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.     

Phylogeny of the Ecnomidae (Insecta: Trichoptera)

Ecnomidae are a family of seven previously accepted extant genera having a typical Gondwanan distribution, except one genus ( Ecnomus ) being widely distributed also in the Oriental and Palearctic regions. We analysed a molecular data set of 3379 characters representing the sum of four different protein-coding genes (COI, CAD, EF-1a and POL-II). Six equally most parsimonious trees were generated from the combined data set, distributed into two distinct islands. In all maximum parsimony (MP) trees the Ecnomidae is monophyletic when the genus Zelandoptila (Psychomyiidae) is included. The sister group to Ecnomidae including Zelandoptila is Pseudoneureclipsis , previously classified in the other families. This sister-group relationship contradicts earlier findings that the Polycentropodidae are the sister group to Ecnomidae. A Bayesian analysis resulted in a monophyletic Ecnomidae when accepting inclusion of Pseudoneureclipsis , which contradicts the results from the MP analysis by leaving Zelandoptila as the sister group to Ecnomidae including Pseudoneureclipsis . In the majority rule tree from this analysis Polycentropodidae form the sister group to Ecnomidae. We were not able to obtain a monophyletic Ecnomus due to the inclusion of Psychomyillodes . We conclude that the genus Zelandoptila or Pseudoneureclipsis probably belongs to the Ecnomidae, and that Psychomyiellodes and Ecnomus are synonyms. Three additional, as yet undescribed monotypic genera from Australia and New Caledonia remain to be erected in Ecnomidae.
© The Willi Hennig Society 2009.     

A multigene phylogeny and timeline for Trichoptera (Insecta)

The Trichoptera, or caddisflies, are traditionally split into two taxonomic subdivisions: the ‘retreat-making’ Annulipalpia and the ‘case-making’ Integripalpia (sensu Ross). The monophyly of these groups is well documented; however, the establishment of a third subdivision, ‘Spicipalpia’, and the positions of the five ‘spicipalpian’ families is much debated. In contrast to previous molecular studies using nuclear ribosomal RNA, a recent trichopteran study (using nuclear protein-coding genes) placed one of these ‘spicipalpian’ families, the free-living predatory Rhyacophilidae, as the sister taxon to the rest of Trichoptera, a result that has significant implications for both the understanding of trichopteran evolution and its timing. This paper sets out to investigate the relationships of Trichoptera using several newly sequenced genes, together with previously published gene sequences. This dataset is the largest trichopteran dataset to date, covering six independent genes and > 10 000 nucleotides, and containing 185 species representing 49 families. With all data included, likelihood and Bayesian analyses support a monophyletic Annulipalpia and a monophyletic Integripalpia, which includes the ‘spicipalpians’ as a paraphyletic grade at the base of this clade. However, an analysis of the protein-coding data alone using similar analytical methods recovers Rhyacophilidae as the most basal taxon in Trichoptera, with low support. A reanalysis correcting for nucleotide composition bias provides support for the placement of the ‘spicipalpian’ taxa as sister to the Integripalpia, consistent with the total data analysis, suggesting that the basal position of Rhyacophilidae in the uncorrected analysis could be (or is probably) an artefact of base composition. We find it likely that ancestral trichopterans made incipient cases and retreats, and these had independent origins as precocious pupal chambers. Molecular dating analysis in beast , using the birth-death model of speciation, with a relaxed-clock model of sequence evolution informed by 37 fossil constraints, suggests that the most recent common ancestor of Trichoptera appeared in the Permian (c. 275 Ma) in line with the first appearance of Trichoptera in the fossil record, and that vicariance explains the distribution of most trichopteran taxa. A new infraordinal name, Phryganides , is introduced for the tube-case-making families of Integripalpia.     

The Micro-caddisflies of Lord Howe Island (Hydroptilidae: Trichoptera: Insecta)

Hydroptilid species are described from Lord Howe Island for the first time. All four species are in the endemic Australian genus Orphninotrichia, and all are endemic to Lord Howe Island. Modifications exhibited by these new species to the normally conservative Orphninotrichia features are suggestive of sympatric speciation and, for several, of a shift in larval niche.     

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.     

The genus Rhyacophila Pictet, 1834 (Insecta: Trichoptera) in Kosovo

The Balkan Peninsula is known for its high species diversity, but it is also one of the least known areas of Europe. Many previously inaccessible regions lack reliable faunistic records. There are especially few data available for the highly diverse aquatic insects of the region. To help fill the gap, we conducted an extensive survey to map the Trichoptera fauna of the Republic of Kosovo, an area where faunistic records are extremely scarce. Adult specimens were collected during 2009 and 2010, with light traps and entomologic nets. Of 84 Trichoptera species identified from the 21 investigated stations, 15 belong to the genus Rhyacophila. This makes the genus the most species-rich in Kosovo. Rhyacophila fasciata and R. tristis are widespread in many streams and rivers all over the country, while R. balcanica, R. trescavicensis, R. obtusa and R. bosnica have limited distribution. Ten Rhyacophila species presented in this paper are recorded for the first time on the area of Kosovo. We also provide new data for the disjunctive distribution of R. palmeni in south-eastern Europe, from Kosovo and Croatia.     

Mortality of Apatania fimbriata (Insecta: Trichoptera) during embryonic, larval and adult life stages

1. Egg masses, oviposition site preferences, and abiotic and biotic factors causing mortality during embryonic, larval and adult life stages of Apatania fimbriata were studied. Laboratory investigations provided information on the temperature dependence of embryonic development, measured as an increase in egg volume.
2. A. fimbriata laid hemispherical egg masses, consisting of a transparent matrix containing a mean of 208 eggs. Egg masses were laid on stones situated just above the water surface in dark cavities in the stream bank.
3. Two hundred egg masses were mapped and individually monitored during embryonic development. There was no significant correlation between mortality during embryonic development and any of the abiotic parameters measured. First-instar larvae of Osmylus fulvicephalus consumed developing eggs, and chironomids preyed on newly hatched larvae.
4. A mean of seventy-two females emerged per metre of stream. Mortality during the 1993/94 life cycle was measured as a percentage of the potential number of eggs laid. Female mortality between emergence and oviposition was ≈ 80%. Eight per cent of individuals were lost during embryonic development. Larval mortality to emergence in 1994 was 11.3%. This indicates that the terrestrial life stage is probably decisive in the regulation of A. fimbriata populations.
5. Duration of embryogenesis at constant temperatures (4–20 °C) in the laboratory was described best by a negative exponential function. This species is cold stenothermal and there was no hatching success at 20 °C.
6. Egg volumes during embryonic development increased sigmoidally over time.     

Mortality of Apatania fimbriata (Insecta: Trichoptera) during embryonic, larval and adult life stages

1. Egg masses, oviposition site preferences, and abiotic and biotic factors causing mortality during embryonic, larval and adult life stages of Apatania fimbriata were studied. Laboratory investigations provided information on the temperature dependence of embryonic development, measured as an increase in egg volume.
2. A. fimbriata laid hemispherical egg masses, consisting of a transparent matrix containing a mean of 208 eggs. Egg masses were laid on stones situated just above the water surface in dark cavities in the stream bank.
3. Two hundred egg masses were mapped and individually monitored during embryonic development. There was no significant correlation between mortality during embryonic development and any of the abiotic parameters measured. First-instar larvae of Osmylus fulvicephalus consumed developing eggs, and chironomids preyed on newly hatched larvae.
4. A mean of seventy-two females emerged per metre of stream. Mortality during the 1993/94 life cycle was measured as a percentage of the potential number of eggs laid. Female mortality between emergence and oviposition was ≈ 80%. Eight per cent of individuals were lost during embryonic development. Larval mortality to emergence in 1994 was 11.3%. This indicates that the terrestrial life stage is probably decisive in the regulation of A. fimbriata populations.
5. Duration of embryogenesis at constant temperatures (4–20 °C) in the laboratory was described best by a negative exponential function. This species is cold stenothermal and there was no hatching success at 20 °C.
6. Egg volumes during embryonic development increased sigmoidally over time.     

The life history of Glyphopsyche irrorata (Trichoptera, Limnephilidae): A caddisfly that overwinters as an adult

Glyphopsyche irrorata has adapted to live in ponds with fluctuating water levels and ponds exhibiting winter drought by passing the winter as an adult in the terrestrial environment.
Larvae, which hatch in May and pupate in late August, are detritivorous shredders. Males and females emerge in September, with a sex ratio of one and commence mating in the autumn. The entire population has mated by May at which time females oviposit in the water. Adult males and females are present in equal numbers in the spring and have similar longevities approaching a maximum of 8 to 9 months.     

New records of polycentropodid caddisflies from Korea (Insecta: Trichoptera)

Four species of the family Polycentropodidae –Plectrocnemia baculifera, P. kusnezovi, Paranyctiophylax hjangsanchonus and Polyplectropus nocturnus– are recorded from Korea. Polyplectropus nocturnus is newly recorded from the Korean peninsula. The other three species are recorded from the Republic of Korea for the first time.     

The First Mitochondrial Genome for Caddisfly (Insecta: Trichoptera) with Phylogenetic Implications

The Trichoptera (caddisflies) is a holometabolous insect order with 14,300 described species forming the second most species-rich monophyletic group of animals in freshwater. Hitherto, there is no mitochondrial genome reported of this order. Herein, we describe the complete mitochondrial (mt) genome of a caddisfly species, Eubasilissa regina (McLachlan, 1871). A phylogenomic analysis was carried out based on the mt genomic sequences of 13 mt protein coding genes (PCGs) and two rRNA genes of 24 species belonging to eight holometabolous orders. Both maximum likelihood and Bayesian inference analyses highly support the sister relationship between Trichoptera and Lepidoptera.     

Larvae of caddisflies (Insecta: Trichoptera) from the Mesozoic of Siberia

Body fossils of caddisfly larvae are described for the first time based on material from the Lower Cretaceous of Siberia (Baissa locality, Neocomian). The material includes a fully grown larva of Baissoplectrum separatum gen. et sp. nov. (Brachycentridae), Creterotesis coprolithica gen. et sp. nov. (Leptoceridae), and Cretolype minuta gen. et sp. nov., a tiny larva probably belonging to an uncertain family of the suborder Annulipalpia (Hydropsychina). C. coprolithica built unusually soft larval cases of pellets; these cases are described as a separate new indusigenus and indusispecies Coprindusia pallida gen. et sp. nov. Bionomics, taxonomy, and evolution of the Mesozoic larvae of Trichoptera are discussed.     

Ultrastructure and central projections of extraocular photoreceptors in caddiesflies (Insecta: Trichoptera)

Summary Retained larval eyes (stemmata) were studied in the imagines of three species of Trichoptera: Phrygania grandis, Agrypnia varia, and Trichostegia minor. At the light-microscopic level the stemmata of all three species appeared to represent different stages of reduction with respect to size, shape and number of lenses. However, in all three species electron-microscopic studies showed units with monolayered rhabdoms, each formed by four retinula cells. By use of immunocytochemistry the presence of S-antigen was demonstrated in the retinula cells and their axons. This method also revealed the central projections of the axons of the retinula cells, which were found (i) to terminate either in the lamina accessoria or (ii) to penetrate this area to join the fibers of the outer chiasma of the optic lobes and then terminate in the medulla accessoria. The lamina accessoria and the medulla accessoria are the assumed remnants of the larval optic lobes. It is suggested that the imaginal stemmata might still be functioning photoreceptors.     

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.     

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.     

Structure and localization of sensilla on antennae of caddisflies (Insecta: Trichoptera)

Structure of antenna segments and ultrastructure of antennal sensillae in representatives of 28 caddisfly families from both modern suborders were studied by electron and light microscopy methods. On Trichoptera antennae, 16 types of sensillae were found, some of them being described for the first time. Morphological peculiarities of cuticular ultrastructures on the antennal surface demonstrate essential differences in structure both at the family level and at the lower taxonomic levels. Specialized sensory fields structurally different from the remaining antennal surface were revealed on antennal flagellae in representatives of the Phryganeina suborder. A new classification of sensillae based on the structure of their cuticular section is proposed.     

Ten new species of caddisflies (Insecta: Trichoptera) from the Baltic amber

Ten new species of caddisflies of the families Polycentropodidae (nine) and Molannidae (one) are described from the Eocene Baltic amber: Holocentropus peregrinator sp. nov., H. proximorepertus sp. nov., H. dugoi sp. nov., H. fundamentalis sp. nov., H. telergon sp. nov., Plectrocnemia varigoria sp. nov., P. synthesia sp. nov., P. novokshonovi sp. nov., Nyctiophylax valideturbidus sp. nov., and Molanna okraina sp. nov.     

A standardized approach to estimate life history tradeoffs in evolutionary ecology

As tradeoffs limit the maximum Darwinian fitness individuals can reach, measuring reliably the strength of tradeoffs using appropriate metrics is of prime importance to understand the evolution of traits under constraints. Tradeoffs involving phenotypic traits and fitness components, however, are difficult to quantify in free‐ranging populations because of confounding effects due to environmental variation and individual heterogeneity. Furthermore, although some methods have been used previously to quantify tradeoffs, these methods cannot be applied with respect to binary traits, which are common to describe life histories (e.g. probability of reproduction, nesting success, offspring survival). Here, we demonstrate how to measure reliably the strength of tradeoffs involving binary traits using (auto)correlation estimates obtained from generalized linear (mixed) models. We first propose a standardized approach that accounts for the variation in the nature of the tradeoffs being compared (e.g. continuous/binary traits, repeated/non‐repeated measures), and then apply this method to longitudinal data from two contrasting species of large herbivores. Empirical estimates of tradeoffs varied among traits, and between‐species comparisons suggested that reproductive tradeoffs between successive breeding attempts might only occur in capital breeders. The empirical results we obtained clearly demonstrate that the method we provide allows measuring reliably the strength of tradeoffs under most circumstances, including tradeoffs on binary traits. Our original approach therefore offers an important first step for comparing the strength and, hence, the relative importance of different tradeoffs, and opens the door to a better understanding of the evolution of life history traits in free‐ranging populations.     

The effect of environmental diversification on species diversification in New Caledonian caddisflies (Insecta: Trichoptera: Hydropsychidae)

Aim To test whether environmental diversification played a role in the diversification of the New Caledonian Hydropsychinae caddisflies. Location New Caledonia, south‐west Pacific. Methods The phylogeny of the New Caledonian Hydropsychinae caddisflies was hypothesized using parsimony and Bayesian methods on molecular characters. The Bayesian analysis was the basis for a comparative analysis of the correlation between phylogeny and three environmental factors: geological substrate (ultrabasic, non‐ultrabasic), elevation and precipitation. Phylogenetic divergence times were estimated using a relaxed clock method, and environmental factors were mapped onto a lineage‐through‐time plot to investigate the timing of environmental diversification in relation to species radiation. The correlation between rainfall and elevation was tested using independent contrasts, and the gamma statistic was calculated to infer the diversification pattern of the group. Results The diversification of extant Orthopsyche–Caledopsyche species began in the Middle–Late Oligocene, when much of the island of New Caledonia was covered by ultrabasic substrate and mountain forming was prevalent. Most lineages originated in the Middle–Late Miocene, a period associated with long‐term climate oscillation. Optimization of environmental factors on the phylogeny demonstrated that the New Caledonian Hydropsychinae group adapted to ultrabasic substrate early in its evolutionary history. The clade living mostly on ultrabasic substrate was far more species‐rich than the clade living mostly on non‐ultrabasic substrate. Elevation and rainfall were significantly correlated with each other. The lineage‐through‐time plot revealed that the main environmental diversification preceded species diversification. A constant speciation through time was rejected, and the negative gamma indicates that most of the diversification occurred early in the history of the clade. According to the inferred phylogeny, the genus Orthopsyche McFarlane is a synonym under Caledopsyche Kimmins, and Abacaria caledona Oláh & Barnard should also be included in Caledopsyche. Main conclusions The age of the radiation does not support a vicariance origin of New Caledonian Hydropsychinae caddisflies. Environmental diversification pre‐dates lineage diversification, and thus environmental heterogeneity potentially played a role in the diversification of the group, by providing a variety of fragmented habitats to disperse into, promoting speciation. The negative gamma indicates that the speciation rate slowed as niches started to fill.