Artificial light at night reverses monthly foraging pattern under simulated moonlight

Mounting evidence shows that artificial light at night (ALAN) alters biological processes across levels of organization, from cells to communities. Yet, the combined impacts of ALAN and natural sources of night-time illumination remain little explored. This is in part due the lack of accurate simulations of the complex changes moonlight intensity, timing and spectra throughout a single night and lunar cycles in laboratory experiments. We custom-built a novel system to simulate natural patterns of moonlight to test how different ALAN intensities affect predator–prey relationships over the full lunar cycle. Exposure to high intensity ALAN (10 and 50 lx) reversed the natural lunar-guided foraging pattern by the gastropod mesopredator Nucella lapillus on its prey Semibalanus balanoides. Foraging decreased during brighter moonlight in naturally lit conditions. When exposed to high intensity ALAN, foraging increased with brighter moonlight. Low intensity ALAN (0.1 and 0.5 lx) had no impact on foraging. Our results show that ALAN alters the foraging pattern guided by changes in moonlight brightness. ALAN impacts on ecosystems can depend on lunar light cycles. Accurate simulations of night-time light cycle will warrant more realistic insights into ALAN impacts and also facilitate advances in fundamental night-time ecology and chronobiology.     

Dietary changes in predators and scavengers in a nocturnally illuminated riparian ecosystem

Aquatic and terrestrial ecosystems are linked by fluxes of carbon and nutrients in riparian areas. Processes that alter these fluxes may therefore change the diet and composition of consumer communities. We used stable carbon isotope (δ¹³C) analyses to test whether the increased abundance of aquatic prey observed in a previous study led to a dietary shift in riparian consumers in areas illuminated by artificial light at night (ALAN). We measured the contribution of aquatic‐derived carbon to diets in riparian arthropods in experimentally lit and unlit sites along an agricultural drainage ditch in northern Germany. The δ¹³C signature of the spider Pachygnatha clercki (Tetragnathidae) was 0.7‰ lower in the ALAN‐illuminated site in summer, indicating a greater assimilation of aquatic prey. Bayesian mixing models also supported higher intake of aquatic prey under ALAN in spring (34% versus 21%). In contrast, isotopic signatures for P. clercki (0.3‰) and Pardosa prativaga (0.7‰) indicated a preference for terrestrial prey in the illuminated site in spring. Terrestrial prey intake increased in spring for P. clercki under ALAN (from 70% to 74%) and in spring and autumn for P. prativaga (from 68% to 77% and from 67% to 72%) and Opiliones (from 68% to 72%; 68% to 75%). This was despite most of the available prey (up to 80%) being aquatic in origin. We conclude that ALAN changed the diet of riparian secondary consumers by increasing the density of both aquatic and terrestrial prey. Dietary changes were species‐ and season‐specific, indicating that the effects of ALAN may interact with phenology and feeding strategy. Because streetlights can occur in high density near freshwaters, ALAN may have widespread effects on aquatic–terrestrial ecosystem linkages.     

Niche differentiation among invasive crayfish and their impacts on ecosystem structure and functioning

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Complex impact of an invasive crayfish on freshwater food webs

Invasive alien species can have complex effects on native ecosystems, and interact with multiple components of food webs, making it difficult a comprehensive quantification of their direct and indirect effects. We evaluated the relationships between the invasive crayfish, Procambarus clarkii, amphibian larvae and predatory insects, to quantify crayfish impacts on multiple levels of food webs, and to evaluate whether crayfish predation of aquatic insects has indirect consequences for their preys. We used pipe sampling to assess the abundance of crayfish, amphibian larvae and their major predators (Ditiscidae, Notonectidae and larvae of Anisoptera) in invaded and uninvaded ponds within a human dominated landscape. We disentangled the multivariate effects of P. clarkii on different components of food web through a series of constrained redundancy analyses. The crayfish had a negative, direct impact on both amphibian communities and their predators. Amphibian abundance was negatively related to both predators. However, the negative, direct effects of crayfish on amphibians were much stronger than predation by native insects. Our results suggest that this crayfish impacts multiple levels of food webs, disrupting natural prey-predator relationships.     

Microbial diversity and community respiration in freshwater sediments influenced by artificial light at night

An increasing proportion of the Earth''s surface is illuminated at night. In aquatic ecosystems, artificial light at night (ALAN) may influence microbial communities living in the sediments. These communities are highly diverse and play an important role in the global carbon cycle. We combined field and laboratory experiments using sediments from an agricultural drainage system to examine how ALAN affects communities and alters carbon mineralization. Two identical light infrastructures were installed parallel to a drainage ditch before the start of the experiment. DNA metabarcoding indicated that both sediment communities were similar. After one was lit for five months (July–December 2012) we observed an increase in photoautotroph abundance (diatoms, Cyanobacteria) in ALAN-exposed sediments. In laboratory incubations mimicking summer and winter (six weeks each), communities in sediments that were exposed to ALAN for 1 year (July 2012–June 2013) showed less overall seasonal change compared with ALAN-naive sediments. Nocturnal community respiration was reduced in ALAN-exposed sediments. In long-term exposed summer-sediments, we observed a shift from negative to positive net ecosystem production. Our results indicate ALAN may alter sediment microbial communities over time, with implications for ecosystem-level functions. It may thus have the potential to transform inland waters to nocturnal carbon sinks.     

Species-specific differences in dynamics of agonistic interactions may contribute to the competitive advantage of the invasive signal crayfish (Pacifastacus leniusculus) over the native narrow-clawed crayfish (Astacus leptodactylus)

Species-specific differences in dynamics of agonistic interactions may influence the outcome of interspecific competition and potentially contribute to competitive advantage of one species over another. In this study, we compared the dynamics of agonistic interactions of one of the most successful crayfish invaders of European freshwater ecosystems, the signal crayfish (Pacifastacus leniusculus) and the widespread native European species currently undergoing range expansion in Croatia and Europe, the narrow-clawed crayfish (Astacus leptodactylus). Comparisons between P. leniusculus pairs and the A. leptodactylus pairs demonstrated significant differences in frequency and duration of agonistic encounters: P. leniusculus dyads engaged in fewer fights, but their duration was significantly longer. In staged interactions between size-matched interspecific pairs, agonistic behaviour of P. leniusculus individuals translated into dominance over their A. leptodactylus counterparts. This indicates that the success of P. leniusculus in agonistic encounters with the native competitor might stem from its readiness to continue fighting for a longer time period, and could lead to ecological advantages during niche competition even when facing a successful native crayfish species.     

Shelter-seeking behavior of crayfish, <Emphasis Type="Italic">Procambarus clarkii,</Emphasis> in elevated carbon dioxide

Carbon dioxide has been found to produce various negative consequences for a number of aquatic species and is projected to rise in the future for many aquatic ecosystems. Crayfish act as indicators of water quality and function as keystone species in aquatic food webs; however, there is a paucity of research on how crayfish may respond to elevated CO₂. This study quantified how shelter-seeking behavior in freshwater crayfish (e.g., family Cambaridae), a behavior critical for survival and fitness, may change following exposure to elevated pCO₂. Red Swamp crayfish (Procambarus clarkii, Girard, 1852) were exposed to one of three potential levels of dissolved CO₂ that could be found in freshwater basins currently or under future climate change conditions: Control (2 levels as the respective initial exposure. The treatment aquarium contained a shelter and was divided into three equal sections based on proximity to that shelter. Crayfish proximity to the shelter (defined by the tank sections) in the treatment aquarium was monitored every 5 s for a 2-min trial. Crayfish spent differing amounts of time in differing zones of the experiment and had different levels of activity, depending on their pCO₂ exposure; crayfish acclimated to High pCO₂ increased their time spent hiding and decreased their overall activity when compared to the Low pCO₂ and Control treatments. Augmented shelter-seeking behavior may affect crayfish social hierarchies, feeding, mating, and mortality, which could generate cascading effects on the ecology of many freshwater ecosystems.     

Bimodal annual reproductive pattern in laboratory-reared marbled crayfish

The marbled crayfish, Procambarus fallax f. virginalis, is a new research model and potent invader of aquatic ecosystems that reproduces by obligatory parthenogenesis. My data show that in captivity, it can reproduce throughout the year. However, when kept at constant 20 °C and natural photoperiod there were two prominent spawning maxima, one before the spring equinox and another one before the fall equinox. If temperature fluctuated between ~15 °C in winter and ~25 °C in summer, the first maximum was shifted beyond the spring equinox and the second maximum was shifted closer towards the fall equinox. These results indicate a bimodal annual rhythm for the reproduction in marbled crayfish that can be modulated by temperature. Comparison with P. fallax suggests that this rhythmicity was inherited from its sexually reproducing mother species. Potential consequences of my laboratory findings for wild populations of marbled crayfish in temperate and tropical regions are discussed. Since marbled crayfish can be kept in captivity for up to four years under a broad range of conditions, they offer the possibility to investigate endogenous circannual rhythms and their entrainment by environmental zeitgebers in decapod crustaceans.     

Towards a mechanistic understanding of individual-level functional responses: Invasive crayfish as model organisms

1. In novel communities, a rising number of new and emerging invasive species interact with resident species, some of which are non-native themselves. We implemented an innovative trophic interaction framework for novel communities and quantified the interaction strength and impact potential of a truly novel species (marbled crayfish Procambarus virginalis) with a resident non-native counterpart (spiny-cheek crayfish Faxonius limosus). As prey, we used Dreissena mussels, which are non-native as well and now hold a key position in many European and North American aquatic ecosystems.
2. For both crayfish species, we predicted functional responses based on a mechanistic model that we parameterised with a set of experimental observations of foraging behaviour and satiation. We compared these predicted functional responses to empirically observed responses. In addition, we incorporated behavioural traits such as aggression, activity, and boldness in the comparisons between the species and individuals to determine their influence on functional responses. We tested individuals from aquarium stocks as well as naturalised individuals from invaded water bodies.
3. Altogether, we performed 1,095 experiments with 26 individual crayfish. We found that per capita predation of spiny-cheek crayfish exceeded that of marbled crayfish from aquaria and naturalised individuals. Functional responses differed between species and were mostly higher for spiny-cheek crayfish males. Marbled crayfish, however, were more voracious and reached satiation more slowly. Consumption rates correlated with aggression for marbled crayfish and with an aggressive threat response for spiny-cheek crayfish.
4. We conclude that spiny-cheek crayfish can reach higher short-term consumption rates than marbled crayfish, but both species probably do not substantially affect Dreissena mussel populations in the field. For marbled crayfish, high long-term consumption, interspecific aggression, and reproduction rates can promote their establishment and spread. Risk assessments of these invaders should be improved by considering numerical responses, and different prey organisms and predators.

     

Differences Found in the Macroinvertebrate Community Composition in the Presence or Absence of the Invasive Alien Crayfish,Orconectes hylas

Introductions of alien species into aquatic ecosystems have been well documented, including invasions of crayfish species; however, little is known about the effects of these introductions on macroinvertebrate communities. The woodland crayfish (Orconectes hylas (Faxon)) has been introduced into the St. Francis River watershed in southeast Missouri and has displaced populations of native crayfish. The effects of O. hylas on macroinvertebrate community composition were investigated in a fourth-order Ozark stream at two locations, one with the presence of O. hylas and one without. Significant differences between sites and across four sampling periods and two habitats were found in five categories of benthic macroinvertebrate metrics: species richness, percent/composition, dominance/diversity, functional feeding groups, and biotic indices. In most seasons and habitat combinations, the invaded site had significantly higher relative abundance of riffle beetles (Coleoptera: Elmidae), and significantly lower Missouri biotic index values, total taxa richness, and both richness and relative abundance of midges (Diptera: Chironomidae). Overall study results indicate that some macroinvertebrate community differences due to the O. hylas invasion were not consistent between seasons and habitats, suggesting that further research on spatial and temporal habitat use and feeding ecology of Ozark crayfish species is needed to improve our understanding of the effects of these invasions on aquatic communities.     

Invasive Crayfish Threaten the Development of Submerged Macrophytes in Lake Restoration

Submerged macrophytes enhance water transparency and aquatic biodiversity in shallow water ecosystems. Therefore, the return of submerged macrophytes is the target of many lake restoration projects. However, at present, north-western European aquatic ecosystems are increasingly invaded by omnivorous exotic crayfish. We hypothesize that invasive crayfish pose a novel constraint on the regeneration of submerged macrophytes in restored lakes and may jeopardize restoration efforts. We experimentally investigated whether the invasive crayfish (Procambarus clarkii Girard) affects submerged macrophyte development in a Dutch peat lake where these crayfish are expanding rapidly. Seemingly favourable abiotic conditions for macrophyte growth existed in two 0.5 ha lake enclosures, which provided shelter and reduced turbidity, and in one lake enclosure iron was added to reduce internal nutrient loading, but macrophytes did not emerge. We transplanted three submerged macrophyte species in a full factorial exclosure experiment, where we separated the effect of crayfish from large vertebrates using different mesh sizes combined with a caging treatment stocked with crayfish only. The three transplanted macrophytes grew rapidly when protected from grazing in both lake enclosures, demonstrating that abiotic conditions for growth were suitable. Crayfish strongly reduced biomass and survival of all three macrophyte species while waterfowl and fish had no additive effects. Gut contents showed that crayfish were mostly carnivorous, but also consumed macrophytes. We show that P. clarkii strongly inhibit macrophyte development once favourable abiotic conditions for macrophyte growth are restored. Therefore, expansion of invasive crayfish poses a novel threat to the restoration of shallow water bodies in north-western Europe. Prevention of introduction and spread of crayfish is urgent, as management of invasive crayfish populations is very difficult.     

Intensive removal of signal crayfish (Pacifastacus leniusculus) from rivers increases numbers and taxon richness of macroinvertebrate species

Invasive species are a major cause of species extinction in freshwater ecosystems, and crayfish species are particularly pervasive. The invasive American signal crayfish Pacifastacus leniusculus has impacts over a range of trophic levels, but particularly on benthic aquatic macroinvertebrates. Our study examined the effect on the macroinvertebrate community of removal trapping of signal crayfish from UK rivers. Crayfish were intensively trapped and removed from two tributaries of the River Thames to test the hypothesis that lowering signal crayfish densities would result in increases in macroinvertebrate numbers and taxon richness. We removed 6181 crayfish over four sessions, resulting in crayfish densities that decreased toward the center of the removal sections. Conversely in control sections (where crayfish were trapped and returned), crayfish density increased toward the center of the section. Macroinvertebrate numbers and taxon richness were inversely correlated with crayfish densities. Multivariate analysis of the abundance of each taxon yielded similar results and indicated that crayfish removals had positive impacts on macroinvertebrate numbers and taxon richness but did not alter the composition of the wider macroinvertebrate community. Synthesis and applications: Our results demonstrate that non‐eradication‐oriented crayfish removal programmes may lead to increases in the total number of macroinvertebrates living in the benthos. This represents the first evidence that removing signal crayfish from riparian systems, at intensities feasible during control attempts or commercial crayfishing, may be beneficial for a range of sympatric aquatic macroinvertebrates.     

Differences in refuge function for prey and tolerance to crayfish among macrophyte species

The invasive crayfish Procambarus clarkii is an omnivore and an ecosystem engineer whose feeding mechanism has reduced the abundance of many native invertebrates and macrophytes. Since macrophytes provide refuges for aquatic insects, macrophyte depletion by crayfish might have indirect negative effects on animal prey in aquatic habitats. We postulated that the prey refuges provided by macrophytes and macrophyte tolerance to crayfish cutting and feeding vary among macrophyte species. We conducted two experiments to (1) investigate differences in macrophyte refuge function for dragonfly larvae against crayfish, and (2) test the tolerance to crayfish cutting and feeding among macrophyte species. Elodea nuttallii (submerged plant), Potamogeton crispus (submerged plant), and Carex idzuroei (emergent plant) had greater refuge effects than Trapa japonica (floating-leaved plant), an effect that might result from the larger total cover of E. nuttallii, P. crispus, and C. idzuroei, and the hardness of C. idzuroei leaves. Tolerance to crayfish cutting and feeding was greater in C. idzuroei than in the other species. As the macro-invertebrate assemblages in submerged vegetation are more abundant and species-rich than those in emergent and floating-leaf vegetation, conservation of E. nuttallii and P. crispus should be prioritized for restoring native aquatic animals in ecosystems invaded by the introduced crayfish.     

An invasive species may be better than none: invasive signal and native noble crayfish have similar community effects

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Rainfall stochasticity controls the distribution of invasive crayfish and its impact on amphibian guilds in Mediterranean temporary waters

Invasive crayfish have severely impacted invaded aquatic ecosystems worldwide. We studied temporal and spatial variation in the range expansion of the red swamp crayfish at one of the first European localities to which it was introduced: Doñana National Park (SW Spain). In contrast to the rapid range expansion witnessed in other areas, this invasive crayfish has not spread across the entire park. Instead, its distribution has expanded during wet periods, but contracted during drought periods. The red swamp crayfish has caused steep amphibian declines in other invaded areas. However, after approximately 35 years of crayfish presence in Doñana National Park, we have yet to detect a reduction in the number or occurrence of amphibian species. Amphibians may thus be protected by the large abundance of temporary ponds in the area, which provides them with an effective refuge network. We show that natural fluctuations in annual rainfall and in the number of ponds filled can temporarily eliminate invasive crayfish from particular areas. This fact should be taken into account when attempting to reduce the impact of crayfish on aquatic communities, intensifying crayfish removal during particularly dry years, when it is most effective.     

Scavenger-driven fish carcass decomposition and phosphorus recycling: Laboratory experiments with freshwater fish and crayfish

1. Fish carcass decomposition can generate fluxes of nutrients to the water column at levels comparable to other major nutrient sources. However, relatively little is known about the biological processes modulating fish decay or the rates at which carcass-bound nutrients are made available to the biota.
2. This study focused on quantifying scavenger-mediated phosphorus (P) recycling, because the availability of this essential element can regulate the trophic state of aquatic ecosystems.
3. To explore the role of some important macroscopic aquatic scavengers in P recycling, laboratory experiments were conducted where carcasses of two fish species (common bleak Alburnus alburnus; pumpkinseed Lepomis gibbosus) were offered to two size classes of black bullhead (Ameiurus melas) and two crustacean species (spiny-cheek crayfish Faxonius limosus; narrow-clawed crayfish Astacus leptodactylus).
4. Our results show that the black bullhead and the two crayfish species are highly efficient macroscopic decomposers as the P contents of scavenged carcasses were reduced at significantly higher rates compared to those of microbially decomposed (control) carcasses.
5. Pumpkinseed carcasses proved to be more resistant to rapid decomposition, as they typically lost lower proportions of their P content than bleak carcasses during the course of the experiments.
6. Scavengers sequestered a relatively large fraction (up to 33% in black bullhead and 36% in crayfish) of total carcass P in their bodies. This suggests that the consumer species used in this study can transfer/return considerable quantities of carcass-derived nutrients directly to higher trophic levels incorporated into their own tissues, and may serve as an additional, short-term sink of these nutrients.

     

Predator and detritivore niche width helps to explain biocomplexity of experimental detritus-based food webs in four aquatic and terrestrial ecosystems

In the study of food webs, the existence and explanation of recurring patterns, such as the scale invariance of linkage density, predator–prey ratios and mean chain length, constitute long-standing issues. Our study focused on litter-associated food webs and explored the influence of detritivore and predator niche width (as δ¹³C range) on web topological structure. To compare patterns within and between aquatic and terrestrial ecosystems and take account of intra-habitat variability, we constructed 42 macroinvertebrate patch-scale webs in four different habitats (lake, lagoon, beech forest and cornfield), using an experimental approach with litterbags. The results suggest that although web differences exist between ecosystems, patterns are more similar within than between aquatic and terrestrial web types. In accordance with optimal foraging theory, we found that the niche width of predators and prey increased with the number of predators and prey taxa as a proportion of total taxa in the community. The tendency was more marked in terrestrial ecosystems and can be explained by a lower per capita food level than in aquatic ecosystems, particularly evident for predators. In accordance with these results, the number of links increased with the number of species but with a significantly sharper regression slope for terrestrial ecosystems. As a consequence, linkage density, which was found to be directly correlated to niche width, increased with the total number of species in terrestrial webs, whereas it did not change significantly in aquatic ones, where connectance scaled negatively with the total number of species. In both types of ecosystem, web robustness to rare species removal increased with connectance and the niche width of predators. In conclusion, although limited to litter-associated macroinvertebrate assemblages, this study highlights structural differences and similarities between aquatic and terrestrial detrital webs, providing field evidence of the central role of niche width in determining the structure of detritus-based food webs and posing foraging optimisation constraints as a general mechanistic explanation of food web complexity differences within and between ecosystem types.     

Seasonal associations with urban light pollution for nocturnally migrating bird populations

The spatial extent and intensity of artificial light at night (ALAN) has increased worldwide through the growth of urban environments. There is evidence that nocturnally migrating birds are attracted to ALAN, and there is evidence that nocturnally migrating bird populations are more likely to occur in urban areas during migration, especially in the autumn. Here, we test if urban sources of ALAN are responsible, at least in part, for these observed urban associations. We use weekly estimates of diurnal occurrence and relative abundance for 40 nocturnally migrating bird species that breed in forested environments in North America to assess how associations with distance to urban areas and ALAN are defined across the annual cycle. Migratory bird populations presented stronger than expected associations with shorter distances to urban areas during migration, and stronger than expected association with higher levels of ALAN outside and especially within urban areas during migration. These patterns were more pronounced during autumn migration, especially within urban areas. Outside of the two migration periods, migratory bird populations presented stronger than expected associations with longer distances to urban areas, especially during the nonbreeding season, and weaker than expected associations with the highest levels of ALAN outside and especially within urban areas. These findings suggest that ALAN is associated with higher levels of diurnal abundance along the boundaries and within the interior of urban areas during migration, especially in the autumn when juveniles are undertaking their first migration journey. These findings support the conclusion that urban sources of ALAN can broadly effect migratory behavior, emphasizing the need to better understand the implications of ALAN for migratory bird populations.     

Role of Life History Strategy in the Colonisation of Western Australian Aquatic Systems by the Introduced Crayfish <Emphasis Type="Italic">Cherax destructor</Emphasis> Clark, 1936

The presence of the yabbie Cherax destructor in a number of wild aquatic systems in the Pilbara and Southwest Coast Drainage Divisions of Western Australia is documented. This is of great concern as all native freshwater crayfishes in Western Australia are endemic and restricted to the southwest, while the Pilbara Division has no native species. An introduced population of C. destructor was sampled monthly from the Hutt River (Pilbara Drainage Division) for determination of life-history and reproductive biology in a wild aquatic system in Western Australia for the first time. Proliferation in that system was attributed to specific traits including: attaining first maturity at the end of its first year of life; a protracted spawning period (July–January); relatively high mean ovarian fecundity of 210.2 (±9.24 S.E.); and a rapid growth rate (curvature parameter K = 0.78 and asymptotic orbital carapace length OCL_∞ = 51.25 mm ascertained from a seasonal von Bertalanffy growth curve) that was comparable to the larger sympatric marron Cherax cainii in this system. The life-history characteristics of C. destructor in the Hutt River were typical of many other invasive crayfish species and it has the potential to impact the unique aquatic ecosystems and the endemic freshwater crayfish species of the region.     

Plasticity in fecundity highlights the females’ importance in the spiny-cheek crayfish invasion mechanism

Invasion is one of the most consequential phenomena affecting the distribution of native species. Few in number of species, European crayfish are losing the competition with introduced North American crayfish. The spiny-cheek crayfish, Orconectes limosus, is an outstanding example, successfully competing against the native narrow-clawed crayfish, Astacus leptodactylus. For four years, we collected data regarding crayfish occurrences, their relative abundance, and the structure of populations in the ongoing colonisation process of O. limosus in the lower Danube. The mature females of both invasive and indigenous crayfish species were analysed with respect to biometry and production of oocytes in relation to the dynamics of invasion. The interspecific comparisons showed no significant differences regarding body size, with an average of approximately 102 mm total length and 31 g wet weight for both species. However, the fecundity of the indigenous species was found to be constant throughout the investigated area, whereas the number of eggs produced by the invasive females was significantly increased at the active front of the invasion. The maximum number of ovarian eggs found was 887 and 1156 in the indigenous species and the invasive species, respectively. We propose the scenario that the invasive species, which carries the deadly crayfish plague, creates an ecological advantage by reducing the populations of indigenous crayfish. Subsequently, the invasive females opportunistically use the available resources to enhance their fecundity, resulting in the acute growth of populations. However, the long-term competitiveness and colonisation success of O. limosus still remain in question.