Potential roles for differential body size and microhabitat complexity in mediating biotic interactions within invasive freshwater amphipod assemblages

Cannibalism and intraguild predation (IGP) are common amongst freshwater amphipod crustacean aswsemblages, particularly between individuals of different body size, with IGP of smaller by larger species. The decline of Gammarus tigrinus Populations in mainland Europe has been accompanied by the arrival of the Ponto-Caspian invader Dikerogammarus villosus and previous studies have implicated IGP of G. tigrinus by the larger D. villosus as the principal driving force in this replacement. We examined how factors such as microhabitat and body size may mediate both cannibalism within G. tigrinus populations and IGP by D. villosus and thus contribute to field patterns of coexistence and exclusion. A field Survey of an invaded Dutch fake indicated that G. tigrinus and D. villosus differed in distribution. with D. villosus being the numerically dominant amphipod (80-96 %) on the rocky boulder Substrate of the shoreline and G. tigrinus being the dominant amphipod (100 %) in the crushed shell/sand matrix immediately adjacent to this. Laboratory microcosm experiments indicated that G. tigrinus cannibalism, particularly of smaller by larger size classes, may be common. In addition, although D. villosus predation of all G. tigrinus size classes was extreme, the smallest size classes Suffered the highest predation. Indeed, when exposed to D. villosus, predation of larger G. tigrinus was lowest when smaller G. tigrinus were also present. Increasing microhabitat complexity from a simple bare substrate littered with Dreissena polymorpha zebra mussels to a Crushed shell/sand matrix significantly reduced both cannibalism and IGP. Our Study emphasizes the need to consider both life history stages and habitat template, when considering the impacts of biotic interactions and it also emphasizes that complex, interacting factors may be mediating the range expansion of D. villosus.

The replacement of a native freshwater amphipod by an invader: roles for environmental degradation and intraguild predation

We assessed the extent to which an invader, Gammarus pulex (Crustacea: Amphipoda), has replaced a native, Gammarus duebeni celticus, over a 13-year period in a European river system and some of the abiotic and biotic factors that could account for this. Between 1988 and 2001, 56% of mixed-species sites had become invader-only sites, whereas no mixed sites had become native only again. The native dominated areas of higher dissolved oxygen and water quality, with the reciprocal true for the invader. Field transplant experiments revealed that native survivorship was lower in areas where it had been replaced than in areas where the invader does not yet occur. In invader-only areas, native survivorship was lower than that of the invader when kept separately and lowest when both species were kept together. We also observed predation of the native by the invader. Laboratory oxygen manipulation experiments revealed that at 30% saturation, the native's survivorship was two thirds that of the invader. We conclude that decreasing water quality favours replacement of the native by the invader.

Lethal and sublethal toxicity of ammonia to native, invasive, and parasitised freshwater amphipods

In lethal and sublethal ammonia toxicity tests, we examined differences in tolerance of three species of freshwater amphipods, one native and two invasive in Ireland. The native Gammarus duebeni celticus was slightly less tolerant to ammonia than the invasive G. pulex (96h LC50 = 1.155 and 1.544 mg l(-1), respectively), while another invader, Crangonyx pseudograeilis, had the lowest tolerance (LC50 = 0.36 mg l(-1)). Parasitism of G. pulex by the acanthocephalan Echinorhynchus truttae greatly reduced the tolerance of the invader to ammonia (LC50 = 0.381 mg l(-1)). Further, precopula pair disruption tests indicated that G. d. celticus was more sensitive to ammonia than G. pulex at sublethal levels. We discuss these results in the context of the ecological replacements of native by invader amphipods. (C) 2004 Elsevier Ltd. All rights reserved.

Differential drift and parasitism in invading and native Gammarus spp. (Crustacea : Amphipoda)

Invading and native species often interact directly, such as by predation, producing patterns of exclusion and coexistence. Less direct factors, such as interactions with the broader abiotic and biotic environment, may also contribute to such patterns, but these have received less recognition. In Northern Ireland, the North American Gammarus tigrinus has invaded freshwaters populated with the native Gammarus duebeni celticus, with intraguild predation between the two implicated in their relative success. However, these species also engage in day and night

Differential physico-chemical tolerances and intraguild predation among native and invasive amphipods (Crustacea); a field study

We used field surveys and transplant experiments to elucidate the relative roles of physico-chemical regime and intraguild predation in determining the generally mutually exclusive distributions of native and invader freshwater amphipod species. Field surveys showed that the native Gammarus duebeni celticus dominates the shoreline of Lough Neagh, N. Ireland, with some co-occurrence with the N. American invader G. tigrinus. However, the latter species dominates the deeper areas of the mid-Lough. Transplant experiments showed no difference in survival of the native and invader in single species 'bioassay tubes' placed along the shoreline. However, there was significantly higher survival of the invader compared with the native in single species tubes placed in the mid-Lough. In mixed species tubes on the shoreline, the native killed and ate the invader, with no reciprocal interaction, leading to significant reductions of the invader. However, the invader had significantly higher survival than the native in mixed species tubes in the mid-Lough, with no evidence. of predation between the two species. These results indicate that, whereas differential intraguild predation may determine domination of the shoreline by the native, differential physico-chemical tolerances may be major determinants of the domination of the mid-Lough by the invader. This study emphasises the need to consider the habitat template in conjunction with biotic interactions before attempting to draw conclusions about mechanisms determining relative distribution patterns of native and invasive species.

Temporal changes in the distribution of native and introduced freshwater amphipods in Lough Neagh, Northern Ireland

Studies of invasion scenarios over long time periods are important to refine explanations and predictions of invasion success and impact. We used data from surveys in 1958 and 1999 of the macroinvertebrates of Lough Neagh, Northern Ireland, to assess changes in the distribution of native and introduced amphipods in relation to the wider assemblage. In 1958, the invader G. tigrinus dominated the shoreline fauna, with the native G. d. celticus present in very low numbers, whereas in 1999 the reverse was evident. In both surveys, G. tigrinus was the only amphipod present in the mid-Lough. G. tigrinus thus seems to have become established within L. Neagh, perhaps overshot and then senesced, with the native species re-establishing on the shoreline, with the invader mostly restricted to the deep mid-Lough. The non-amphipod macroinvertebrate assemblage was similar between the two surveys, in terms of Bray-Curtis community similarity, assemblage diversity, dominance and the taxa based ASPT water quality index. However, the mean density of macroinvertebrates (all taxa combined) was lower in 1999 compared to 1958, largely accounted for by a decline in oligochaete numbers. Since Gammarus species may be predators of other macroinvertebrates and influence their distribution and abundance, we investigated this trophic link in staged laboratory encounters. Both G. tigrinus and G. d. celticus preyed on isopods, alderflies, mayflies, chironomids and mysids, however, the native G. d. celticus had a significantly greater predatory impact on isopods and chironomids than did the invader G. tigrinus. While we cannot definitively ascribe cause and effect in the present scenario, we discuss how replacement of one amphipod species by another may have impacts on the wider macroinvertebrate assemblage.

An acanthocephalan parasite mediates intraguild predation between invasive and native freshwater amphipods (Crustacea)

1. The balance of predation between closely related invasive and native species can be an important determinant of the success or failure of biological invasions. In Irish freshwaters, the introduced amphipod Gammarus pulex has replaced the native G. duebeni celticus, possibly through differential mutual intraguild predation (IGP). Theoretically, parasitism could mediate such predation and hence the invasion outcome. However, this idea remains poorly studied.

Resolution of a taxonomic conundrum: an ultrastructural and molecular description of the life cycle of Pleistophora mulleri (Pfeiffer 1895; Georgevitch 1929)

The classification of a microsporidian parasite observed in the abdominal muscles of amphipod hosts has been repeatedly revised but still remains inconclusive. This parasite has variable spore numbers within a sporophorous vesicle and has been assigned to the genera Glugea, Pleistophora, Stempellia, and Thelohania. We used electron microscopy and molecular evidence to resolve the previous taxonomic confusion and confirm its identification as Pleistophora mulleri. The life cycle of P. mulleri is described from the freshwater amphipod host Gammarus duebeni celticus. Infection appeared as white tubular masses within the abdominal muscle of the host. Light and transmission electron microscope examination revealed the presence of an active microsporidian infection that was diffuse within the muscle block with no evidence of xenoma formation. Paucinucleate merogonial plasmodia were surrounded by an amorphous coat immediately external to the plasmalemma. The amorphous coat developed into a merontogenetic sporophorous vesicle that was present throughout sporulation. Sporogony was polysporous resulting in uninucleate spores, with a bipartite polaroplast, an anisofilar polar filament and a large posterior vacuole. SSU rDNA analysis supported the ultrastructural evidence clearly placing this parasite within the genus Pleistophora. This paper indicates that Pleistophora species are not restricted to vertebrate hosts.

Differential physico-chemical tolerances of amphipod species revealed by field transplantations

Physico-chemical regimes of river systems are major determinants of the distributions and relative abundances of macroinvertebrate taxa. Other factors, however, such as biotic interactions, may co-vary with changes in physico-chemistry and concomitant changes in community composition. Thus, direct cause and effect relationships may not always be established from field surveys. Equally, however, laboratory studies may suffer from lack of realism in extrapolation to the field. Here, we use balanced field transplantation experiments to elucidate the role of physico-chemical regime in determining the generally mutually exclusive distributions of two amphipod taxa, Gammarus (two species) and Crangonyx pseudogracilis. Within two river systems in Ireland, the former species dominate stretches of well oxygenated, high-quality water, whereas the latter dominates stretches of poorly oxygenated, low-quality water. G. pulex and G. duebeni celticus did not survive in bioassay tubes in areas dominated by C. pseudogracilis, which itself survived in tubes in such areas. However, both C. pseudogracilis and Gammarus spp. survived equally well in tubes in areas dominated by Gammarus spp. Physicochemical regime thus limits the movement of Gammarus spp. into C. pseudogracilis areas, but some other factor excludes C. pseudogracilis from Gammarus spp. areas. Since previous laboratory experiments showed high predation rates of Gammarus spp. on C. pseudogracilis, we propose that predation by the former causes exclusion of the latter. Hence, presumed effects of physico-chemical regime on macroinvertebrate presence/abundance may often require experimental field testing and appreciation of alternative explanations.

Predator-prey interactions between brown trout Salmo trutta and native and introduced amphipods; their implications for fish diets

An attempt to improve the food base for brown trout Salmo trutta in Northern Ireland was made in 1958.59 by deliberately introducing English Gammarus pulex into several Irish rivers. In addition. another amphipod Crangonyx pseudogracilis, was later accidently introduced into II ish waters. Our study represents the first attempt to examine the trophic interactions between a native fish predator (S. trutta) and an array of these native (Gammarus duebeni celticus) and introduced (G. pulex and C. pseudogracilis) amphipods. Feeding experiments, involving young brown trout predators and ampiphod prey, revealed that the fish actively selected C. pseudogracilis relative to two alternative Gammarus prey species. Although the trout encountered the Gammarus species more than C. pseudogracilis, they were eaten less than Crangonyx. Difficulties in handling and ingestion of Gammarus by trout may be a. key component of the preference fbr the smaller, more easily handled Crangonyx. The microdistribution of the species was altered by the fish, due to predation being greater in particular microhabitats, Our study showed that the introduction of the herbivorous C. pseudogracilis into Irish freshwaters may represent a useful addition to fish diets. particularly for small and/or juvenile fish. The reprecussions of the deliberate introduction of G. pulex are less clear. It may improve feeding for fish. but only if it can coexist with indigenous macroinvertebrates and thus ultimately improve the range and quantity of possible food items in predator diets. Alternatively, being highly predatory towards other macroinvertebrates including G. d. celticus and C. pseudogracilis. G. pulex may be deleterious to the diversity of the resident benthic community and hence reduce the diversity of prey available to fish predators.