Updated checklist, habitat affinities, and changes over time of the Indiana (USA) caddisfly fauna (Insecta, Trichoptera).

Based on recent collecting and a synthesis of ~100 years of historical data, 219 caddisfly species are reported from the state of Indiana. Seventeen species are reported herein from the state for the first time, including two previously thought to be endemic to the southeastern USA. Species records are also presented herein organized by drainage basin, ecoregion, glacial history, and waterbody type for two distinct time periods: before 1983 and after 2005. More species were reported from the state before 1983 than after 2005, despite collecting almost 3× the number of occurrence records during the latter period. Species occurrence records were greater for most families and functional feeding groups (FFGs) for the post-2005 time period, although the Limnephilidae, Phryganeidae, Molannidae, and Lepidostomatidae, particularly those in the shredder FFG, instead had greater records before 1983. This loss of shredders probably reflected the ongoing habitat degradation within the state. While species rarefaction predicts only a few more species to be found in Indiana, many regions still remain under-sampled and 44 species have not been collected in >40 years.

Distribution patterns of Ohio stoneflies, with an emphasis on rare and uncommon species.

Presently, 102 stonefly species (Plecoptera) have been reported from Ohio. All 9 Nearctic families are represented. Over 90% of the fauna exhibit a combination of broad Nearctic-widespread, eastern Nearctic-widespread, Appalachian, and eastern Nearctic-unglaciated distributions. In contrast, only 2 species display a central Nearctic-Prairie distribution. Seven species of Perlidae are likely no longer present (Acroneuria evoluta Klapálek, A. perplexa Frison, Attaneuria ruralis (Hagen), and Neoperla mainensis Banks) or have experienced marked range reductions (Acroneuria abnormis (Newman), A. frisoni Stark and Brown, and A. filicis Frison). Another nearly 31% of the fauna (32 species) are rare, uncommon, or have highly-limited distributions within the state. Twelve of these species have Appalachian distributions, and an additional 8 have eastern Nearctic-unglaciated distributions. The distributional status for each of the 32 rare/uncommon species is discussed.

Checklist of the caddisflies (Insecta, Trichoptera) of the Upper Midwest region of the United States.

Five hundred and fifty-two caddisfly species are reported from the Upper Midwest region of the United States, an area that includes 13 states and ~ 2 million km2. Of these, 62 species are reported for the first time from the state of Iowa, 25 from Wisconsin, 18 from South Dakota, 12 from Illinois, five from Indiana, four from North Dakota, four from Minnesota, and one from Nebraska. The Upper Midwest fauna contains nearly 40% of all species known from the United States and Canada, as well as 22 species endemic to the region. Overall species richness was highest in Michigan (319 species), Kentucky (296), Minnesota (292), and Wisconsin (284). Differences in state species assemblages within the region largely followed a geographic pattern, with species richness declining in the western prairie states. There are almost certainly further species remaining to be found in this large region.

Towards a comprehensive barcode library for arctic life - Ephemeroptera, Plecoptera, and Trichoptera of Churchill, Manitoba, Canada.

BACKGROUND: This study reports progress in assembling a DNA barcode reference library for Ephemeroptera, Plecoptera, and Trichoptera ("EPTs") from a Canadian subarctic site, which is the focus of a comprehensive biodiversity inventory using DNA barcoding. These three groups of aquatic insects exhibit a moderate level of species diversity, making them ideal for testing the feasibility of DNA barcoding for routine biotic surveys. We explore the correlation between the morphological species delineations, DNA barcode-based haplotype clusters delimited by a sequence threshold (2%), and a threshold-free approach to biodiversity quantification--phylogenetic diversity. RESULTS: A DNA barcode reference library is built for 112 EPT species for the focal region, consisting of 2277 COI sequences. Close correspondence was found between EPT morphospecies and haplotype clusters as designated using a standard threshold value. Similarly, the shapes of taxon accumulation curves based upon haplotype clusters were very similar to those generated using phylogenetic diversity accumulation curves, but were much more computationally efficient. CONCLUSION: The results of this study will facilitate other lines of research on northern EPTs and also bode well for rapidly conducting initial biodiversity assessments in unknown EPT faunas.

Ecosystem Services, Global Diversity, and Rate of Stonefly Species Descriptions (Insecta: Plecoptera).

Stoneflies (Insecta: Plecoptera) provide ecosystem services as indicators of water quality, as food for predators, as mediators of energy flow and nutrient cycling, and through cultural services related to recreation and artistic creativity. The Plecoptera Species File (PSF) aggregates stonefly nomenclature, distribution, and literature to help society and scientists understand the value of services stoneflies provide. Using PSF data, we examined global and regional diversity, compared species description rates, and predicted future species description numbers through the year 2100. Through 2018, extant species totaled 3,718 with Temperate Asia having the greatest regional diversity at 1178 species. The Perlidae was the most species-rich of the 16 families at 1120 species. The recent global rate of species description was 43.6 species/yr, with Temperate Asia having the highest regional rate at 13.7 species/yr, followed by China and South America adding approximately 9.0 species/yr. We predicted that 1140 ± 130 new species would be described globally by 2050, and 2130 ± 330 by the year 2100, most of the increase occurring in China and South America. We discuss the possibility of reaching these predicted values.

Plecoptera of Canada.

Currently, a total of 267 stonefly species are known for Canada. The biodiversity hotspot of Canadian stoneflies is British Columbia with at least 138 species, nearly 52% of all species known from Canada. Four families, the Perlodidae, Capniidae, Chloroperlidae, and Nemouridae, contain nearly 75% of all species known to occur in Canada. The family with the fewest species represented in Canada is the Peltoperlidae. The stonefly fauna of Canada consists of two major faunal assemblages, west and east. The western clade consists of those species inhabiting Manitoba, all provinces to the west, and the three territories. The eastern clade consists of species from Ontario eastward. The two clades share only 29 species (10.9% of the Canadian total), suggesting a separate origin for each clade. The available taxonomic literature for the stoneflies of Canada is reviewed.

A new stonefly species (Plecoptera, Perlidae) from the Interior Highlands USA, with morphological and molecular comparison to other congeneric species.

Thirty-one species of Nearctic Perlesta Banks, 1906 (Plecoptera: Perlidae) are recognized. A new species is described from western Arkansas and eastern Oklahoma, USA, Perlestasublobata South & DeWalt, sp. nov., from the adult male, adult female, and egg. Perlestasublobata males are differentiated from other congeners by a combination of a prominent ventral caecum and a distinct dorsal extension of the lateral sclerites of the aedeagus. A preliminary molecular phylogenetic hypothesis is proposed for Perlesta based on 17 congeners and three outgroup taxa using partial mitochondrial cytochrome c oxidase subunit I sequence data. Illustrations, stereomicroscope images, and scanning electron micrographs support the description and comparison to other Perlesta.

Effects of Stream Temperature and Substrate Type on Emergence Patterns of Plecoptera and Trichoptera From Northeastern United States Headwater Streams.

The timing and spatial distribution of aquatic insect emergence is linked to the abiotic and biotic environment in streams. Studies of aquatic insect emergence are needed to generate baseline data to identify potential shifts in phenology and habitat-related emergence with global change. The purpose of this study was to 1) compare the timing of Plecoptera (stonefly) species emergence between two streams with different thermal regimes and 2) characterize the distribution of emerging Plecoptera and Trichoptera (caddisflies) from wood, rock, gravel, and sand substrates in five forested, headwater streams. Emergence timing and duration varied among Plecoptera species, with Ostrocerca albidipennis (Walker) (Plecoptera: Nemouridae) emerging only in May and four species in the genus Leuctra (Plecoptera: Leuctridae) collectively emerging throughout the summer (May to September). We observed earlier emergence of Amphinemura nigritta (Provancher) (Plecoptera: Nemouridae) and a longer total emergence period for Leuctra ferruginea (Walker) (Plecoptera: Leuctridae) in the stream with ~1.5°C warmer temperatures, which suggested that some insects may experience phenological shifts in streams with subtle differences in temperature. The abundance of plecopteran and trichopteran taxa emerging from wood was generally greater than for gravel or sand, and sand was the least preferred emergence substrate. The results suggest that human actions that decrease large wood and increase fine sedimentation may decrease habitat quality for many insect larvae and limit preferred emergence substrates.

Perlesta armitagei n. sp. (Plecoptera: Perlidae): More cryptic diversity in darkly pigmented Perlesta from the eastern Nearctic.

Perlesta Banks, 1906 (Plecoptera: Perlidae) is a genus of small, summer-emergent stoneflies known primarily from the eastern Nearctic. Thirty-two species are currently recognized, including two from China and nymphs have been reported from Costa Rica. We report here on some cryptic diversity within a small group of Perlesta with dark wings and bodies. Perlesta armitagei sp. nov. is described from the adult male, adult female, and egg. Diagnostic characters are presented with light microscope and scanning electron microscope (SEM) photomicrographs. Perlesta armitagei sp. nov. most closely resembles P. browni Stark, 1989 and P. cinctipes (Banks, 1905), two species distributed mainly within the Interior Highland Region. Perlesta armitagei sp. nov. is known currently within the Ohio River drainage from Indiana eastward to western Pennsylvania and southward into central Kentucky. Comparative light microscope and SEM images are also provided for P. adena Stark, 1989, P. browni, P. cinctipes, and P. xube Stark and Rhodes, 1997 in an effort to better define the morphological concepts of these dark colored species.

Wet collections accession: a workflow based on a large stonefly (Insecta, Plecoptera) donation.

This study details a workflow used to accession a large stonefly (Plecoptera) collection resulting from several donations. The eastern North American material of Kenneth W. Stewart (deceased, University of North Texas), the entire collection of Stanley W. Szczytko (deceased, University of Wisconsin, Stevens Point), and a small portion of the Barry C. Poulton collection (active, United States Geological Survey, Columbia, Missouri) were donated to the Illinois Natural History Survey in 2013. These 5,767 vials of specimens were processed to help preserve the specimen legacy of these world renowned Plecoptera researchers. The workflow used an industrialized approach to organize the specimens taxonomically, image the specimens and labels, and place the specimens into new storage. Utilizing the images as a verbatim data source, we transcribed labels in iterative steps that yielded more information with each pass. The data were normalized, locations georeferenced, all specimen data formatted to meet Darwin Core Archive format for occurrence data, and a data set created using Pensoft's Integrated Publishing Toolkit. This is the first time that any of the specimen data has been made available electronically. We also provide two important electronic supplements that include the Bill P. Stark (active, Mississippi College) Oklahoma field notebook for 1971 and 1972 detailing locations for many coded stonefly specimens in the Stewart collection, and the coded locations of B. C. Poulton's Arkansas and Missouri study. Again, we have linked coded labels in vials to normalized and georefenced site data. We confirmed 243 stonefly species were contained within the collections, and the potential for many more species exists among the specimens identified to family and genus level. Twenty-one new state, province, and other significant stonefly records are reported herein with all identifications verified by the senior author, often through consultation with other stonefly taxonomists. Researchers are encouraged to utilize the specimen data, form collaborations with the authors, and borrow specimens for research.

Updated checklist of the Michigan (USA) caddisflies, with regional and habitat affinities.

Based on examination of ~180,000 specimens from 695 collections of 443 localities collected from the 1930s to 2015 we report 295 species of caddisflies from Michigan. Of these, 41 are reported from the state for the first time. Another 18 species previously reported from Michigan are listed as doubtful. The 11 most abundant species collectively represented over half of all specimens collected. Conversely, 80 species were known from <10 specimens, and 27 species from a single specimen. The Michigan fauna is similar to those of Minnesota and Ohio, adjacent states with comparable recent collecting effort. Regional and habitat affinities for each Michigan species are reported herein. Due to the high level of species discovery over the last few years, despite a >80-year collecting history, it is likely that additional species remain undiscovered in the state.

Mayflies, stoneflies, and caddisflies of streams and marshes of Indiana Dunes National Lakeshore, USA.

United States National Parks have protected natural communities for one hundred years. Indiana Dunes National Lakeshore (INDU) is a park unit along the southern boundary of Lake Michigan in Indiana, USA. An inventory of 19 sites, consisting of a seep, 12 streams, four marshes, a bog, and a fen were examined for mayflies (Ephemeroptera), stoneflies (Plecoptera), and caddisflies (Trichoptera) (EPT taxa). Volunteers and authors collect 35 ultraviolet light traps during summer 2013 and supplementary benthic and adult sampling added species not attracted by lights or that were only present in colder months. Seventy-eight EPT species were recovered: 12 mayflies, two stoneflies, and 64 caddisflies. The EPT richness found at INDU was a low proportion of the number of species known from Indiana: caddisflies contributed only 32.7% of known state fauna, mayflies and stoneflies contributed 8.4% and 2.3%, respectively. Site EPT richness ranged from one for a seep to 34 for an 8 m-wide stream. Richness in streams generally increased with stream size. Seven new state records and rare species are reported. The number of EPT species at INDU is slightly larger than that found at Isle Royale National Park in 2013, and the community composition and evenness between orders were different.

Atlas of Ohio Aquatic Insects: Volume II, Plecoptera.

BACKGROUND: We provide volume II of a distributional atlas of aquatic insects for the eastern USA state of Ohio. This treatment of stoneflies (Plecoptera) is companion to Armitage et al. (2011) on caddisflies (Trichoptera). We build on a recent analysis of Ohio stonefly diversity patterns based on large drainages (DeWalt et al. 2012), but add 3717 new records to the data set. We base most analyses on the United States Geological Survey Hierarchical Unit Code eight (HUC8) drainage scale. In addition to distributional maps for each species, we provide analyses of species richness versus HUC8 drainage area and the number of unique locations in a HUC8 drainage, species richness versus Ohio counties, analyze adult presence phenology throughout the year, and demonstrate stream size range affiliation for each species. NEW INFORMATION: This work is based on a total of 7797 specimen records gathered from 21 regional museums, agency data, personal collections, and from the literature Table 1. To our knowledge this is the largest stonefly data set available for a similarly sized geopolitical area anywhere in the world. These data are made available as a Darwin Core Archive supported by the Pensoft Integrated Publishing Toolkit (DeWalt et al. 2016b). All known published papers reporting stoneflies from Ohio are detailed in Suppl. material 1. We recovered 102 species from Ohio, including all nine Nearctic families Table 2​. Two species were removed from the DeWalt et al. (2012) list and two new state records added. Perlidae (32 spp.) was most speciose, compared to the low diversity Pteronarcyidae (2 spp.) and Peltoperlidae (1 sp.). The richest HUC8 drainages occurred in northeastern, south-central, and southern regions of the state where drainages were heavily forested, had the highest slopes, and were contained within or adjacent to the unglaciated Allegheny and Appalachian Plateaus. Species poor drainages occurred mainly in the northwestern region where Wisconsinan aged lake plains climaxed to an expansive wooded wetland, the Black Swamp. The unglaciated Lower Scioto drainage (72 spp.) in south-central Ohio supported the greatest species richness. There was no relationship between species richness and HUC8 drainage size, but the number of unique locations in a drainage strongly related to species richness. All Ohio counties were represented in the data set with Hocking County (59 spp.) of the Lower Scioto drainage being the richest and most heavily sampled. Adult presence phenology was influenced by phylogenetic relationships such that the superfamily Nemouroidea (Capniidae, Leuctridae, Nemouridae, and Taeniopterygidae) generally emerged in winter and spring while the superfamilies Pteronarcyoidea (Pteronarcyidae, Peltoperlidae) and Perloidea (Chloroperlidae, Perlidae, Perlodidae) emerged later, some species continuing emergence through summer months. Species often occupied specific stream size ranges, while others were generalists. Two species once histrorically abundant in the western Lake Erie Bass Islands no longer reside there. Each of the 102 species is discussed in detail, including several that require additional collecting efforts to confirm their identities, presence, and distribution in Ohio.

Ephemeroptera, Plecoptera, and Trichoptera on Isle Royale National Park, USA, compared to mainland species pool and size distribution.

Extensive sampling for aquatic insects was conducted in the orders Ephemeroptera (mayflies), Plecoptera (stoneflies), and Trichoptera (caddisflies) (EPT) of Isle Royale National Park (ISRO), Michigan, United States of America, during summer 2013. The island was ice covered until 8,000 to 10,000 years ago and is isolated by 22-70 km distance from the mainland. Two hypotheses were examined: that ISRO EPT richness would be much reduced from the mainland, and that the species colonizing ISRO would be of smaller size than mainland, adults presumably using updrafts to bridge the distance from mainland sources. Data sets were developed for known mainland EPT species and size for those species. The first hypothesis was confirmed with the mainland species pool consisting of 417 EPT, while ISRO is known to support 73 species. Richness of EPT is directly related to the number of specimens examined. Small streams supported five EPT species, while 15-25 species were found in larger streams. Lakeshores had intermediate diversity. The second hypothesis was substantiated for stoneflies, but not for mayflies or caddisflies. Stoneflies apparently are poorer fliers than either of the other two orders.

A review of the Nearctic genus Prostoia (Ricker) (Plecoptera, Nemouridae), with the description of a new species and a surprising range extension for P. hallasi Kondratieff & Kirchner.

The Nearctic genus Prostoia (Plecoptera: Nemouridae) is reviewed. Prostoia ozarkensis sp. n. is described from the male and female adult stages mainly from the Interior Highland region encompassing portions of Arkansas, Missouri, and Oklahoma. Prostoia ozarkensis sp. n. appears most closely related to two species, one distributed broadly across the western Nearctic region, P. besametsa (Ricker), and one found widely throughout the central and eastern Nearctic regions, P. completa (Walker). A surprising range extension is noted for P. hallasi Kondratieff & Kirchner, a species once known only from the Great Dismal Swamp, from small upland streams in southern Illinois. Additional new state records are documented for P. besametsa, P. completa, P. hallasi and P. similis (Hagen). Taxonomic keys to Prostoia males and females are provided, and scanning electron micrographs of adult genitalia of all species are given.

A review of the Nearctic genus Zealeuctra Ricker (Plecoptera, Leuctridae), with the description of a new species from the Cumberland Plateau region of eastern North America.

The stonefly genus Zealeuctra (Plecoptera: Leuctridae) is endemic to the central and eastern Nearctic regions and is presently comprised of 10 species. Scanning electron microscopy (SEM) was used to examine and redescribe two important diagnostic features typically used to identify and define the adult male stage: the large, anteriorly-recurved epiproct and the medial cleft of the ninth abdominal tergite. SEM was also employed to depict the posteromedial portion of female 7(th) sternum. A new species, Z. ukayodi sp. n., is described from the Cumberland Plateau region of northeastern Alabama and Tennessee. The new species appears superficially similar to Z. talladega Grubbs, but is easily differentiated by characteristics of the male medial cleft. An updated taxonomic key to the males of Zealeuctra is provided.

Perlesta ephelida, a new Nearctic stonefly species (Plecoptera, Perlidae).

A new Nearctic species of Perlidae (Insecta, Plecoptera), Perlesta ephelidasp. n., is described from the male, female, and egg stages. This species has been previously reported as, or confused with, Perlesta shubuta Stark from several central and eastern U.S. states. Perlesta ephelida is distinctive from Perlesta shubuta and other regional Nearctic congeners mainly according to male genitalic and egg characteristics. Perlesta ephelida is a widely-distributed eastern Nearctic species, whereas Perlesta shubuta appears to be restricted to a narrow latitudinal belt in the Gulf Coast region from Louisiana east conservatively to the Florida panhandle. The egg of Perlesta shubuta is depicted with scanning electron microscopy for the first time.

Ohio USA stoneflies (Insecta, Plecoptera): species richness estimation, distribution of functional niche traits, drainage affiliations, and relationships to other states.

Ohio is an eastern USA state that historically was >70% covered in upland and mixed coniferous forest; about 60% of it glaciated by the Wisconsinan glacial episode. Its stonefly fauna has been studied in piecemeal fashion until now. The assemblage of Ohio stoneflies was assessed from over 4,000 records accumulated from 18 institutions, new collections, and trusted literature sources. Species richness totaled 102 with estimators Chao2 and ICE Mean predicting 105.6 and 106.4, respectively. Singletons and doubletons totaled 18 species. All North American families were represented with Perlidae accounted for the highest number of species at 34. The family Peltoperlidae contributed a single species. Most species had univoltine-fast life cycles with the vast majority emerging in summer, although there was a significant component of winter stoneflies. Nine United States Geological Survey hierarchical drainage units level 6 (HUC6) were used to stratify specimen data. Species richness was significantly related to the number of unique HUC6 locations, but there was no relationship with HUC6 drainage area. A nonparametric multidimensional scaling analysis found that larger HUC6s in the western part of the state had similar assemblages with lower species richness that were found to align with more savanna and wetland habitat. Other drainages having richer assemblages were aligned with upland deciduous and mixed coniferous forests of the east and south where slopes were higher. The Ohio assemblage was most similar to the well-studied fauna of Indiana (88 spp.) and Kentucky (108 spp.), two neighboring states. Many rare species and several high quality stream reaches should be considered for greater protection.

A DNA barcode library for North American Ephemeroptera: progress and prospects.

DNA barcoding of aquatic macroinvertebrates holds much promise as a tool for taxonomic research and for providing the reliable identifications needed for water quality assessment programs. A prerequisite for identification using barcodes is a reliable reference library. We gathered 4165 sequences from the barcode region of the mitochondrial cytochrome c oxidase subunit I gene representing 264 nominal and 90 provisional species of mayflies (Insecta: Ephemeroptera) from Canada, Mexico, and the United States. No species shared barcode sequences and all can be identified with barcodes with the possible exception of some Caenis. Minimum interspecific distances ranged from 0.3-24.7% (mean: 12.5%), while the average intraspecific divergence was 1.97%. The latter value was inflated by the presence of very high divergences in some taxa. In fact, nearly 20% of the species included two or three haplotype clusters showing greater than 5.0% sequence divergence and some values are as high as 26.7%. Many of the species with high divergences are polyphyletic and likely represent species complexes. Indeed, many of these polyphyletic species have numerous synonyms and individuals in some barcode clusters show morphological attributes characteristic of the synonymized species. In light of our findings, it is imperative that type or topotype specimens be sequenced to correctly associate barcode clusters with morphological species concepts and to determine the status of currently synonymized species.