The brackishwater hyporheic zone: invertebrate community structure across a novel ecotone

This study describes a hyporheic zone that exists beneath a river-dominated estuary in North Wales, and which spans the freshwater/saltwater boundary. A series of 72 cores was taken from the Aber Estuary, in September, at depths from 10–60 cm below the bed surface. Site 1 was above the extreme high water mark and therefore was never inundated by salt water. Site 5 was the furthest downstream and was inundated twice daily by incoming tides. Substrates contained more gravel at the upstream sites and closer to the bed surface, with more sand and silt at downstream sites and at depth. Salinity of interstitial water increased both with distance away from Site 1 and with increasing depth into the zone, reaching a maximum of 15‰ at 60 cm at Site 5, as measured at low tide. Longitudinal and vertical zonations of the invertebrate taxa were evident, and densities were greater in the top 40 cm than below. Nematodes were especially abundant from 10–60 cm, and genera differed along the estuary. Oligochaetes were well represented at most sites, with the less saline sites yielding Naididae, Enchytraeidae, Aeolosomatidae and Tubificidae; Site 5 was dominated by Tubifex costatus. Chironomid larvae were most abundant to 40 cm at Sites 1 and 2 (primarily Brillia modesta, Corynoneura sp., Tanytarsus sp., and unidentified Tanypodinae), but they were also found at Site 3 (to 60 cm), and at 10 cm at Site 5 (Orthocladius sp.). Nymphs of mayflies and caddisfly larvae were found primarily at Site 1, but small stonefly nymphs and elmid beetle larvae were taken at Site 2 depths where the interstitial water was brackish. The most saline sediments were populated by the triclad Uteriporus vulgaris, the polychaete Nereis sp., and snails (Hydrobia sp.). Site, salinity, pH, and sand and silt contents appeared to influence species distribution most; together, these explained 37.8% of the variance (CANOCO). There was an overall negative relationship between silt content of the sediments and total invertebrate density. The hyporheic zone of the Aber Estuary is thus not only influenced by surface water and, presumably, estuarine groundwater exchanges, but is further complicated, biologically, by having a salinity gradient running through it. The term `brackishwater hyporheic zone' (BHZ) is proposed to describe this and similar systems.     

Fish use of an inundated saltmarsh flat in a temperate Australian estuary

Abstract Fish associated with Australian saltmarshes have previously been sampled from within creeks or semi-permanent pools on the marsh; this is the first record of fish use of marsh flats inundated on high tides but emergent on low tides. Buoyant pop nets (9 m², 1 mm mesh) were used to collect fish from two areas of the marsh flat on Torrens Island in the Barker Inlet-Port River estuary. South Australia. Fyke nets of the same mesh size were used to collect fish from small creeks draining the marsh flat. Fish were caught in only four of 48 pop net releases, for a total of 19 fish of two species (families Gobiidae and Atherinidae). A total of 1109 fish from six species were caught in creeks draining the same areas, and the two species caught on the flats were the most common species in the creeks. Economically important species were poorly represented, with just four specimens of the mugilid Aldrichetta forsteri (Valenciennes).     

Biodiversity loss in Appalachian headwater streams (Kentucky,USA): Plecoptera and Trichoptera communities

Government and academic studies indicate that many streams in the Appalachian Mountains have degraded biological communities stemming from a variety of regional landuses. Headwater stonefly (Plecoptera) and caddisfly (Trichoptera) assemblages were assessed between 1999 and 2004 in relation to pervasive landuse disturbances (coal mining and residential) in mountainous areas of eastern Kentucky, USA. Indicator metrics (richness, abundance, tolerance, and an observed/expected (O/E) null model) were compared among 94 sites with different land use pressures including least disturbed reference, residential, mining, and mixed mining and residential categories. Thirty-three stonefly species from 26 genera and 9 families were identified; Leuctra, Acroneuria, Haploperla, and Isoperla comprised the core genera that commonly decreased with disturbance. Caddisflies were represented by 48 species, 32 genera, and 14 families. Core caddisfly genera (Neophylax, Pycnopsyche, Rhyacophila, Lepidostoma, and Wormaldia) were extirpated from most disturbed sites. Species richness was significantly higher at reference sites and reference site mean tolerance value was lowest compared to all other categories; relative abundance of both orders was variable between disturbance groups. Non-metric multidimensional scaling (for riffle-dwelling stonefly and caddisfly genera) clustered reference sites distinctly from most other sites. The O/E index was highly correlated with individual habitat and chemical stressors (pH, conductivity) and on average, it estimated ~ 70% loss of common stoneflies and caddisflies across all disturbed landuse categories. Expected plecopteran and trichopteran communities were radically altered in streams draining mining and residential disturbance. Long-term impacts incurred by both landuses will continue to depress these vulnerable indigenous fauna.     

Insect community organisation in estuaries: the role of the physical environment

Insects are reportedly uncommon in marine habitats and, from a spatial/temporal intercomparison perspective, estuaries are among the least studied. We examined the natural variability seen among insect community organisation in estuaries on both sides of the North Atlantic, and evaluated the role of their physical environments. Community composition was found to be strongly influenced by three physical factors: estuary size, the degree of inundation by incoming tides, and substrate size/stability. Insects formed a significant proportion (17–54%, by numbers) of the benthic community of coarse-grained-substratum estuaries, and species richness increased with estuary size. Nymphs/larvae of mayflies, stoneflies, caddisflies, elmid beetles and chironomids dominated channel sites inundated by up to 25% of all incoming tides, but a gradual loss in species richness occurred downstream. However, even the most seaward sites supported high insect densities (up to 25 016 and 5433 m⁻², supporting 26 and 4 species, at sites inundated by 75 and 100% of all incoming tides, respectively). Sites covered by tides for between 3 and 5 h twice daily were dominated by orthocladine chironomids, especially of the genus Orthocladius . Chironomid larvae contribute significantly to the diets of some coastal fish species, particularly juvenile flounder and sticklebacks. We present a schematic model summarising the relationships between estuary size, degree of inundation by salt water and insect community structure.     

Life cycle and production of Skwala parallela (Frison) (Plecoptera: Perlodidae) in a Colorado montane stream

The life cycle and production of Skwala parallela, a perlodid stonefly, was investigated in a third-order Colorado montane stream. The species exhibited a univoltine life cycle with a distinct cohort. Small nymphs appeared in May. Rapid growth was exhibited throughout summer and autumn. During winter, growth slowed somewhat but was continuous until April. Maximum density of 34 nymphs/m² occurred in July. Based upon the instantaneous growth method, annual production was 395.3 mg/m² or 3.95 kg/ha dry weight with a P/B ratio of 4.4     

Trematodes and nematodes parasitizing the benthic insect community of an Andean Patagonian stream,with emphasis on plagiorchiid metacercariae

In freshwater systems, parasitological studies have mainly been carried out on vertebrates and molluscs, but little is known about parasites of aquatic insects. We describe the trematodes and nematodes parasitizing the benthic insects of an Andean Patagonian stream and the presence of parasites in the terrestrial adult stages. Members of 3 of 20 insect taxa were found to be parasitized by larval nematodes, and members of six taxa harbored metacercariae of digeneans. In benthic samples, chironomids, simuliids (Order Diptera), and baetids (Order Ephemeroptera) harbored mermithid larvae (Nematoda). The stonefly Antarctoperla michaelseni (Order Plecoptera), the caddisfly Smicridea annulicornis (Order Trichoptera), a watersnipe fly (Order Diptera: Athericidae), and three species of leptophlebiid mayflies (Order Ephemeroptera) were parasitized by encysted plagiorchiid metacercariae (Order Plagiorchiida). Most metacercariae were found in the three species of mayflies with prevalences ranging 15–63% and mean intensities ranging 1.2–4.9. Prevalence declined from summer to early winter, probably because of the emergence of infected nymphs and the recruitment of uninfected new cohorts. The imagos had live metacercariae with higher prevalences and intensities of infection than nymphs. We suggest that these plagiorchiids have an allogenic life cycle, involving a terrestrial definitive host.     

Effects of a forest disturbance on shredder production in southern Appalachian headwater streams

• 1 The effects of a forest disturbance were investigated by comparing production of leaf-shredding aquatic insects in three streams draining a mature hardwood forest and three streams draining an 11-year-old, cable-logged clearcut.
• 2 Reference streams contained significantly greater mean annual standing crop of leaf material and significantly more slow-processing leaf material than disturbed streams. Disturbed streams had a significantly higher mean annual standing crop of fast-processing leaf material than the reference streams.
• 3 Leaf-shredding cranefly (Tipula abdominalis), caddisfly (Pycnopsyche gentilis), and stonefly (Tallaperla maria) larvae comprised over 95% of shredder biomass in all streams. Total shredder production was significantly greater (P<0.05) in disturbed versus reference streams, but individual production rates were not significantly different between stream types.
• 4 Pycnopsyche gentilis larvae were present at higher densities and achieved significantly greater annual biomass in disturbed versus references streams, Biomass of P. gentilis was significantly correlated with the standing crop of fast-processing, early successional leaf material in samples, whereas biomass of other shredders was correlated significantly with medium or slow-processing leaf species characteristic of later stages of forest succession.

     

The biology of Bothrometopus elongatus (Coleoptera, Curculionidae) in a mid-altitude fellfield on sub-Antarctic Marion Island

Bothrometopus elongatus is one of four Ectemnorhinus-group species restricted to the epilithic biotope on the Prince Edward Islands. Here we examine the biology of this species over a full year at Kerguelen Rise, a mid-altitude fellfield site on Marion Island. B. elongatus adults eclose from April onwards, reaching maximum densities (ca. 17 individuals m⁻²) in September. Females mature approximately three eggs at a time and these commence hatching in July. Larval eclosion reaches a peak in November, during which time larval densities are also highest (ca. 153 individuals m⁻²). The larvae develop through six instars, which is within the range found for other Ectemnorhinus-group species. The high densities of B. elongatus in fellfield habitats, and its single, virtually discrete annual generation, make this species unusual among insects indigenous to the sub-Antarctic. The latter generally have low densities compared to other micro-arthropods, prolonged life-cycles, and flexible life-histories. We suggest that the diversity of life-histories found amongst the indigenous insects at Marion Island presents considerable potential for testing environmental effects on insect life-histories. An overview of sub-Antarctic insect life-history data suggests that the indigenous species, with generally prolonged life-cycles, are at a disadvantage relative to introduced species that have more rapid cycles and often complete several generations per year. This is reason for concern given rapid climate change at these islands. Accepted: 14 November 1999     

Food consumption by the larvae of Sericostoma vittatum (Trichoptera), an endemic species from the Iberian Peninsula

The caddisfly Sericostoma vittatum Rambur (Trichoptera: Sericostomatidae) is an endemic species of the Iberian Peninsula. Under laboratory conditions, larvae of S. vittatum had a higher activity and metabolism during the night. Besides consuming particulate allochthonous organic matter, young stages are also able to feed and grow on faecal pellets from adults. Daily growth rates varied from 0.02 mg (0.8–3.7 mg size class animals) to 0.31 mg dry mass (10.6–22.8 mg size class animals). Due to the high densities of this species (annual mean of 25 individuals m^–2; maximum of 96 individuals m^–2) and high consumption rates (0.47 mg leaf dry mass mg animal^–1 d^–1for small larvae), this species has a potential key role on the fragmentation of allochthonous organic matter of streams in central Portugal.     

Occurrence of Gobiidae larvae in a tropical Brazilian estuary,with particular emphasis on the use of size classes to categorize species guilds

The structure and seasonal dynamics of larvae of the Gobiidae family in the Mucuri Estuary (Bahia, Brazil) were studied for nine consecutive years. Sampling was conducted at three stations in the lower estuary channel, between 2002 and 2010, in relation to season, day and night and tidal variations. A total of 5802 Gobiidae larvae, representing 15 taxa (12 species and three morphotypes), were collected in the Mucuri Estuary during this time. The highest mean ± s.d. density of fish larvae, 54·7 ± 79·8 larvae 100 m^?3, was recorded during the flood tide and night sampling. Ctenogobius boleosoma was the most abundant species (68%), being dominant in the rainy and dry seasons and had a long reproductive period. This species was classed as a marine estuarine‐opportunist because it was observed at high frequencies and active larvae entering the estuary between 6·1 and 12·0 mm standard length (L_S). Gobionellus oceanicus, second in abundance (12%), occurred only in later larval stages but did not use the estuary as a preferred location for spawning, being classed a marine estuarine dependent. Microgobius carri (11%) was represented in all L_S classes and was resident in the estuary for spawning, remaining there throughout their life cycle. The other species were considered rare due to their low densities and could not be classified in any guild.     

Maternal behaviours may explain riffle‐scale variations in some stream insect populations

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Hatching rhythms and dispersion of decapod crustacean larvae in a brackish coastal lagoon in Argentina

Mar Chiquita, a brackish coastal lagoon in central Argentina, is inhabited by dense populations of two intertidal grapsid crab species,Cyrtograpsus angulatus andChasmagnathus granulata. During a preliminary one-year study and a subsequent intensive sampling programme (November–December 1992), the physical properties and the occurrence of decapod crustacean larvae in the surface water of the lagoon were investigated. The lagoon is characterized by highly variable physical conditions, with oligohaline waters frequently predominating over extended periods. The adjacent coastal waters show a complex pattern of semidiurnal tides that often do not influence the lagoon, due to the existence of a sandbar across its entrance. Besides frequently occurring larvae (exclusively freshly hatched zoeae and a few megalopae) of the two dominating crab species, those of three other brachyurans (Plathyxanthus crenulatus, Uca uruguayensis, Pinnixa patagonica) and of one anomuran (the porcellanidPachycheles haigae) were also found occasionally. Caridean shrimp (Palaemonetes argentinus) larvae occurred in a moderate number of samples, with a maximum density of 800·m⁻³. The highest larval abundance was recorded inC. angulatus, with almost 8000°m⁻³. Significantly moreC. angulatus andC. granulata zoeae occurred at night than during daylight conditions, and more larvae (statistically significant only in the former species) during ebb (outflowing) than during flood (inflowing) tides. In consequence, most crab zoeae were observed during nocturnal ebb, the least with diurnal flood tides. Our data suggest that crab larvae do not develop in the lagoon, where the adult populations live, but exhibit an export strategy, probably based upon exogenously coordinated egg hatching rhythms. Zoeal development must take place in coastal marine waters, from where the megalopa eventually returns for settlement and metamorphosis in the lagoon. Significantly higher larval frequency ofC. granulata in low salinities (≤12‰) and at a particular sampling site may be related to local distribution patterns of the reproducing adult population. Unlike crab larvae, those of shrimp (P. argentinus) are retained inside the lagoon, where they develop from hatching through metamorphosis. They significantly prefer low salinity and occur at the lagoon surface more often at night. These patterns cannot be explained by larval release rhythms like those in brachyuran crabs, but may reflect diel vertical migrations to the bottom. It is concluded that osmotic stress as well as predation pressure exerted by visually directed predators (small species or life-cycle stages of estuarine fishes) may be the principal selection factors for the evolution of hatching and migration rhythms in decapod larvae, and that these are characteristics of export or retention mechanisms, respectively.     

Ecology and production of invertebrates in a Canadian coldwater spring-springbrook system

Valley Spring is a hardwater rheocrene consisting of a single point issue and a springbrook some 60 m long. The fauna consists of > 60 taxa, with insects and mites predominating. During 1985–86, benthic densities varied both along the springbrook and at single stations over time (range 58 to 756 animals 900 cm⁻²). A clear longitudinal zonation of many of the major taxa occurred with the Diptera tending to be more abundant downstream, the Plecoptera and Acari more abundant upstream, and the Copepoda and Bivalvia most abundant in mid- and downstream sections. The Gastropoda and Trichoptera occurred throughout the system although, in the case of the latter, there were distributional differences at the species level (e.g. Diplectrona modesta downstream, Parapsyche apicalis and Lepidostoma vernale upstream). Life histories varied between major species and were grouped into: (A) those species in which there was a wide range of larval sizes for most of the year, with no easily discernible temporal succession of cohorts, e.g. L. vernale, P. apicalis, Sialis sp., Dixa sp., Metriocnemus sp., Trissopelopia sp., Micropsectra sp. and Mucronothrus nasalis (the generation time of species in this group ranged from < 1 year to > 2 years); and (B) those species in which there was a clear pattern of larval growth, with obvious temporal succession of cohorts, e.g. D. modesta, Nemoura trispinosa and Hyalella azteca (all of these species were univoltine). Differences between the “adaptive strategies” of these two life history types are discussed. Total secondary production in Valley Spring in 1985–86 was estimated to be 913.2 g wet wt yr⁻¹. Production upstream was more than twice that downstream (11.21 and 4.01 g wet wt m⁻² yr⁻¹, respectively). The three most abundant species of caddisfly together contributed > 30% of the total production, the stonefly N. trispinosa contributed > 17%, while the chironomids accounted for >16% of production at downstream stations but only 3% upstream.     

Flight activity of insects along a mountain stream: is directional flight adaptive?

1. Flight activity of Trichoptera, Plecoptera and Ephemeroptera was studied by sticky trapping for 12 months at five sites along a New Zealand mountain stream. Over 19 000 insects were captured by the traps, which were located in forest and grassland reaches, including a reach with intermittent flow.
2. Most species occurred predominantly in forest or grassland, although some were trapped throughout the stream. Longitudinal distributions of adults and their larvae were strongly correlated.
3. Flight periods of 24 caddisflies, three mayflies and four stoneflies ranged from 2 to 12 months. Six species were trapped in all months and 17 (55%) in more than 5 months.
4. The most abundant forest-dwelling caddisfly species were over-represented on the downstream sides of sticky traps located in, and immediately below, forest indicating a majority was flying upstream. Upstream flight compensates for downstream drift of larvae and should maximize the likelihood that forest-dwelling species will locate preferred habitat for egg, larval and/or adult development. Unlike the caddisflies, the stonefly Spaniocerca zelandica was over-represented on the upstream sides of traps, suggesting that some adults may float or fly downstream following emergence.
5. In contrast to forest-dwelling species, only one common caddisfly ( Oxyethira albiceps ) was over-represented on the downstream sides of traps at grassland sites. Unlike the forest-dwelling species, most species taken at the downstream sites probably came from a variety of sources, including a nearby stream.     

Copepod community structure and abundance in a tropical mangrove estuary,with comparisons to coastal waters

Zooplankton, sampled at five stations from the upper Sangga estuary (7 km upstream) in Matang Mangrove Forest Reserve (MMFR), Malaysia, to 16 km offshore, comprised more than 47% copepod. Copepod abundance was highest at nearshore waters (20,311 ind m⁻³), but decreased toward both upstream (15,572 ind m⁻³) and offshore waters (12,330 ind m⁻³). Copepod abundance was also higher during the wetter NE monsoon period as compared to the drier SW monsoon period, but vice versa for copepod species diversity. Redundancy analysis (RDA) shows that copepod community structure in the upper estuary, nearshore and offshore waters differed, being influenced by spatial and seasonal variations in environmental conditions. The copepods could generally be grouped into estuarine species (dominantly Acartia spinicauda Mori, Acartia sp1, Oithona aruensis Früchtl, and Oithona dissimilis Lindberg), stenohaline species (Acartia erythraea Giesbrecht, Acrocalanus gibber Giesbrecht, Paracalanus aculateus Giesbrecht, and Corycaeus andrewsi Farran) and euryhaline species (Parvocalanus crassirostris Dahl, Oithona simplex Farran, and Bestiolina similis (Sewell)). Shifts in copepod community structure due to monsoonal effects on water parameters occurred at the lower estuary. Copepod peak abundance in mangrove waters could be associated with the peak chlorophyll a concentration prior to it. Evidence of copepod consumption by many species of young fish and shrimp larvae in the MMFR estuary implies the considerable impact of phytoplankton and microphytobenthos on mangrove trophodynamics.     

Use of 13C/12C to trace dissolved and particulate organic matter utilization by populations of an aquatic invertebrate

Summary The ¹³C/¹²C of limnephilid caddisfly larvae (Clistoronia magnifica) grown in the presence of red alder leaf particulate organic matter (POM) and alder leaf leachate was found to be identical to the ¹³C/¹²C of larvae grown on alder POM, but with isotopically contrasting grass leachate substituted for the above alder leachate. The isotopic similarity between these insects and the alder POM common in both treatments indicates that grass leachate was not a source of insect carbon. In contrast to these results, the ¹³C/¹²C of larvae grown on grass POM and leachate was markedly different from the ¹³C/¹²C of larvae raised on grass POM and alder leachate. The relative ¹³C depletion found in larvae from the latter treatment could be accounted for only if a significant amount of carbon derived from alder leachate had been incorporated into insect biomass. Significant differences in yield of insect biomass among the above treatments also were found.     

Carbon flux between an estuary and the ocean: a case for outwelling

The classical outwelling hypothesis states that small coastal embayments (e.g. estuaries, wetlands) export their excess production to inshore marine waters. In line with this notion, the present study tested whether the Swartkops estuary acts as source or sink for carbon. To this end, concentrations of dissolved inorganic carbon (DIC), dissolved organic carbon (DOC) and particulate organic carbon (POC) were determined hourly during the first monthly spring and neap tides over one year in the tidal waters entering and leaving the estuary. Each sampling session spanned a full tidal cycle, yielding a total of 936 concentration estimates. Carbon fluxes were calculated by integrating concentrations with water flow rates derived from a hydrodynamic model calibrated for each sampling datum. Over the year, exports to marine waters markedly exceeded imports to the estuary for all carbon species: on the basis of total spring tidal drainage area, 1083 g m^–2 of DIC, 103 g m^–2 of DOC, and 123 g m^–2 of POC left the estuary annually. Total carbon export from the estuary to the ocean amounted to 4755 tonnes, of which 83% was in the inorganic form (DIC). Thus, the bulk of carbon moving in the water column is inorganic - yet, DIC seems to be measured only rarely in most flux studies of this nature. Salt marshes cover extensive areas in this estuary and produce some carbon, particularly DOC, but productivity of the local Spartina species is low (P:B=1.1). Consequently, the bulk of carbon exported from the estuary appears to originate from the highly productive macroinvertebrate and the phytoplankton component and not from the salt marsh plants.     

Predator defense along a permanence gradient: roles of case structure, behavior, and developmental phenology in caddisflies

Species replacements along freshwater permanence gradients are well documented, but underlying mechanisms are poorly understood for most taxa. In subalpine wetlands in Colorado, the relative abundance of caddisfly larvae shifts from temporary to permanent basins. Predators on caddisflies also shift along this gradient; salamanders (Ambystoma tigrinum nebulosum) in permanent ponds are replaced by predaceous diving beetles (Dytiscus dauricus) in temporary habitats. We conducted laboratory and field experiments to determine the effectiveness of caddisfly cases in reducing vulnerability to these predators. We found that larvae of a temporary-habitat caddisfly (Asynarchus nigriculus) were the most vulnerable to salamanders. Two relatively invulnerable species (Limnephilus externus, L. picturatus) exhibited behaviors that reduced the likelihood of detection and attack, whereas the least vulnerable species (Agrypnia deflata) was frequently detected and attacked, but rarely captured because cases provided an effective refuge. Vulnerability to beetle predation was also affected by cases. The stout cases of L. externus larvae frequently deterred beetle larvae, whereas the tubular cases of the other species were relatively ineffective. Two of these vulnerable species (A. nigriculus and L. picturatus) often co-occur with beetles; thus, case construction alone is insufficient to explain patterns of caddisfly coexistence along the permanence gradient. One explanation for the coexistence of these two species with beetles is that they develop rapidly during early summer and pupate before beetle larvae become abundant. One species (L. picturatus) pupates by burying into soft substrates that serve as a refuge. The other (A. nigriculus) builds stone pupal cases, which in field experiments, more than doubles survival compared to organic pupal cases. The combined results of these experiments suggest that caddisfly distributions along permanence gradients depend on a suite of primary and secondary predator defenses that include larval and pupal case structure, predator-specific escape behaviors, and the phenology of larval development.     

A comparison of alternative methods for estimating population density of the fiddler crab Uca annulipes at Saco Mangrove, Inhaca Island (Mozambique)

Visual counts of surface-active crabs both by binocular and burrow counting methods have been used in many studies to estimate population density. However, their reliability has not yet been assessed comparatively. Three methods for estimating the abundance of fiddler crabs Uca annulipes in a mangrove forest (Inhaca Island, Mozambique) were compared from three different sub-areas: two sub-areas inundated only during spring tides and one sub-area inundated in both spring and neap tides. Burrow, binocular and direct (excavation) counting methods were performed by plotting ten 0.25 m² quadrats in each sub-area over the four moon phases. Overall densities (per 0.25 m²) differed according to method, sub-area and lunar phase. Burrow count overestimated crab density by up to 20%, while binocular count underestimated density by up to 41%. Correlation coefficient estimated for both counting methods showed that burrow count gives better density estimates than binocular count (0.91 and 0.56, respectively). Sex ratios were also investigated within the three sub-areas and at the moon phases. Males are dominant throughout the studied period except during new moon and first quarter, indicating that when the number of gravid females is low, sex ratio bias for binocular count is minimal.     

Group size of feeding stream case-bearing caddisfly grazers and resource abundance

Several herbivorous insects are known to benefit from feeding in groups; however, little is known about (1) the resource conditions under which herbivorous insects perform group feeding and (2) the optimum population size to get any benefits by group feeding, for example, in terms of growth performance. To test the hypotheses that the benefits from group feeding change with resource level and population size, we performed field investigations and an enclosure experiment using the grazer caddisfly larva Micrasema quadriloba. The field investigations revealed aggregated distributions of larvae (indicator of aggregation, I_δ=4.1±1.55, aggregated density: 12.7±5.3 individuals per 3.1×3.1 cm² (mean±1 SD)) when periphyton was abundant on stream cobbles and random distributions (I_δ=1.0±0.11) when periphyton was scarce. In the enclosure experiment, the relative growth rate (RGR) of the larvae at each population size showed different tendencies at high and low periphyton abundance levels; RGR with abundant periphyton had a convex curve with a peak at intermediate population size, whereas RGR with scarce periphyton decreased linearly with increasing population size. The benefits from group feeding thus changed with resource level; larvae obtained high growth performance by group feeding behavior only when the resource was sufficiently abundant. The present study revealed not only that the optimum group size of larvae increased their growth performance, but also that this optimum group size occurred frequently in the field. We also discuss the mechanisms and benefits of group feeding by case-bearing caddisfly grazers.