Interactions between invasive and native crustaceans: differential functional responses of intraguild predators towards juvenile hetero-specifics

Intraguild predation (IGP) between invasive and native species can lead to species exclusions or co-existence, dependent on the direction and strength of the interaction. Recently, derivation of ??functional responses?? has been identified as a means of comparing the community impacts of invasive and native species. Here, we employ a novel use of this functional response methodology to evaluate any IGP asymmetries between the invasive Ponto-Caspian amphipod Echinogammarus ischnus and the North American native Gammarus fasciatus. The direction and magnitude of intraguild predation of adult males on hetero-specific adult females has previously been shown to reverse across a water conductivity gradient. This partially explains field patterns, but does not predict the co-existence of the two species observed in many habitats and locations. Here, we compared intraguild predation by both species on each other??s juveniles in high- and low- conductivity water. G. fasciatus has a higher type II functional response towards E. ischnus juveniles compared to the reciprocal interaction. Conductivity did not influence the predation rate on juveniles of either E. ischnus or G. fasciatus. Thus, the male/female IGP advantage to the native G. fasciatus in low conductivity water is compounded by a juvenile IGP asymmetry, which also counteracts the male/female IGP advantage to E. ischnus in high conductivity waters, helping to explain field patterns of exclusion and co-existence. Thus, complex asymmetries in mutual IGP associated with inherent species differences, environmental modulation, and life-history effects can help us understand and predict the population and community level outcomes of species invasions.     

Differential infection of exotic and native freshwater amphipods by a parasitic water mold in the St. Lawrence River

A parasitic water mold (oomycete) of unknown origin was discovered infecting live native and exotic freshwater gammarid amphipods (Gammarus fasciatus and Echinogammarus ischnus, respectively) in the upper St. Lawrence River. Infections were associated with rapid die-offs of natural populations of amphipods, especially the exotic E. ischnus. Analysis of sequences of three different segments of the rRNA gene cluster indicated that the parasite was in the Saprolegniaceae family, and is related to other crustacean-associated Saprolegniaceae. Specific primers were designed based on the SSU rRNA gene and utilized for semi-quantitative analysis of parasite presence in live and dead amphipods. In laboratory experiments, infection prevalence was higher in E. ischnus individuals than in native amphipods. In addition, dead E. ischnus individuals exhibited more intense infections than G. fasciatus individuals. In contrast to the Great Lakes where E. ischnus has replaced G. fasciatus at many locations, the native species remains abundant in the St. Lawrence River more than a decade after invasion by E. ischnus in the late 1990s. We hypothesize that the parasite is facilitating the co-existence of the two amphipod species by reducing the abundance of E. ischnus in environments in which it might otherwise be dominant.     

Interference competition between alien invasive gammaridean species

The relative abundances of gammaridean species in the river Rhine have profoundly changed since the invasion of Dikerogammarus villosus in 1994/1995. This study tested whether these changes in gammaridean dominance could have been determined by interspecific competition and unequal mortality, for example by intraguild predation (IGP). Single and two species tests have been carried out in aquariums provided with all substrata present in the main channel of the Rhine. Changes in substratum choice, increased swimming activity and increased mortality of a species were used as indicators of interspecific competition during interaction between gammaridean species. Interspecific competition and mortality between the most abundant invasive gammaridean species in the Rhine, viz. Gammarus tigrinus, Echinogammarus ischnus and Dikerogammarus villosus were tested. In single-species experiments, G. tigrinus and D. villosus showed similar preferences for a stony substratum, whereas E. ischnus mostly occupied the water column. The two-species aquarium experiments indicated direct interference competition for substratum and unequal mortality between G. tigrinus and D. villosus, with D. villosus being the stronger competitor. Competitive stress was influenced by population density, was size-dependent and varied between the different types of substratum due to substratum choice. G. tigrinus did not show any behaviour indicative of interference competition in the presence of E. ischnus, and neither did E. ischnus or D. villosus in the presence of any of the other gammarideans. Swimming in the water layer may already enable E. ischnus to minimise its encounters with the stone-dwelling D. villosus and G. tigrinus. To maximise the encounters between E. ischnus and D. villosus, a fish (Lepomis gibbosus) was added to occupy the water layer during the aquarium experiments. E. ischnus showed a higher mortality in the presence of both D. villosus and fish, probably due to increased stress, as shelter opportunities to escape the predators had been minimised. The study shows that interference competition between gammaridean species can explain the replacement of the North American invader G. tigrinus by D. villosus in the river Rhine. E. ischnus and D. villosus both Ponto-Caspian invaders did not show interference competition in our experiments and co-exist in the Rhine.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

Establishment of the Baikalian endemic amphipod Gmelinoides fasciatus Stebb. in Lake Ladoga

Gmelinoides fasciatus Stebb., a small amphipod from the Lake Baikal basin, was discovered in July 1988 in Lake Ladoga, the largest European lake. G. fasciatus likely invaded Lake Ladoga as a consequence of its intentional introduction, aimed at enhancing fish production, in some Karelian Isthmus lakes close to Lake Ladoga's western shore in the early 1970's. Benthos studies conducted in 1989 and 1990 revealed that G. fasciatus was well established in littoral communities along the western and northern shores of Lake Ladoga. G. fasciatus was the dominant species in these littoral communities and contributed over 70% of the macroinvertebrate biomass. The species was abundant in different macrophyte beds and stony littoral areas, both in heavily polluted and undisturbed sites. The maximum abundance and biomass of G. fasciatus was about 54,000 ind m^–2 and 160 g (wet wt.) m^–2. Negative impacts of G. fasciatus on native species have been observed. Studies are needed to identify the effects of G. fasciatus on the functioning of littoral communities and to predict and control the spread of this amphipod.     

Environmental mediation of intraguild predation between the freshwater invader Gammarus pulex and the native G. duebeni celticus

Invasive species and environmental change often occur simultaneously across a habitat and therefore our understanding of their relative roles in the decline of native species is often poor. Here, the environmental mediation of a critical interspecific interaction, intraguild predation (IGP), was examined between invasive (Gammarus pulex) and native (G. d. celticus) freshwater amphipods. In the laboratory, IGP asymmetries (males preying on congeneric females) were examined in river water sourced from zones where: (1) the invader has completely displaced the native; (2) the two species currently co-exist, and (3) the native currently persists uninvaded. The invader was always a more effective IG predator, but this asymmetry was significantly weaker moving from ‘invader-only water’ through ‘co-existence water’ to ‘native-only water’. The constituent of the water that drives this mediation of IGP was not identified. However, balancing the rigour of laboratory experiments with field derived ‘environment’ has advanced understanding of known patterns in a native species decline, and its co-existence and persistence in the face of an invader.     

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. 2. In a field survey, we show that the acanthocephalan parasite Echinorynchus truttae is present in more G. pulex populations than G. d. celticus populations. In addition, within parasitised populations, E. truttae is more prevalent in the invader than in the native. 3. We show for the first time that an acanthocephalan parasite mediates predation between its intermediate macroinvertebrate hosts. In a field experiment, E. truttae parasitism of the invader lowered IGP upon the unparasitised native. In laboratory experiments, parasitism of G. pulex significantly reduced their predatory impact on recently moulted female G. d. celticus. Parasitism also appeared to cause reduction in predatory behaviour, such as attacks per contact on precopula guarded female natives. 4. We conclude that higher parasite prevalence in invaders as compared with natives, by mediation of interspecific interactions, could promote species coexistence, or at least slow species replacements, in this particular biological invasion.     

Recent drastic changes in the gammarid fauna (Crustacea, Amphipoda) of the Vistula River deltaic system in Poland caused by alien invaders

During the last decade of 20th century, the nonindigenous gammarid species Gammarus tigrinus, Dikerogammarus haemobaphes, Pontogammarus robustoides and Obesogammarus crassus invaded the lower Vistula River and its deltaic, partly brackish regions. G. tigrinus, an oligohaline North‐American species, was introduced to western Europe in the 1950s; the remaining three species are oligohaline/freshwater Ponto‐Caspian species. All these species are now invading central and western Europe using the network of man‐made canals connecting different European river systems. In the Vistula River, the native European freshwater gammarid species Gammarus pulex and G. varsoviensis were replaced in the 1920s by the Ponto‐Caspian Chaetogammarus ischnus (syn. Echinogammarus ischnus), which in turn has been outnumbered by the more recent invasions of D. haemobaphes and P. robustoides. In brackish waters, the native Atlantic‐boreal species Gammarus zaddachi and Gammarus duebeni are replaced or at least outnumbered by G. tigrinus, P. robustoides and O. crassus. Possible invasion routes are discussed.     

Intraguild predation and species exclusions in amphipods: the interaction of behaviour, physiology and environment

1. Data from field surveys, laboratory experiments and computer simulations of community dynamics revealed that a novel interaction among intraguild predation, physiological adaptation and environment may explain the complex distributions of two putatively competing aquatic amphipods. 2. Gammarus pulex and G. tigrinus both thrive in fresh and oligohaline waters in western Europe. However, the native European G. pulex excludes the invading North American G. tigrinus from freshwaters of relatively low conductivity, whereas the reverse occurs at higher conductivities. Additionally, there is much spatio-temporal fluctuation in the patterns of coexistence of these species. 3. Laboratory experiments in The Netherlands and Ireland revealed that mutual predation of moulting individuals occurred frequently between these species. However, predation frequencies were differentially in favour of G. pulex under the ionic conditions to which this species is physiologically adapted (freshwater). On the other hand, predation was not differential under the ionic conditions to which G. tigrinus is physiologically adapted (oligohaline water). 4. A mathematical model, which extends the logistic equation to include mutual intraguild predation, simulated interactions over a range of values of relevant population parameters. This indicated that G. pulex would be excluded when balanced instantaneous rates of mutual predation were combined with the known greater reproductive output of G. tigrinus. However, this reproductive advantage is overcome by any relatively small bias in the instantaneous rate of predation favouring G. pulex, leading to the exclusion of G. tigrinus. This occurs even when the reproductive advantage to G. tigrinus is relatively large. Moreover, the model generated ‘switches’ in species dominance that are determined by the relative values of reproductive rate and mutual predation. The time taken to ‘switch’ may explain the transient periods of apparent coexistence of these species observed in the field. 5. The complex community dynamics of such species may thus be understood in terms of variation in the intensity of species interactions mediated by behavioural, physiological and environmental factors.     

Interspecific differences in drift behaviour between the native <Emphasis Type="Italic">Gammarus pulex</Emphasis> and the exotic <Emphasis Type="Italic">Gammarus roeseli</Emphasis> and possible implications for the invader’s success

“Drifting” is known to subject aquatic invertebrates to intense predation by drift feeding fish. Consequently, interspecific variations in drifting behaviour could lead to differences in predation pressure between coexisting prey species. Predation being an important factor determining the success of invaders, differences in drift patterns could advantage either native or exotic invertebrates through differential predation by native fish predators. The exotic freshwater amphipod (Gammarus roeseli) has now largely colonized Western Europe where it is often found in sympatry with a native species (Gammarus pulex). Here we documented interspecific differences in drifting behaviour that might have favored the invader’s success through differential predation. Benthic and drifting amphipods were sampled three times at the same site to compare the proportion of each species within and between sample types (benthos or drift) across time. Compared with the benthos, where the invader (G. roeseli) was significantly less abundant than the native (G. pulex), G. roeseli was proportionally overrepresented in the drift but displayed a very different drifting pattern. While G. pulex drift rates remained roughly constant over a 24 h period, G. roeseli showed a marked diel periodicity with low diurnal and high nocturnal drift rates. Such drifting behaviour could procure this species with a competitive advantage regarding predation as most drift feeding fish are diurnal. As a result, the native appears more disadvantaged with respect to drift. This may partly explain the ability of G. roeseli to coexist with G. pulex in a habitat more suitable to the native.     

Differential predatory and interference interactions between native and invasive freshwater amphipods and a co-occurring mysid (Crustacea)

Animal introductions can often have unexpected and complex consequences for both native and invader species. Freshwater crustaceans such as Gammarus spp. (amphipods) and Mysis relicta (an ‘opossum shrimp’) have frequently come into contact because of deliberate and accidental introductions. However, there remains poor understanding of mechanisms leading to the co-existence and/or exclusion among amphipods and mysids. Here, we examined predatory and interference interactions between native (Gammarus duebeni celticus) and invasive (Gammarus tigrinus and Crangonyx pseudogracilis) amphipods and the native M. relicta in Britain’s largest expanse of freshwater, Lough Neagh. Laboratory mesocosm experiments simulating near-shore/mid-lough benthic habitats showed that both Gammarus species, but not C. pseudogracilis, preyed on M. relicta, which itself preyed on C. pseudogracilis. Further, M. relicta micro-distribution and habitat use changed because of interference from G. d. celticus and to a lesser extent G. tigrinus, with C. pseudogracilis having no such impact. In smaller microcosms, predation of M. relicta adults and juveniles by Gammarus spp. was significant. Although predation of Gammarus spp. by M. relicta was low, adult C. pseudogracilis were killed by M. relicta and its predation of juvenile C. pseudogracilis was high. We discuss the concurrence of these laboratory interactions with known field patterns of co-existence amongst these species.     

Mutual predation between and cannibalism within several freshwater gammarids: Dikerogammarus villosus versus one native and three invasives

Dikerogammarus villosus, a Ponto-Caspian species, is known to be a particularly successful invader, and is currently the prevailing invasive gammarid in Southern German large waters. Field observations suggest that D. villosus has replaced the native G. pulex and the invasive D. haemobaphes, also a Ponto-Caspian species, in some reaches of the German Danube. Dikerogammarus villosus is also believed to be the reason why Echinogammarus berilloni, a Mediterranean species, and Dikerogammarus bispinosus, a further Ponto-Caspian gammaridean species, could not build significant populations in Southern German rivers. Because intraguild predation (IGP) is regarded as a major force leading to species displacements, we hypothesized that superiority in predation by D. villosus is responsible for the disappearance or limited occurrence of several native and invasive gammarideans in many reaches of Southern German waters. To test this hypothesis, we conducted a series of laboratory IGP experiments with 1:1 combinations in which D. villosus was allowed to prey on other gammarids and vice versa. We also studied the extent of cannibalism within species. Dikerogammarus villosus was a stronger predator than G. pulex and E. berilloni. However, D. villosus was not stronger in predation than its relatives D. haemobaphes and D. bispinosus, although results with the combinations of D. villosus and D. bispinosus were less conclusive, especially in the female–female combination. Cannibalism rate was high in D. haemobaphes compared with other species. We conclude that superiority in IGP experiments could explain the disappearance of G. pulex and the missing or limited occurrence of E. berilloni since the arrival of D. villosus in Southern German rivers. However, the results of IGP experiments do not explain why D. haemobaphes was displaced by D. villosus in many places and why D. bispinosus could not build significant populations in Southern German waters. Possibly specific combinations between mutual predation and cannibalism or factors other than predation could have affected displacement and occurrence patterns of Dikerogammarus species in Southern German waters.     

Testing the differential predation hypothesis for the invasion of rusty crayfish in a stream community: laboratory and field experiments

1. We tested the hypothesis that the non‐native rusty crayfish (Orconectes rusticus) is less vulnerable to predators than two native species (O. propinquus and O. obscurus) it is replacing in streams of the upper Susquehanna River catchment (New York, U.S.A.). 2. We used laboratory experiments to compare species‐specific predation rates by smallmouth bass (Micropterus dolomieu) on crayfish of equal size and field tethering experiments to compare relative predation rates between native O. propinquus and non‐native O. rusticus by the suite of crayfish predators in our system. We predicted that crayfish size would affect predation rate but that predation rates would be equal among species when size was controlled. 3. We also tested for two potential artefacts of tethering. We tethered crayfish in cages to test whether the ability to escape from tethers is size specific, and we tested whether tethering alters differential predation among crayfish species by the smallmouth bass. 4. In the laboratory, smallmouth bass predation on rusty crayfish was lower than on either of the native species. In the field, predation risk for tethered crayfish was inversely related to size but did not differ among species when size was taken into account. Because rusty crayfish in the field experiment were slightly larger than the native species, as in nature, mortality was overall lower for the rusty crayfish. 5. In cages, smaller crayfish were more probably to escape from tethers than larger ones, an artefact that may partially confound results from our tethering experiments. Unexpectedly, tethering nearly eliminated predation by smallmouth bass. This artefact prevented us from testing for an interaction of tethering with differential predation and means that the results of field tethering experiments do not include any contribution from smallmouth bass predation. 6. Our experiments highlight the importance of explicitly considering potential artefacts that could confound results. 7. Our results indicate that differential predation contributes to the rusty crayfish's invasion of a stream community. In our study system, predation rates on rusty crayfish are lower than for native species mostly because of selection by predators for smaller crayfish; species‐specific characteristics such as behaviour that further reduce predation may also contribute, especially where smallmouth bass predation is important.     

How to be an invasive gammarid (Amphipoda: Gammaroidea)–comparison of life history traits

About six life history and two ecological traits of gammarid species occurring in Central European waters were compared in order to identify the characters of successful invader. The species were (1) natives: Gammarus fossarum, G. pulex, G. lacustris, G. varsoviensis, G. balcanicus, G. leopoliensis, G. roeselii, and (2) aliens: Gammarus tigrinus, Chaetogammarus ischnus (=Echinogammarus ischnus), Pontogammarus robustoides, Obesogammarus crassus, Dikerogammarus haemobaphes, D. villosus. Generally the alien species were characterised by a combination of large brood size, high partial fecundity, early maturation and by appearance of higher number of generations per year. Also, these species presented higher tolerance towards severe environmental conditions, i.e. elevated salinity and human degradation of the environment. The above features seem to facilitate the colonisation of new areas and competition with native species -- a phenomenon that has been currently observed in various parts of Europe.     

Ethological Isolation,Sexual Behavior and Pheromones in the Fasciatus Species Group of the Lizard Genus Eumeces

Ethological isolation was found among North American members of the fasciatus species group of Eumeces. Ethological isolation was investigated by staging a series of interspecific and intraspecific heterosexual encounters. No male E. laticeps or E. fasciatus courted heterospecific females. In both species, males courted and copulated with conspecific females in a significantly higher frequency of trials than with heterospecific females. However, male E. inexpectatus courted females of all three species, courting conspecific females and female E. laticeps at similar frequencies, but female E. fasciatus at a significantly lower frequency. No females of any of the three species were sexually receptive to heterospecific males, but forced copulation occurred in two of nine courtships of female E. fasciatus by male E. inexpectatus. Thus, ethological isolation in the fasciatus group appears to be complete with the possible exception of occasional forced copulation between male E. inexpectatus and female E. fasciatus. Preliminary evidence on the role of chemical stimuli in maintaining ethological isolation was obtained from experiments involving interspecific transfers of female odors. Male E. inexpectatus courted female E. fasciatus labelled with the odor of female E. inexpectatus in a significantly greater proportion of trials than they courted such females lacking the conspecific female odor. In a similar experiment, male E. fasciatus did not court female E. inexpectatus even if the females bore odors of female fasciatus.     

Cannibalism and Interspecific Predation in a Phytoseiid Predator Guild from Cassava Fields in Africa: Evidence from the Laboratory

Interspecific predation and cannibalism are common types of interaction in phytoseiid predator guilds, but the extent and nature of these interactions have not been determined yet in phytoseiid guilds composed of African native and neotropical exotic phytoseiid predators found in cassava habitat in southern Africa. We determined in laboratory experiments the level of cannibalism and interspecific predation among the three phytoseiid mite species Euseius fustis, Iphiseius degenerans, and Typhlodromalus aripo in the absence of food and in the presence of limited or abundant quantities of two food types – Mononychellus tanajoa and maize pollen – commonly found on cassava in Africa. When confined without food, only two T. aripo females laid each two eggs within 5 days, and this species survived longer than I. degenerans and E. fustis. In the presence of con- or hetero-specific larvae or protonymphs, the three species fed more on the former than on the latter, and more on hetero-specifics than on con-specifics. Oviposition rates of the three species did not exceed 0.7 egg/female/day on con- and hetero-specific immatures. Typhlodromalus aripo and E. fustis survived longer on con-specific and hetero-specific larvae and on hetero-specific protonymphs than in the absence of any food, while T. aripo survived longer than the two other species on the same diets. Provision of limited quantity of food decreased interspecific predation rate by I. degenerans and T. aripo, but not by E. fustis, and increased oviposition rate and longevity of all three species. Provision of abundant food, however, eliminated cannibalism by all three species and further reduced interspecific predation rates, but their oviposition and longevity remained relatively unchanged compared with limited food provision. Potential consequences of cannibalism and interspecific predation among phytoseiid mites on cassava for the biological control of M. tanajoa are discussed.     

A long-term study (1949–2005) of experimental introductions to an island; freshwater amphipods (Crustacea) in the Isle of Man (British Isles)

Aim Using data from field introduction experiments with Gammarus spp. conducted in the rivers of a small island, commencing in 1949, with resampling in the 60s, 70s, 80s, 90s and finally in 2005, we aimed to examine the long‐term interaction of the native freshwater amphipod Gammarus duebeni celticus with the introduced G. pulex. Using physico‐chemical data from a 2005 island‐wide survey, we also aimed to find what environmental factors could influence the distribution of the two species. Location The Isle of Man, British Isles. Methods We used species distribution data from 10 river sites from a field introduction study in 1949–69, resampling of these sites in 1974/5, 1988, 1995 and 2005. We sampled a further 82 sites in 2005, where physico‐chemical characteristics of each site were obtained. ANOVA was used to ascertain significant differences in environmental variables between native and introduced amphipod assemblages. Multiple discriminant analysis was used to relate Gammarus distribution to environmental gradients in rivers. Results Field introductions of G. pulex into previously G. d. celticus dominated sites failed, despite the large propagule pressure in terms of numbers of individuals released and repeated introductions. The island‐wide survey revealed that G. d. celticus was found in areas of higher water quality than G. pulex. Sites where the two species co‐occurred also tended to be of lower water quality than native only sites. Continuing metal pollution from past mining activity may be a major factor excluding all amphipods from many rivers. The North American amphipod Crangonyx pseudogracilis, which was first discovered on the Isle of Man in one river site in 1995, was found in a further five sites. Main conclusions This ecological experiment, through long‐term monitoring, thus continues to elucidate factors influencing the distributions of introduced and native species. Under some physicochemical regimes, G. d. celticus is able to withstand its usual displacement by G. pulex, whereas increased river pollution could promote replacement of the native by the introduced species.     

Ranging Behavior of Proboscis Monkeys in a Riverine Forest with Special Reference to Ranging in Inland Forest

We observed a unimale group (BE-Group) of proboscis monkeys (Nasalis larvatus) comprising an α-male, 6 adult females, and several immatures from May 2005 to May 2006. We followed the group for 2014 h along the Menanggul River, Sabah, Malaysia (118°30′E, 5°30′N). Observations focused mainly on ranging behavior. We determined availability and seasonal changes in plant species consumed by the members of the group by vegetation surveys in a 2.15-ha area along 200–500 m trails in the riverine forest. During the observation period, the group ranged ≤800 m from the riverbank, within a total range of 138.3 ha. The daily path length of the group ranged from 220 to 1734 m (mean, 799 m), and daily path length correlates negatively with fruit availability. The monkeys were apt to remain within a small range in fruit-abundant seasons. Because the monkeys preferred to feed on fruits of dominant plant species in the study area, their daily path length may decrease on days when they feed on fruits. The core areas of the group’s home range were along the river because the monkeys typically returned to riverside trees to sleep. The group most often used areas that were nearer the riverbank and where the availability of fruits was higher. The most frequently used grids were the ones where the group often had sleeping sites and crossed the river. Avoiding predation may be the main reason for river crossing and selecting particular sleeping sites; hence not only food availability but also the risk of predation appears to influence the ranging of the BE-Group.     

Distribution,population structure and salinity tolerance of the invasive amphipod Gmelinoides fasciatus (Stebbing) in the Neva Estuary (Gulf of Finland,Baltic Sea)

In the early 1970s, the Baikalian amphipod Gmelinoides fasciatus (Stebbing) was intentionally introduced into several lakes in the Gulf of Finland basin in order to enhance fish production. By 1996, G. fasciatus successfully colonized the littoral zone of Lake Ladoga and, via the Neva River, invaded the Neva Bay, the freshwater part of the Neva Estuary. In 1999, G. fasciatus was first registered in the inner Neva Estuary, the very first record of the Baikalian amphipod in brackish waters of the Baltic Sea. Distribution, abundance, reproduction and population structure of G. fasciatus in the Neva Estuary were studied during 1998–2000. In some locations of the Neva Estuary, maximum densities of G. fasciatus reached 3500 ind. m⁻². In general, density and biomass of G. fasciatus in the freshwater part of the Neva Estuary were higher (around 1.5 fold) than in the brackish-water part. Fecundity of this amphipod averaged 10–20 eggs per female, depending on body size of females and season. In order to assess the possibility of further spread of G. fasciatus in the Baltic Sea, the salinity tolerance of this species was determined in a series of laboratory experiments. Our results showed that the invasive amphipod G. fasciatus is potentially able to colonize shallow coastal habitats of, for example, the Gulf of Bothnia, Gulf of Riga and other parts of the Baltic Sea with water salinities ranging from 1 to 5 psu.

     

Microhabitat selection by the invasive amphipod Echinogammarus ischnus and native Gammarus fasciatus in laboratory experiments and in Lake Erie

1. The amphipod Echinogammarus ischnus was first reported in the Laurentian Great Lakes during 1995. However, analysis of archived samples revealed the presence of the species from western Lake Erie in 1994 and possibly as early as 1993. 2. Surveys conducted in Lake Erie between 1995 and 1998 revealed that Echinogammarus was the dominant amphipod on rocks covered by Dreissena molluscs compared with those fouled by the filamentous alga Cladophora, while Gammarus fasciatus used both Dreissena and Cladophora substrata extensively. 3. In laboratory habitat selection studies, Echinogammarus chose Dreissena‐ over Cladophora‐encrusted rocks and bare rocks, while Gammarus occupied the more complex substrata equally. 4. Field colonisation experiments demonstrated that the densities of Echinogammarus and Gammarus were positively correlated when the total density of the species was low, in contrast to the large‐scale natural distribution of the species that revealed a strong inverse relationship. 5. The on‐going replacement of Gammarus by Echinogammarus in the Laurentian Great Lakes may be related to the stronger affinity of the latter for substrata fouled by Dreissena, a genus with which it co‐evolved.