Changes in the distribution of native fishes in response to introduced species and other anthropogenic effects

Aim Globally, one of the major threats to the integrity of native faunas is the loss of biodiversity that can result from the introduction of exotics. Here we document recent changes in the distribution of five common fish species that are linked to introductions in Chile. Location Chile from 28° S to 54° S. Methods We assess the extent of changes in distribution of galaxiid species by comparing their historical and current distributions based on the results of the most extensive survey of freshwater fishes in Chile to date, a range that encompasses the full latitudinal and elevational range of the Galaxiidae in Chile. We test for relationships of the distributions and abundances of native fishes with the incidence of introduced species. Results The latitudinal range of Galaxias maculatus has declined by 26%, and most of this reduction has occurred in the northern part of its range. Aplochiton taeniatus and Brachygalaxias bullocki have experienced reductions (8–17% loss) in total drainage area occupied, and they have disappeared from, or are now extremely difficult to find, in latitudes 36° to 41° S, coincidently with areas of urban growth and intense economic activities. The distribution of Galaxias platei has, instead, increased considerably. In northern basins, G. maculatus has apparently been replaced by an introduced poeciliid Gambusia sp. High‐elevation systems remain dominated by native Galaxias platei, whereas systems at intermediate elevations, especially rivers, are now dominated by introduced salmonids. Within drainages, native galaxiids remain abundant where exotic salmonid abundance is low. Main conclusions We suggest that negative interactions between introduced and native fish are responsible for some of the range reductions among Galaxiidae in Chile. The severity of the impacts varies with latitude and altitude and is probably related to temperature. The effects of Gambusia are restricted to warmer systems. Native fish also appear to have found temperature refugia from salmonids; impacts are low in the warmer northern and coastal systems, as well as in high‐altitude relatively cold systems. Native fish also appear less vulnerable to salmonids in lakes than in rivers. This study identifies watersheds critical for the conservation of biodiversity within the Galaxiidae.     

Molecular systematics and adaptive radiation of Hawaii's endemic Damselfly genus Megalagrion (Odonata: Coenagrionidae)

Damselflies of the endemic Hawaiian genus Megalagrion have radiated into a wide variety of habitats and are an excellent model group for the study of adaptive radiation. Past phylogenetic analysis based on morphological characters has been problematic. Here, we examine relationships among 56 individuals from 20 of the 23 described species using maximum likelihood (ML) and Bayesian phylogenetic analysis of mitochondrial (1287 bp) and nuclear (1039 bp) DNA sequence data. Models of evolution were chosen using the Akaike information criterion. Problems with distant outgroups were accommodated by constraining the best ML ingroup topology but allowing the outgroups to attach to any ingroup branch in a bootstrap analysis. No strong contradictions were obtained between either data partition and the combined data set. Areas of disagreement are mainly confined to clades that are strongly supported by the mitochondrial DNA and weakly supported by the elongation factor 1alpha data because of lack of changes. However, the combined analysis resulted in a unique tree. Correlation between Bayesian posterior probabilities and bootstrap percentages decreased in concert with decreasing information in the data partitions. In cases where nodes were supported by single characters bootstrap proportions were dramatically reduced compared with posterior probabilities. Two speciation patterns were evident from the phylogenetic analysis. First, most speciation is interisland and occurred as members of established ecological guilds colonized new volcanoes after they emerged from the sea. Second, there are several instances of rapid radiation into a variety of specialized habitats, in one case entirely within the island of Kauai. Application of a local clock procedure to the mitochondrial DNA topology suggests that two of these radiations correspond to the development of habitat on the islands of Kauai and Oahu. About 4.0 million years ago, species simultaneously moved into fast streams and plant leaf axils on Kauai, and about 1.5 million years later another group moved simultaneously to seeps and terrestrial habitats on Oahu. Results from the local clock analysis also strongly suggest that Megalagrion arrived in Hawaii about 10 million years ago, well before the emergence of Kauai. Date estimates were more sensitive to the particular node that was fixed in time than to the model of local branch evolution used. We propose a general model for the development of endemic damselfly species on Hawaiian Islands and document five potential cases of hybridization (M. xanthomelas x M. pacificum, M. eudytum x M. vagabundum, M. orobates x M. oresitrophum, M. nesiotes x M. oahuense, and M. mauka x M. paludicola).     

Adaptive evolution to novel predators facilitates the evolution of damselfly species range shifts

Most species have evolved adaptations to reduce the chances of predation. In many cases, adaptations to coexist with one predator generate tradeoffs in the ability to live with other predators. Consequently, the ability to live with one predator may limit the geographic distributions of species, such that adaptive evolution to coexist with novel predators may facilitate range shifts. In a case study with Enallagma damselflies, we used a comparative phylogenetic approach to test the hypothesis that adaptive evolution to live with a novel predator facilitates range size shifts. Our results suggest that the evolution of Enallagma shifting from living in ancestral lakes with fish as top predators, to living in lakes with dragonflies as predators, may have facilitated an increase in their range sizes. This increased range size likely arose because lakes with dragonflies were widespread, but unavailable as a habitat throughout much of the evolutionary history of Enallagma because they were historically maladapted to coexist with dragonfly predators. Additionally, the traits that have evolved as defenses against dragonflies also likely enhanced damselfly dispersal abilities. While many factors underlie the evolutionary history of species ranges, these results suggest a role for the evolution of predator‐prey interactions.     

Phylogeographic patterns of Hawaiian Megalagrion damselflies (Odonata: Coenagrionidae) correlate with Pleistocene island boundaries

The Pleistocene geological history of the Hawaiian Islands is becoming well understood. Numerous predictions about the influence of this history on the genetic diversity of Hawaiian organisms have been made, including the idea that changing sea levels would lead to the genetic differentiation of populations isolated on individual volcanoes during high sea stands. Here, we analyse DNA sequence data from two closely related, endemic Hawaiian damselfly species in order to test these predictions, and generate novel insights into the effects of Pleistocene glaciation and climate change on island organisms. Megalagrion xanthomelas and Megalagrion pacificum are currently restricted to five islands, including three islands of the Maui Nui super-island complex (Molokai, Lanai, and Maui) that were connected during periods of Pleistocene glaciation, and Hawaii island, which has never been subdivided. Maui Nui and Hawaii are effectively a controlled, natural experiment on the genetic effects of Pleistocene sea level change. We confirm well-defined morphological species boundaries using data from the nuclear EF-1alpha gene and show that the species are reciprocally monophyletic. We perform phylogeographic analyses of 663 base pairs (bp) of cytochrome oxidase subunit II (COII) gene sequence data from 157 individuals representing 25 populations. Our results point to the importance of Pleistocene land bridges and historical island habitat availability in maintaining inter-island gene flow. We also propose that repeated bottlenecks on Maui Nui caused by sea level change and restricted habitat availability are likely responsible for low genetic diversity there. An island analogue to northern genetic purity and southern diversity is proposed, whereby islands with little suitable habitat exhibit genetic purity while islands with more exhibit genetic diversity.     

Isolation and characterization of 16 polymorphic microsatellite loci in the endemic Hawaiian damselfly Megalagrion xanthomelas (Odonata: Coenagrionidae)

Microsatellite loci have been isolated from two species of endemic Hawaiian damselflies, Megalagrion xanthomelas and M. eudytum, that are of conservation concern. Sixteen polymorphic loci were characterized in 32 M. xanthomelas from one population on Molokai and one on Hawaii Island. The total number of alleles per locus ranged from two to 16 and observed population heterozygosity ranged from 0.0 to 0.963. Eleven of these loci amplified successfully in M. eudytum as well. These loci will be used to further conservation efforts and infer genetic consequences of Pleistocene glaciations.     

Global diversity of dragonflies (Odonata) in freshwater

Larvae of almost all of the 5,680 species of the insect order Odonata (dragonflies and damselflies) are dependent on freshwater habitats. Both larvae and adults are predators. The order is relatively well studied, and the actual number of species may be close to 7,000. Many species have small distributional ranges, and are habitat specialists, including inhabitants of alpine mountain bogs, seepage areas in tropical rain forests, and waterfalls. They are often successfully used as indicators for environmental health and conservation management. The highest diversity is found in flowing waters in rain forests of the tropics, the Oriental and Neotropical regions being the most speciose. This paper discusses diversity, summarises the biogeography of dragonflies in the different biogeographical regions and gives the total number of species and genera per family per biogeographical region. Examples are given of areas of particular diversity, in terms of areas of endemism, presence of ancient lineages or remarkable recent radiations but no well-based review of areas with high endemism of dragonflies is available so far. The conservation status of dragonflies is briefly discussed. Species confined to small remnants of forest in the tropics are most under threat of extinction by human activities. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

A reptilian smoking gun: first record of invasive Jackson’s chameleon (<Emphasis Type="Italic">Chamaeleo jacksonii</Emphasis>) predation on native Hawaiian species

Here we report the first conclusive evidence of an introduced reptile (Chamaeleo jacksonii) feeding on Hawaiian taxa, including 11 snails in four endemic genera from two families, including four individuals of an endangered species (Achatinella mustelina), and native insects in five genera. Native Hawaiian invertebrates were discovered in the dissected stomachs of wild caught Jackson’s chameleons collected from June to November 2009 on the island of Oahu. Although Jackson’s chameleons were introduced to the Hawaiian Islands in the early 1970s, ecological impacts have never been documented. Of particular concern is the fact that chameleons have previously only rarely been found in native Hawaiian habitat, although 12 were recently collected in a mid-elevation native forest, an area that is not likely to be suitable for their long-term persistence, but that is adjacent to higher elevation pristine forest where endemic prey are abundant and favorable climatic conditions exist for chameleon persistence. One concern is that Jacksons’s chameleons may be undergoing a range expansion into upper elevation pristine forests. If chameleons reach and establish populations in these areas, devastating impacts to the native ecosystem are possible. A thorough understanding of the impacts of chameleons on Hawaiian fauna will require additional evaluation and sampling, but dissemination of this discovery in a timely fashion is important as it provides new information regarding this threat. Monitoring and collection of chameleons is ongoing, particularly in native Hawaiian forest habitats at mid and upper elevations (600–1,300 m).     

Sexual selection, sexual size dimorphism and Rensch's rule in Odonata

Odonata (dragonflies and damselflies) exhibit a range of sexual size dimorphism (SSD) that includes species with male-biased (males > females) or female-biased SSD (males < females) and species exhibiting nonterritorial or territorial mating strategies. Here, we use phylogenetic comparative analyses to investigate the influence of sexual selection on SSD in both suborders: dragonflies (Anisoptera) and damselflies (Zygoptera). First, we show that damselflies have male-biased SSD, and exhibit an allometric relationship between body size and SSD, that is consistent with Rensch's rule. Second, SSD of dragonflies is not different from unit, and this suborder does not exhibit Rensch's rule. Third, we test the influence of sexual selection on SSD using proxy variables of territorial mating strategy and male agility. Using generalized least squares to account for phylogenetic relationships between species, we show that male-biased SSD increases with territoriality in damselflies, but not in dragonflies. Finally, we show that nonagile territorial odonates exhibit male-biased SSD, whereas male agility is not related to SSD in nonterritorial odonates. These results suggest that sexual selection acting on male sizes influences SSD in Odonata. Taken together, our results, along with avian studies (bustards and shorebirds), suggest that male agility influences SSD, although this influence is modulated by territorial mating strategy and thus the likely advantage of being large. Other evolutionary processes, such as fecundity selection and viability selection, however, need further investigation.     

Ecology and conservation of Pseudolestes mirabilis (Odonata: Zygoptera), a damselfly endemic to Hainan Island of China

Although some efforts have addressed oriental dragonfly conservation, knowledge on the ecology and geographic distribution of such dragonflies remains scant. The phoenix damselfly Pseudolestes mirabilis Kirby, 1900 is the single species in the family Pseudolestidae that is endemic to Hainan Island of China. This damselfly was recommended by the International Union for the Conservation of Nature to be of priority for further study and conservation. In this work, we use ecological niche modeling techniques to estimate the dimensions of the realized niches of this damselfly and to predict its potential distribution. Our findings suggest that the phoenix damselfly possessed a small climate space characterized by low temperature and high precipitation. Highly suitable areas are mainly distributed in the low‐altitude regions of southern central tree‐covered mountains in Hainan. Caution is warranted when considering the potential habitat loss attributed to human activity and climate change.     

The significance of littoral and shoreline habitat integrity to the conservation of lacustrine damselflies (Odonata)

Human development of pond and lake shorelines may significantly impact native lacustrine biota including a variety of aquatic macroinvertebrate groups. In an effort to better understand the habitat associations and sensitivities of lacustrine damselflies (Odonata: Zygoptera), we sampled adults in littoral macrophyte habitat during two flight periods at 35 randomly selected pond and lake sites in southern Maine during 2000 and 2001. Data were also collected to help characterize water body, shoreline disturbance, and aquatic vegetation at each study site. Nonmetric multidimensional scaling was used for ordination of damselfly assemblages, and coordinates from the most stable solution were related to site variables using forward stepwise multiple regression. Our results suggest that the diversity and composition of damselfly assemblages is related to the abundance and richness of littoral zone macrophytes, extent of riparian disturbance, benthic substrate granularity, and lake productivity; all variables subject to anthropogenic degradation on excessively developed waterbodies. Additionally, we developed a Habitat Tolerance Index useful for distinguishing between relative habitat specialists and generalists from among a diverse assemblage of 19 lacustrine species. Finally, species-specific damselfly associations with multiple genera of floating and emergent macrophytes were assessed using both nonparametric correlation and multiplicative regression yielding significant relationships for 17 species, including two damselflies of global conservation concern (Enallagma laterale and E. pictum). We conclude that the protection of littoral and shoreline habitat integrity, with special emphasis on emergent and floating macrophytes, is critical to the conservation of lacustrine biodiversity.     

Invasion of the Hawaiian Islands by a parasite infecting imperiled stream fishes

Points of origin and pathways of spread are often poorly understood for introduced parasites that drive disease emergence in imperiled native species. Co‐introduction of parasites with non‐native hosts is of particular concern in remote areas like the Hawaiian Islands, where the introduced nematode Camallanus cotti has become the most prevalent parasite of at‐risk native stream fishes. In this study, we evaluated the prevailing hypothesis that C. cotti entered the Hawaiian Islands with poeciliid fishes from the Americas, and spread by translocation of poeciliid hosts across the archipelago for mosquito control. We also considered the alternative hypothesis of multiple independent co‐introductions with host fishes originating from Asia. We inferred conduits of introduction and spread of C. cotti across the archipelago from geographic patterns of mtDNA sequence variation and allelic variation across 11 newly developed microsatellite markers. The distribution of haplotypes suggests that C. cotti spread across the archipelago following an initial introduction on O'ahu. Approximate Bayesian Computation modeling and allelic variation also indicate that O'ahu is the most likely location of introduction, from which C. cotti dispersed to Maui followed by spread to the other islands in the archipelago. Evidence of significant genetic structure across islands indicates that contemporary dispersal is limited. Our findings parallel historical records of non‐native poeciliid introductions and suggest that remediating invasion hotspots could reduce the risk of infection in native stream fishes, which illustrates how inferences on parasite co‐introductions can improve conservation efforts by guiding responses to emerging infectious disease in species of concern.     

Determinants of fine‐scale homogenization and differentiation of native freshwater fish faunas in a Mediterranean Basin: implications for conservation

Aim Increasing threats to freshwater biodiversity are rapidly changing the distinctiveness of regional species pools and local assemblages. Biotic homogenization/differentiation processes are threatening the integrity and persistence of native biodiversity patterns at a range of spatial scales and pose a challenge for effective conservation planning. Here, we evaluate the extent and determinants of fine‐scale alteration in native freshwater fish assemblages among stream reaches throughout a large river basin and consider the implications of these changes for the long‐term conservation of native fishes. Location Guadiana River basin (South‐Western Iberian Peninsula). Methods We quantified the magnitude of change in compositional similarity between observed and reference assemblages and its potential effect on natural patterns of compositional distinctiveness. Reference assemblages were defined as the native species expected to occur naturally (in absence of anthropogenic alterations) and were reconstructed using a multivariate adaptive regression splines predictive model. We also evaluated the role of habitat degradation and introduced species as determinants of biotic homogenization/differentiation. Results We found a significant trend towards homogenization for native fish assemblages. Changes in native fish distributions led to the loss of distinctiveness patterns along natural environmental gradients. Introduced species were the most important factor explaining the homogenization process. Homogenization of native assemblages was stronger in areas close to reservoirs and in lowland reaches where introduced species were more abundant. Main conclusions The implementation of efficient conservation for the maintenance of native fish diversity is seriously threatened by the homogenization processes. The identification of priority areas for conservation is hindered by the fact that the most diverse communities are vanishing, which would require the selection of broader areas to adequately protect all the species. Given the principal role that introduced species play in the homogenization process and their relation with reservoirs, special attention must be paid to mitigating or preventing these threats.     

A northward shift of range margins in British Odonata

Many species are predicted to shift their ranges to higher latitudes and altitudes in response to climate warming. This study presents evidence for 37 species of nonmigratory British dragonflies and damselflies shifting northwards at their range margins over the past 40 years, seemingly as a result of climate change. This response by an exemplar group of insects associated with fresh water, parallels polewards range changes observed in terrestrial invertebrates and other taxa.     

Impact of exotic fish removal on native communities in farm ponds

Introduced largemouth bass (Micropterus salmoides spp.) and bluegill (Lepomis macrochirus spp.) are thought to threaten native aquatic organisms worldwide and hence their eradication has recently begun in Japan. Our previous studies suggested that the removal of largemouth bass increases native fish, shrimp, dragonflies, and exotic crayfish, but decreases macrophytes. To test this prediction, we removed the exotic fishes by draining farm ponds and compared the numbers of these organisms before and after the drain, as well as between drained and undrained ponds. The number of dragonfly Pseudothemis zonata, crayfish, shrimp, and goby increased rapidly after the drain, but the coverage of macrophyte declined. The reduction in macrophyte is assumed to be caused by increased herbivory by crayfish. The number of exuviae of damselfly Cercion calamorum and the total number of species of odonate also decreased after the drain. These decreases can be due to the reduction of macrophyte because reduced odonate species are known to use macrophytes as oviposition sites. Therefore, the removal of largemouth bass has a potential to cause negative effects on some native organisms. We propose that reduction of exotic crayfish should be considered when eradicating the exotic fishes.     

First Baltic amber megapodagrionid damselfly (Odonata: Zygoptera)

Electropodagrion szwedoi n. gen., n. sp., first Baltic amber megapodagrionid damselfly, is described. The European and North American fossils document a very high diversity and a much wider distribution of this group of damselflies during the Cenozoic than today. A checklist of described fossil species of damselflies of the family Megapodagrionidae is given.     

Impact of predation by spiders on a territorial damselfly (Odonata: Calopterygidae)

Summary Predation by orb-weaving spiders and crab spiders on the damselfly Calopteryx haemorrhoidalis was studied at a small stream in Southern France. One species of orb-weaving spider, Larinioides folium, caught 76% of the damselflies which fell prey to spiders. Displacement experiments on spiders on sections of bank and the positioning of webs in male territories show that the density and distribution of damselflies is not influenced by orbweb density or by the position of webs. Predation rates corresponded to orb-web density, but neither for sex nor for stage was there a relationship with damselfly density. Mean daily predation rates ranged between 0.9% for females and 4.1% for adult males. Predation risk to adult damselflies by orb-weaving spiders was male biased, whereas among tenerals there was no bias. Males were captured more frequently at territories near the water. Captures show a maximum at noon when territorial disputes of adult males were most frequent. After orbwebs were placed within territories predation rate of males was strongly increased. Predation risk to adult females in the direct vicinity of the stream was less than in the bank vegetation where they perch close to orbwebs. The risk of predation by crab spiders, which catch damselflies at their perching sites, was not sex-biased.     

Modeling the impacts of global warming on predation and biotic resistance: mosquitoes, damselflies and avian malaria in Hawaii

Biotic resistance from native predators can play an important role in regulating or limiting exotic prey. We investigate how global warming potentially alters the strength and spatial extent of these predator–prey interactions in aquatic insect ecosystems. As a simple model system, we use rock pools in streams of rainforests of Hawaii, which contain the beautiful Hawaiian damselfly Megalagrion calliphya as predator and the invasive southern house mosquito Culex quinquefasciatus as prey. This abundant mosquito is the major vector of avian malaria transmission to native forest birds. We use mathematical modeling to evaluate the potential impacts of damselfly predation and temperature on mosquito population dynamics. We model this predator–prey system along an elevational gradient (749-1952 m elevation) and assess the effect of 1°C and 2°C climate warming scenarios as well as the effects of El Niño and La Niña oscillations, on predator–prey dynamics. Our results indicate that the strength of biotic resistance of native predators on invasive prey may decrease with increasing temperature because demographic rates of predator and prey are differentially affected by temperature. Future warming could therefore increase the abundance of invasive species by releasing them from predation pressure. If the invasive species is a disease vector, these shifts could increase the impact of disease on both humans and wildlife.     

Beyond latitude: Temperature,productivity and thermal niche conservatism drive global body size variation in Odonata

Aim

So far, latitudinal body size clines have been discussed primarily in the context of thermoregulation, sensu Bergmann. However, body size patterns are ambiguous in ectotherms, and this heterogeneity remains poorly understood. We tested whether Bergmann's rule and the resource availability rule, which states that energetic requirements determine species body size, apply to damselflies and dragonflies (Odonata). Furthermore, we hypothesized that the contrasting effects of thermoregulation and resource availability (e.g., productivity) can obscure the overall gradient in body size variation.

Location

Global.

Time period

Contemporary.

Major taxa studied

Odonata.

Methods

Using data for 43% of all odonate species described so far, we tested our hypotheses in phylogenetically and spatially comparative analyses at assemblage and species levels. For the distribution data, we integrated expert range maps and ecoregional ranges based on all available occurrence records. To distinguish between long-term and evolutionarily recent responses of environmental drivers in body size, we constructed a phylogenetically informed classification of all odonate species and decomposed the body size into its phylogenetic and specific components for our subset of species.

Results

We documented a weak positive relationship between body length and latitude but found strong and contrasting effects for temperature between dragonflies and damselflies and consistent positive effects for productivity that explained 35–57% of body size variation. Moreover, we showed a strong phylogenetic signal in sized-based thermoregulation that shaped the distribution of dragonflies, but not of damselflies.

Main conclusions

We concluded that temperature, productivity and conservatism in size-based thermoregulation synergistically determine the distribution of ectotherms, while the taxon-specific importance of these factors can lead to contrasting and weak latitude–size relationships. Our results reinforce the importance of body size as a determinant of species distributions and responses to climate change.     

Protozoal and epitheliocystis-like infections in the introduced bluestripe snapper Lutjanus kasmira in Hawaii

The bluestripe snapper, or taape, was introduced into Hawaii in the 1950s and has since become very abundant throughout the archipelago. As part of a health survey of reef fish in Hawaii, we necropsied 120 taape collected from various coastal areas south of Oahu and examined fish histology for extraintestinal organisms. Forty-seven percent of taape were infected with an apicomplexan protozoan compatible with a coccidian. Infection was evident mainly in the spleen and, less commonly, the kidney. Prevalence of this coccidian increased with size of fish, and we saw no significant pathology associated with the organism. Twenty-six percent of taape were also infected with an epitheliocystis-like organism that occurred mainly in the kidney and, less commonly, the spleen. In contrast to the coccidian, fish mounted a notable inflammatory response to the epitheliocystis-like organism, and this inflammation appeared to increase in severity with age. Prevalence of the epitheliocystis-like organism infection increased with age, but infection was not seen in fish greater than 26.5 cm fork length. The high prevalence of coccidial infection in introduced taape prompts the concern that these organisms, along with the epitheliocystis-like organism, have the potential to be transmitted to native reef fish. Given the impact of other introduced microbial organisms on native Hawaiian fauna, there is a clear need to assess whether protozoa and bacteria are endemic to Hawaii, and whether they negatively impact native reef fish that closely associate with taape.