Population dynamics of the glacial relict amphipods in a subarctic lake: role of density‐dependent and density‐independent factors

Relative role of intrinsic density‐dependent factors (such as inter‐ and intraspecific competition, predation) and extrinsic density‐independent factors (environmental changes) in population dynamics is a key issue in ecology. Density‐dependent mechanisms are considered as important drivers of population dynamics in many vertebrate and insect species; however, their influence on the population dynamics of freshwater invertebrates is not clearly understood. In this study, I examined interannual variations in the abundance of the glacial relict amphipod Monoporeia affinis in a small subarctic lake based on long‐term (2002–2019) monitoring data. The results suggest that the population dynamics of amphipods in the lake is influenced by the combined effects of both intrinsic and extrinsic factors. The reproductive success of amphipod cohorts was inversely related to its initial abundance, indicating it is influenced by density‐dependent factors. M. affinis recruitment was negatively correlated with population density and near‐bottom temperature but positively correlated with food availability, which is defined as the concentration of chlorophyll a. Multiple regression with chlorophyll, temperature, and abundance of parent cohort as independent factors explained about 80% of the variation in the reproductive success of amphipods. The negative correlation between amphipod recruitment and water temperature indicates that the current climate conditions adversely affect the populations of glacial relict amphipods even in cold‐water lakes of the subarctic zone. Results of this study can be useful in environmental assessments to separate population oscillations connected with density‐dependent mechanisms from human‐mediated changes.     

Algal epiphytes of Zostera marina: Variation in assemblage structure from individual leaves to regional scale

Data from a hierarchical study of four Zostera marina beds in Wales were used to identify the spatial scales of variation in epiphyte assemblages. There were significant within and among bed differences in assemblage structure. The differences in assemblage structure with spatial scale generally persisted when species identifications were aggregated into functional groups. There was also significant within and among bed variability in Zostera density and average length. Local variations in Zostera canopy variables at the quadrat scale (total leaf length, average leaf length and leaf density per quadrat) were not related to epiphyte species richness nor to the structure of the assemblage. In contrast, individual leaf length was significantly related to species richness in two of the beds and the structure of epiphyte assemblages was always related to individual leaf lengths. The absence of links between quadrat scale measurements of canopy variables and assemblage structure may reflect the high turnover of individual Zostera leaves. Experimental work is required to discriminate further between the potential causes of epiphyte assemblage variation within and between beds. No bed represented a refuge where a rare species was abundant. If a species was uncommon at the bed scale, it was also uncommon in beds where it occurred. The heterogeneous assemblages found in this study suggest that a precautionary approach to conservation is advisable. As much of a bed as possible should be retained, both to protect the integrity of local assemblages and to retain rare species at regional scales.     

Community structure of amphipods on shallow <Emphasis Type="Italic">Posidonia oceanica</Emphasis> meadows off Tunisian coasts

The structure, diversity and spatial distribution of the amphipod fauna associated with Posidonia oceanica meadows were studied along the Tunisian coasts in 2007. Samples were collected in nine different meadows at 2 m depth. A total of 44 species belonging to 12 families were collected. The most common species in terms of abundance were Ampithoe helleri, Hyale camptonyx and Ericthonius punctatus. The highest values of abundance and species richness and the lowest values of diversity and equitability were found in meadows with high epiphyte biomass. Multivariate analyses of data indicated that epiphyte biomass and geographical position were major determinants of the distribution and composition of amphipod assemblages along Tunisian coasts. The presence of two lessepsian amphipod species in one of the southern Posidonia oceanica meadows modified the structure of assemblage.     

EFFECT OF ROCKY INTERTIDAL AMPHIPODS ON ALGAL RECRUITMENT: A FIELD STUDY1

Evidence about the potential of mobile marine invertebrates to act as algal spore dispersal agents is presently circumstantial. Using a field correlational and experimental protocol, our study tested the hypothesis that amphipods can increase the spore recruitment of the red alga Iridaea laminarioides Bory. Iridaea laminarioides spore recruitment onto glass slides was measured at a site with high amphipod abundance and a site with low density of amphipods. To evaluate the effect of an Ulva canopy on recruitment, replicated glass slides with and without a surrounding Ulva canopy were installed at both sites. The number of I. laminarioides spores recruited on the glass slides was four to eight times higher at the high amphipod abundance site than at the low density site. However, the presence of an Ulva canopy covering the glass slides did not significantly increase the recruitment of I. laminarioides. Because the abundance of I. laminarioides, the proportion of cystocarpic plants, and the percentage of open cystocarps only differed slightly between the low and high abundance amphipod sites, we suggest that the variation in recruitment between the sites is due to the differences in amphipod abundance (and their movements) and not to differences in spore production. Moreover, the presence of I. laminarioides cystocarps showing amphipod grazing scars was significantly higher at the high amphipod density site than at the low density site.     

The importance of shore height and host identity for amphipod assemblages

The spatial distribution of organisms associated with marine intertidal macroalgae may be a direct result of their tolerance to air exposure or an indirect consequence of the distribution of their host. We compared amphipod assemblages from five intertidal macroalgae to investigate their relationship with algal identity. To test the effect of height regardless of algal characteristics, we transplanted coralline algal turfs to three different levels within the intertidal zone and compared amphipod assemblages after 1 and 14 days. Interstitial volume was positively correlated to the abundance of amphipods, suggesting that this attribute may correspond better to the potential space for their occupation when compared to algal biomass, thallus volume or the ratio between thallus and interstitial volume. Algal level determined the structure of the amphipod assemblages. Upper-level (Acanthophora spicifera and Caulerpa racemosa) and intermediate-level (coralline) algae host similar amphipod assemblages dominated by Apohyale media, but different from lower-level algae (Padina gymnospora and Sargassum cymosum), which were dominated by Hyale niger. Ten of the 15 amphipod species reported from natural communities were found in the transplanted plots. Distinct pools of amphipod species colonized coralline transplants at upper and lower levels after 1 day. However, regardless of the position on the shore, transplanted coralline turfs supported similar assemblages after 14 days, indicating that algal identity is also important for species assemblages. Our results suggest that both height on the shore and host identity combine to determine the vertical structure of amphipod assemblages in the rocky intertidal.     

Amphipods on seaweeds: partners or pests?

Summary Herbivorous marine amphipods have been implicated as important grazers on filamentous and ephemeral algae, and thus as beneficial to macrophytes in reducing overgrowth by epiphytic competitors. In North Carolina, USA, amphipods comprise 97% of all macroscopic animals inhabiting the abundant brown seaweed Sargassum filipendula, and peak in abundance between late winter and early summer. I used outdoor tank experiments to test the species-specific impact of common phytal amphipods on the growth of Sargassum and its epiphytes. The results show that seaweed-associated amphipods are a trophically diverse group that could either increase or decrease host fitness depending on their feeding preferences. The amphipods Ampithoe marcuzii, Caprella penantis, and Jassa falcata each significantly reduced growth of epiphytes on Sargassum plants relative to amphipod-free controls, while Ericthonius brasiliensis had no significant effect on Sargassum or its epiphytes. However, amphipod grazing was not necessarily beneficial to Sargassum. A. marcuzii consumed Sargassum in one outdoor tank experiment, reducing its mass by 11%, while Sargassum plants without amphipods grew by 81%. Epiphytes (mostly diatoms and the filamentous brown alga Ectocarpus siliculosus) and detritus remained abundant on these plants suggesting that A. marcuzii preferred the host to its epiphytes. Similarly, when given simultaneous access to Sargassum and to several common foliose and filamentous epiphytes in the lab, A. marcuzii ate Sargassum almost exclusively. The other three amphipods ate no macroalgae. In contrast to A. marcuzii, C. penantis consistently reduced epiphytes with no negative effect on Sargassum. Thus the species composition of the amphipod fauna can determine whether these animals increase or decrease seaweed fitness.     

Amphipod (Gammarus minus) responses to predators and predator impact on amphipod density

Recent theoretical work suggests that predator impact on local prey density will be the result of interactions between prey emigration responses to predators and predator consumption of prey. Whether prey increase or decrease their movement rates in response to predators will greatly influence the impact that predators have on prey density. In stream systems the type of predator, benthic versus water-column, is expected to influence whether prey increase or decrease their movement rates. Experiments were conducted to examine the response of amphipods (Gammarus minus) to benthic and water-column predators and to examine the interplay between amphipod response to predators and predator consumption of prey in determining prey density. Amphipods did not respond to nor were they consumed by the benthic predator. Thus, this predator had no impact on amphipod density. In contrast, amphipods did respond to two species of water-column predators (the predatory fish bluegills, Lepomis macrochirus, and striped shiners, Luxilus chrysocephalus) by decreasing their activity rates. This response led to similar positive effects on amphipod density at night by both species of predatory fish. However, striped shiners did not consume many amphipods, suggesting their impact on the whole amphipod “population” was zero. In contrast, bluegills consumed a significant number of amphipods, and thus had a negative impact on the amphipod “population”. These results lend support to theoretical work which suggests that prey behavioral responses to predators can mask the true impact that predators have on prey populations when experiments are conducted at small scales. Received: 21 March 1997 / Accepted: 15 December 1997     

Turbidity and kelp holdfast amphipoda. I. Wales and S. W. England

Analyses are given of a collection of over 65,000 amphipods derived from kelp holdfasts surveyed around Wales and S.W. England. Faunal affinity is predominantly biased latitudinally, warmwater elements being confined to the extreme southwest. Total amphipod density is related to holdfast weight. There are no significant differences in amphipod density with differences in wave exposure or turbidity. Standardized density is, however, higher in west coast than in east coast holdfasts. Diversity is lower in turbid waters due to the increased dominance of a few species. The distribution, population structure, and fecundity of the previously implicated turbidity-susceptible species, Lembos websteri Bate and Corophium bonnellii Milne Edwards are examined in the light of new ethological information. Supporting and conflicting data are examined and mechanisms are suggested to show how suspended inorganic particles could be directly detrimental to both species. In addition, indirect effects involving climatic fluctuation and competitive interactions, particularly with C. sextonae Crawford emend. Hurley and Jassa falcata (Montagu) cannot be excluded.     

Water current velocity as an environmental factor regulating the distribution of amphipod species in Gamo lagoon, Japan

The role of current velocity as an environmental factor affecting the distribution of amphipod species was investigated in the brackish Gamo lagoon, Japan. Two gammaridean amphipod species, Eogammarus possjeticus and Melita setiflagella, were found inhabiting stations of different water current with same substrates. E. possjeticus and M. setiflagella selected substrates at low and high water current, respectively. The lifestyles and anatomical features of these two free-living amphipods contributed to their adaptation to habitats with different water currents. The results of field and experimental observations suggest the existence of some type of interspecific competition between the two species. The presence of E. possjeticus caused a displacement of the fundamental niche of M. setiflagella. Received: October 10, 2000 / Accepted: August 17, 2001     

Structural variation in the brown alga Sargassum cymosum and its effects on associated amphipod assemblages.

The presence of diverse biological substrates adds complexity to coastal landscapes and increases the number of ecological niches that can be used by the mobile epifauna. Studies on the influence of structural complexity have focused mainly on algal host species, but there is little information about the influence of intraspecific structural variation on the associated mobile epifauna. In this work, we examined whether intraspecific variation in the brown alga Sargassum cymosum influenced the structure of amphipod assemblages on two shores with different wave exposure. At least 15 fronds were randomly sampled at Fortaleza and Perequê-Mirim beaches, on the Atlantic coast of S?o Paulo state, southeastern Brazil, and 12 variables were measured for each alga. The amphipods were identified and counted. The greatest structural variation in S. cymosum occurred within shores, whereas the differences between shores were mainly related to algal size. These characteristics influenced amphipod assemblages differently on each shore, with the greatest effects being associated with variables related to morphological complexity, such as holdfast size, the number and size of branches, and the extent of cover by sessile colonial animals. These findings show that monospecific algal banks are not homogeneous, and that morphological differences and interactions with other biological substrates can influence the mobile epifaunal assemblages.     

Long-term change in limnology and invertebrates in Alaskan boreal wetlands

Climate change is more pronounced at high northern latitudes, and may be affecting the physical, chemical, and biological attributes of the abundant wetlands in boreal forests. On the Yukon Flats, located in the boreal forest of northeast Alaska, wetlands originally sampled during 1985–1989 were re-sampled for water chemistry and macroinvertebrates in summer 2001–2003. Wetlands sampled lost on average 19% surface water area between these periods. Total nitrogen and most metal cations (Na, Mg, and Ca, but not K) increased between these periods, whereas total phosphorus and chlorophyll a (Chl a) declined. These changes were greater in wetlands that had experienced more drying (decreased surface area). Compared with 1985–1989, densities of cladocerans, copepods, and ostracods in both June and August were much higher in 2002–2003, whereas densities of amphipods, gastropods, and chironomid larvae were generally lower. In comparisons among wetlands in 2002–2003 only, amphipod biomass was lower in wetlands with lower Chl a, which might help explain the decline of amphipods since the late 1980s when Chl a was higher. The decline in Chl a corresponded to greatly increased zooplankton density in June, suggesting a shift in carbon flow from scrapers and deposit-feeders to water-column grazers. Declines in benthic and epibenthic deposit-feeding invertebrates suggest important food web effects of climate change in otherwise pristine wetlands of the boreal forest. Handling editor: R. Bailey     

The effect of depth and sediment type on the spatial distribution of shallow soft-bottom amphipods along the southern Portuguese coast

The present study provides updated information on amphipod assemblages of the southern Portuguese coast, the most south-western part of continental Europe. The influence of depth and sediment type on the spatial distribution of shallow soft-bottom amphipods was tested by means of univariate and multivariate methods. Samples were collected down to 40?m depth within areas ranging from very fine sand to gravel. The ratio between density and diversity was found to increase with depth and from coarser to finer sediments. Both physical variables tested were found to play a major role in structuring amphipod assemblages. However, some dominance of depth was evident as samples were mainly separated according to this variable. Moreover, while only 15 taxa were common to all depth levels, 35 were collected from all sediment types. Both depth and sediment type may override the potential effects of geographical location, as samples from different areas along the southern Portuguese coast were biologically related. A peculiar habitat was sampled in this study: a maerl bed, which was confirmed to be a hotspot of density. Nevertheless, this habitat did not harbor a distinct amphipod assemblage but some taxa such as Caprella spp., Lembos websteri, lysianassids and Gammarella fucicola were preferentially collected there. The biogeographical analysis pointed out the occurrence of 8% of Mediterranean endemic species, confirming the influence of the Mediterranean Sea beyond the Strait of Gibraltar, and the Atlantic-Mediterranean feature of the amphipod fauna of this geographically relevant area. The present results support the need to interpret the spatial distribution patterns of marine species in relation to the combined effect of depth and sediment characteristics.     

Prey selection by an intertidal beetle: field test of an optimal diet model

Summary Adult beetles Thinopinus pictus LeConte (Staphlyinidae) live on sand beaches in temporary burrows from which they emerge at night to prey on amphipods Orchestoidea calforniana (Brandt). I constructed models of amphipod size selection by beetles, using the size distributions of amphipods measured on the beach, and the results of laboratory experiments on capture success, reaction distance and feeding rates. Capture success decreased and the probability that an amphipod was detected increased with increasing amphipod size. Beetles observed during beach searches selected larger sizes of amphipods than predicted from availability and vulnerability of different sizes. To apply an optimal foraging model, I estimated the profitability of different sizes of amphipods from the number of amphipods of a given size required to satiate a beetle in the laboratory. Profitability was highest for large amphipods and lowest for small amphipods and isopods. However, amphipod abundance on the beach was always below the threshold at which specialization on larger sizes was predicted to occur.     

Responses of hyporheic invertebrate assemblages to large-scale variation in flow permanence and surface–subsurface exchange

1. Flow permanence (the proportion of time that flowing water is present) strongly influences benthic invertebrate assemblages in ephemeral and intermittent river reaches. Effects of varying flow permanence on hyporheic invertebrate assemblages are not well understood, and have not previously been studied at large spatial scales. 2. We used a 52‐km long flow‐permanence gradient in the alluvial Selwyn River, New Zealand to assess hyporheic assemblage responses to variation in flow permanence and surface–subsurface exchange. The Selwyn mainstem consists of perennial and temporary reaches embedded in longer downwelling (losing) and upwelling (gaining) sections. 3. We predicted that hyporheic invertebrate diversity, density and assemblage stability would increase with increasing flow permanence. We further predicted that assemblage structure would be influenced by the relative contribution of downwelling and upwelling water at the reach‐scale. 4. Hyporheic invertebrates were collected at 15 river cross‐sections over a 13‐month period. As predicted, hyporheic taxon richness, density and assemblage stability varied directly with flow permanence. The distribution of taxa along the flow permanence gradient appeared to be related to desiccation resistance. However, it is possible that proximity to colonist sources also contributed to distribution patterns. 5. Taxon richness was significantly higher at sites in the gaining section compared with the losing section. Sites with high flow permanence in the gaining and losing sections supported distinct hyporheic assemblages, characterised by amphipods and isopods in the gaining section, and ostracods, Hydra sp. and the mayfly Deleatidium spp. in the losing section. 6. Results of the study suggest an expansion of the scope of the Hyporheic Corridor Concept to include large hyporheic flowpaths associated with unbounded alluvial plains rivers. Hyporheic assemblages in alluvial rivers are strongly influenced by large‐scale flow permanence gradients, large‐scale surface water–groundwater exchange, and their interactions.     

Contrasting patterns of amphipod dispersion in a seagrass meadow between day and night: consistency through a lunar cycle

Changes in light intensity, typically over the course of a day, affect the dispersion of aquatic organisms at short temporal scales. Amphipods, for example, have strong behavioural responses to light conditions. In this study, we used amphipod assemblages inhabiting a Cymodocea nodosa seagrass meadow on the east coast of Gran Canaria Island (eastern Atlantic) to test whether short-term dispersion of seagrass-associated amphipods differed between day and night, testing the consistency throughout an entire lunar cycle. Replicated artificial seagrass units were deployed, and subsequently retrieved, during the day (from 8:00 am to 18:00 pm) and the night (from 18:00 pm to 8:00 am) on three consecutive days within each of the four moon phases of a complete lunar cycle. We collected 13,467 amphipods corresponding to 32 species and 17 families. Significantly larger abundances of amphipods were collected during the night through the entire moon cycle. The total abundance of amphipods was also affected by the moon phases; under full moon, larger abundances of amphipods dispersed into the artificial seagrass units followed by the third quarter, the new moon and the first quarter. The species density of amphipods per unit followed the same pattern. In conclusion, the short-term dispersion of amphipods living in a seagrass meadow was considerably greater during the night than the day, while dispersion of amphipods was more intense under full moon.     

Molecular relationships of gammaridean amphipods from Arctic sea ice

The information on the biology and ecology of the Arctic sea ice-associated amphipods (Apherusa glacialis, Gammarus wilkitzkii, Onisimus glacialis, and O. nanseni) has increased, but their molecular taxonomic information still remains undisclosed. In the present study, we investigated long-range DNA sequences spanning 18S to 28S rDNA of these four sea ice-associated amphipods and analyzed their genetic relationships with other amphipod taxa. Variations of rDNA within the individuals of the same species were not detected. Phylogenetic analyses showed that each ice amphipod was separated, forming clusters with other conspecifics. Pairwise comparisons led to similar phylogenetic results, showing that the molecular taxonomy of the ice amphipods was in accordance with morphological systematics. In addition, these findings suggest that all four amphipods have little genetic variation compared with their morphologically defined conspecifics from temperate regions. Based on DNA taxonomy, G. wilkitzkii was supported as a species in good standing, refuting a recent synonymization with Gammarus duebeni. Considerably low genetic divergences of O. glacialis and O. nanseni in 18S, ITS, and 28S rDNA suggest the presence of population distinctions within species.     

Nutrient impacts on epifaunal density and species composition in a subtropical seagrass bed

The capacity of epifauna to control algal proliferation following nutrient input depends on responses of both grazers and upper trophic level consumers to enrichment. We examined the responses of Thalassia testudinum (turtle grass) epifaunal assemblages to nutrient enrichment at two sites in Florida Bay with varying levels of phosphorus limitation. We compared epifaunal density, biomass, and species diversity in 2 m² plots that had either ambient nutrient concentrations or had been enriched with nitrogen and phosphorus for 6 months. At the severely P-limited site, total epifaunal density and biomass were two times higher in enriched than in unenriched plots. Caridean shrimp, grazing isopods, and gammarid amphipods accounted for much of the increase in density; brachyuran crabs, primary predatory fish, and detritivorous sea cucumbers accounted for most of the increase in biomass. At the less P-limited site, total epifaunal density and biomass were not affected by nutrient addition, although there were more caridean shrimp and higher brachyuran crab and pink shrimp biomass in enriched plots. At both sites, some variation in epifaunal density and biomass was explained by features of the macrophyte canopy, such as T. testudinum and Halodule wrightii percent cover, suggesting that enrichment may change the refuge value of the macrophyte canopy for epifauna. Additional variation in epifaunal density and biomass was explained by epiphyte pigment concentrations, suggesting that enrichment may change the microalgal food resources that support grazing epifauna. Increased epifaunal density in enriched plots suggests that grazers may be able to control epiphytic algal proliferation following moderate nutrient input to Florida Bay. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

PHYTOPLANKTON PIGMENTS IN COASTAL LAKE MICHIGAN: DISTRIBUTIONS DURING THE SPRING ISOTHERMAL PERIOD AND RELATION WITH EPISODIC SEDIMENT RESUSPENSION1

Phytoplankton pigment distributions during the spring isothermal periods of 1998 and 1999 and their association with episodic sediment resuspension were characterized in coastal waters of southern Lake Michigan. Total and phylogenetic group chl a concentrations (derived using chemical taxonomy matrix factorization of diagnostic carotenoids) corresponded with assemblage and group biovolumes estimated from microscopic enumeration (P≤ 0.001). Diatoms and cryptophytes dominated assemblages and together typically comprised greater than 85% of relative chl a. Total chl a concentrations and both fucoxanthin·chl a ^{? 1} and alloxanthin·chl a ^{? 1} ratios were similar across depths (P> 0.05), indicating uniform distributions of and photophysiological states for assemblages and diatoms and cryptophytes, respectively, throughout the mixed water column. Total chl a concentrations were not always spatially uniform from near‐shore to offshore waters, with the greatest variability reflecting the influence of tributary inflows upon coastal assemblages. Sediment resuspension strongly influenced water column particle density and light climate; however, total and group chl a concentrations did not correspond with coefficients of K_d and suspended particulate matter concentrations (P> 0.05). The correspondence of both light attenuation and suspended particulate matter concentration with relative diatom chl a (P≤ 0.001) indicated an apparent association between sediment resuspension and diatoms. This, and the negative association (P≤ 0.0001) between relative diatom and cryptophyte chl a, corresponded with the spatial dominance of diatom and cryptophyte chl a in near‐shore and offshore waters, respectively. The presence of viable chl a and fucoxanthin within the surficial sediment layer, established this layer as a potential source of meroplanktonic diatoms for near‐shore assemblages.     

Effect of grazing by isopods and amphipods on growth of Ulva spp. (Chlorophyta)

Eutrophication of shallow coastal waters often leads to blooms of macroalgae. Grazing by crustaceans, such as amphipods and isopods, can reduce macroalgal biomass accumulation. At the same time, growth of the macroalgae can be stimulated by epiphyte removal. The role of grazing by isopods and amphipods on Ulva spp. biomass development was investigated in the Veerse Meer, a brackish lagoon situated in the southwest Netherlands. Exclusion of grazing in the field did not stimulate Ulva spp. growth. In fact, growth rates were higher in exclosures that allowed grazers to enter. Edibility tests identified the amphipod Gammarus locusta, and the isopods Idotea chelipes and Sphaeroma hookeri as potential grazers on Ulva spp. However, when epiphytic diatoms were present on the Ulva spp. thalli, Gammarus and Sphaeroma grazed on ephiphytes and not on Ulva tissue. Only Idotea continued to graze on Ulva spp. A laboratory growth experiment revealed a positive effect of Gammarus presence on Ulva spp. growth, probably caused by preferential removal of epiphytic diatoms from the Ulva spp. thalli. The growth stimulation by epiphyte removing grazers such as Gammarus may explain the higher growth rates in the presence of grazers observed in the field. When determining the potential role of invertebrate grazers in controlling macroalgal biomass accumulation, it is important to include an assessment of the epiphyte abundance on the macroalgae, as preferential removal of epiphytes may stimulate growth and thus have the opposite effect.