John C. Morse

Freshwater biomonitoring with macroinvertebrates in East Asia

This paper summarizes the history and current status of efforts to implement macroinvertebrate biomonitoring protocols for surface water pollution in China, Japan, South Korea, Malaysia, Mongolia, Russia (Far East), and Thailand. Impediments to biomonitoring in some of these countries include: (1) lack of knowledge about macroinvertebrate fauna and their tolerance values, especially during the aquatic, immature stages; (2) the scarcity of research programs and formal training opportunities for biomonitoring offered in universities; (3) the shortage of high-quality microscopes and other necessary equipment; and (4) limited government understanding and support for biomonitoring, few skilled regulatory staff, and the persistence of old and unusable biomonitoring protocols. A recently established regional network, the Aquatic Entomological Society of East Asia (AESEA), and several major recent publications are helping to coordinate and promote science and technology in East Asia.

Associating larvae and adults of Chinese Hydropsychidae caddisflies (Insecta:Trichoptera) using DNA sequences

Abstract The utility of hydropsychid (Trichoptera:Hydropsychidae) caddisfly larvae for freshwater biomonitoring has been demonstrated, but the major impediment to its implementation has been the lack of species-level larval descriptions and illustrations. A rapid and reliable molecular protocol that also uses morphology is proposed because conventional approaches to associating undescribed larvae with adults have been slow and problematic. Male adults were identified before DNA sequence analyses were used. These identifications established morphospecies boundaries that were mapped on phylograms constructed from 2 independent gene fragments: mitochondrial cytochrome c oxidase subunit I (COI) and large subunit (28S) nuclear ribosomal DNA expansion fragment D2 (D2). Species boundaries were confirmed if they were monophyletic on both molecular phylograms. Larval associations were made with reference to the phylogenetic analyses under 2 criteria: sequence identity across both genes or nested placement within a reference species boundary. A total of 133 individuals belonging to Chinese Hydropsyche sensu lato group (including Hydropsyche [Hydropsyche], Hydropsyche [Occutanspsyche], Ceratopsyche, Mexipsyche, Hydatomanicus, and Herbertorossia) were included in our study to test the new protocol. D2 sequences (all individuals) and COI sequences (101 individuals) were obtained, and 2 independent phylograms were constructed using neighbor joining. Both fragments provided enough nucleotide changes to differentiate independently most Hydropsyche sensu lato species, with ambiguity in only a few species that eventually could be resolved with additional sequences and specimens. COI diverges significantly within some species, suggesting a need for caution when applying typical genetic divergence thresholds in species diagnoses. The study enabled us to establish a procedure for delimiting species boundaries and associating larvae and adults using DNA sequences and morphological evidence. Ideal sampling strategies for larval–adult association are suggested. Associating larvae and adults of hydropsychids using DNA sequences appears to be promising in terms of both reliability and speed.

Evolution of feeding and case-making behavior in Trichoptera

A phylogeny of the families of Trichoptera is reviewed to provide a basis for understanding the probable evolution of feeding tactics and case or retreat constructions by larvae. At least 48 hierarchically inclusive homologues are known, mostly from larval, pupal, and adult morphology. Their resulting phylogeny indicates that Rhyacophilidae, Hydrobiosidae, Glossosomatidae, and Hydroptilidae are more closely related to Philopotamidae, Hydropsychidae, and their allies than to Limnephilidae, Leptoceridae, and their allies. This phylogeny implies that the ancestral caddisfly larva was probably a tube-dwelling detritivore, inhabiting humus and detrital mats near the shores of lentic or lotic-depositional habitats. This ancestor evolved into a tube-case-making detritivore and scraper in the ancestor of Integripalpia and into a retreat-making collector-gatherer in the ancestor of Annulipalpia. All other larval feeding and case-making tactics evolved from these ancestral habits.

A Phylogeny and Classification of Family-Group Taxa of Leptoceridae (Trichoptera)

Until now only four tribes have been defined in Leptoceridae, all in the subfamily Leptocerinae. Genera in Triplectidinae and most genera described since 1955 in Leptocerinae have never been assigned to tribes. Based on observations by Ulmer and on recent phylogenetic studies two of these four tribes (Mystacidini and Leptocerini) are paraphyletic groups. Consequently, as a conceptual framework for predictive biology and for future generic revisions, a new phylogeny and tribal classification are proposed for the family and its 45 currently recognized genera. The resulting eleven tribes in two subfamilies include six new family-group names.

The Trichoptera barcode initiative: a strategy for generating a species-level Tree of Life.

DNA barcoding was intended as a means to provide species-level identifications through associating DNA sequences from unknown specimens to those from curated reference specimens. Although barcodes were not designed for phylogenetics, they can be beneficial to the completion of the Tree of Life. The barcode database for Trichoptera is relatively comprehensive, with data from every family, approximately two-thirds of the genera, and one-third of the described species. Most Trichoptera, as with most of lifes species, have never been subjected to any formal phylogenetic analysis. Here, we present a phylogeny with over 16 000 unique haplotypes as a working hypothesis that can be updated as our estimates improve. We suggest a strategy of implementing constrained tree searches, which allow larger datasets to dictate the backbone phylogeny, while the barcode data fill out the tips of the tree. We also discuss how this phylogeny could be used to focus taxonomic attention on ambiguous species boundaries and hidden biodiversity. We suggest that systematists continue to differentiate between ‘Barcode Index Numbers’ (BINs) and ‘species’ that have been formally described. Each has utility, but they are not synonyms. We highlight examples of integrative taxonomy, using both barcodes and morphology for species description. This article is part of the themed issue ‘From DNA barcodes to biomes’.

The aberrant spermatozoa of Hydropsychidae caddisflies (Trichoptera): an electron microscope analysis on spermiogenesis.

The spermatozoon of caddisflies in the family Hydropsychidae lacks acrosome and centriole adjunct. The basal body and nucleus are separated. The axoneme is cone shaped and its microtubular doublets diverge, forming separate filaments which trail backward from the cell surface. The mitochondria do not fuse, conserve their cristae, form no derivatives, and appear within the lumen of the aberrant axoneme. Some of those may be secondarily derived characteristics since they are not present in early spermatids. Spermatozoa isolated into a Ringer solution from ejaculatory ducts or inseminated females show no autonomous movements. In the female, the spermatozoa may be moved by peristaltic contractions of the genital duct. The direction of their movement may be determined by tubular projections of the duct wall which engage the trailing microtubular doublets, thus impeding backward movement of the spermatozoa. No apyrene (anucleated) spermatozoa, similar to those characterizing Lepidoptera, could be found.

Defining the genus Hydropsyche (Trichoptera:Hydropsychidae) based on DNA and morphological evidence

Abstract In this paper, we review the history of Hydropsychinae genus-level classification and nomenclature and present new molecular evidence from mitochondrial cytochrome c oxidase subunit I (COI) and nuclear large subunit ribosomal ribonucleic acid (28S) markers supporting the monophyly of the genus Hydropsyche. Both molecular and morphological characters support a broad conservative definition of Hydropsyche. Caledopsyche, Hydatomanicus, and Occutanspsyche are synonymized with Hydropsyche. The following species groups are established: Hydropsyche bronta Group (generally corresponding with Ceratopsyche and Hydropsyche morosa and newae Groups), Hydropsyche colonica Group (generally corresponding with Orthopsyche), Hydropsyche instabilis Group (generally corresponding with Hydropsyche s.s.), and Hydropsyche naumanni Group (generally corresponding with Occutanspsyche). Molecular data recovered Hydromanicus as paraphyletic, and Cheumatopsyche and Potamyia as sister taxa. The genus names Plectropsyche and Streptopsyche are reinstated.

Pseudoneureclipsinae in Dipseudopsidae (Trichoptera: Hydropsychoidea), with Descriptions of Two New Species of Pseudoneureclipsis from East Asia

A phylogenetic analysis of all known life history stages of Hydropsychoidea implies that Pseudoneureclipsinae should be removed from Polycentropodidae and placed in Dipseudopsidae. Two synapomorphies especially informative for this conclusion are the fused condition of female sternum VIII sclerites and the presence of a long larval spinneret without labial palpi. A new species, Pseudoneureclipsis tiani Li, sp. n., is described, representing the first report of this genus from China. A male lectotype and two male paralectotypes of P. ussuriensis Martynov are designated from the Primorye Region of Far East Russia and distinguished from specimens described but not named by Botosaneanu in 1970 from Korea. These latter are herein named Pseudoneureclipsis botosaneanui Morse, sp. n.

Higher Classification of Triplectidinae (Trichoptera : Leptoceridae)

A phylogenetic analysis of 29 morphological characters of larvae, pupae, and adults for the 22 nominal genus-group taxa of Triplectidinae implies that the three tribes Grumichellini (including Atanatolica, Grumichella, and Triplexa), Hudsonemini (including Condocerus, Hudsonema, and Notalina) and Triplectidini (including Notoperata, Triplectides, and Westriplectes) are each monophyletic and the latter two are sister tribes. The other 13 nominal genus-group taxa are probably synonyms of these genera. At least 69 of the 124 known species of the subfamily are included in Triplectides, suggesting a need to recognize 17 or more subgenera within it for conceptual and predictive purposes.

Two new species of the genus Plectrocnemia Stephens (Trichoptera: Polycentropodidae) from the Russian Far East

Two new species of the genus Plectrocnemia Stephens, Plectrocnemia levanidovae, sp. n., from Sakhalin Island and the southern Kuril Islands, and Plectrocnemia martynovae, sp. n., from southern Primorye, are described and illustrated. The new species belong to the P. tortosa Group and are the sister species of P. tortosa Banks, from China, and P. appensata Mey, from Vietnam.