Two new species of the stonefly genus Amphinemura (Insecta,Plecoptera, Nemouridae) from China

Two new species of the genus Amphinemura Ris from China are described and illustrated, i.e. A. annulata Du & Ji, sp. n. from Zhejiang, Shanxi, Shaanxi and Guizhou Province, and A. lingulata Du & Wang, sp. n. from Shaanxi and Sichuan Province. A. annulata is similar to A. tricintusidens Wang & Zhu in having an apical cavity of the epiproct, but the epiproct ventral sclerite and the median paraproct lobe of the two species are different. A. lingulata is related to A. didyma Zhu & Yang in having the similar epiproct, but they differ mostly in paraproct median and outer lobes.     

Molecular systematics and biogeography of the western Mediterranean stonefly genus Tyrrhenoleuctra (Insecta, Plecoptera)

The stonefly genus Tyrrhenoleuctra includes species living in western Mediterranean temporary freshwater streams, sometimes also at sea level, a very unusual habitat for most Plecoptera. Traditional morphological approaches proved unsuccessful in drawing both taxonomic and phylogenetic patterns, thus hampering discussion of biogeographical patterns for this interesting group. We aimed at: (a) assessing the taxonomic status of populations of Tyrrhenoleuctra covering the geographic range of the genus; (b) studying the phylogenetic relationships among the recognized species; and (c) describing biogeographic patterns. We used phylogenetic analyses to infer the phylogenetic history of this group of stoneflies based on a combined data set of 1666 bp including fragments of the 12S ribosomal (12S) and cytochrome oxidase I (CO‐I) mtDNA genes, with maximum likelihood and Bayesian methods. Two main clades have been identified: a Sardo‐Corsican one, including Tyrrhenoleuctra zavattarii, and an Ibero‐Maghrebian one including four lineages of unkown taxonomic rank from the Balearic Islands (Maiorca), from northern Africa (Ceuta) and southernmost Spain (Algeciras), and a complex preliminarily referred herein to T. minuta (Klapálek, 1901), which includes two lineages, one from Cordoba, and one from Sierra de Grazalema (El Cerro) and Portugal (Tellhares) respectively. Dating the nodes by fixing the split of the Ibero‐Maghrebian clade from the Sardo‐Corsican one at 29 million years ago (Mya), yielded dates referring to the major geological events in the Mediterranean basin. Estimated molecular evolutionary rates ranged from 0.02–0.09% per million years (my) in the T. zavattarii lineages, to 0.2–0.7% per MY in the Ibero‐Maghrebian clade. The phylogenetic pattern emerged from the present study is congruent with the known paleo‐history of the western Mediterranean basin, with the divergence of the two main Tyrrhenoleuctra lineages corresponding to the split of the Sardo‐Corsican microplate from the Iberian block. Vicariance events have characterized the history of this stonefly group along its entire biogeographical history. Surprisingly low evolutionary rates, previously supposed by Fochetti (1991, 1994) and Fochetti et al. (2004) based on nuclear markers (allozymes), have been herein found also in mitochondrial markers.     

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.     

Molecular systematics,evolution and zoogeography of the stonefly genus Siphonoperla (Insecta: Plecoptera,Chloroperlidae)

The stonefly genus Siphonoperla Zwick, 1967 (Chloroperlidae) ranges from the Atlas Mountains of North Africa, throughout Europe and east to the Lesser Caucasus Mountains in Armenia. Systematic relationships within the genus are unknown. We provide the first molecular phylogenetic reconstruction of the genus, based on sequence variation of 1348 bp of the mitochondrial cytochrome c oxidase subunit I gene and relate this phylogeny to a few key morphological features as well as zoogeographical perspectives. A total of 15 taxa (32 individuals), including four outgroup species (Chloroperla Newman, 1836 and Xanthoperla Zwick, 1967 ) were screened. Monophyly of Siphonoperla, as well as each purported taxon was supported, with net pairwise distances ranging from 0.7% to 13.7%. Within‐taxon variability ranged from 0% to 6.2%, whereby the upper value relates to Siphonoperla torrentium Pictet, 1841 represented in our sample by three allopatric subspecific taxa. These taxa, displayed as a polytomy with an estimated time to the most recent common ancestor of about 5.4 Myr, are also shown to have distinct genitalia. These data could be used to support a species‐level distinction for the three subspecific taxa. Siphonoperla ranged from 13% (Xanthoperla) to 16.2% (Chloroperla) divergent from the outgroup genera. The recently described Siphonoperla ottomoogi, Graf, 2008 from Austria is not closely related to the sympatric Siphonoperla montana Pictet, 1841 and is a minimum of 7.8% divergent from congenerics, supporting its status as a micro‐endemic relict surviving on the edge of Alpine glaciation throughout the Pleistocene. Overall, the genus shows the highest levels of diversification in the Mediterranean and Southeast regions whereby at least some of the species found in Central Europe today may have persisted there in peri‐glacial refugia throughout the Pleistocene.     

Digestive enzyme activity and trophic behavior in two predator aquatic insects (Plecoptera, Perlidae): a comparative study

Plecoptera (Perlidae) are among the major macroinvertebrate predators in stream ecosystems and one of the insect families with lower tolerance to environmental alterations, being usually employed as bioindicators of high water ecological quality. The differences in the trophic roles of the coexisting species have been exclusively studied from their gut contents, while no data are available on the comparative digestive capacity. In the present paper, we make a comparative study of the activity of several digestive enzymes, namely proteases (at different pH), amylase, lipase, trypsin and chymotrypsin, in two species of stoneflies, Perla bipunctata and Dinocras cephalotes, which cohabit in the same stream. The study of digestive enzyme activity together with the analysis of gut contents can contribute to a better understanding of the ecology of these aquatic insects and their role in freshwater food webs. Thus, our results show that the two studied predator species inhabiting in the same stream present specializations on their feeding behaviors, facilitating their coexistence, and also differences in their capacity of use the resources. One of the main findings of this study is that D. cephalotes is able to assimilate a wider trophic resource spectrum and this could be one of the reasons why this species has a wider global distribution in all its geographical range.     

Impact of human activities on stonefly (Insecta,Plecoptera) ecological metrics in the Hron River (Slovakia)

A total of 57 stonefly species have been recorded in the Hron River. The natural gradient (slope and stream width) and pollution gradient of the river were defined using CCA based on physical, chemical and stonefly data. Stonefly metrics (abundance, richness/diversity, sensitivy/tolerance and functional metrics) were used to estimate the quality of the Hron River and the degree of proximity to its natural state. Similar results were obtained using two different methods. The first method was based on the homogeneity of variance and the interquartile range of different groups of stretches of the Hron River and the second was based on deviations from the expected values of biological metrics in a given stretch of the river. These values continuously decreased with increasing distance from the spring area, with the exception of the saprobic index, which increased in a downstream direction, and the stonefly average score, which did not change significantly along the whole river flow. The Stonefly Average Score (SAS) metric is universal for a variety of habitats such as the Hron River upstream and downstream, and is a reliable indicator of water quality and the natural course of a stream.     

Agnetina den, a new species of perlid stonefly (Plecoptera: Perlidae) from Vietnam

A new species of perlid stonefly, Agnetina den, is described from Vietnam. Male adults of this species can be easily distinguished by their black body color pattern, submedian patches of sensilla basiconica on hemitergites 7–8, and clusters of long dense fine setae on the inner sides the abdominal tergites. Illustrations of habitat and diagnostic characters are presented.     

Study of antioxidant defense in four species of Perloidea (Insecta, Plecoptera)

The aim of the present work is to conduct a comparative study of oxidative states in the nymphs of four species of Plecoptera belonging to the superfamily Perloidea: Perla marginata (Panzer, 1799) (family Perlidae), Guadalgenus franzi (Aubert, 1963), Isoperla curtate Navás, 1924, and lsoperla grammatica (Poda, 1761) (family Perlodidae) in relation to their ecological and biological characteristics. For this, the activity of the following antioxidant enzymes was determined: superoxide dismutase (SOD), catalase (CAT), glutathione reductase (GR), glutathione peroxidase (GPX), glutathione transferase (GST), and DT-diaphorase (DTD), together with lipid peroxidation. Glucose 6-phosphate dehydrogenase (G6PDH) was also determined. The four species studied were selected based on significant ecological and biological differences. The results obtained when studying different indicative parameters of the oxidative state of the nymph of different species showed that each has an important enzymatic antioxidant potential, and that differences among species are conditioned by the duration of the nymphal development period more than by whether they come from permanent or temporary habitats. Thus, Plecoptera, although traditionally considered as typical inhabitants of permanent waters, seem to have sufficient variability in physiological mechanisms, together with behavioral and ecological adaptations, to cope with potentially unfavorable conditions that may occur in temporary waters.     

Two new species of genus Rhopalopsole (Insecta, Plecoptera, Leuctridae) from China

Two new species of Rhopalopsole Klapálek from China are described: Rhopalopsole exiguspina Du & Qian, sp. n. and Rhopalopsole ampulla Du & Qian, sp. n., which were collected in Guizhou province, China.     

Life history and larval feeding of some species of Ephemeroptera and Plecoptera (Insecta) in the Sierra Nevada (Southern Iberian Peninsula)

We monitored two high mountain streams from the Sierra Nevada (Southern Spain) monthly throughout an annual cycle, measuring and analyzing physical and chemical parameters. Temperature was hourly registered. We studied life cycles and larval feeding of four mayfly species [Serratella ignita (Poda, 1761), Serratella spinosa nevadensis (Alba-Tercedor, 1981), Alainites muticus (Linnaeus, 1758), and Baetis alpinus Pictet, 1843] and two stonefly species [Amphinemura triangularis (Ris, 1902) and Capnioneura mitis Despax, 1932]. Some species changed their strategies (growth rate, life cycle duration, and/or voltinism) in accordance with different conditions (depending on the altitude and temperature). The gut content data showed that species fed mainly on detritus (as collector-gatherers or as shredders), except B. alpinus that behaved as a scraper in one of the streams. Differences in the larval diet of the same species in different streams, and even through its life, support the idea of not using general models to extrapolate the function of a species from a higher taxonomical level. Handling editor: D. Dudgeon     

Diel food habits of two species of setipalpian stoneflies (Plecoptera) in tributaries of the Clearwater River, Idaho

SUMMARY. The foreguts of two setipalpian stonefly nymphs, Skwala parallela and Hesperoperla pacifica , from tributaries of the Clearwater River, Idaho, were examined in terms of composition and fullness. A total of 736 nymphs were collected at 06.00,12.00,18.00 and 24.00 hours during a 1-week period in August in order to examine diel variation in both diet composition and feeding periodicity. Substantial diel variation in diet composition and Electivity Indices (E) occurred. No single prey taxon was consistently the principal component in the diet of either stonefly species during all 6-h sampling intervals. Overall, hydropsychids (15.3–60.6%) were the major food of H. pacifica in Mission Creek whereas chironomid larvae were the chief prey of S. parallela in Mission (10.0–47.2%) and Lapwai (14.9–28.0%) Creeks. Nymphs were nocturnal feeders with feeding increasing from 18.00 to 06.00 hours and with peak feeding occurring from 24.00 to 06.00 hours.     

Life cycle and nymphal feeding of Besdolus ravizzarum (Plecoptera: Perlodidae), a threatened stonefly

Abstract We have studied some aspects of Besdolus ravizzarum biology in the Apennines. This stonefly has a very localized distribution in southeastern France and northern Italy. This species is considered threatened. For this reason it is especially relevant to improve knowledge about its natural history, previously unknown. Despite the large size of nymphs, this species shows a univoltine life cycle. The growth is slow but uniform from October to April or May when the short flight period occurs. The analysis of the gut contents indicates this species is a predator, feeding mainly on Baetidae and Chironomidae. It is notable that the development of this carnivorous species through cold periods coincides with the season with higher richness and density of potential prey in Apennine streams. Further studies are needed to improve knowledge of the distribution of the species and to start planning effective conservation efforts.     

New species and records of the stonefly genus Neoperla (Plecoptera,Perlidae) from Jinhuacha Nature Reserve,Guangxi of China

Two new Neoperla species (Neoperla mesospina, Neoperla latispina) are described from the adult male stage from the Jinhuacha Nature Reserve, Guangxi of China. The new species are compared with similar taxa. Taxonomic remarks are also provided for N. transversprojecta Du & Sivec and N. yao Stark. The latter species is newly recorded for Guangxi.     

Alimentary canal formation in the stonefly,Kamimuria tibialis (pictet) (Plecoptera : Perlidae)

The alimentary canal formation in the stonefly, Kamimuria tibialis (Plecoptera : Perlidae) is described. The stomodaeum is formed as in other insect embryos. The proctodaeum is derived from the ectodermal fold an the caudal end of the embryo without the contribution of the amnion. The 3 Malpighian tubules develop from the blind end of the proctodaeum. The rectal pad is formed by the thickening of the dorsal wall of the proctodaeum. The midgut epithelium rudiment arises only from the blind end of the proctodaeum, i.e. it is completed by unipolar formation instead of bipolar. The yolk cells do not contribute to the formation of the midgut epithelium. The alimentary canal is transformed during the 1st nymphal instar and becomes functional in the next instar. The stomodaeum is differentiated into 3 parts: pharynx, oesophagus, and proventriculus. The midgut becomes shortened and its epithelium is well developed. Gastric caeca with tapering processes are formed.     

Interspecific variation in temperature dependence of egg development of five congeneric stonefly species (Protonemura Kempny, 1898, Nemouridae,Plecoptera)

Embryonal development of the five congeners Protonemura auberti Illies, 1954, P. hrabei Rauser, 1956, P. meyeri (Pictet, 1841), P. nitida (Stephens, 1835), and P. praecox (Morton, 1894) was studied under various laboratory temperatures and different photoperiods.Mean number of eggs in field collected batches was between 470 (P. praecox) and 1211 (P. auberti). Spring species had smaller egg batches than autumn species (Table 1). Mean hatching success in the laboratory was 50–100% at 2–18 °C. In most species hatching success decreased slightly with increasing temperature (Figs. 1a-e). None of the eggs incubated at 24 °C developed. Hatching pattern followed an asymmetric frequency distribution. In general, the hatching periods were the shorter the higher the incubation temperature.Embryonic development of all five species was inversely temperature dependent (Figs. 2a-e), and well described by a power law relationship (Figs. 3a-e). Interspecific differences in incubation periods were notable at nearly all temperatures (Fig. 4). There was a distinct interspecific sequence in length of incubation period (with steps of about 4 days), which was the same as can be seen in the flight periods: The later the species flies the longer the incubation period. Temperature fluctuations and variations in photoperiod had no influence on incubation and hatching periods or hatching success.The thermal demand of the egg stage neither explains the recent geographical distribution of the Protonemura species, nor does it directly correspond to the field temperatures common during their egg development. However, it is optimal in respect to resource partitioning between the five species, with the consequence of temporal displacement of life cycles.Derived from Brittain's (in press) proposal to compare the two constants a and b of the regressions describing the temperature dependence of embryonal development, a new index (Integral Development Time, IDT) indicating the thermal demand was created for easier comparison of numerous species (Table 2). Evaluation of the IDT for various species of Plecoptera (Fig. 5) suggests that species belonging to the family group Systellognatha generally have higher thermal requirements in the egg stage than species belonging to the Euholognatha.     

A revision of the Oriental stonefly genus Phanoperla (Plecoptera: Perlidae)

The genus Phanoperla (=Dyaperla Banks, 1939) (Plecoptera: Perlidae: Perlini) is revised and generic diagnoses are provided for adults and larvae. Diagnostic and constitutive characters of the tribe Neoperlini are discussed, and Chinoperla Zwick, 1980, is shown to be the closest relative of Phanoperla. Many past misidentifications of Phanoperla species have been corrected by the use of characters recently recognized as important, namely the structure of the internal genitalia of male and female specimens and particularly the complex pattern of spines on the male penial sac made visible by eversion of this structure, and details of sculpturing of the egg chorion.
The following species of Phanoperla are recognized: amorpha sp.n.; anomala (Banks, 1939); bakeri (Banks, 1924); ceylonica Kawai, 1975; comuta sp.n.; flaveola (Klapálek, 1921), comb.n. ( =clarissa (Banks, 1913), syn.n.), (=Neoperla hageni Banks, 1920, syn.n.), (= N.consimilis Banks, 1924, syn.n.); guttata sp.n.; himalayana Zwick, 1977 (= siwalika Harper, 1977); limosa (Hagen, 1858); maculata sp.n.; maindroni (Navas, 1926), comb.n.; malayana sp.n.; minutissima (Enderlein, 1909); nana sp.n.; nervosa Banks, 1939; nuwara Kawai, 1975; omega sp.n.; pallipennis (Banks, 1938); parva sp.n.; pumilio (Klapálek, 1921), comb.n.; peniculus Kawai, 1969a; schmidi sp.n.; sertispina Jewett, 1975; srilanka sp.n.; sumatrae sp.n.; testacea (Hagen, 1858); wedda sp.n. P.claggi (Banks, 1938) is a nomen nudum.
All species are (re-)described and figured; all primary types have been examined. A key to species is provided. Most species can be assigned to one of seven species groups which are defined.
Phanoperla is endemic to the Oriental Region. Species groups are generally widespread, but individual species are in most cases known only from restricted areas.     

Variability In the rate of egg development of the stonefly, Nemoura cinerea (Plecoptera)

SUMMARY. 1. The egg development of the widely distributed European stonefly. Nemoura cinerea Retzius. was investigated in the laboratory. There was a significant relationship between water temperature (T°C) and incubation period (Ydays), expressed by the regression equation: Y = 239 T^−0.85 (r²= 0.85. P<0.001).
2. The number of day-degrees above 0°C required for hatching showed a curvilinear relationship with water temperature, with a minimum requirement around 12°C.
3. Nemoura cinerea shows more variation in the rate of egg development than most other stoneflies. This, in part, explains the wide size range in nymphal populations and the species' extended emergence period. Flexibility in life cycle and asynchrony in egg development enable this species to colonize a wide range of freshwater habitats.     

On the stonefly (Plecoptera) fauna of the transfrontier Selenga River basin

Based on the literature and new data, a list of 48 species belonging to 26 genera of 8 stonefly families from the Selenga River basin is presented. Isoperla kozlovi Zhiltzova, I. lunigera Klapálek, and I. obscura (Zetterstedt) are recorded from the Russian part of the Selenga River basin for the first time; Filchneria mongolica Klapálek is new to the Russian fauna.