Seasonal variation in the diet of Redshank Tringa totanus in the Odiel Marshes,southwest Spain: a comparison of faecal and pellet analysis

Capsule Redshank diet from southern Europe during migration shows spatial and seasonal variations.

Aims To assess seasonal variation in Redshank diet at a major passage site, and to compare data derived from analysing pellets or faeces.

Methods At the Odiel Marshes in 2001, pellets from spring migration (39), autumn migration (121) and midwinter (15) were analysed, together with faecal samples from autumn (84).

Results The abundance of different invertebrate groups in pellets varied between seasons. In spring, Chironomus salinarius pupae and larvae dominated by volume, followed by Ephydridae larvae and the beetle Paracymus aenus. Polychaetes and molluscs dominated in autumn, and isopods in midwinter. In autumn, chironomid larvae, Mesembryanthemum nodiflorum seeds and Artemia cysts were relatively more abundant in faeces, whereas polychaetes, isopods, molluscs and cestode cysticercoids were more abundant in pellets. Harder and/or larger items were thus relatively more abundant in pellets than faeces. Pellet analysis gave more emphasis to mudflat prey, and faeces to saltpan prey.

Conclusion Pellet and faecal analysis give different results for wader diet, and it is useful to combine the two methods. However, they show significant correlations both in diet range and rank abundance of prey items. Redshank diet shows much seasonal and spatial variation in southern Europe.     

Passive internal dispersal of insect larvae by migratory birds

It has long been assumed that the resistant eggs of many zooplankton are able to survive passage through the gut of migratory waterbirds, thus facilitating their dispersal between isolated aquatic habitats. We present the first evidence that such passive internal transport within birds may be relevant for insect populations. In three out of six faecal samples from black-tailed Godwits on autumn migration in southwest Spain, we found larvae of the chironomid Chironomus salinarius which had survived gut passage. Although adult chironomids can fly, they are likely to disperse greater distances when transported as larvae via birds. In insects with discrete generations, such passive transport also enables colonization of new habitats at times when flight by adults is not an option.     

Herbivory in an acid stream

• 1 Spatial and temporal variation in the distribution and feeding of non‐predatory macroinvertebrates was investigated in a first‐order, acid stream in the Ashdown Forest, southern England.
• 2 Stonefly (Nemouridae) and chironomid (Orthocladiinae) larvae were abundant on the upper surfaces of mineral substrata of three sizes (small stones, large stones, bedrock). The density of larvae in each taxonomic group did not vary among substrata of different sizes, although strong seasonal variation existed.
• 3 Nemourids and chironomids (H. marcidus) collected from the upper surfaces of substrata exhibited generalist feeding habits, consuming algae (diatoms, coccoid and filamentous green algae), detritus (biofilm matrix material and fine particulate organic matter (FPOM)) and inorganic debris.
• 4 There was spatial variation in the gut contents of nemourids. The proportion of algae in the guts of larvae often increased with the size of the substratum from which they were collected. Strong temporal variation in the composition of the diet also existed. Nemourids ingested a large quantity of attached algae and biofilm matrix from the biofilm in spring and winter, but consumed loose FPOM and associated microflora in summer and autumn.
• 5 We conclude that, in this acid stream, the trophic linkage between algae and grazers is maintained by ‘detritivorous’ stonefly and chironomid species. The relationship between the feeding habits of these larvae and other life‐history attributes, such as mouthpart morphology and mobility, is discussed.

     

Variation in prey-specific consumption rates and patterns of field co-occurrence for two larval predaceous diving beetles

Freshwater predatory insects can exert strong effects on prey, although how multiple similar predators may coexist is not well understood. Larval predaceous diving beetles are often numerically and taxonomically abundant predators in lentic systems, but the proximate mechanisms that explain their high abundance remain unknown. Field surveys were conducted twice in June in ponds in Alberta, Canada to assess the associations between larvae of two genera (Graphoderus, Rhantus), their spatial locations, and correlations with potential prey. Both larvae were common and positively correlated within wetlands although neither varied with pond depth nor distance from edge. Laboratory trials indicated that Graphoderus consumed more prey (corixids) at the surface, whereas Rhantus killed benthic prey (chironomids) and corixids at an equal rate; damselflies were the least consumed prey. Predation also varied with depth, with both larvae feeding at higher rates in the shallowest environments compared to Graphoderus at an intermediate depth. Predator–prey correlations from ponds were mostly congruent with predation trials; Graphoderus was positively correlated with corixids, Rhantus was positively correlated with corixids and chironomids; beetles were uncorrelated with damselflies. Reliance on different prey in different microhabitats may be an important mechanism for the maintenance of high abundance of dytiscid larvae.     

The effect of tubificids and chironomids on particle redistribution of lake sediment

The effects of chironomid larvae,Chironomus plumosus, and tubificid worms,Limnodrilus spp., on particle redistribution in lake sediment were investigated experimentally using pots containing sediments obtained from Lake Suwa, Japan. The chironomids and tubificids increased the water content of surface sediment. The chironomid larvae had no effect on particle size distribution, while tubificids continuously accumulated small particles on the surface sediment through their selective feeding activity. Particles larger than 0.125 mm were buried at a sediment depth of 6 cm. In Lake Suwa, long diatom frustules, large plant debris and blue-green algal flocs were found to accumulate in the deeper layer of the lake sediment inhabited by tubificids at high density.     

Patterns of macroinvertebrate abundance in inland saline wetlands: a trophic analysis

We used stable isotopes and gut-content analysis to compare trophic relations of macroinvertebrates between two types of saline wetlands in the Laramie Basin, Wyoming, USA. Amphipods (Hyallela azteca), chironomid larvae, and predatory insects (mainly Zygopteran larvae) occurred in both wetland types. However, in oligosaline wetlands (0.5–5‰ total dissolved solids) amphipods were dominant, whereas in mesosaline wetlands (5–18‰) amphipods were scarce and chironomid larvae and predatory insects were much more abundant. Salinity alone seemed inadequate to explain these differences, so we examined trophic interactions to address three main questions: (1) why are predatory insects more abundant in mesosaline wetlands, (2) why are chironomid larvae less abundant in oligosaline wetlands, and (3) why are amphipods uncommon in mesosaline wetlands? Zygopteran larvae ate mainly chironomid larvae and zooplankton, and did not eat amphipods; chironomid larvae were an average 93 dry mass of zygopteran intake. Guts of amphipods contained no animal parts and little vascular plant tissue, but rather mainly amorphous detritus. Chironomid larvae ate amorphous detritus as well as diatoms. There is potential for competition between amphipods and chironomids, but the nature of amorphous detritus in these wetlands needs further study. Because amphipods appear unavailable as prey for predatory insects, top-down impacts on chironomid larvae may be greater in oligosaline than in mesosaline wetlands. Our results suggest that (1) predatory zygopteran larvae and hemipterans are more abundant in mesosaline wetlands due to more abundant chironomid prey, (2) chironomids are less common in oligosaline wetlands because of both competition with amphipods and greater per capita consumption by predatory insects, and (3) amphipods are scarce in mesosaline wetlands because, lacking a resting stage and mode of direct dispersal, they do not cope well with extremes created by hydrologic instability in mesosaline wetlands. These mechanisms may apply to wetlands in other regions where similar patterns of invertebrate community structure have been reported.     

Interactions ofMonoporeia affinis (Lindström) (Amphipoda) with sedentary Chironomidae

Interannual population oscillations ofMonoporeia affinis and sedentary chironomids are negatively correlated in Lake Mälaren. Sedentary chironomid abundance regressed againstM. affinis density was highly significant at a lag response of one year (adj. R²=0.54, P=0.0001). The inverse correlation between Tanytarsini (Micropsectra sp. andTanytarsus sp.) andM. affinis densities supports a recruitment limitation hypothesis,viz. that the growth or survival of early instar chironomids is low at high amphipod densities. Microcosm studies showed density-dependent effects on the growth and survival of second instarCh. riparius larvae across an amphipod density gradient. No significant effects were found on growth or survival of third instars. These findings support the chironomid recruitment limitation hypothesis. Amphipod predation on early instar chironomid larvae is suggested as a population controlling mechanism.     

Feeding ecology of larval and juvenile American shad (Alosa sapidissima) in a small pond

Release ponds are used as part of a multifacet effort to restore American shad ( Alosa sapidissima ) in the upper Susquehanna River basin. Little information exists, however, on the feeding ecology of young shad in small ponds. Consequently, we examined feeding ecology and prey selection of 299 larval and 299 juvenile American shad in a small pond during spring and summer. Larval shad mainly consumed copepods (37.7%) and cladocerans (37.4%) whereas juvenile shad ate chironomids (43.1%) and ostracods (28.4%). Larval and juvenile shad exhibited diel variation in diet composition and feeding periodicity. Food consumption by shad was minimal at night; feeding activity was highest during the day, peaking at 2000 h for both larvae and juveniles. Electivity values of shad larvae for prey taxa were highest for cladocerans (+0.27) and lowest for ostracods (−0.07). Electivity values of juvenile shad were highest for chironomids (+ 0.21) and ostracods (+ 0.09), and lowest for copepods (− 0.08) and baetids (− 0.14). Our data indicate differences in diet composition, prey preference and, to a lesser extent, feeding patterns between larval and juvenile American shad in small pond environments.     

Influence of competing and predatory invertebrate taxa on larval populations of mosquitoes in temporary ponds of wetland areas in Germany

Abundances of mosquito larvae and associated invertebrate communities were assessed in 27 temporary ponds during the spring season in wetland areas of Germany. Four genera of mosquitoes were identified: Aedes, Anopheles, Culex, and Culiseta. We focused our analyses on Aedes spp. because this genus was the most abundant (92% of total abundance) and frequently encountered mosquito (present in 65% of investigated sites). The abundance of Aedes spp. was negatively associated with the abundance of competitors for food, and to a lesser extent with those of intraguild predators and strict predators. The influence of these natural antagonists on larvae of Aedes was stronger in ponds with higher levels of dissolved oxygen (53 ± 4%) than in ponds with lower levels (16 ± 1%). The overall abundance of antagonists explained 42% of the variation in abundance of Aedes spp. at sites with higher levels of dissolved oxygen. Of this explained variation, competitors accounted for 34.7%, whereas the abundance of intraguild predators and strict predators accounted for only 6.8 and 0.5%, respectively. Therefore, the promotion of competing species might be an appropriate ecological approach for the control of Aedes spp. in temporary ponds in these areas.     

Just add water: sources of chironomid drift in a large river floodplain

Although seasonal floodplains represent one of the most dynamic and productive of aquatic ecosystems, the sources of this productivity are poorly understood. We examined composition and sources of chironomid drift in the Yolo Bypass, the primary floodplain of the Sacramento River. We found that invertebrate drift during winter floodplain inundation is dominated by a single species, the newly identified chironomid Hydrobaenus saetheri (Diptera: Chironomidae). In order to determine sources of chironomids in the Yolo Bypass, invertebrates were sampled from several potential sources prior to and during initial floodplain inundation. Rehydration of dried floodplain sediments from several locations showed that H. saetheri dominated insect emergence from this colonization pathway. By contrast, H. saetheri was not a substantial component of inundated floodplain ponds or of tributary inputs to the floodplain. We conclude that the initial pulse of invertebrate abundance in Yolo Bypass floodwaters is dominated by chironomid emergence from sediments in multiple regions of the floodplain. Handling editor: S. Declerck     

Temporal and spatial distribution of chironomid larvae and oligochaetes in two Dutch storage reservoirs

Long-term changes in distribution and taxonomic composition of chironomid larvae and oligochaetes in two water storage reservoirs in The Netherlands were studied. A succession among the chironomid species was observed. In the last 10–15 years chironomid densities varied. Compared with other lakes chironomid densities were high. Densities of chironomids were higher in the littoral zone than in the profundal zone. The opposite was found for oligochaetes. Densities and composition of the chironomid fauna in the two reservoirs were similar.Procladius, Tanytarsus andHarnischia dominated at all depths. However, pupal exuviae samples showed some differences in chironomid taxonomic composition between the two reservoirs. Orthocladiinae, rarely found in bottom samples, abounded in pupal exuviae samples.     

A trophic pathway from biogenic methane supports fish biomass in a temperate lake ecosystem

Although some primary consumers such as chironomid larvae are known to exploit methane‐derived carbon via microbial consortia within aquatic food webs, few studies have traced the onward transfer of such carbon to their predators. The ruffe Gymnocephalus cernuus is a widespread benthivorous fish which feeds predominantly on chironomid larvae and is well adapted for foraging at lower depths than other percids. Therefore, any transfer of methanogenic carbon to higher trophic levels might be particularly evident in ruffe. We sampled ruffe and chironomid larvae from the littoral, sub‐littoral and profundal areas of Jyväsjärvi, Finland, a lake which has previously been shown to contain chironomid larvae exhibiting the very low stable carbon isotope ratios indicative of methane exploitation. A combination of fish gut content examination and stable isotope analysis was used to determine trophic linkages between fish and their putative prey. Irrespective of the depth from which the ruffe were caught, their diet was dominated by chironomids and pupae although the proportions of taxa changed. Zooplankton made a negligible contribution to ruffe diet. A progressive decrease in δ¹³C and δ¹⁵N values with increasing water column depth was observed for both chironomid larvae and ruffe, but not for other species of benthivorous fish. Furthermore, ruffe feeding at greater depths were significantly larger than those feeding in the littoral, suggesting an ontogenetic shift in habitat use, rather than diet, as chironomids remained the predominant prey item. The outputs from isotope mixing models suggested that the incorporation of methane‐derived carbon to larval chironomid biomass through feeding on methanotrophic bacteria increased at greater depth, varying from 0% in the littoral to 28% in the profundal. Using these outputs and the proportions of littoral, sub‐littoral or profundal chironomids contributing to ruffe biomass, we estimated that 17% of ruffe biomass in this lake was ultimately derived from chemoautotrophic sources. Methanogenic carbon thus supports considerable production of higher trophic levels in lakes.     

Temporal distribution of larval Chironomidae (Diptera) in experimental rice fields in Penang,Malaysia

The distribution and abundance of chironomid larvae were investigated in experimental rice fields at the Bukit Merah Agricultural Experimental Research Station, Penang, Northern Malaysia. The mean density of chironomid larvae during the period of the study (including two cultivation seasons) was 121 larvae m^?2. Chironomus kiiensis (48.6%) was the most abundant species, while four other species, Polypedilum trigonus (4.8%), Tanytarsus formosanus (22.6%), Tanypus punctipennis (13.9%), and Clinotanypus sp. (10.1%) occurred less frequently, although they were present during all wet phases of rice cultivation. Generally, water depth, rice plant height, conductivity, and nitrate–nitrogen content of the water only weakly influenced the abundance of the larvae. Other environmental parameters such as pH, dissolved oxygen, daytime water temperature, total suspended solids, phosphate, and sediment total organic matter did not significantly affect larval populations. The community structure of chironomids in this rice agroecosystem followed the dynamic changes of the field including agronomic practices, patterns of water availability, and phases of rice plant growth.     

Seasonal dynamics of moss-dwelling chironomid communities

Chironomid communities of mosses in a small upland stream in central Germany were highly dynamic across the year with respect to their abundance, biomass and dominant taxa. During 1988 semi-submersed mosses near a main spring and those occurring some 700 m downstream were compared with permanently submersed mosses in immediate vicinity of the downstream site. All the chironomids sampled were conspicuously small, with nearly 98% being less than 5 mm in length. A total of 65 chironomid species from 26 genera were found, with a higher diversity occurring near the source and a change in dominant taxa along the upper stream section. The mean abundance in permanently submersed mosses (250 larvae/10 cm², n = 125) was about five times higher than in semi-submersed mosses. The maximum value of 830 larvae/10 cm² (n = 1) is the highest chironomid density ever reported, which is explained by the sampling method used. The mean standing crop was also highest in permanently submersed mosses (1.5 mg AFDW/10 cm² (n = 125)), even though the highest individual value was recorded in semi-submersed mosses near the spring (10.4 mg AFDW/10 cm²). The evidence suggested that the dominance of chironomid taxa depended mainly on the location of the moss along the stream, whereas abundance and biomass were determined mainly by constancy in the ambient discharge as well as the factors influenced by this (e.g. temperature, detritus deposition). A trend was seen towards a seasonal succession among the chironomid taxa colonizing lotic mosses.     

Phenology of fleshy fruits in a wet sclerophyll forest in southeastern Australia: are birds an important influence?

Summary The hypothesis that seasonality in the production of fleshy fruits in temperate regions is the result of selection by avian seed dispersal agents or avian seed predators was tested in a temperate wet sclerophyll forest in south-eastern Australia. I determined whether ten common fleshy-fruited species produced fruits when potential avian seed dispersers were most abundant or avian seed predators were least abundant. The season of fruit production was not correlated with avian disperser abundance nor with seasonal changes in avian seed predators. Peak fruiting occurred during autumn but fruit-eating birds were equally abundant from spring through to autumn. Avian seed predators (parrots) did not show any seasonal variation in abundance. If dispersers are influencing fruiting seasonality in wet sclerophyll forest, it is likely that changes in social behaviour and feeding patterns of dispersers during autumn, rather than increases in abundance, will be an important influence on fruiting patterns in wet sclerophyll forest. However, environmental and life history factors may also influence phenology.     

The effect of biomanipulation of the zooplankton on the growth, feeding and survival of pikeperch (Stizostedion lucioperca) in nursing ponds

The following treatments were evaluated for use in rearing pikeperch larvae (Stizostedion lucioperca L.) in nursing ponds: treatment with Dipterex (Bayer); concurrent stocking with common carp (Cyprinus carpio L.); short flooding periods and prefeeding fish larvae. Treatment with Dipterex (active ingredient trichloorfon, i.e. 00-dimethyl-2,2,2, trichlorohydroxyethyl phosphonate) gave zooplankton populations dominated by rotatoria and nauplii at the onset of exogenous feeding and dominated by copepods later on. Cladocerans dominated the plankton in untreated ponds but were never prominent in treated ones. Stomach analysis revealed a fast changing feeding preference in pikeperch from nauplii and copepodites towards cladocerans and chironomids. The excellent food conditions for first feeding pikeperch larvae in Dipterex treated ponds resulted in a significant higher number of harvested fingerlings. In spite of the more favourable food conditions in untreated ponds during the second half of the culture period, a significant growth difference was not found. Concurrent carp stocking did not affect significantly the survival and growth of the pikeperch, although it decreased the accessability of chironomids for pikeperch. Short flooding periods and prefeeding pikeperch larvae failed as useful management strategies for pikeperch nursing. Pikeperch fingerlings cultured in ponds are larger than those found in natural populations at the same time, giving them a high chance to become piscivorous and to increase the year class strength when stocked in natural waters.     

Seasonally varying marine influences on the coastal ecosystem detected through molecular gut analysis

Terrestrial predators on marine shores benefit from the inflow of organisms and matter from the marine ecosystem, often causing very high predator densities and indirectly affecting the abundance of other prey species on shores. This indirect effect may be particularly strong if predators shift diets between seasons. We therefore quantified the seasonal variation in diet of two wolf spider species that dominate the shoreline predator community, using molecular gut content analyses with general primers to detect the full prey range. Across the season, spider diets changed, with predominantly terrestrial prey from May until July and predominantly marine prey (mainly chironomids) from August until October. This pattern coincided with a change in the spider age and size structure, and prey abundance data and resource selection analyses suggest that the higher consumption of chironomids during autumn is due to an ontogenetic diet shift rather than to variation in prey abundance. The analyses suggested that small dipterans with a weak flight capacity, such as Chironomidae, Sphaeroceridae, Scatopsidae and Ephydridae, were overrepresented in the gut of small juvenile spiders during autumn, whereas larger, more robust prey, such as Lepidoptera, Anthomyidae and Dolichopodidae, were overrepresented in the diet of adult spiders during spring. The effect of the inflow may be that the survival and growth of juvenile spiders is higher in areas with high chironomid abundances, leading to higher densities of adult spiders and higher predation rates on the terrestrial prey next spring.     

Loop migration in adult marsh harriers Circus aeruginosus,as revealed by satellite telemetry

Loop migration among birds is characterized by the spring route lying consistently west or east of the autumn route. The existence of loops has been explained by general wind conditions or seasonal differences in habitat distribution. Loop migration has predominantly been studied at the population level, for example by analysing ring recoveries. Here we study loop migration of individual marsh harriers Circus aeruginosus tracked by satellite telemetry. We show that despite a generally narrow migration corridor the harriers travelled in a distinct clockwise loop through Africa and southern Europe, following more westerly routes in spring than in autumn. We used the Normalized Difference Vegetation Index (NDVI) to identify potential feeding habitat in Africa. Suitable habitat seemed always more abundant along the western route, both in spring and autumn, and no important stopover site was found along the eastern route. Observed routes did thus not coincide with seasonal variation in habitat availability. However, favourable habitat might be more important during spring migration, when the crossing of the Sahara seems more challenging, and thus habitat availability might play an indirect role in the harriers’ route choice. Grid‐based wind data were used to reconstruct general wind patterns, and in qualitative agreement with the observed loop marsh harriers predominantly encountered westerly winds in Europe and easterly winds in Africa, both in autumn and in spring. By correlating tail‐ and crosswinds with forward and perpendicular movement rates, respectively, we show that marsh harriers are partially drifted by wind. Thus, we tentatively conclude that wind rather than habitat seems to have an overriding effect on the shape of the migration routes of marsh harriers. General wind conditions seem to play an important role also in the evolution of narrow migratory loops as demonstrated for individual marsh harriers.     

Saline rivers provide arid landscapes with a considerable amount of biochemically valuable production of chironomid (Diptera) larvae

Saline rivers are supposed to be ‘hot spots’ of high biological productivity in arid landscapes. To test this, we quantified the production of chironomid larvae, because river production is known to be transferred to arid landscapes primarily by birds fed on these larvae. In addition, we studied the potential biochemical quality of the larvae for birds based on the essential highly unsaturated fatty acid (HUFA) contents in their biomass. We studied species composition and measured production of chironomid larvae in two saline rivers (Volgograd region, Russia). We also evaluated the fatty acid composition and contents of the dominant taxa and estimated the flux of HUFA from the studied saline rivers to land via chironomid potential emergence. Average monthly production of chironomids measured for only 1 month, August, was quite comparable to annual production in some freshwater rivers. All the dominant chironomid larvae had comparatively high essential eicosapentaenoic acid contents, especially Cricotopus salinophilus, which showed the highest value, reported for Chironomidae. The monthly flux of HUFA from the studied rivers to land due to the chironomid potential emergence was roughly comparable to the global average estimation of annual water–land HUFA export via emerging insects.     

Population dynamic aspects of chironomid larvae of the littoral zone of Lake Maarsseveen I

Population dynamics of chironomid larvae in the littoral zone of Lake Maarsseveen I were investigated to a depth of 6 m. No essential differences in the composition and density between various sampling stations in the shallow sand flats were found. Two species,Cladotanytarsus mancus andStictochironomus sticticus, were found to be the main components of the chironomid population to a depth of 4 m. The density ofCladotanytarsus, which occurred predominantly to a depth of 1 m, increased from June to a maximum in December. In May and June, the entireCladotanytarsus population emerged.Stictochironomus had its peak of abundance in June. From this cohort, a part, differing in size for each of the various years, emerged in August. The remainder of this population, together with the offspring of the August adults, emerged in May. The size of the August midges, and the number of its eggs was smaller than that of the May midges. At depths greater than 4 m,Tanytarsus bathophillus was found to be the dominant chironomid.In summer the abundance of the small (second and third instars)Stictochironomus was primarily affected by water mite predation. In autumn and winter,Cladotanytarsus was the main food source for water mites. In autumn, whenStictochironomus larvae have molted into the fourth instar, predation by bream decreased the density of these larvae. In winter, the impact of this fish is negligible.