Angela L. Strecker

The Aquarium Trade as an Invasion Pathway in the Pacific Northwest

Abstract The aquarium trade moves thousands of species around the globe, and unwanted organisms may be released into freshwaters, with adverse ecological and economic effects. We report on the first investigation of the ornamental pet trade as an invasion pathway in the Pacific Northwest region of the United States, where a moderate climate and a large human population present ample opportunities for the introduction and establishment of aquarium trade species. Results from a regional survey of pet stores found that the number of fish (n=400) and plant (n=124) species currently in the aquarium trade is vast. Pet stores import thousands of fish every month, the majority of which (58%) are considered to pose an ecological threat to native ecosystems. Our propagule pressure model suggests that approximately 2,500 fish (maximum 21,000 individuals) are likely released annually to the Puget Sound region by aquarists, and that water temperatures in many parts of Washington are suitable for establishment of popul...

Defining conservation priorities for freshwater fishes according to taxonomic, functional, and phylogenetic diversity

To date, the predominant use of systematic conservation planning has been to evaluate and conserve areas of high terrestrial biodiversity. Although studies in freshwater ecosystems have received recent attention, research has rarely considered the potential trade-offs between protecting different dimensions of biodiversity and the ecological processes that maintain diversity. We provide the first systematic prioritization for freshwaters (focusing on the highly threatened and globally distinct fish fauna of the Lower Colorado River Basin, USA) simultaneously considering scenarios of: taxonomic, functional, and phylogenetic diversity; contemporary threats to biodiversity (including interactions with nonnative species); and future climate change and human population growth. There was 75% congruence between areas of highest conservation priority for different aspects of biodiversity, suggesting that conservation efforts can concurrently achieve strong complementarity among all types of diversity. However, si...

A multi-scale comparison of trait linkages to environmental and spatial variables in fish communities across a large freshwater lake

Species present in communities are affected by the prevailing environmental conditions, and the traits that these species display may be sensitive indicators of community responses to environmental change. However, interpretation of community responses may be confounded by environmental variation at different spatial scales. Using a hierarchical approach, we assessed the spatial and temporal variation of traits in coastal fish communities in Lake Huron over a 5-year time period (2001–2005) in response to biotic and abiotic environmental factors. The association of environmental and spatial variables with trophic, life-history, and thermal traits at two spatial scales (regional basin-scale, local site-scale) was quantified using multivariate statistics and variation partitioning. We defined these two scales (regional, local) on which to measure variation and then applied this measurement framework identically in all 5 study years. With this framework, we found that there was no change in the spatial scales of fish community traits over the course of the study, although there were small inter-annual shifts in the importance of regional basin- and local site-scale variables in determining community trait composition (e.g., life-history, trophic, and thermal). The overriding effects of regional-scale variables may be related to inter-annual variation in average summer temperature. Additionally, drivers of fish community traits were highly variable among study years, with some years dominated by environmental variation and others dominated by spatially structured variation. The influence of spatial factors on trait composition was dynamic, which suggests that spatial patterns in fish communities over large landscapes are transient. Air temperature and vegetation were significant variables in most years, underscoring the importance of future climate change and shoreline development as drivers of fish community structure. Overall, a trait-based hierarchical framework may be a useful conservation tool, as it highlights the multi-scaled interactive effect of variables over a large landscape.

Increased abundance of the non-indigenous zooplanktivore, Bythotrephes longimanus, is strongly correlated with greater spring prey availability in Canadian Shield lakes

The non-indigenous zooplanktivore, Bythotrephes longimanus, is a large Palaearctic cladoceran that is spreading rapidly in the Great Lakes watershed in North America. As a voracious predator, Bythotrephes can reduce herbivorous cladoceran abundance and diversity; however, the variables that affect its abundance are not well understood. To determine what bottom-up factors are associated with the abundance and seasonal dynamics of established Bythotrephes populations, two Bythotrephes datasets from lakes in south-central Ontario, Canada, were analysed using multiple regression and multivariate analyses: a multi-lake dataset of nine lakes sampled in 2003 and a multi-year dataset of one of these lakes, Harp Lake, sampled from 1994–1998 and 2001–2004. Bottom-up variables tested were Secchi disk depth, epilimnetic temperature, cladoceran (prey) density, total phosphorus, dissolved organic carbon and Chlorophyll a, as well as maximum depth for the multi-lake dataset. In both analyses and datasets, springtime abundance of herbivorous cladocerans was consistently found to be a significant factor associated with Bythotrephes (June–September) abundance; Bythotrephes annual abundance was significantly and positively associated with mean May and June prey abundance, along with mean Secchi disk depth for the multi-lake dataset, and groups of lakes or years with similar Bythotrephes seasonal abundance patterns were predicted by June prey abundance. Additionally, prey availability was the dominant contributor towards changes in weekly Bythotrephes birth rates calculated for two of the study lakes. Our study suggests that prey availability influences Bythotrephes abundance, which provides evidence that Bythotrephes establishment success is affected by the abundance of its prey.

Complex interactions between regional dispersal of native taxa and an invasive species

In the event of an environmental disturbance, dispersal of native taxa may provide species and genetic diversity to ecosystems, increasing the likelihood that there will be species and genotypes present that are less vulnerable to the disturbance. This may allow communities to maintain functioning during a disturbance and may be particularly important when the perturbation is novel to the system, such as the establishment of an invasive species. We examined how dispersal of native species may influence crustacean zooplankton communities in freshwater lakes invaded by the invertebrate predator, Bythotrephes longimanus. Using large enclosures, we experimentally tested the effect of dispersal on zooplankton community abundance, richness, and composition in (1) a community invaded by Bythotrephes, (2) the same community with the invader removed, and (3) a community that was never invaded. Dispersal increased zooplankton community abundance and richness; however, these effects were usually only significant in the invader-removed treatment. Dispersal tended to make the invader-removed communities more similar to never-invaded communities in abundance, richness, and composition. Dispersal had little effect on zooplankton abundance in the invaded community; however, richness significantly increased, and the community composition changed to resemble a never-invaded community by the end of the experiment. Our results have implications for understanding the role of dispersal during transitory states in communities. Dispersal of native taxa may be particularly important during the period between the arrival and broad-scale establishment of Bythotrephes, as dispersal through space or time (i.e., from resting eggs) may rapidly increase zooplankton abundance when the invader is absent or in low abundances. Overall, our results suggest that communities with strong local predatory and competitive interactions may be closed to immigration from colonists, but that invasive species may alter the conditions under which species can establish. These results have implications for the interaction of native and invasive species across broad spatial scales, as regional dispersal of native taxa may forestall the local extirpation of native species. In particular, transient phases that result from variable persistence of invaders within habitats or across a region may permit native colonists to successfully establish, and thus increase local and regional resistance to future disturbance.

Bythotrephes invasion elevates trophic position of zooplankton and fish: implications for contaminant biomagnification

We estimated the effects of Bythotrephes longimanus invasion on the trophic position (TP) of zooplankton communities and lake herring, Coregonus artedi. Temporal changes in lacustrine zooplankton communities following Bythotrephes invasion were contrasted with non-invaded reference lakes, and along with published information on zooplankton and herring diets, formed the basis of estimated changes in TP. The TP of zooplankton communities and lake herring increased significantly following the invasion of Bythotrephes, whereas TP in reference lakes decreased (zooplankton) or did not change significantly (lake herring) over a similar time frame. Elevated TP following Bythotrephes invasion was most prominent in lakes that also supported the glacial relict, Mysis diluvania, suggesting a possible synergistic interaction between these two species on zooplankton community composition. Our analysis indicated that elevated TPs of zooplankton communities and lake herring are not simply due to the presence of Bythotrephes, but rather reflect changes in the zooplankton community induced by Bythotrephes; namely, a major reduction in the proportion of herbivorous cladoceran biomass and a concomitant increase in the proportion of omnivorous and/or predatory copepod biomass in invaded lakes. We demonstrated that increases in TP of the magnitude reported here can lead to substantial increases in fish contaminant concentrations. In light of these results, we discuss potential mechanisms that may be responsible for the disconnect between empirical and theoretical evidence that mid-trophic level species invasions (e.g., Bythotrephes) elevate contaminant burdens of consumer species, and provide testable hypotheses to evaluate these mechanisms.

Fish species introductions provide novel insights into the patterns and drivers of phylogenetic structure in freshwaters

Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. We quantify phylogenetic clustering and overdispersion of native and non-native fishes of a large river basin in the American Southwest to test for the mechanisms (environmental filtering versus competitive exclusion) and spatial scales influencing community structure. Contrary to expectations, non-native species were phylogenetically clustered and related to natural environmental conditions, whereas native species were not phylogenetically structured, likely reflecting human-related changes to the basin. The species that are most invasive (in terms of ecological impacts) tended to be the most phylogenetically divergent from natives across watersheds, but not within watersheds, supporting the hypothesis that Darwins naturalization conundrum is driven by the spatial scale. Phylogenetic distinctiveness may facilitate non-native establishment at regional scales, but environmental filtering restricts local membership to closely related species with physiological tolerances for current environments. By contrast, native species may have been phylogenetically clustered in historical times, but species loss from contemporary populations by anthropogenic activities has likely shaped the phylogenetic signal. Our study implies that fundamental mechanisms of community assembly have changed, with fundamental consequences for the biogeography of both native and non-native species.

Differential sensitivity of planktonic trophic levels to extreme summer temperatures in boreal lakes

The stress–size hypothesis predicts that smaller organisms will be less sensitive to stress. Consequently, climate warming is expected to favour smaller taxa from lower trophic levels and smaller individuals within populations. To test these hypotheses, we surveyed zooplankton communities in 20 boreal lakes in Killarney Provincial Park, Canada during 2005 (an anomalously warm summer) and 2006 (a normal summer). Higher trophic levels had larger responses to warm temperatures supporting the stress–size hypothesis; however, rather than imposing negative effects, higher density and biomass were observed under warmer temperatures. As a result, larger taxa from higher trophic levels were disproportionately favoured with warming, precluding an expected shift towards smaller species. Proportionately greater increases in metabolic rates of larger organisms or altered biotic interactions (e.g. predation and competition) are possible explanations for shifts in biomass distribution. Warmer temperatures also favoured smaller individuals of the two most common species, in agreement with the stress–size hypothesis. Despite this, these populations had higher biomass in the warm summer. Therefore, reduced adult survivorship may have triggered these species to invest in reproduction over growth. Hence, warmer epilimnions, higher zooplankton biomass and smaller individuals within zooplankton populations may function as sensitive indicators of climate warming in boreal lakes.

Spatial and Temporal Variability of Contaminants Within Estuarine Sediments and Native Olympia Oysters: A Contrast Between a Developed and an Undeveloped Estuary

Chemical contaminants can be introduced into estuarine and marine ecosystems from a variety of sources including wastewater, agriculture and forestry practices, point and non-point discharges, runoff from industrial, municipal, and urban lands, accidental spills, and atmospheric deposition. The diversity of potential sources contributes to the likelihood of contaminated marine waters and sediments and increases the probability of uptake by marine organisms. Despite widespread recognition of direct and indirect pathways for contaminant deposition and organismal exposure in coastal systems, spatial and temporal variability in contaminant composition, deposition, and uptake patterns are still poorly known. We investigated these patterns for a suite of persistent legacy contaminants including polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers (PBDEs) and chemicals of emerging concern including pharmaceuticals within two Oregon coastal estuaries (Coos and Netarts Bays). In the more urbanized Coos Bay, native Olympia oyster (Ostrea lurida) tissue had approximately twice the number of PCB congeners at over seven times the total concentration, yet fewer PBDEs at one-tenth the concentration as compared to the more rural Netarts Bay. Different pharmaceutical suites were detected during each sampling season. Variability in contaminant types and concentrations across seasons and between species and media (organisms versus sediment) indicates the limitation of using indicator species and/or sampling annually to determine contaminant loads at a site or for specific species. The results indicate the prevalence of legacy contaminants and CECs in relatively undeveloped coastal environments highlighting the need to improve policy and management actions to reduce contaminant releases into estuarine and marine waters and to deal with legacy compounds that remain long after prohibition of use. Our results point to the need for better understanding of the ecological and human health risks of exposure to the diverse cocktail of pollutants and harmful compounds that will continue to leach from estuarine sediments over time.

Effects of Bythotrephes longimanus (Crustacea, Cladocera) on the abundance, morphology, and prey community of Leptodora kindtii (Crustacea, Cladocera)

We hypothesized that native Leptodora kindtii would be shorter and have smaller feeding baskets in central Ontario lakes with greater abundances of small-bodied zooplankton prey, and that differences in zooplankton size among lakes could be attributed to the invasive cladoceran Bythotrephes longimanus. We evaluated these conjectures by comparing size metrics of Leptodora and the size of their preferred cladoceran prey in lakes invaded or not by Bythotrephes. Leptodora was less abundant in invaded lakes, but were smaller bodied with smaller feeding baskets only in lakes with long invasion histories. Small cladoceran abundance was greater in non-invaded lakes and was directly related to Leptodora abundance although not to Leptodora size. Mean Leptodora body size declined with increasing abundance of Bythotrephes. We evaluated three possible explanations for these patterns in Leptodora—(a) competition with Bythotrephes for zooplankton prey, (b) direct predation by Bythotrephes, and (c) size-selective predation by fish. While we were unable to unequivocally distinguish among these hypotheses, our observations are most consistent with predation by Bythotrephes changing zooplankton community composition and size structure in a manner that is detrimental to Leptodora. Our results indicate that Bythotrephes invasion may trigger more complex and subtle changes in food webs than previously thought.