Emergence of some caddisflies (Trichoptera) from a wooded stream in southern Ontario

Twenty-eight species of nine families of caddisflies (Trichoptera) were identified in 170 samples taken over an 8-month period from five emergence traps placed on a second-order, forested, cold-stenothermal stream on the Niagara Escarpment, Ontario, Canada.A mean of 980.9 caddisflies m^–2 of streambed was obtained over the entire sampling period. Eleven common species accounted for 92.8% of the total emergence with specific proportions ranging from 23.8% (Wormaldia moesta) to 0.11 % (Rhyacophila sp.). The use of various kinds of traps in other studies and their effects on the detection of species composition and abundance are discussed and compared with the present study.Although the distributions of all the common species were invariant over time, four species showed low to high degrees of patchiness in the streambed; the other seven common species were uniformly distributed. However, a large residual variance suggested a subtle mechanism of microhabitat selection by the larvae and (or) pupae, not detectable by even the small emergence traps used.Both sexes of 15 species, only males of 4 and only females of 9 species were collected. Eight of the eleven common species showed significant departures from a balanced sex ratio and five exhibited a protandry of from 1 to 3 weeks. Neither this study nor others have been able to establish a predictable pattern of sex ratios in Trichoptera.The emergence periods and patterns of the eleven common species are described and compared with other studies. Of these common species, one emerged in the spring, seven during the summer and three during the late summer or early fall. Ten species had a short emergence period with a distinct peak and a significantly skewed pattern. One species,Parapsyche apicalis, exhibited a prolonged emergence period, no distinct peak and a significantly platykurtotic² pattern. With the exception ofLepidostoma sp. A, the emergence patterns of the common species were unimodal.     

Distribution of filter-feeding caddisflies (Trichoptera) and plankton drift in a Swedish lake-outlet stream

The objectives of this investigation were to relate the longitudinal distribution of filter-feeding caddisflies to quality and quantity of seston, and to document factors facilitating coexistence among filter-feeding species, in a 4th-order lake-outlet stream in central Sweden. A sharp decline in abundance and biomass of filtering caddisflies was evident within 1 km from the lake outlet following negative power functions. In contrast to particulate organic carbon, significant reductions of both phytoplankton and zooplankton biomass, especially some large species, were recorded between the lake-outlet and 1.9 km downstream. Differences in longitudinal and temporal distribution provide evidence of resource partitioning between filtering species included.     

Phylogeny of caddisflies (Insecta, Trichoptera)

Trichoptera are holometabolous insects with aquatic larvae that, together with the Lepidoptera, comprise the Amphiesmenoptera. Previous phylogenetic hypotheses and progress on our ongoing data collection are summarized. Fragments of the large and small subunit nuclear ribosomal RNAs (D1, D3, V4–5), the nuclear elongation factor 1 alpha gene and a fragment of mitochondrial cytochrome oxidase 1 (COI) were sequenced, and molecular data were combined with previously published morphological data. Equally and differentially weighted parsimony analyses were conducted in order to present a phylogeny of Trichoptera, including 43 of 45 families. Our phylogeny closely resembles that proposed by Herbert Ross with respect to the relationships among suborders, with a monophyletic Annulipalpia at the base of the tree, and a clade consisting of Spicipalpia plus a monophyletic Integripalpia. The monophyly of Spicipalpia is weakly supported in the combined equally weighted analysis, and Spicipalpia is paraphyletic in the differentially weighted analysis. Within Integripalpia, our phylogeny recovered monophyletic Plenitentoria, Brevitentoria and Sericostomatoidea. Leptoceroidea was unresolved in the equally weighted analysis and monophyletic in the differentially weighted analysis. Within Annulipalpia, we recovered a basal but paraphyletic Philopotamoidea and a monophyletic Hydropsychoidea.     

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.     

Factors influencing the longitudinal distribution of larval Hydropsychidae (Trichoptera) in a southern Appalachian stream system (U.S.A.)

The influence of physical habitat variables and suspended particulate organic matter (seston) on the distribution and production of eight species of larval Hydropsychidae was studied along a 6.4 km section of a southern Appalachian stream. Samples were collected at six stations encompassing stream orders 1–4 and an elevation range of 610 m. Multivariate analysis of covariance (using time as the covariable) and discriminant function analysis were used to examine habitat differences between the sampling stations due to the following variables: current velocity; coarse benthic detritus; substrate composition (by particle size); substrate heterogeneity; degree-days; and diel temperature fluctuation. The associations of these variables with the abundance (and production) of larval hydropsychids was also examined using the same statistical procedures. The six sampling stations represented three or four distinct habitats based on patterns of change in the variables along the stream continuum. Diel temperature fluctuation, median substrate particle size, the proportion of sand substrate, and substrate heterogeneity were most closely associated with the overall difference between the sampling stations. Hydropsychid species distribution along the stream system followed subfamily lines, i.e., Arctopsychinae and Diplectroninae were more abundant and productive in the upper 4.5 km of the stream, while Hydropsychinae were dominant in the lower 1.9 km. Diel temperature fluctuation was the habitat variable most highly correlated with patterns of hydropsychid abundance and production. The longitudinal pattern of species distribution, i.e., larger particle feeding Arctopsychinae being replaced downstream by smaller particle feeding Hydropsychinae, also coincided with the distribution of seston particle size classes along the stream. Mean seston particle size generally declined downstream, as particles <42 µm increased in relative abundance while those between 43 µm and 5 mm decreased. Temperature, seston, and substrate composition all undoubtedly exerted an important influence on the distribution and production of Hydropsychidae and other filter feeding insects in this stream. The validity and generality of these results depend upon the scope of the sampling effort. Conclusions drawn from data collected over the entire range (and limits) of a species' distribution are more sound than those based on data from a limited area.     

Restricted movement by mottled sculpin (pisces: cottidae) in a southern Appalachian stream

1. We used direct observation and mark‐recapture techniques to quantify movements by mottled sculpins (Cottus bairdi) in a 1 km segment of Shope Fork in western North Carolina. Our objectives were to: (i) quantify the overall rate of sculpin movement, (ii) assess variation in movement among years, individuals, and sculpin size classes, (iii) relate movement to variation in stream flow and population size structure, and (iv) quantify relationships between movement and individual growth rates. 2. Movements were very restricted: median and mean movement distances for all sculpin size classes over a 45 day period were 1.3 and 4.4 m respectively. Nevertheless, there was a high degree of intrapopulation and temporal variation in sculpin movement. Movement of juveniles increased with discharge and with the density of large adults. Movement by small and large adults was not influenced by stream flow, but large adults where more mobile when their own density was high. Finally, there were differences in the growth rates of mobile and sedentary sculpins. Mobile juveniles grew faster than sedentary individuals under conditions of low flow and high density of large adults, whereas adults exhibited the opposite pattern. 3. Our results support the hypothesis that juvenile movement and growth is influenced by both intraspecific interactions with adults and stream flow. In contrast, adult movement appears to be influenced by competitive interactions among residents for suitable space. The relationship between movement and growth may provide a negative feedback mechanism regulating mottled sculpin populations in this system.     

Patch selection by mottled sculpin (Pisces: Cottidae) in a southern Appalachian stream

1. We examined the effects of prey abundance on patch selection by a benthic fish, the mottled sculpin ( Cottus bairdi ), in a fourth order, southern Appalachian stream (North Carolina, U.S.A.). This habitat is a mosaic of small (i.e. < 0.5 m²), relatively discrete patches.
2. Patches were characterized in terms of physical habitat variables, detritus (coarse particulate organic matter, CPOM) and macroinvertebrate abundance (number and biomass). We quantified patch selection by comparing the characteristics of patches utilized by sculpin with those of locally available patches. Locally available patches were selected using a constrained random sampling design (i.e. randomly selected within a 2 m radius from each fish). We also examined the relationship between macroinvertebrate abundance, CPOM and the physical characteristics of available patches.
3. Patches selected by sculpin contained significantly higher macroinvertebrate abundances (both number and biomass) than locally available patches in five out of six seasonal samples. Sculpin also occupied patches with significantly higher amounts of CPOM in three out of five seasonal samples. Patches utilized by sculpin, however, could not be consistently differentiated from locally available patches on the basis of physical variables. In addition, macroinvertebrate abundance was not consistently related to physical habitat variables or CPOM during the course of the study.
4. Our results suggest that sculpin are able to assess patch quality on the basis of prey abundance and select patches that potentially maximize energy gain. This behaviour may produce an increase in individual fitness, especially when prey distributions are not consistently related to habitat variables. Quantifying patch use in relation to prey abundance may help elucidate the causal factors determining habitat use by benthic fishes in other lotic systems.     

Patch selection by mottled sculpin (Pisces: Cottidae) in a southern Appalachian stream

1. We examined the effects of prey abundance on patch selection by a benthic fish, the mottled sculpin ( Cottus bairdi ), in a fourth order, southern Appalachian stream (North Carolina, U.S.A.). This habitat is a mosaic of small (i.e. < 0.5 m²), relatively discrete patches.
2. Patches were characterized in terms of physical habitat variables, detritus (coarse particulate organic matter, CPOM) and macroinvertebrate abundance (number and biomass). We quantified patch selection by comparing the characteristics of patches utilized by sculpin with those of locally available patches. Locally available patches were selected using a constrained random sampling design (i.e. randomly selected within a 2 m radius from each fish). We also examined the relationship between macroinvertebrate abundance, CPOM and the physical characteristics of available patches.
3. Patches selected by sculpin contained significantly higher macroinvertebrate abundances (both number and biomass) than locally available patches in five out of six seasonal samples. Sculpin also occupied patches with significantly higher amounts of CPOM in three out of five seasonal samples. Patches utilized by sculpin, however, could not be consistently differentiated from locally available patches on the basis of physical variables. In addition, macroinvertebrate abundance was not consistently related to physical habitat variables or CPOM during the course of the study.
4. Our results suggest that sculpin are able to assess patch quality on the basis of prey abundance and select patches that potentially maximize energy gain. This behaviour may produce an increase in individual fitness, especially when prey distributions are not consistently related to habitat variables. Quantifying patch use in relation to prey abundance may help elucidate the causal factors determining habitat use by benthic fishes in other lotic systems.     

Studies on production and food consumption by the larval simuliidae (Diptera) of a chalk stream

Summary Large populations of S. ornatum and S. equinum inhabit Ranunculus plants in the Bere Stream at Bere Heath. A weekly sampling programme obtained both larvae and pupae. The results of weekly sampling indicated four annual generations of S. ornatum and three of S. equinum. Quantitative samples of larvae, obtained at least three times in each generation gave estimates of production ranging from 0.26 g to 2.75 g dry weight m-² per generation of larvae of S. equinum and 0.15 g to 1.19 g dry weight m-² per generation of larvae of S. ornatum.Retention times of food by larvae, under field conditions were in the order of twenty to thirty minutes. Following determination of the weight of the gut contents for each size class of larvae, the activity of feeding larvae was calculated to achieve complete theoretical clearance of suspended material in a distance of 0.6 km, at peak population densities in the summer months.Dept. of Applied Biology, Bath University of Technology     

Species assemblages and community structure of adult caddisflies along a headwater stream in southeastern Ghana (Insecta: Trichoptera)

Adult caddisflies were collected at 12 sites along a 5 km stretch of a forested headwater stream in southeastern Ghana in March and November 1993 and the catches are related to environmental variables. Some 34,000 specimens, belonging to 178 morphospecies in 43 genera and 11 families, were caught with Malaise traps and light traps. Many species demonstrate differences between sampling method, seasonal occurrence, and sex-ratios, with the light trap in the second wet sunny season (November) being most efficient with respect to the diversity of the catches. A Moran index analysis for global structures indicates a major transition in the fauna from the ravine with waterfalls to the forest, and secondly, a gradient through the forest. A Geary index analysis for local structures indicates further faunal turnovers in the lower reaches of the stream. Species-environment relationships were analyzed through correspondence analysis and co-inertia analysis, allowing ordination of both species and the environment into three zones. The first co-inertia axis reveals a transition between the waterfalls in the open ravine (sites 1–3), and the stream through the riverine forest (sites 4–12), while the second co-inertia axis reveals a gradient from the lotic stream in the moist semi-deciduous forest (sites 4–8) to more lentic stretches of the stream in a gradually more depleted forest (sites 9–12). The most important environmental factors related to the species transition are: riparian moss growth (r=0.94), leaves in the stream substratum (r=0.91), forest type (r=0.90), hygropetric surfaces (r=0.88), bedrock (r=0.87), lotic [riffle] (r=0.85) and lentic [pool] (r=0.85) stretches. The most important environmental factors related to the species gradient are: lotic [riffle] stretches (r=0.92), gravel (r=0.88) and leaves (r=0.85) in the stream substratum, forest type (r=0.81), and agricultural use (r=0.76). Three clusters representing three major environmental zones (zone I, II and III) along the stream are identified by projecting the average species positions on the co-inertia plane. By analyzing the Malaise trap samples significant indicator values are found for 29 species in zone I, 16 species in zone II, and 18 species in zone III. By analyzing the light trap samples significant indicator values are found for 17 species in zone I, 57 species in zone II, and 38 species in zone III. The high diversity of caddisflies in the sanctuary might be reflected both in the geological and climatological history of West Africa, as well as in the recognition of three major environmental zones, and the clean water with high diversity of stream microhabitats favorable for Trichoptera larvae. The closed forest seems to play an important role for the Trichoptera community, and the study demonstrates the importance of protecting forested headwater streams in order to maintain a sustainable aquatic biodiversity in tropical Africa.     

The Neotropical species of Notalina,a southern group of long-horned caddisflies (Trichoptera: Leptoceridae)*

The leptocerid trichopteran genus Notalina is recorded from the South American continent for the first time. Seven new species are described in it and placed in a new subgenus, Neonotalina: N.brasiliana, N.morsei, N.cipo and N.hamiltoni from southeastern Brazil, comprise the brasiliana species-group, and N.roraima from Venezuela, N.nanay from Peru and N.matthiasi from Colombia form the roraima species-group. Notalina (Neonotalina) is identifiable by the presence of a long, slender baso-mesal process on the male inferior appendage. The two species-groups may be distinguished by the morphology of the male phallus. The historical biogeography of the genus in South America is discussed.     

New long-horned caddisflies in Eocene Baltic amber (Insecta,Trichoptera)

We describe two new extinct caddisflies, Electroadicella kuenowi sp. nov. and Triaenodes simoni sp. nov., embedded in Baltic amber. Both species belong to the tribe Triaenodini and enlarge the family Leptoceridae to a remarkable number of 18 extinct Eocene species from Baltic amber. The extinct Electroadicella and the extant Triaenodes are possibly closely related, by a distinct synapomorphy, a recurved process from the basal plate of the inferior appendages of the ninth abdominal segment in the male genitalia.     

Biological diversity of the Minnesota caddisflies (Insecta, Trichoptera)

The caddisfly fauna of Minnesota contains at least 277 species within 21 families and 75 genera. These species are based on examination of 312,884 specimens from 2,166 collections of 937 Minnesota aquatic habitats from 1890 to 2007. Included in these totals is my own quantitative sampling of 4 representative habitat types: small streams, medium rivers, large rivers, and lakes, from each of the 58 major Minnesota watersheds from June through September during 1999-2001. All species are illustrated herein, and their known Minnesota abundances, distributions, adult flight periodicities, and habitat affinities presented. Four species: Lepidostoma griseum (Lepidostomatidae), Psilotreta indecisa (Odontoceridae), and Phryganea sayi and Ptilostomis angustipennis (Phryganeidae) are added to the known fauna. An additional 31 dubious species records are removed for various reasons. Of the 5 determined caddisfly regions of the state, species richness per watershed was highest in the Lake Superior and Northern Regions, intermediate in the Southeastern, and lowest in the Northwestern and Southern. Of the 48 individual collections that yielded >40 species, all but 1 were from the Northern Region. Many species, especially within the families Limnephilidae and Phryganeidae, have appeared to decrease in distribution and abundance during the past 75 years, particularly those once common within the Northwestern and Southern Regions. Many species now appear regionally extirpated, and a few have disappeared from the entire state. The loss of species in the Northwestern and Southern Regions, and probably elsewhere, is almost certainly related to the conversion of many habitats to large-scale agriculture during the mid-20th century.     

Global diversity of caddisflies (Trichoptera: Insecta) in freshwater

The not yet uploaded Trichoptera World Checklist (TWC) [], as at July 2006, recorded 12,627 species, 610 genera and 46 families of extant and in addition 488 species, 78 genera and 7 families of fossil Trichoptera. An analysis of the 2001 TWC list of present-day Trichoptera diversity at species, generic/subgeneric and family level along the selected Afrotropical, Neotropical, Australian, Oriental, Nearctic and Palaearctic (as a unit or assessed as Eastern and Western) regions reveals uneven distribution patterns. The Oriental and Neotropical are the two most species diverse with 47–77% of the species in widespread genera being recorded in these two regions. Five Trichoptera families comprise 55% of the world’s species and 19 families contain fewer than 30 species per family. Ten out of 620 genera contain 29% of the world’s known species. Considerable underestimates of Trichoptera diversity for certain regions are recognised. Historical processes in Trichoptera evolution dating back to the middle and late Triassic reveal that the major phylogenetic differentiation in Trichoptera had occurred during the Jurrasic and early Cretaceous. The breakup of Gondwana in the Cretaceous led to further isolation and diversification of Trichoptera. High species endemism is noted to be in tropical or mountainous regions correlated with humid or high rainfall conditions. Repetitive patterns of shared taxa between biogeographical regions suggest possible centres of origin, vicariant events or distribution routes. Related taxa associations between different regions suggest that an alternative biogeographical map reflecting Trichoptera distribution patterns different from the Wallace (The Geographical Distribution of Animals: With a Study of the Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Earth’s Surface, Vol. 1, 503 pp., Vol. 2, 607 pp., Macmillan, London, 1876) proposed biogeography patterns should be considered. Anthropogenic development threatens biodiversity and the value of Trichoptera as important functional components of aquatic ecosystems, indicator species of deteriorating conditions and custodians of environmental protection are realised.     

Sexual dimorphism in relation to swarming and pair formation patterns in leptocerid caddisflies (Trichoptera: Leptoceridae)

North European Leptoceridae (Trichoptera) perform three types of swarming flight patterns: (1) swarming males of Athripsodesand Ceracleafly in horizontal zigzag patterns over the water surface, (2) the Mystacidesspp. perform vertical zigzag movements, and (3) the flight of males of Triaenodes unanimisMcLach. is a mixture of the horizontal and vertical zigzagging. Also three groups of pair formation behavior can be distinguished. In the first group, of Athripsodesand Ceraclea,the females fly into the male swarms, where they are grasped and carried to the riparian vegetation by the flying males with the females hanging upside-down in genitalia coupling. In the second group, a Mystacidesfemale is caught by a male, when approaching a swarm and both use their wings to fly in tandem to the shore where they copulate. In the third group, of Triaenodes bicolor(Curt.) and Oecetis lacustris(Curt.), the males fly searching for females sitting on aquatic plants and when a female is found the male lands and they copulate immediately while clinging to the plant. The different swarming and mating behaviors might have favored selection for three types of sexual dimorphism: (1) longer forewings in males than females in species which fly in copula, (2) larger eyes in males of the vertically zigzagging species, and (3) much smaller males in the group where males search for females sitting on aquatic plants. In the second group approaching females are detected by males before reaching the swarm and in the third group the female almost always mates with the male which is the first to find her. In conclusion, we suggest that females of Athripsodesand Ceracleahave a greater choice among swarming males than do females of Mystacides, T. bicolor,and O. Lacustris.     

Invertebrate drift,discharge, and sediment relations in a southern Appalachian headwater stream

Drifting invertebrates and suspended sediments were collected at monthly intervals from June 1977 to May 1978. The numbers and biomass of drifting organisms reflected the seasonal cycles of aquatic insects. Some aquatic organisms showed behavioral drift either during a sample day or during some portion of their life cycle. Parapsyche cardis Ross and Diplectrona modesta Banks (Trichoptera: Hydropsychidae) dispersed as first instar larvae; few later instars of these two net-spinning caddisflies drifted. The drift of nymphal Peltoperla maria Needham et Smith (Plecoptera: Peltoperlidae) was apparently related more to detritus transport than to benthic densities or discharge alone. Power law relations between the magnitude of daily invertebrate drift and discharge or sediment variables are demonstrated for some taxa in Hugh White Creek. The general level of stream invertebrate drift appears to be related to detritus transport, and drift during storms is also related to detritus transport. During storms, terrestrial invertebrate drift was related to rainfall intensity, canopy washing, and channel expansion. Drift density of aquatic invertebrates in Hugh White Creek was within the range of previously reported values for other streams, but the estimate of yearly export (aquatic invertebrates = 134 g · y^?1; terrestrial invertebrates = 23 g · y^?1) is lower reflecting the smaller size of Hugh White Creek in comparison with those other streams.     

Community structure of Trichoptera in a mountain stream: spatial patterns of production and functional organization

SUMMARY. 1. Annual production was estimated for Trichoptera occurring in each of three distinct habitats of a mountain stream: bedrock-outcrops, riffles and pools. Production was greatest on bedrock-outcrops (2608 mg ash-free dry weight m^?2), followed by riffles (1038) and pools (950). 2. Annual production in bedrock-outcrops and pools was dominated by single functional groups, with collector-filterers and shredders contributing 73% and 75% of the annual production, respectively. Production in riffles was due primarily to shredders (46%), followed by collector-filterers (27%). 3. Taking account of the amount of stream area occupied by each habitat type, total annual production was estimated at 1336 mg AFDW m^?2. 53% of this production was attributable to four taxa: Parapsyche cardis Ross (25%), Pycnopsyche gentilis (MacLachlan) (10%), Neophylax mitchelli Carpenter (9%) and Rhyacophila nigrita Banks (9%). 4. Habitat-weighted production was distributed among functional groups as follows: collector-filterers (41%), shredders (29%), engulfing-predators (15%), scrapers (13%) and collector-gatherers (2%). 5. The distinct taxonomic and functional structures of trichopteran sub-communities were shaped by the distinct physical characteristics of their principal habitats. Bedrock-outcrops were characterized by low roughness and high current and were sites of low deposition or organic matter; thus the predominance of collector-filterers. In contrast, the other habitats of greater roughness (riffles) and/or lower current (pools) were sites of deposition of food (e.g. leaf litter) and greatest shredder production. 6. By distinguishing discrete mesoscale habitats, each with a functionally distinct caddisfly sub-community, we speculate that small mountain streams provided the diverse physical templates essential for the evolution of the major feeding tactics (e.g. scraping, shredding, filter-feeding) of the. Trichoptera.     

Life history and production of Agapetus quadratus (Trichoptera: Glossosomatidae) in a temporary, spring-fed stream

1. The life history and trophic basis of production of the caddisfly grazer Agapetus quadratus were studied in the torrent Gorg Blau, a spring‐fed stream on the island of Majorca that dries annually during summer. 2. Quantitative random samples were taken every 2–3 weeks during an annual surficial flow period, from November 2000 to mid‐July 2001. Instars of field‐collected larvae were determined by measurements of head width and pronotum length, and the sex of all pupae was determined to study sexual dimorphism and sex ratio. 3. Stage‐frequency histograms suggested a trivoltine population, with an average cohort time of 4 months. Larval development was asynchronous, with continuous growth and overlapping generations. Recruitment peaks were identified in mid‐November, early March and late June, indicative of winter, spring and summer generations. On average, females were larger than males and the mean sex ratio was 2 : 3 (females : males). Population densities and biomasses derived from the field data were used to calculate production and turnover rate. 4. Annual production of A. quadratus in the torrent Gorg Blau (4.80 g dry mass m^?2 year^?1) was the highest ever reported for the genus, being comparable with that estimated for some insects with rapid development and multiple cohorts. 5. Estimates of production of A. quadratus were combined with foregut content analysis to estimate the fraction of total production derived from the principal food sources: algae and organic detritus. Algae supported a major proportion of the production of this grazer. 6. The low density of predators characteristic of many temporary streams, and the small amplitudes in discharge and temperature during most of the wet period that characterise the spring habitats might allow high levels of grazer production in this particular Mediterranean stream.     

Phylogenetic position of rhyacophiloid caddisflies (Insecta, Trichoptera): a spermatological analysis of Rhyacophilidae and Glossosomatidae

Most caddisflies (Insecta, Trichoptera) are classified into two suborders, Annulipalpia and Integripalpia. However, the use of the derived characters that are regularly applied in systematic and phylogenetic analyses of Trichoptera is insufficient to determine with certainty the position of the families belonging to Rhyacophiloidea, which are considered by different authors to be either Annulipalpia, or Integripalpia, or even a separate suborder. Rhyacophiloidea comprise four overall similar families: free-living Rhyacophilidae and Hydrobiosidae, saddle-case making Glossosomatidae, and purse-case making Hydroptilidae. It was previously found that Annulipalpia spermatozoa have aberrant axonemes while Integripalpia spermatozoa display the plesiomorph 9 + 2 axoneme. The present spermatological analysis of the families Rhyacophilidae and Glossosomatidae shows that both have spermatozoa with aberrant axonemes lacking the two central microtubules found in the typical axoneme of insect spermatozoa. This is an apomorphic character shared with the superfamily Hydropsychoidea, indicating that from this point of view, Rhyacophiloidae are more closely related to Annulipalpia than to Integripalpia.