Linking aquatic and terrestrial food webs – Odonata in boreal systems

1. It is increasingly realised that aquatic and terrestrial systems are closely linked. We investigated stable isotope variations in Odonata species, putative prey and basal resources of aquatic and terrestrial systems of northern Mongolia during summer. 2. In permanent ponds, δ¹³C values of Odonata larvae were distinctly lower than those of putative prey, suggesting that body tissue comprised largely of carbon originating from isotopically light carbon sources. Presumably, prey consumed during autumn and winter when carbon is internally recycled and/or methanotrophic bacteria form an important basal resource of the food web. In contrast, in a temporary pond, δ¹³C values of Odonata larvae were similar to those of putative prey, indicating that their body carbon originated mainly from prey species present. 3. Changes in δ¹⁵N and δ¹³C values between larvae and adults were species specific and reflected differential replacement of the larval isotopic signature by the terrestrial diet of adult Odonata. The replacement was more pronounced in Odonata species of permanent ponds than in those of the temporary pond, where larvae hatched later in the year. Replacement of larval carbon varied between tissues, with wings representing the larval isotopic signature whereas thoracic muscles and eggs reflected the δ¹⁵N and δ¹³C values of the terrestrial diet of adults. 4. The results suggest that because of their long larval development, Odonata species of permanent ponds carry the larval signature, which is partly replaced during their terrestrial life. Terrestrial prey forms the basis for egg production and thus the next generation of aquatic larvae. In temporary ponds, in contrast, Odonata species rely on prey from a single season, engage in a prolonged aquatic phase and hatch later, leaving less time to acquire terrestrial prey resources for offspring production. Stable isotope analysis provided important insights into the food webs of the waterbodies and their relationship to the terrestrial system.     

Experimental study on the effect of diet on fatty acid and stable isotope profiles of the squid Lolliguncula brevis

Fatty acid and stable isotope analyses have previously been used to investigate foraging patterns of fish, birds, marine mammals and most recently cephalopod species. To evaluate the application of these methods for dietary studies in squid, it is important to understand the degree to which fatty acid and stable isotope signatures of prey species are reflected in the squids' tissue. Four groups of Lolliguncula brevis were fed on prey species with distinctly different fatty acid and stable isotope profiles over 30 consecutive days. One group of squid were fed fish for fifteen days, followed by crustaceans for a further fifteen days. A second and third group were fed exclusively on fish or crustaceans for thirty days. And a fourth group was fed on a mixture of fish and crustaceans for thirty days. Analysis of squid tissue showed that, after 10 days of feeding, fatty acid profiles of squid tended to reflect those of their prey. Squid that fed on a single prey type, i.e. fish or crustacean, showed only minor modifications in fatty acid proportions after the initial change and fatty acid profiles were clearly distinguishable between the two feeding groups. Shifts in fatty acid proportions towards respective prey profiles could clearly be observed in squid the diet of which was swapped after 15 days. Clear differences could also be seen in fatty acid profiles of squid feeding on a mixed diet with trends towards either fish or crustacean fatty acid signatures. Stable isotope signatures of squid tissues clearly distinguished between animals feeding on different diets and supported findings from fatty acid analysis, thus indicating both methods to be viable tools in feeding studies on squid species.     

Aquatic Terrestrial Linkages Along a Braided-River: Riparian Arthropods Feeding on Aquatic Insects

Rivers can provide important sources of energy for riparian biota. Stable isotope analysis (δ¹³C, δ¹⁵N) together with linear mixing models, were used to quantify the importance of aquatic insects as a food source for a riparian arthropod assemblage inhabiting the shore of the braided Tagliamento River (NE Italy). Proportional aquatic prey contributions to riparian arthropod diets differed considerable among taxa. Carabid beetles of the genus Bembidion and Nebria picicornis fed entirely on aquatic insects. Aquatic insects made up 80% of the diet of the dominant staphylinid beetle Paederidus rubrothoracicus. The diets of the dominant lycosid spiders Arctosa cinerea and Pardosa wagleri consisted of 56 and 48% aquatic insects, respectively. In contrast, the ant Manica rubida fed mainly on terrestrial sources. The proportion of aquatic insects in the diet of lycosid spiders changed seasonally, being related to the seasonal abundance of lycosid spiders along the stream edge. The degree of spatial and seasonal aggregation of riparian arthropods at the river edge coincided with their proportional use of aquatic subsidies. The results suggest that predation by riparian arthropods is a quantitatively important process in the transfer of aquatic secondary production to the riparian food web.     

Nutritional composition of the preferred prey of insectivorous birds: popularity reflects quality

Food availability is emerging as a key determinant of avian occurrence and habitat use in a variety of systems, but insectivores have received less attention than other groups and the potential influence of nutritional quality has rarely been considered. Rather than a uniform food source, arthropods vary greatly in terms of nutritional composition, but does this variation translate into differential consumption? Building on previous work that demonstrated clear preference for some arthropod groups by 13 species of ground‐foraging insectivores, we compare the nutritional composition of these arthropod groups with other groups commonly encountered but seldom consumed in the same habitat types. Using samples of arthropods collected from a eucalypt woodland in southern Australia, we found the high frequency prey groups (Coleoptera, Lepidoptera, Orthoptera and Araneae) consistently contained higher fractions of crude protein and total fat than the low frequency groups (Diptera, Hymenoptera and Odonata). Even more clear‐cut differences were noted in terms of micronutrients; high frequency prey containing significantly greater concentrations of seven elements than low frequency prey and significantly greater amounts per individual arthropod for all eleven elements measured. These results indicate that the nutritional quality plays an important role in prey selection in insectivores and suggests that micronutrients may be more important determinants of prey choice than previously recognized. Integrating these findings with previous work suggesting food limitation may constrain distribution patterns of birds in fragmented landscapes, we contend that variation in nutritional quality helps explain observed patterns in insectivore diets and occurrence. In addition to explaining why smaller and more disturbed habitats are unable to support resident insectivore populations, this bottom‐up mechanism may underlie the disproportionate sensitivity of insectivores to land‐use intensification.     

Resource partitioning between Pacific walruses and bearded seals in the Alaska Arctic and sub-Arctic

Climate-mediated changes in the phenology of Arctic sea ice and primary production may alter benthic food webs that sustain populations of Pacific walruses (Odobenus rosmarus divergens) and bearded seals (Erignathus barbatus). Interspecific resource competition could place an additional strain on ice-associated marine mammals already facing loss of sea ice habitat. Using fatty acid (FA) profiles, FA trophic markers, and FA stable carbon isotope analyses, we found that walruses and bearded seals partitioned food resources in 2009–2011. Interspecific differences in FA profiles were largely driven by variation in non-methylene FAs, which are markers of benthic invertebrate prey taxa, indicating varying consumption of specific benthic prey. We used Bayesian multi-source FA stable isotope mixing models to estimate the proportional contributions of particulate organic matter (POM) from sympagic (ice algal), pelagic, and benthic sources to these apex predators. Proportional contributions of FAs to walruses and bearded seals from benthic POM sources were high [44 (17–67)% and 62 (38–83)%, respectively] relative to other sources of POM. Walruses also obtained considerable contributions of FAs from pelagic POM sources [51 (32–73)%]. Comparison of δ¹³C values of algal FAs from walruses and bearded seals to those from benthic prey from different feeding groups from the Chukchi and Bering seas revealed that different trophic pathways sustained walruses and bearded seals. Our findings suggest that (1) resource partitioning may mitigate interspecific competition, and (2) climate change impacts on Arctic food webs may elicit species-specific responses in these high trophic level consumers.     

Integrating stomach content and stable isotope analyses to elucidate the feeding habits of non-native sharptooth catfish <Emphasis Type="Italic">Clarias gariepinus</Emphasis>

Sharptooth catfish Clarias gariepinus was introduced into the Eastern Cape Province, South Africa, in 1976 and there are concerns about its possible negative impacts on native biota. This study investigated its trophic impact by examining its feeding habits. Stomach content and stable isotope analyses were compared from three localities—the Great Fish River, Sundays River and Glen Melville Dam. Stomach content analysis indicated a catholic diet dominated by fish particularly in all localities. Spatially, however, the diets revealed differences based on the dominance of macrophytes that were only present in the rivers, and aquatic invertebrates that appeared more diverse within the Great Fish River compared to other localities. By contrast, stable isotopes revealed a more generalised feeding pattern with no clear dominance of particular prey. Stable isotopes further showed that the catfish was a complex predator, with large catfish being top predators whereas smaller size groups appeared to feed lower in the food chain. An ontogenetic shift in diet was evident, with small fish predominantly consuming aquatic invertebrates and shifting towards fish with increasing size. High dietary overlap suggests the potential risk associated catfish feeding, especially the potential of piscivory by small catfish that are more likely to persist in shallow and marginal where endangered indigenous minnows occur. The alteration of environmental conditions, especially flow by inter-basin water transfer (IBWT) schemes, was inferred to have had a probable influence its invasion success. Occurrence of other invaders, which was facilitated by the IBWT together with the catfish, posits the risk of invasion meltdown within the study systems.     

Terrestrial contributions to Afrotropical aquatic food webs: The Congo River case

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Carbon Sources in Riverine Food Webs: New Evidence from Amino Acid Isotope Techniques

A nearly 40-year debate on the origins of carbon supporting animal production in lotic systems has spawned numerous conceptual theories emphasizing the importance of autochthonous carbon, terrestrial carbon, or both (depending on river stage height). Testing theories has been hampered by lack of adequate analytical methods to distinguish in consumer tissue between ultimate autochthonous and allochthonous carbon. Investigators initially relied on assimilation efficiencies of gut contents and later on bulk tissue stable isotope analysis or fatty acid methods. The newest technique in amino acid, compound specific, stable isotope analysis (AA-CSIA), however, enables investigators to link consumers to food sources by tracing essential amino acids from producers to consumers. We used AA-CSIA to evaluate nutrient sources for 5 invertivorous and 6 piscivorous species in 2 hydrogeomorphically contrasting large rivers: the anastomosing Upper Mississippi River (UMR) and the mostly constricted lower Ohio River (LOR). Museum specimens we analyzed isotopically had been collected by other investigators over many decades (UMR: 1900–1969; LOR: 1931–1970). Our results demonstrate that on average algae contributed 58.5% (LOR) to 75.6% (UMR) of fish diets. The next highest estimated contributions of food sources were from C₃ terrestrial plants (21.1 and 11.5% for the LOR and UMR, respectively). Moreover, results from 11 individually examined species consistently demonstrated the importance of algae for most fish species in these trophic guilds. Differences among rivers in relative food source availability resulting from contrasting hydrogeomorphic complexity may account for relative proportions of amino acids derived from algae.     

The importance of aquatic vegetation in beaver diets and the seasonal and habitat specificity of aquatic‐terrestrial ecosystem linkages in a subarctic environment

The extent and ecological significance of trophic linkages across ecosystem boundaries have been the subject of considerable recent research attention. North American beavers Castor canadensis engineer terrestrial influences in aquatic ecosystems by constructing terrestrial food caches near their lodges and aquatic influences in terrestrial ecosystems by building dams and flooding low lying areas. However, it is poorly resolved to what extent beavers rely on aquatic food sources and whether this reliance is greater during winter when ice cover physically confines beavers to aquatic habitats or during summer when warm, ice free water promotes the growth and accessibility of aquatic vegetation. Working in a subarctic region, we surveyed the abundance of aquatic and terrestrial food sources in and around lotic and lentic environments and estimated their contributions to beaver diets during open water and ice covered periods using carbon and nitrogen stable isotope analysis of hair samples. Ponds had four times more aquatic vegetation than streams, but terrestrial habitats around ponds had less than half as much shrub cover as habitats adjacent to streams. Beaver diets in this subarctic environment are estimated to be comprised of 60 to 80% aquatic vegetation, with beavers occupying ponds consuming more aquatic vegetation in winter than beavers occupying streams, which rely more on terrestrial shrubs cached near their lodge. Collectively, these results show how the influence of physical barriers on ecosystem linkages can be modified by habitat‐ and season‐specific abundances of preferred resources and the potential for animals to consume food in ecosystems and seasons different from where and when the food was harvested.     

Diverse diet compositions among harpaline ground beetle species revealed by mixing model analyses of stable isotope ratios

1. Species diversities of some insect lineages have been attributed to differentiation of feeding habits among species. Our objective was to determine variation in diet composition among harpaline ground beetle species occurring in a riverside grassland. 2. We examined the diet compositions of 14 species from six genera in the spring and 10 species from two genera in the autumn. We performed measurements of nitrogen and carbon stable isotope ratios in consumers and in their potential food items, and estimated relative contributions of different food items with two mixing models, IsoSource and MixSIR. 3. IsoSource and MixSIR software gave similar results, but IsoSource tended to calculate higher contributions of principal food items and smaller percentile ranges than MixSIR. Among harparine beetle species, there were diverse food utilisation patterns among four food categories (detritivorous invertebrates, herbivorous invertebrates, C3 plants, and C4 plants). Detritivores comprised the main diets of abundant harpaline species in the spring, whereas abundant harpaline species in the autumn were primarily herbivores feeding on C4 plants, or omnivores feeding on herbivorous invertebrates and C3 plants. Seasonal changes in food use were related to seasonal changes in the abundance of each food resource. 4. Mixing model analysis of stable isotope ratios is a convenient and effective method for roughly estimating diets of many species with diverse food habits (such as ground beetles). This method can contribute to determining the trophic relationships of related insects in one ecosystem.     

高营养级捕食者在浅水湖泊沿岸带与敞水区能量耦合的维持作用

研究利用保安湖沿岸带与敞水区两种生境中高营养级捕食者(达氏鲌与红鳍原鲌)的碳(δ13C)、氮(δ15N)稳定同位素数据,通过稳定同位素质量平衡混合模型、非度量多维尺度分析(NMDS)等方法,分析了两种生境中鲌类食物来源的差异.结果表明,两种生境中鲌类食物来源基本一致,且食物来源较为广泛,包括沿岸带饵料鱼类、敞水区饵料鱼...     

Simulated eutrophication and browning alters zooplankton nutritional quality and determines juvenile fish growth and survival

The first few months of life is the most vulnerable period for fish and their optimal hatching time with zooplankton prey is favored by natural selection. Traditionally, however, prey abundance (i.e., zooplankton density) has been considered important, whereas prey nutritional composition has been largely neglected in natural settings. High‐quality zooplankton, rich in both essential amino acids (EAAs) and fatty acids (FAs), are required as starting prey to initiate development and fast juvenile growth. Prey quality is dependent on environmental conditions, and, for example, eutrophication and browning are two major factors defining primary producer community structures that will directly determine the nutritional quality of the basal food sources (algae, bacteria, terrestrial matter) for zooplankton. We experimentally tested how eutrophication and browning affect the growth and survival of juvenile rainbow trout (Oncorhynchus mykiss) by changing the quality of basal resources. We fed the fish on herbivorous zooplankton (Daphnia) grown with foods of different nutritional quality (algae, bacteria, terrestrial matter), and used GC‐MS, stable isotope labeling as well as bulk and compound‐specific stable isotope analyses for detecting the effects of different diets on the nutritional status of fish. The content of EAAs and omega‐3 (ω‐3) polyunsaturated FAs (PUFAs) in basal foods and zooplankton decreased in both eutrophication and browning treatments. The decrease in ω‐3 PUFA and especially docosahexaenoic acid (DHA) was reflected to fish juveniles, but they were able to compensate for low availability of EAAs in their food. Therefore, the reduced growth and survival of the juvenile fish was linked to the low availability of DHA. Fish showed very low ability to convert alpha‐linolenic acid (ALA) to DHA. We conclude that eutrophication and browning decrease the availability of the originally phytoplankton‐derived DHA for zooplankton and juvenile fish, suggesting bottom‐up regulation of food web quality.     

A comparison of digestive efficiency in birds of prey

Differences in how raptors hunt and what they catch are correlated with size-independent differences in length of the small intestine, the region of the digestive tract responsible for food absorption. The study examined the functional significance of these differences among ten raptor species. Dry matter apparent digestive efficiency was calculated for each species fed a diet of day-old cockerels. For Falconiformes and Strigiformes, efficiencies varied between 75% and 82%, digestive efficiency being positively correlated with intestine length.
Generalist species, with a wide prey spectrum and feeding on relatively easily caught prey and carrion, had long intestines and high digestive efficiencies. Raptors specializing on fast-moving, avian prey had short intestines and reduced digestive efficiency. The Peregrine Falco peregrinus and the Common Buzzard Buteo buteo were used as examples of specialist and generalist feeders, respectively. Rabbit and pigeon were fed to both. Buzzards digested both diets more efficiently than Peregrines. Body-mass changes were used to examine the nutritional value of the two diets to each species. Buzzards gained body-mass when eating rabbit, Peregrines lost mass. Both species gained mass when feeding on pigeon. It seems that consideration of diet quality, not just quantity, is essential in understanding raptor food requirements. Generalist raptors have high efficiencies on several diets, specialists compensate for their reduced efficiency by eating food of high nutritional quality. Various aspects of prey quality are examined.     

Food sources and lipid retention of zooplankton in subarctic ponds

1. Subarctic ponds are seasonal aquatic habitats subject to short summers but often have surprisingly numerous planktonic consumers relative to phytoplankton productivity. Because subarctic ponds have low pelagic productivity but a high biomass of benthic algae, we hypothesised that benthic mats provide a complementary and important food source for the zooplankton. To test this, we used a combination of fatty acid and stable isotope analyses to evaluate the nutritional content of benthic and pelagic food and their contributions to the diets of crustacean zooplankton in 10 Finnish subarctic ponds. 2. Benthic mats and seston differed significantly in total lipids, with seston (62.5 μg mg^?1) having approximately eight times higher total lipid concentrations than benthic mats (7.0 μg mg^?1). Moreover, the two potential food sources differed in their lipid quality, with benthic organic matter completely lacking some nutritionally important polyunsaturated fatty acids (PUFA), most notably docosahexaenoic acid and arachidonic acid. 3. Zooplankton had higher PUFA concentrations (27–67 μg mg^?1) than either of the food sources (mean benthic mats: 1.2 μg mg^?1; mean seston: 9.9 μg mg^?1), indicating that zooplankton metabolically regulate their accumulation of PUFA. In addition, when each pond was evaluated independently, the zooplankton was consistently more ¹³C‐depleted (δ¹³C ?20 to ?33‰) than seston (?23 to ?29‰) or benthic (?15 to ?27‰) food sources. In three ponds, a subset of the zooplankton (Eudiaptomus graciloides, Bosmina sp., Daphnia sp. and Branchinecta paludosa) showed evidence of feeding on both benthic and planktonic resources, whereas in most (seven out of 10) ponds the zooplankton appeared to feed primarily on plankton. 4. Our results indicate that pelagic primary production was consistently the principal food resource of most metazoans. While benthic mats were highly productive, they did not appear to be a major food source for zooplankton. The pond zooplankton, faced by strong seasonal food limitation, acquires particular dietary elements selectively.     

Trophic structure of coastal freshwater stream fishes from an Atlantic rainforest: evidence of the importance of protected and forest-covered areas to fish diet

The role of riparian forests in the functioning of aquatic ecosystems is well known, and they are recognized as an important food source for riverine fauna. This study investigates the trophic structure of coastal freshwater stream fishes from a large conservation area in an Atlantic rainforest using stomach content and food availability analyses. Four samples were collected from 19 sample sites. Fishes were caught with electrofishing. Prey were sampled with trays, Surber, traps, and electrofishing to evaluate the availability of food resources. The diets of 20 fish species were determined from the stomach contents of 1691 individuals. Terrestrial and aquatic insects and detritus were the most consumed items. Fish diet and prey availability were not seasonally dependent. A cluster analysis showed five trophic functional groups: terrestrial insectivores, aquatic insectivores, detritivores, carnivores, and omnivores. Insectivores predominated in species richness (60%), abundance (47%) and biomass (39%). Allochthonous and autochthonous items were found in similar proportions in the environment; however, allochthonous items were representative for insectivores and detritivores, whereas autochthonous items were important for primarily aquatic insectivores. The preference for certain insects by insectivorous fishes was associated with food selectivity rather than the availability of the resource and demonstrated the strong relationship between feeding behavior and food preference. The absence of seasonal variation in the diets of the fishes was possibly related to the consistent food supply. Our results confirm the role of the forest as a food provider for stream fishes, such as terrestrial insects and plant debris/detritus (also consumed by aquatic insects, which subsequently serve as food for fish), highlighting the importance of conserving the Brazilian Atlantic rainforests.     

Diet diversity among five co-existing fish species in a tropical river: integration of dietary and stable isotope data

Diets were estimated from stomach contents and the MixSIR model using stable isotope values for five co-existing and abundant benthic fishes in relation to potential prey from a riffle habitat in a tropical river in eastern Thailand. Collectively, aquatic insects were of greatest dietary importance based on stomach contents and, except for one fish species, predicted from the MixSIR. The most prominent functional feeding insect groups in fish diets were collector-filterers and scrapers and, to a lesser extent, predators. MixSIR predicted shrimp to be the most important single dietary constituent for all fishes in contrast to stomach contents, which indicated they are a major item for only one species. MixSIR predicted plant material to be more important in fish diets than stomach contents where the composition of detritus was a concern. Differences in temporal feeding schedules, prey availability, species adaptations and others are important in understanding diet and in the construction of food webs. Stable isotope and dietary analysis provides a more accurate assessment of the food web structure and dynamics of tropical river ecosystems than either method alone.     

Feeding differences among common littoral mysids, <Emphasis Type="Italic">Neomysis integer,Praunus flexuosus</Emphasis> and <Emphasis Type="Italic">P. inermis</Emphasis>

The energy flow in aquatic food webs and their structures are largely determined by food utilisation of predators. Mysid shrimps are important predators in various aquatic ecosystems. We studied the stomach contents of three common littoral mysids from the Baltic Sea. The aim was to study whether the diets differ between species and size classes inhabiting shallow coastal areas. The effects of season (spring, summer, autumn) and habitat were also explored. Results showed that all species were highly omnivorous, utilising various phyto- and zooplankton prey, algal filaments and dead organic material through the growing season. No ontogenetic diet shifts were observed although different size classes preferred slightly different prey. The amount of detritus increased in the diets during growth. In addition, large mysids ate more macro- and less microzooplankton compared with the small ones. There were also species-specific differences in the food utilisation. Neomysis integer ate more benthic material, Praunus flexuosus more macrozooplankton and P. inermis more phytoplankton compared with the others. These differences reflect microhabitat differences and probably also size differences of the studied species. Seasonal variation was also observed in the diets. Food utilisation followed the changes in the food availability, e.g. phytoplankton spring bloom and zooplankton peak abundances in late summer. Results confirm the omnivorous nature of mysids showing the importance of a diversity of prey as energy sources during growth.     

Feeding Responses of Hatchery-Reared Gilthead Sea Bream (Sparus aurata L.) to a Commercial Diet and Natural Prey Items

Many fish species have evolved feeding mechanisms and behaviours enabling them to feed on specific prey. However, such mechanisms may not be optimal for feeding on commercial-pelleted diets in aquaculture. Gilthead sea bream chew and occasionally eject pellets or parts of pellets from the mouth when feeding on commercial diets. This may result in an increase in nutritional waste from the intensive culture of this species. In this study we examined the prevalence of this food processing behaviour in two sizes of sea bream, feeding on three types of natural prey items in comparison to a commercial pellet, to give an insight into the circumstances in which excess chewing and ejection of food items from the mouth occurred. These included two hard-textured food items (commercial pellet and hard-shelled prey) and two soft-textured food items (larvae and small crustacean). Both sizes of sea bream frequently consumed the soft-textured food types, however large sea bream also frequently consumed hard-textured pellets. Hard-textured food required longer handling times and elicited more chewing and the ejection of food items from the mouth. These results suggest that future investigations on the food processing behaviour and consequent waste when fed commercial diets differing in texture could give an insight into improving diets and feeding efficiency for intensively cultivated gilthead sea bream.     

Feeding ecology and basal food sources that sustain the Paradoxal frog Pseudis minuta: a multiple approach combining stomach content,prey availability,and stable isotopes

In the present study, we investigated ontogenetic diet shifts, feeding strategy, prey preferences, and basal food sources that sustain the Paradoxal frog (Pseudis minuta) based on stomach content, prey availability, and stable isotope (δ¹³C, δ¹⁵N) approaches. The feeding strategy analysis showed that the population can be considered a generalist species with each individual displaying a marked opportunism for different preys. Trophic positions estimated using nitrogen isotopic ratio (δ¹⁵N) revealed that tadpoles are primary consumers, but post-metamorphic individuals shifted to secondary and tertiary trophic levels as they increase in body size. A stable isotopic mixing model revealed that most of the carbon (0.61–0.72) sustaining the post-metamorphic P. minuta is derived from the aquatic rather than the adjacent terrestrial environment. This finding suggests that the post-metamorphic P. minuta is strongly dependent on carbon sources that primarily originate in aquatic systems, regardless of the terrestrial or aquatic origins of the arthropods in its diet. Our results indicated that this species is a generalist-opportunistic predator that derives most of their carbon sources from the aquatic environment where it shows preference for aquatic preys with higher individual biomasses.