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Plastocerus angulosus (Germar, 1844) is one of the only two species of genus Plastocerus Schaum, 1852 within the monogeneric click beetle tribe Plastocerini. It is distributed in the area comprising Greece, Turkey, Syria, Israel, and Lebanon (first record for Lebanon published here). Due to the slightly modified morphology of P. angulosus, this taxon has a convoluted taxonomic history and was earlier classified in various families and even superfamilies. However, recent phylogenies place it in Elateridae: Dendrometrinae. In this study, we review the morphology, intraspecific morphological and genetic variability, sexual dimorphism, systematics, bibliography, and distribution of P. angulosus. Our results show rather low morphological and relatively high genetic variability in this species. Females, which are larger than males and differ mainly in the antennae and abdominal ventrites, are not so rare as previously thought. Further field research should focus on the discovery of immature stages to describe their morphology and understand their biology and ecology.

Crop production is affected by drought duration and severity, which become more frequent with climate change. Several studies reported the positive effect of Silicate (Si) fertilization on soil and plant water balance. However, the relation between soil type and the impact of Silicate (Si) fertilization on plant performance, as well as the underlying mechanism for water stress tolerance, is poorly understood. To investigate the effect of Si fertilization on soil-plant-water relation in different soil types, we set up a Si fertilization experiment in an arable landscape of Northeast Germany (Uckermark region, 53° 23' N, 13° 47' E) using Barley (Hordeum vulgare) at three different sites with different soil types and erosion stages: 1) Haplic luvisol (non-eroded), 2) Haplic Regesol (extremely eroded) and 3) Endogleyic colluvic regosol (deposition). 3 Kg "Aerosil 300" are applied to the soil (~1% ASi in topsoil) compared to control plots (no fertilization), with four replicates per treatment at each site. A campaign of 2-3 consecutive days was conducted throughout the experiment period (from April to July 2022) every two weeks. During these campaigns, we measured leaf water potential, gas fluxes (CO2 and Evapotranspiration), NDVI (normalized difference vegetation index) and biomass sampling. Soil water content and temperature were continuously monitored by soil sensors planted in situ at each plot. We harvested Barley at the end of the growing season and measured each plot's plant biomass and seed production. Our data showed that Si fertilization significantly increased the soil water content in the different soil erosion stages by 2-3%. At the plant's early growth stage, the increase in the soil water content related to Si fertilization significantly affected drought mitigation, balancing leaf water potential decrease during drought. Moreover, while plant development was not generally affected by Si fertilization, germination was delayed in non-fertilized plots. However, the vegetation period in 2022 was rather wet and a drought occurred only during the early phenological development of the plant, and no significant effects of Si fertilization on plant performance were visible (leaf water potential, net ecosystem exchange, evapotranspiration, NDVI and yield) after the early stages. Thus, no lasting effect of Si fertilization on drought mitigation on Barley could be detected, as Barley recovered quickly from drought during the early vegetation stage, irrespective of Si fertilization. All things considered, Si fertilization as an approach to enhance plant tolerance during drought is more complex than previously expected. Our results suggest that the timing and duration of drought, as well as soil type, are important factors to consider.Keywords: Si fertilization, Drought, Soil erosion, Water Stress, Soil water content, Leaf water potential, NDVI, Gas fluxes, Plant performance.

Context Plant populations in agricultural landscapes are mostly fragmented and their functional connectivity often depends on seed and pollen dispersal by animals. However, little is known about how the interactions of seed and pollen dispersers with the agricultural matrix translate into gene flow among plant populations. Objectives We aimed to identify effects of the landscape structure on the genetic diversity within, and the genetic differentiation among, spatially isolated populations of three temperate forest herbs. We asked, whether different arable crops have different effects, and whether the orientation of linear landscape elements relative to the gene dispersal direction matters. Methods We analysed the species’ population genetic structures in seven agricultural landscapes across temperate Europe using microsatellite markers. These were modelled as a function of landscape composition and configuration, which we quantified in buffer zones around, and in rectangular landscape strips between, plant populations. Results Landscape effects were diverse and often contrasting between species, reflecting their association with different pollen- or seed dispersal vectors. Differentiating crop types rather than lumping them together yielded higher proportions of explained variation. Some linear landscape elements had both a channelling and hampering effect on gene flow, depending on their orientation. Conclusions Landscape structure is a more important determinant of the species’ population genetic structure than habitat loss and fragmentation per se. Landscape planning with the aim to enhance the functional connectivity among spatially isolated plant populations should consider that even species of the same ecological guild might show distinct responses to the landscape structure.

Context Evidence for effects of habitat loss and fragmentation on the viability of temperate forest herb populations in agricultural landscapes is so far based on population genetic studies of single species in single landscapes. However, forest herbs differ in their life histories, and landscapes have different environments, structures and histories, making generalizations difficult. Objectives We compare the response of three slow-colonizing forest herbs to habitat loss and fragmentation and set this in relation to differences in life-history traits, in particular their mating system and associated pollinators. Methods We analysed the herbs’ landscape-scale population genetic structure based on microsatellite markers from replicate forest fragments across seven European agricultural landscapes. Results All species responded to reductions in population size with a decrease in allelic richness and an increase in genetic differentiation among populations. Genetic differentiation also increased with enhanced spatial isolation. In addition, each species showed unique responses. Heterozygosity in the self-compatible Oxalis acetosella was reduced in smaller populations. The genetic diversity of Anemone nemorosa, whose main pollinators are less mobile, decreased with increasing spatial isolation, but not that of the bumblebee-pollinated Polygonatum multiflorum. Conclusions Our study indicates that habitat loss and fragmentation compromise the long-term viability of slow-colonizing forest herbs despite their ability to persist for many decades by clonal propagation. The distinct responses of the three species studied within the same landscapes confirm the need of multi-species approaches. The mobility of associated pollinators should be considered an important determinant of forest herbs’ sensitivity to habitat loss and fragmentation.

The authors present the results of morphological, anatomic and genetic studies on Hypena helenae Berio, 1972. They revealed the synonymy of this species with Hypena commixtalis Zeller, 1852 (Hypena helenae Berio, 1972 syn. nov.).

Genetic diversity is of crucial importance for population fitness and the potential of populations to adapt to environmental change. Population-level genetic diversity is expected to be reduced in specialists having small and isolated populations. We analysed genetic structure and diversity in (pre-)Alpine populations of four Copper butterfly species, differing in the degree of habitat specialisation. Despite substantial variation among genetic markers and molecular indices, genetic diversity and thus evolutionary potential tended to be highest in Lycaena hippothoe and lowest in L. tityrus. Microsatellite analyses revealed that genetic diversity tended to be higher in L. hippothoe and L. virgaureae than in L. helle and L. tityrus. MDH sequencing indicated only few differences among species, while GAPDH sequencing showed higher genetic diversity in L. hippothoe and L. tityrus than in L. virgaureae and L. helle. In L. helle, populations showed strong genetic differentiation but without clear spatial structure, while the other species showed genetic differentiation among clusters according to cardinal directions. These results do not match our a priori predictions of a high genetic diversity in L. tityrus and a particularly low one in the glacial relict L. helle. We thus document that evolutionary potential is not necessarily constrained in relict taxa, and that predicting genetic diversity is challenging. Conservation status does not necessarily reflect genetic diversity at the species level. It thus seems to be more meaningful to focus on specific populations of concern to assess genetic vulnerability to environmental change.

On arable land, C dynamics and storage are significantly influenced by tillage and N fertilization. Therefore, new practices such as the combination of topsoil dilution (e.g., through fractional deep tillage) and organic N fertilization may not only ameliorate soil's physical and chemical properties and promote root development but might also enhance soil organic carbon (ΔSOC) stocks. However, the impact of these practices depends on site-specific conditions as agricultural landscapes are often characterized by distinct small-scale soil heterogeneities. To upscale and evaluate the effects or benefits of these new farming practices, accurate and precise information on the complex spatio-temporal C flux pattern and their drivers are thus needed.To investigate the impact of topsoil dilution and organic N fertilization on SOC storage, we performed a study in the strongly erosion affected arable landscape of NE Germany (Uckermark region, 53° 23' N, 13° 47' E; ~50-60 m a.s.l). The study area consisted of 36 measurement plots, of which each 12 covered one out of three erosion induced soil types; Calcic Luvisol (non-eroded), Nudiargic Luvisol (strongly eroded) and Calcaric Regosol (extremely eroded). During July 2020, a two factorial experimental design was established (topsoil dilution vs no topsoil dilution and mineral N fertilization vs organic N fertilization) through implementing topsoil dilution and organic N fertilization on three replicates of each of the three measured soil types. Topsoil dilution was achieved by removing the upper 6 cm of the topsoil layer adding/mixing equivalent weight of subsoil into it.Subsequently, relevant C fluxes, especially the CO2 exchange, were measured using a new robotic chamber system. C in plant biomass was measured by weekly biomass sampling on a nearby reference site and related to plot measurements of CO2 through NDVI (normalized difference vegetation index) and RVI (ratio vegetation index) measurements. Here, we present our first results on the effect of soil type, topsoil dilution, and N-fertilization form on CO2 and C exchange of winter rye. Our results show that there are not only differences between the distinct soil types but also differences between the non-diluted and diluted topsoil treatments. The latter show lower cumulated ecosystem respiration and gross primary productivity, as well as a lower RVI/NDVI and above-ground biomass production, compared to the non-diluted soil. No substantial difference, however, was detected in the case of net ecosystem exchange. As a result, net ecosystem carbon balance was lower for diluted topsoil compared to the non diluted treatments.

Improved agricultural practices are considered as one of the potential solutions for mitigating global climate change. However, agricultural used landscapes are complex and their function as source and sink of greenhouse gases like CO2, CH4, and N2O might differ substantially in time and space. Hence, accurate and precise information on the complex spatio-temporal gas flux pattern is needed to evaluate the effects/benefits of new agricultural practices aiming towards increasing soil organic carbon. Automatic chamber measurements are increasingly used in agricultural systems to determine emissions of greenhouse gases as well as the net ecosystem C balance (NECB). While the eddy covariance (EC) technique remains to be the most common method at field scale, automated chamber measurements might close a gap, by detecting small-scale spatial emission patterns, while still compromising a sufficient temporal resolution. Infrared gas analysers (IRGAs) have been available for decades and helped to facilitate CO2 measurements substantially. In addition, further technical progress resulted in the development of multigas analysers, which are able to measure not only CO2, but also CH4, N2O, as well as their isotopes. However, most of these analysers are rather cost-intensive and many of them are primary designed for use in the laboratory.Here, we compare CO2 fluxes and derived emission estimates, obtained using a widely applied IRGA (LI-850 CO2/H2O, Licor, Germany) with results of a new, medium cost, CO2, CH4, and N2O gas analyser (ProCeas GENERAL, AP2E, France). Two of both sensors were mounted on a novel robotic chamber system (“CarboCrane”), which was installed in 2019 at an undulating summit position of the hummocky ground moraine landscape of NE Germany. The system is comprised of a gantry crane mounted on two tracks (110 m) transporting the sensors and two transparent closed chambers. Measurements of the net CO2 exchange were performed by moving the system along the tracks with each chamber along one half of the covered area. Altogether, 36 measurement plots have been established. On each of these plots, an area for net CO2 exchange measurement has been set up by inserting round iron frames (diameter=1.59 m) 5 cm deep into the soil on which the transparent chambers were deployed for measurements. CO2 fluxes were determined by measuring the development of chamber headspace CO2 concentrations (4 sec frequency; measurements of both sensors in parallel) over chamber deployment time (7 min; see 2.5) in a flow-through non-steady-state (FT-NSS) mode (Livingston and Hutchinson, 1995). CO2 fluxes and emission estimates were derived for all four sensors for a test period of three month (April – June 2021) at six plots, covered with winter rye situated at a mineral fertilized, non-eroded Calcic Luvisol. To guarantee an enhanced variability in measured CO2 fluxes, the six measured plots divide into topsoil diluted and non-diluted treatments. Our results show in general a great consistency between the results delivered by both sensors and support the assumption of a rather small error fraction of the sensor type for both, the calculated CO2 flux and the emission estimates based on it.

Beetles are the most species-rich animal radiation and are among the historically most intensively studied insect groups. Consequently, the vast majority of their higher-level taxa had already been described about a century ago. In the 21st century, thus far, only three beetle families have been described de novo based on newly collected material. Here, we report the discovery of a completely new lineage of soft-bodied neotenic beetles from the Brazilian Atlantic rainforest, which is one of the most diverse and also most endangered biomes on the planet. We identified three species in two genera, which differ in morphology of all life stages and exhibit different degrees of neoteny in females. We provide a formal description of this lineage for which we propose the new family Jurasaidae. Molecular phylogeny recovered Jurasaidae within the basal grade in Elateroidea, sister to the well-sclerotized rare click beetles, Cerophytidae. This placement is supported by several larval characters including the modified mouthparts. The discovery of a new beetle family, which is due to the limited dispersal capability and cryptic lifestyle of its wingless females bound to long-term stable habitats, highlights the importance of the Brazilian Atlantic rainforest as a top priority area for nature conservation.

Cydistinae are a rare monogeneric beetle lineage from Asia with a convoluted history of classification, historically placed in various groups within the series Elateriformia. However, their position has never been rigorously tested. To resolve this long-standing puzzle, we are the first to present sequences of two nuclear and two mitochondrial markers for four species of Cydistinae to determine their phylogenetic position. We included these sequences in two rounds of analyses: one including a broad Elateriformia dataset to test placement at the superfamily/family level, and a second, including a richer, targeted sampling of presumed close relatives. Our results strongly support Cydistinae as sister to Phengodidae in a clade with Rhagophthalmidae. Based on our molecular phylogenetic results and examination of morphological characters, we hereby transfer the formerly unplaced Cydistinae into Phengodidae and provide diagnoses for the newly circumscribed Phengodidae, Cydistinae and Cydistus. Since both Phengodidae and Rhagophthalmidae have bioluminescent larvae and strongly neotenic females, similar features can be hypothesized for Cydistinae. Additionally, Cydistus minor is transferred to the new genus Microcydistus.

Keys to adults and larvae of the genera of West Palaearctic nematine sawflies are presented. Species of some of the smaller genera are keyed, and their taxonomy, distribution, and host plants reviewed, with a geographic focus on north-western Europe, particularly Sweden.DinematusLacourt, 2006 is a new junior subjective synonym ofPristiphoraLatreille, 1810, resulting in the new combinationPristiphora krausi(Lacourt, 2006) for the type species ofDinematus.Hemichroa monticolaErmolenko, 1960 is a new junior subjective synonym ofHemichroa australis(Serville, 1823). Lectotypes are designated forTenthredo opacaFabricius, 1775, Mesoneura opaca var. nigerrima Enslin, 1914, Mesoneura opaca var. obscuriventris Enslin, 1914,Nematus hypogastricusHartig, 1837,Nematus alnivorusHartig, 1840,Leptopus rufipesFörster, 1854,Nematus protensusFörster, 1854, and Platycampus luridiventris var. pleuritica Enslin, 1915. A phylogenetic analysis based on four genes (mitochondrial COI and nuclear NaK, POL2, and TPI) supports the current generic classification.

Beetles are the most species-rich animal radiation and are among the historically most intensively studied insect groups. Consequently, the vast majority of their higher-level taxa had already been described about a century ago. In the 21st century, thus far, only three beetle families have been described de novo based on newly collected material. Here, we report the discovery of a completely new lineage of soft-bodied neotenic beetles from the Brazilian Atlantic rainforest, which is one of the most diverse and also most endangered biomes on the planet. We identified three species in two genera, which differ in morphology of all life stages and exhibit different degrees of neoteny in females. We provide a formal description of this lineage for which we propose the new family Jurasaidae. Molecular phylogeny recovered Jurasaidae within the basal grade in Elateroidea, sister to the well-sclerotized rare click beetles, Cerophytidae. This placement is supported by several larval characters including the modified mouthparts. The discovery of a new beetle family, which is due to the limited dispersal capability and cryptic lifestyle of its wingless females bound to long-term stable habitats, highlights the importance of the Brazilian Atlantic rainforest as a top priority area for nature conservation.

Many riparian (Tugai) forests growing along rivers in arid and hyper-arid regions of Central Asia are dominated by the Euphrates poplar (Populus euphratica). Besides generative reproduction, which is only possible upon flooding events and at a distance to the groundwater of less than 2 m, this phreatophytic tree species also reproduces vegetatively by forming clones that can cover land surface areas of several hectares. Along a gradient of groundwater distances, we investigated whether the fraction of clones in P. euphratica stands (1) increases with increasing distance to the water table; (2) is higher if supplied with water via river cut-offs; and (3) approaches 100% at a short distance to the groundwater, but at high salt concentrations in the upper soil layers, which would prevent germination and establishment of seedlings. AFLP (Amplified Fragment Length Polymorphism) analyses were conducted on leaf samples taken from mature P. euphratica trees growing at the fringes of the Taklimakan Desert in stands with different distances (2–12 m) to the groundwater at two plots at the middle and the lower reaches of the Tarim River and in a stand close to Ebinur Lake, Xinjiang, China. Genetic diversity was large among plots, but considerably smaller within plots. We found the highest genetic diversity (caused by regeneration from seeds) at plots that have a short distance to the groundwater or are supplied with additional water. There was no significant relationship between groundwater distance and clonal fraction. All investigated trees at the saline Ebinur Lake site belonged to one single clone. Our results demonstrate that the genetic pattern of this widespread species is not easily predictable even over small distances as it is a result of a complex interplay of stand history and dispersal of propagules (pollen, seeds, and vegetative diaspores) by wind and water. In conservation and restoration schemes, P. euphratica stands with a high genetic diversity and stands that grow at short distances to the water table and are regularly subjected to flooding (which favors generative over clonal reproduction) should be prioritized.

The neotenic elaterid genus Drilus Olivier, 1790 (Elateridae: Agrypninae: Drilini) contains more than 40 species distributed primarily in the Mediterranean region. The precise number of species remains ambiguous due to the insufficient taxonomic investigation in several areas. Herein, we study Drilus diversity in Cyprus and we recognize the following species: D. humeralis Pic, 1931, D. rectus Schaufuss, 1867, and D. rufipes ( Baudi di Selve, 1871 ). Drilus testaceipes Pic, 1933 syn. nov. is synonymized under D. rectus, and D. truquii ( Baudi di Selve, 1871 ) syn. nov. is synonymized under D. rufipes. As consequence of these taxonomic changes, Drilus humeralis remains the only species endemic to Cyprus. Females of D. rectus and D. rufipes are described and figured for the first time, as well as the larvae of D. rufipes. A key to the males of Drilus species in Cyprus is given. Drilus rectus is recorded in Syria for the first time and D. rufipes is recorded in Rhodes. To examine the phylogenetic relationships of Drilus in Cyprus, we produced the cox1-3ʹ, cox1-5′ and rrnL mtDNA sequences for available specimens of D. rectus and D. rufipes and merged them with data from GenBank. Results of the maximum likelihood analysis and Bayesian inference suggest a close relationship between D. rectus and Drilus cf. mertliki from Turkey, and a sister position of D. rufipes to a clade formed by the species from the Ionian Islands, Peloponnese, and Crete. Using the multigene barcoding approach including the above mentioned mitochondrial markers and the nuclear arginine kinase, we have been able to associate the different larval stages from Cyprus with the adult males of D. rufipes, and confirm the intraspecific variability in male body size and antennal coloration. Moreover, the low genetic difference in the cox1-3′ fragments indicates the close relationship between D. rectus populations from Cyprus and southern Turkey.

Tianmuthredo Wei, 1997 is recorded for the first time in the Palearctic realm, and Tianmuthredo davei Taeger and Kramp, new species is described from the Primorskij Kraj in the Russian Far East. The placement of the genus within Tenthredininae is discussed.

Xyela davidsmithi Blank and Kramp sp. n. is described from the Sierra Nevada of California. Morphological characters place it into the X. alpigena group. Unlike other members of this group, whose larvae feed inside staminate cones of species of Pinus (Strobus), larvae of X. davidsmithi identified by COI barcoding have been extracted from staminate cones of P. (Pinus) jeffreyi. Field observations on imagines of X. brunneiceps Rohwer, 1913 indicate that this species is oligophagous on P. (S.) flexilis in Colorado and on P. (S.) strobiformis in Arizona. A key to the Nearctic species of the X. alpigena group is provided.

North-Western Palaearctic species ofPristiphoraLatreille, 1810 are revised. Altogether, 90 species are treated, two of which are described as new:P.caraganaeVikberg & Prous,sp. n.from Finland andP.dedearaListon & Prous,sp. n.from Germany. Host plant ofP.caraganaeisCaraganaarborescensLam.Pristiphoradasiphorae(Zinovjev, 1993) (previously known from East Palaearctic) andP.cadmaWong & Ross, 1960 (previously known from North America) are recorded for the first time from Europe.NematusnigricansEversmann, 1847 [=Pristiphoranigricans(Eversmann, 1847),comb. n.],N.breviusculusEversmann, 1847 [=Euuramelanocephalus(Hartig, 1837)], andN.caudalisEversmann, 1847 [=E.caudalis(Eversmann, 1847),comb. n.] are removed from synonymy withP.pallidiventris(Fallén, 1808),N.paralellusHartig, 1840 [=P.paralella(Hartig, 1840),comb. n.] is removed from synonymy withP.bufo(Brischke, 1883), andP.mesatlanticaLacourt, 1976 is removed from synonymy withP.insularisRohwer, 1910. The following 29 new synonymies are proposed:P.nigropuncticepsHaris, 2002,syn. n.withP.albitibia(Costa, 1859);LygaeonematuskarvoneniLindqvist, 1952,syn. n.withP.alpestris(Konow, 1903); P. (P.) anivskiensis Haris, 2006,syn. n.withP.appendiculata(Hartig, 1837);NematuscanaliculatusHartig, 1840,syn. nwithP.carinata(Hartig, 1837);P.nigrogroenblomiHaris, 2002,syn. n.withP.cinctaNewman, 1837;TenthredoflavipesZetterstedt, 1838,syn. n.,NematuscongenerW.F. Kirby, 1882,syn. n., andP.thomsoniLindqvist, 1953,syn. n.withP.dochmocera(Thomson, 1871);P.atrataLindqvist, 1975,syn. n.withP.friesei(Konow, 1904);P.gelidaWong, 1968,syn. n.withP.frigida(Boheman, 1865);PachynematusnigricorpusTakagi, 1931,syn. n.withP.laricis(Hartig, 1837); Nematus (Pikonema) piceae Zhelochovtsev in Zhelochovtsev and Zinovjev, 1988,syn. n.and P. (P.) hoverlaensis Haris, 2001,syn. n.withP.leucopodia(Hartig, 1837);MesoneuraarcticaLindqvist, 1959,syn. n.,PachynematusincisusLindqvist, 1970,syn. n.,PachynematusintermediusVerzhutskii, 1974,syn. n., andP.mongololaricisHaris, 2003,syn. n.withP.malaisei(Lindqvist, 1952);NematusanderschiZaddach, 1876,syn. n.,P.inocreataKonow, 1902,syn. n., andP.discolorLindqvist, 1975,syn. n.withP.nigricans(Eversmann, 1847);LygaeonematustenuicornisLindqvist, 1955,syn. n.withP.paralella(Hartig, 1840);LygaeonematusconcolorLindqvist, 1952,syn. n.withP.pseudocoactula(Lindqvist, 1952);P.flavipictaLindqvist, 1975,syn. n.,P.flavopleuraHaris, 2002,syn. n.,P.mongoloexiguaHaris, 2002,syn. n., andP.mongolofaustaHaris, 2003,syn. n.withP.punctifrons(Thomson, 1871);P.listoniLacourt, 1998,syn. n.withP.sootryeniLindqvist, 1955;P.gaunitziLindqvist, 1968,syn. n.withP.testacea(Jurine, 1807); andNematusbreviusculusEversmann, 1847,syn. n.withEuuramelanocephalus(Hartig, 1837). The valid name of Pachynematus (Pikonema) carpathiensis Haris, 2001 isNematinuscarpathiensis(Haris, 2001)comb. n.Lectotypes are designated for 43 taxa. An illustrated electronic key made with Lucid and a traditional dichotomous key are provided to facilitate identification of the species. Species belonging to thecarinata(previouslyLygaeotus),micronematica(previouslyLygaeophora), andrufipes(also known asthalictrioraquilegiae) groups are not keyed to the species level, because additional research is needed to delimit the species more reliably in these groups. Phylogeny ofPristiphorais reconstructed based on one mitochondrial (COI) and two nuclear (NaK and TPI) genes. Remarkably, around 50–60% (depending on the exclusion or inclusion of thecarinata,micronematica, andrufipesgroups) of the species cannot be reliably identified based on COI barcodes. Limited data from nuclear genes indicate a better identification potential (about 20% remain problematic).

The ‘Symphyta’ is a paraphyletic assemblage at the base of the order Hymenoptera, comprising 14 families and about 8750 species. All have phytophagous larvae, except for the Orussidae, which are parasitoids. This study presents and evaluates the results of DNA barcoding of approximately 5360 specimens of ‘Symphyta’, mainly adults, and 4362 sequences covering 1037 species were deemed of suitable quality for inclusion in the analysis. All extant families are represented, except for the Anaxyelidae. The majority of species and specimens are from Europe, but approximately 38% of the species and 13% of the specimens are of non‐European origin. The utility of barcoding for species identification and taxonomy of ‘Symphyta’ is discussed on the basis of examples from each of the included families. A significant level of cryptic species diversity was apparent in many groups. Other attractive applications include the identification of immature stages without the need to rear them, community analyses based on metabarcoding of bulk samples and association of the sexes of adults.

The boreo-montane wetland butterfly species Colias palaeno has a European distribution from the Alps to northern Fennoscandia. Within its European range, the species’ populations have shrunk dramatically in recent historical times. Therefore, detailed baseline knowledge of the genetic makeup of the species is pivotal in planning potential conservation strategies. We collected 523 individuals from 21 populations across the entire European range and analyzed nuclear (20 allozyme loci) and mitochondrial (600 bp of the cytochrome c oxidase subunit I gene) genetic markers. The markers revealed contrasting levels of genetic diversity and divergence: higher in allozymes and lower in mitochondrial sequences. Five main groups were identified by allozymes: Alps, two Czech groups, Baltic countries, Fennoscandia, and Poland. The haplotype mitochondrial network indicates a recent range expansion. The most parsimonious interpretation for our results is the existence of a continuous Wurm glacial distribution in Central Europe, with secondary disjunction during the Last Glacial Maximum into a south-western and a north-eastern fragment and subsequent moderate differentiation. Both groups present signs of postglacial intermixing in the Czech Republic. However, even a complete extinction in this region would not considerably affect the species’ genetic basis, as long as the source populations in the Alps and in northern Europe, comprising the most relevant evolutionary units for conservation, are surviving.

Megaxyela gigantea Mocs��ry, 1909 Figs 5A���H, 11C, 12E, 13C, G���H Megaxyela gigantea Mocs��ry, 1909: 39 (♀, type locality: Russia, Khabarovskiy Kray, Kazakevichevo). Megaxyela mikado Sato, 1930: 4���5, figs a���b (♀, type locality: South Korea, Suwon, Kazan). Megaxyela mikado ��� Takeuchi 1937: 25 (synonymy with M. gigantea). Megaxyela gigantea ��� Shinohara 1992: 785���789 (description, distribution, hosts, references), 794���795 (key), figs 1A���B, 2A���B, 2E, 3A���C, 4A, 4F���G. Type material Lectotype (here designated) RUSSIA: ♀, lectotype of Megaxyela gigantea, ��� Ussuri Kasakewitsh [= Khabarovskiy Kray, Kazakevichevo, 48.27�� N, 134.77�� E] 1907 Korb ���; ��� Megaxyela gigantea det. Mocs��ry���; [rectangular red label without inscription]; ��� Megaxyela gigantea Mocs. ���; [red:] ��� Lectotype Megaxyela gigantea Mocs��ry, 1909 Det. A. Shinohara 1997 ���, in perfect condition (HNHM). The lectotype corresponds well with Shinohara���s (1992) characterization of the species. Paralectotypes RUSSIA: 1 ♀, same collecting data as lectotype (HNHM), Mocs��ry (1909) described M. gigantea from an unknown number of females. SOUTH KOREA: 1 ♀, holotype of Megaxyela mikado, ��� Suigen [= Suwon, 37.28�� N, 127.02�� E], Chosen [= Korea], Apr. 24., 1927, coll. K. Sato ���; ��� 270���, ���Type of Megaxyela mikado Sato���, ��� Megaxyela gigantea Mocs. det. K. Sato, XI���1957 ���, ��� DEI-GISHym 22350 ���, left wings missing (NSMT). The holotype of M. mikado is a comparatively dark female of M. gigantea with a large dark brown spot on ocellar area and vertex. Terga 2���4 bear white stripes laterally, which are of equal width (see figure in Sato 1930; Shinohara & Smith 1979: fig. 13, in synonymy with M. gigantea). The synonymy first proposed by Takeuchi (1937) is here confirmed. Additional material examined or reported in literature NORTH KOREA: 1 ♀, P���y��ngan-pukto [���Pyongah Prov. N���], Mt. Myohyang, 40.02�� N, 126.33�� E, 23 May 1985, Vojnits & Zombori leg. (HNHM). RUSSIA: 1 ♀, Primorskiy Kray, Voroshilovskiy rayon, southern slopes of Sikhote Alin, on river Suputinka [���Fluss Suputinka, S. Abh��nge von Sichota-Alin���], 43.77�� N, 131.90�� E, 20 Jun. 1937, T. Samoylov leg. (NHRS). SOUTH KOREA: 4 ♀♀, 51 ♂♂ (including DEI-GISHym 708), Chollanam-do, Mt. Nogodan, Mts. Chirisan, 35.30�� N, 127.53�� E, 1220 m alt., 4���5 Jun. 1996, A. Shinohara leg. (NSMT); 1 ♀, Kangwondo, Mirugam (Pugdaesa), Mt. Odaesan, 37.80�� N, 128.57�� E, 1300 m alt., 30 May 1991, A. Shinohara leg. (NSMT) (Shinohara 1992); 1 ♀, same locality, 27 May 1998, A. Shinohara leg. (NSMT); 1 ♂ (specimen DEI-GISHym 18507 used for barcoding), same locality, collected from Juglans ? mandshurica, 1 Jun. 2002, A. Shinohara leg. (NSMT); 5 ♀♀, 5 ♂♂, Kyonggi-do, Suwon [��� Suigen ���], 37.283�� N, 127.017�� E, 12 May 1932, S. Fujii leg. (CIS, NSMT, USNM) (Shinohara 1992); 2 ♀♀, 4 ♂♂, same locality, 14 May 1932, S. Fujii leg. (NSMT, USNM) (Shinohara 1992); 5 ♀♀, Seoul [��� Keijo ���], 37.57�� N, 127.00�� E, 11 May 1935, K. Sato leg. (NSMT) (Shinohara 1992); 2 ♀♀, Seoul, Ch���ongnyangni [��� Seiryori ���], 37.58�� N, 127.05�� E, 6 May 1934, H. Doi leg. (UOP) (Shinohara 1992). Description See Shinohara (1992) and Figs 5A���B, E, 11C (ovipositor sheath). Female Ovipositor about 2.7 mm long, weakly curved ventrally, valvula 1 and valvula 2 bulbous distal of middle, without ctenidia (Fig. 12E). Pulvilli absent on article 1 of metatarsus, present on article 4 and 50 ��m long, presence ambiguous for articles 2���3 (Fig. 5F���G). Male See Figs 5C���D, H, 13G���H. Valviceps of penis valve distally parabolically rounded, basal quarter of upper side expanded to a round lobe coiled laterally, medial 0.3���0.8 of upper edge with one or two shallow elevations and numerous small teeth. Distal 0.7 of valviceps with long setae, most dense in medial lower portion (Fig. 13C). Host plant Saito (1941), Ko (1969) and Xiao (2006) reported Juglans ailanthifolia Carri��re (cited as J. sieboldiana Maxim.), J. mandshurica Maxim., J. regia L. (including the synonym J. sinensis [C. DC.] Dode), Pterocarya rhoifolia Sieb. & Zucc., and P. stenoptera C. DC. All recorded hostplant associations of M. gigantea need confirmation, because they can also refer to the new species M. pulchra sp. nov. In South Korea, A. Shinohara captured a male of M. gigantea together with 3 ♀♀, 2 ♂♂ M. pulchra sp. nov. from the same J. ? mandshurica tree at end of May and beginning of June 2002. Remarks Megaxyela gigantea is similar to species with the terga 2���3 or 2���4 laterally extensively white (dorsally black in M. euchroma sp. nov. and in the Nearctic species except for M. major). It is unique among the East Asian species in the predominantly red brown head (predominantly black in M. parki, pale yellow color and less extensive in M. togashii) bearing a large infuscate spot on face and vertex (completely red brown in M. fulvago sp. nov.), which does not extend to the antennal toruli ventrally (spot extending to toruli in M. pulchra sp. nov.). As in M. fulvago sp. nov., the surface of the face is minutely areolate and bears scattered, shallow, 50 ��m large pits (face rugose in other East Asian species at least laterally). The dorsal side of terga 2���4 is white laterally for 0.15���0.20 �� the tergal width in M. gigantea and M. fulvago sp. nov., whereas the white marks are generally narrower in M. parki or narrower at least on tergum 4 in M. pulchra sp. nov. and M. togashii. The single specimen studied genetically is placed within the group, which otherwise comprises M. fulvago sp. nov., but we interpret this placement as an artefact caused by the very short sequence length of M. gigantea. Morphological results indicate two separate species., Published as part of Stephan M. Blank, Katja Kramp, David R. Smith, Yuri N. Sundukov, Meicai Wei & Akihiko Shinohara, 2017, Big and beautiful: the Megaxyela species (Hymenoptera, Xyelidae) of East Asia and North America, pp. 1-46 in European Journal of Taxonomy 348 on pages 16-19, DOI: 10.5852/ejt.2017.348, http://zenodo.org/record/1042663, {"references":["Mocsary A. 1909. Chalastogastra nova in collectione musei nationalis Hungarici. Annales historiconaturales Musei Nationalis Hungarici 7: 1 - 39.","SatoK. 1930. Anewspeciesofthegenus Megaxyela AshmeadfromKorea (Hymenoptera; Tenthredinidae). Journal of Chosen Natural History Society 3: 4 - 5.","Takeuchi K. 1937. [Megaxyela gigantea Mocsary found in Honshu, Japan.] (In Japanese.) Akitu 1: 25.","Shinohara A. 1992. The sawfly genus Megaxyela (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology 60: 783 - 796.","Shinohara A. & Smith D. R. 1979. The Sato types of sawflies (Hymenoptera: Symphyta). Bulletin of the National Science Museum, Series A, Zoology 5: 281 - 288.","Saito K. 1941. [A dendro-entomological contribution.] Bulletin of the Agricultural and Forestry College Suigen (Suwon) (6): 1 - 235. [In Japanese.]","Ko J. - H. 1969. A List of Forest Insect Pests in Korea. Forest Research Institute, Seoul.","Xiao G. - R. 2006. Suborder Chalastogastra (Symphyta). In: Hua L. - Z. (ed.) List of Chinese Insects. Vol. IV: 187 - 200 + 331 - 335. Sun Yat-sen University Press, Guangzhou."]}

Megaxyela fulvago Blank, Shinohara & Wei sp. nov. urn:lsid:zoobank.org:act:DF04584F-B33D-42A7-836D-E758A66F4081 Figs 4A–G, 11B, 12B–C, 13B Megaxyela gigantea – Takeuchi 1940: 484 (misidentification). ― Maa 1949: 30 (misidentification). Megaxyela sp. 2 – Shinohara et al. 2017: fig. 15 (phylogenetic analysis). — Blank et al. 2017: 115. Diagnosis The red brown head, prothorax and mesothorax are unique to this species. Etymology The Latin noun fulvago indicates the predominantly pale brown color of this species. Type material Holotype CHINA: ♂: “[CHINA: Hunan] Mt. Yunshan 1250 m [alt.] 26°39′ N, 110°37′ E nr. Wugang 12.IV.2012 [leg.] A. Shinohara ”; “ DEI-GISHym 5236 ”; [red:] “Holotype Megaxyela fulvago det. SMBlank & AShinohara 2015” (CSCS). Paratypes (6 ♀♀, 6 ♂♂) CHINA: 1 ♀, 5 ♂♂, same collecting data as holotype, DEI-GISHym 5235, 5237–5239, 5751–5752 (5237, 5239, 5751–5752 used for barcoding) (SDEI, NSMT); 1 ♀, Jiangsu Sheng, Nanjing, 32.06° N, 118.78° E, 1 May 1985, Jinniang leg. (CSCS); 2 ♀♀, 1 ♂, Jiangsu Sheng [“Prov. Kiangsu”], Zhenjiang [“Chinkiang”], 32.20° N, 119.43° E, 24 Apr.–1 May 1918 (IZCAS) (Takeuchi 1940; Maa 1949; images for re-identification provided by Yang Ganyan); 1 ♀, Zhejiang Province, Kaishan Laodian, Tianmushan, Lin’an, 30.343° N, 119.435° E, 1140 m alt., 10 Apr. 2014, Liu Ting and Yu Xinjie leg., CSCS14012, DEI- GISHym 30882 (specimen used for barcoding) (CSCS); 1 ♀, same locality, 1110 m alt., 14 Apr. 2016, Li Zejian, Liu Mengmeng and Chen Zhiwei leg., CSCS16143, DEI-GISHym 30883 (specimen used for barcoding) (CSCS). Description Female COLOR. Body red brown with black-and-white pattern, black parts partly with bronze tinge (Fig. 4A). Head and thorax red brown (specimen 30882 with diffuse transverse stripe on vertex – artifact?), posterior half of metanotum and ventral half of metepisternum infuscate. Scape red brown, pedicel black, flagellum dark brown. Abdomen in dorsal view black with continuous white bands laterally on terga 2–4, each 0.15–0.20 × as wide as tergal width, tergum 8 with narrower lateral white band, ventral portions of terga 2–4 and 8 completely white, terga 5–6 narrowly and tergum 7 broadly white along ventral margin, terga 9+10 brown with small dark spot dorsolaterally. Sterna white. Valvifer 2 and valvula 3 red brown, membrane between valvifer 2 and valvula 3 pale (Fig. 11B). Fore and mid legs pale red brown. On hind leg, coxa red brown, infuscated laterally in middle and dorsally, trochanter and trochantellus pale red brown, femur brown dorsally and red brown ventrally, tibia and tarsus brown. Wings weakly yellow stained, venation and pterostigma yellow brown. MORPHOLOGY. Body 11.4–13.0 mm, fore wing 14.9–15.4 mm long. POL: OOL: OCL = 1.0: 2.2–2.4: 2.2–2.3. Interantennal distance 1.0–1.1 × as wide as distance between torulus and eye margin, malar space 0.4–0.5 × as wide as interantennal distance. Eye 1.3–1.4 × as long as wide, frons at dorsal edge of antennal toruli 1.7–1.8 × as wide as maximum diameter of eye. Synantennomere 3 4.5–4.9 mm, filament 1.1 mm, with 8–9 articles.Article 3 of maxillary palp 0.5–0.6 mm long. Ovipositor sheath 3.30–3.50 mm long, valvula 3 1.9–2.3 × as long as wide. On hind leg, femur 4.9–5.0 mm, tibia 5.6–6.0 mm, tarsus 4.1–4.3 mm, tarsomere 1 4.5–5.1 × as long as wide, longer distal spur of tibia 0.70 × as long as tarsomere 1, subapical tooth of claw stout and little shorter than apical tooth (Fig. 4B; similar to Shinohara 1992: fig. 4D). Face and vertex with areolate surface microsculpture, dull, interantennal area and frons above antennal toruli with scattered, shallow, 50 µm large pits (similar to Fig. 4G), gena weakly coriarious, shining. Mesoscutum and mesoscutellum homogeneously areolate. Mesepisternum weakly coriarious and shining, with scattered pits. Metatarsomere 1 dorsally and dorsolaterally very sparsely setose, ventrally and ventrolaterally densely setose, setae 150–200 µm long ventrally (Fig. 4B). Pulvilli absent on article 1 of metatarsus, present on articles 3–4, presence ambiguous for article 2, on article 4 pulvillus 50 µm long. Basal and medial sections of valvula 3 of ovipositor sheath convex dorsally, almost straight ventrally, preapical section concave on dorsal and ventral edge, medial to preapical section of dorsal margin with setae up to 200 µm long, ventral margin with setae up to 100 μm long (Fig. 11B). Ovipositor about 2.4 mm long, weakly curved ventrally, valvula 1 and valvula 2 with bulbous areas in middle, without ctenidia (Fig. 12B–C). Valvula 1 in distal 0.7 with very narrowly spaced, subparallel, straight, vertical annuli, ventral margin in distal 0.3 bearing small serrulae. Valvula 2 in distal 0.6 with narrowly spaced annuli, distal 0.3 of dorsal edge of valvula 1 with small denticles. Male COLOR. Similar to female (Fig. 4E–G). Metepimeron red brown or predominantly infuscate. Ventral portions of terga 5–7 predominantly white. Subgenital plate and genitalia red brown. On hind leg, coxa sometimes completely black laterally, trochanter and trochantellus red brown, femur black dorsally and medially, dark red brown ventrally and laterally, tibia and tarsus black (Fig. 4C–D). MORPHOLOGY. Body 8.8–10.7 mm, fore wing 11.3–12.8 mm long. POL: OOL: OCL = 1.0: 1.6–1.7(– 1.9): 1.7–1.9(–2.0). Interantennal distance 0.9–1.1 × as long as distance between torulus and eye margin, malar space 0.5 × as wide as interantennal distance. Eye 1.3–1.4 × as long as wide, frons at dorsal edge of antennal toruli 1.6–1.8 × as wide as maximum diameter of eye. Synantennomere 3 (3.6–)4.0– 4.6 mm, filament 1.0– 1.1 mm and with 8–9 articles (one male with left filament 0.8 mm long, 6 articles, right filament 1.0 mm long, 8 articles). Article 3 of maxillary palp 0.4–0.5 μm long. On hind leg, femur 3.7–4.2 mm, tibia 4.5–5.2 mm, tarsus (3.8–) 4.1–4.4 mm, tarsomere 1 4.9–5.5 × as long as wide, longer distal spur of tibia 0.60 × as long as tarsomere 1. Claws and microsculpture similar to female (Fig. 4D). Subgenital plate bluntly pointed at apex. Valviceps of penis valve distally parabolically rounded, basal 0.3 of upper side expanded to a round lobe coiled laterally, medial 0.3–0.8 of upper edge shallowly convex, with numerous small teeth. Distal 0.7 of valviceps with long setae, most dense in medial lower portion of valviceps (Fig. 13B). Remarks Megaxyela fulvago sp. nov. is most similar to M. gigantea regarding color and structure. In both species, terga 2–4 are extensively white on the dorsolateral and ventral portion. This pattern is absent in M. euchroma sp. nov. and in most of the Nearctic species. White is present on the dorsal side of only terga 2–3 in M. parki, M. pulchra sp. nov. and M. togashii, as well as in the Nearctic M. major. Megaxyela fulvago sp. nov. and M. gigantea share the dull, minutely areolate and shallowly pitted face. A rugose surface structure is absent from the frons, which has been observed in the other East Asian species. Megaxyela fulvago sp. nov. and M. gigantea are primarily discriminated by color: head, prothorax and mesothorax are completely red brown in M. fulvago sp. nov., but these parts bear an extensive dark brown to black pattern in M. gigantea. The antennal filament of females is longer in M. fulvago sp. nov. (1.1 mm) than in M. gigantea (0.8 mm), and sometimes it is subdivided into fewer articles in M. fulvago sp. nov. (♀: 8–9; ♂: usually 8–9) than in M. gigantea (♀: 9; ♂: 9–10). The relative distance of POL: OOL: OCL differs weakly in males (1.0: 1.6–1.7[–1.9]: 1.7–1.9[–2.0] and 1.0: 1.8–2.0: 2.0–2.2, respectively). The metatarsomere 1 of males is 4.9–5.5 × as long as wide in M. fulvago sp. nov. but 4.6–5.0 × in M. gigantea. The female of M. fulvago sp. nov. falls into the range of variability of M. gigantea regarding these two characters. The COI sequences group the specimens identified as M. fulvago sp. nov. by morphology into a clade which is supported by a bootstrap of 97%. This clade additionally includes a specimen of M. gigantea. Supposedly this placement is an artefact caused by the short sequence length (261 bp) of this specimen. The maximum intraspecific variation within M. fulvago sp. nov. is 0.8%. Megaxyela togashii is placed at a minimum pairwise distance of 7.6% (Fig. 2). The specimens from Mount Yunshan were collected on a sunny day flying around birch trees with lots of catkins shedding pollen. Pollen feeding of adults is well known for Xyela species, which bear distally modified and usually extended maxillary palps (Burdick 1961). A similar behavior is unknown for adult Macroxyelinae. Although M. major has been caught from catkins of willow (Bridwell 1906) and pollen has been identified from the intestine of two fossil Macroxyelinae species (Krasilov & Rasnitsyn 1982), Megaxyela is not specialized in pollen feeding but is facultatively palynivorous, because the labiomaxillar complex exhibits no particular modifications for the uptake of pollen as in Xyelinae. Takeuchi (1940) and Maa (1949) referred to the specimens of M. gigantea collected in “Chinkiang [= Zhenjiang], Prov. Kiangsu [= Jangsu Sheng]” deposited in the collection of the Musée Heude, Shanghai at that time. Major parts of this collection have been incorporated into IZCAS, Beijing (Yang Ganyan, personal communication). The two females and one male collected in “Chinkiang” in 1918 could be identified as M. fulvago sp. nov. with help of photos kindly provided by Yang Ganyan. An additional female from “Chinkian” (3 May 1936, E. Suenson leg.), which we identified as M. gigantea prior to the recognition of M. fulvago sp. nov., is kept at INHS.

Megaxyela euchroma Blank, Shinohara & Wei sp. nov. urn:lsid:zoobank.org:act:63EF6069-E728-4107-8DC2-F02C9081880A Figs 1F, 3A–3K, 11A, 12A, 13A Megaxyela sp. 3 – Shinohara et al. 2017: fig. 15 (phylogenetic analysis). Diagnosis This species is unique upon the black head bearing narrow yellow lines along inner and outer orbits and a linear yellow spot on vertex (Fig. 3A–B, D, F–G), and the presence of ctenidia along the annuli of the ovipositor (Figs 1F, 12A). It is separated from other Eurasian species by the metallic blue shine of dark colored body parts (Fig. 3A–G), the laterally black terga 2–4 (Fig. 3A–E; only the ventral margins of the terga bear a narrow longitudinal white line ventrally, Fig. 3C, E), the almost completely black valvula 3 of the ovipositor sheath (Fig. 11A), the black hypopygium of the male (Fig. 3E), and the large and irregular teeth of the upper edge of the valviceps (Fig. 13A). Etymology The species name is a noun derived from ancient Greek ευ- (eu-, beautiful) and χρωμα (chroma, color). Type material Holotype CHINA: ♀: “ CSCS14007 [..., Chinese characters for Kaishan Laodian, Mt. Tianmushan, Linan, Zhejiang Province], E119°26′05″ N30°20′33″ 1142 m 2014. IV.8 [..., Chinese characters for the collectors Nie Haiyan & Hu Ping] CH3COOC2H5”, [red:] “Holotype Megaxyela euchroma sp.n. det. SMBlank & AShinohara 2015”; “ DEI-DISHym 22554 ” (CSCS). Paratypes CHINA: 2 ♀♀, same site as holotype, 4–5 Apr. 2015, Li Zejian leg. (LSAF); 1 ♀, same site as holotype, 8 Apr. 2014, Liu Ting and Yu Xingjie leg., CSCS14008 / DEI-DISHym 22555 (CSCS); 1 ♀, same site, 9 Apr. 2014, Liu Ting and Yu Xingjie leg., CSCS14010 / DEI-DISHym 22556 (CSCS); 1 ♂, same site, 10 Apr. 2014, Liu Ting and Yu Xingjie leg., CSCS14012 / DEI-DISHym 22557 (CSCS); 1 ♀, 2 ♂♂, same site as holotype, 11 Apr. 2015, Li Zejian leg., CSCS); 3 ♀♀, same site, 16 Apr. 2014, Nie Haiyan and Hu Ping leg., CSCS14026 / DEI-DISHym 22558–22560 (CSCS, SDEI, YUIC); 1 ♀, 1 ♂, same site, 16 Apr. 2014, Liu Ting and Yu Xingjie leg., CSCS14027 / DEI-DISHym 22561–22562, (CSCS); 1 ♂, same site, 1150 m, 4 Apr. 2015, A. Shinohara leg., DEI-DISHym 22515 (SDEI) (specimen used for barcoding); 1 ♀, same site, 8 Apr. 2014, A. Shinohara leg., DEI-DISHym 22512 (NSMT); 1 ♂, same site, 9 Apr. 2014, A. Shinohara leg., DEI-DISHym 22513 (NSMT) (specimen used for barcoding); 1 ♂, 1 ♀, same site, 11 Apr. 2015, A. Shinohara leg., DEI-DISHym 22514, DEI-DISHym 22516 (NSMT); 1 ♀, same site, 15 Apr. 2014, A. Shinohara leg., DEI-DISHym 22511 (NSMT); 3 ♀♀, same site, 16 Apr. 2014, A. Shinohara leg., DEI-DISHym 22509–22510 and 22355 (latter specimen used for barcoding) (NSMT, SDEI). Description Female COLOR. Body black with white, yellow white and red brown pattern, black parts with blue tinge (Fig. 3A– B). Head black with three yellow white lines, one on vertex posterior of eye, one along dorsal half of inner orbits, one along ventral half of outer orbits and along ventral edge of eye; clypeus along anterior margin, labrum, mandibles and most of other mouthparts red brown to white (similar to Fig. 3F–G).Antenna black. Tegula, dorsolateral corner and narrow lateroventral margin of pronotum yellow white. Abdominal terga 1–8 with narrowly white distal margins, lateroventral portions of terga 2–7 with continuous 50–100 µm wide white stripe along ventral margin (Fig. 3C). Abdominal sterna 2–7 broadly white along lateral and distal margins (Fig. 3C). Valvifer 2 of ovipositor red brown in basal half, black in distal half, valvula 3 black with brown tip, membrane in between valvifer 2 and valvula 3 white (Fig. 11A). Fore and mid legs red brown, coxae predominantly black from bases, trochanters partly white. On hind leg, basal third of coxa laterally black, distal portion red brown, trochanter and trochantellus white, femur predominantly red brown in basal half, predominantly black in distal half, tibia black, tarsus white, tarsomere 1 weakly infuscate in basal third (Fig. 3A–C, H–I). Wings weakly brown stained, venation brown, pterostigma black (Fig. 3D–E), sometimes brown (Fig. 3A–B, immature specimens?). MORPHOLOGY. Body 11.3–13.1 mm, fore wing 14.1–14.6 mm long. POL: OOL: OCL = 1.0: 1.6–1.8: 1.8–2.0. Interantennal distance 0.9–1.0 × as wide as distance between torulus and eye margin, malar space 0.4–0.5 × as wide as interantennal distance. Eye 1.3–1.4 × as long as wide, frons at dorsal edge of antennal toruli 1.6 × as wide as maximum diameter of eye. Synantennomere 3 (4.0–) 4.5–4.9 mm, filament 0.9–1.1 mm, with 7–8 articles (9 in one specimen). Article 3 of maxillary palp 0.5–0.6 mm. Ovipositor sheath 2.5–2.7 mm, valvula 3 1.4–1.6 × as long as valvifer 2, valvula 3 2.0–2.1(–2.2) × as long as wide. On hind leg, femur (4.5–) 4.9–5.2 mm, tibia (5.4–) 5.9–6.5 mm, tarsus (4.3–) 4.6–5.2 mm, tarsomere 1 4.2–4.4 × as long as wide, longer distal spur of tibia 0.4–0.5 × as long as tarsomere 1, subapical tooth of claw stout and little shorter than apical tooth. Head dull, medial part of frons and vertex minutely areolate, lateral parts of frons rugose, gena minutely areolate (Fig. 3F and 3G). Inner orbit and ventral half of gena with carina, on inner orbit partly obscured by generally rugose structure. Mesonotum minutely areolate, posterio-medial part of medial lobes of mesoscutum and lateral parts of mesoscutellum rugulose. Mesepisternum minutely areolate, dull, with scattered 20 µm large pits. Metatarsomere 1 dorsally almost glabrous, laterally sparsely setose, lateroventrally with some stout setae up to 150 µm, ventrally densely setose, setae about 100 µm long (Fig. 3H–I). Pulvilli present on metatarsomeres 1–4, on tarsomere 1 pulvillus 70–100 µm long, on tarsomere 4 200–230 µm (Fig. 3H– J). Valvula 3 of ovipositor sheath convex dorsally and ventrally in basal and medial sections, preapically weakly concave, setae up to 250 μm long scattered mainly along dorsal and ventral margins (Fig. 11A). Ovipositor about 2.1 mm long, weakly curved ventrally, valvula 1 and valvula 2 without bulbous areas (Fig. 12A). Valvula 1 in distal 0.6 with annuli, basal annuli sigmoid, distal annuli weakly oblique, annuli with ctenidia composed of minute denticles, ventral edge in distal 0.3 bearing serrulae, cypsellae between distal 6 serrulae deep (Figs 1F, 11A). Valvula 2 in distal 0.4 with annuli, basal annuli weakly curved, distal annuli straight, annuli with ctenidia, denticles larger than those on valvula 1, dorsal edge of valvula 2 indented between annuli. Male COLOR. Similar to female (Fig. 3D–G, K). Subgenital plate and genitalia black. MORPHOLOGY. Body 9.4–10.2 mm, fore wing 11.9–12.3 mm long. POL: OOL: OCL = 1.0: 1.5–1.8: 1.6–1.9. Interantennal distance 0.9–1.0 × as wide as distance between torulus and eye margin, malar space 0.4–0.5 × as wide as interantennal distance. Eye 1.3–1.4 × as long as wide, frons at dorsal edge of antennal toruli 1.6–1.7 × as wide as maximum diameter of eye. Synantennomere 3 4.0– 4.4 mm, filament (0.8–) 0.9–1.1 mm, with 7–8 articles. Article 3 of maxillary palp 0.5 mm long. On hind leg, femur 4.1–4.4 mm, tibia 5.1–5.3 mm, tarsus 4.2–4.5 mm, tarsomere 1 4.7–5.3 × as long as wide, longer distal spur of tibia 0.40 × as long as tarsomere 1. Claws and microsculpture similar to female. Setation of metatarsus less dense than in female (Fig. 3K). Subgenital plate bluntly pointed at apex. Valviceps of penis valve distally evenly rounded, basal 0.4 of upper side expanded to a roundly angular lobe, upper edge of valviceps distal of lobe weakly concave, with large, irregular teeth. Distal half of valviceps in upper and medial portion with long setae, distal of middle in lower portion with dense assemblage of short, conical setae (Fig. 13A). Remarks Megaxyela euchroma sp. nov. is characterized by a number of unique characters (see Diagnosis). All other East Asian species are completely white on the ventral portion and partly on the dorsolateral portion of terga 2–3 or 2–4. In these cases, the white band on terga 2–3 or 2–4 is much wider than on the subsequent terga. The head is more extensively yellow or red brown, at least on the gena in M. gigantea, M. pulchra sp. nov. and M. togashii, or the eye is completely surrounded by black except for the malar space in M. parki. Megaxyela euchroma sp. nov. and M. parki share the white metatarsus, which is black in the other East Asian species. With respect to the white pattern of the terga, M. euchroma sp. nov. is similar to the North American M. alisonae, M. bicoloripes, M. inversa sp. nov. and M. tricolor (Smith & Schiff 1998). Megaxyela alisonae and M. bicoloripes have a white metatarsus similar to M. euchroma sp. nov., while M. tricolor has a black metatarsus. In the Nearctic species, a yellow white pattern on the orbits and the vertex, similar to that of M. euchroma sp. nov., is absent. Megaxyela alisonae and M. bicoloripes bear an extensive red brown pattern on thorax and abdomen, which is absent in M. euchroma sp. nov. Megaxyela inversa sp. nov. differs in the shape of the ovipositor sheath, which is straight dorsally in the basal and the medial sections, vs convex in M. euchroma sp. nov. The analysis of the COI sequences supports the monophyly of the studied specimens by a bootstrap of 100%. The maximum intraspecific variation is 0.3%. The nearest neighbor, M. fulvago sp. nov., is placed at a distance of 12.7% (Fig. 2). All the available adults were found on the undergrowth of open forests during cloudy weather. There were Juglans trees nearby and new shoots of leaflets were beginning to grow.

Megaxyela langstoni Ross, 1936 sp. rev. Figs 7A���F, 11E Megaxyela langstoni Ross, 1936: 131 ���132 (♀ ♂, type locality: USA, Mississippi, Starkville, Mississippi State University). Megaxyela langstoni ��� Smith 1978: 25 (listed in synonymy with M. major). ― Smith 1979: 10 (synonymized with M. major). Megaxyela major ��� Smith & Schiff 1998: 644���648 (misidentification, partim). ― Ree 2012: [1] (damage to pecan). ― Ree 2014: [2] (damage to pecan). ― Ree 2016: [2] (damage to pecan). Type material Holotype USA: ♀: ���State College Miss. 4/7/32 ���; ��� J. M. Langston Collector���; ��� Pecan 283813 ���; [red:] ��� Holotype Megaxyela langstoni Ross ♀���; ���INHS Type #1071���; ��� DEI-GISHym 30821 ��� (INHS), left posterior leg missing. The locality data correspond with the present-day Mississippi State University of Agriculture and Applied Science in Starkville. Additional material examined USA: 1 ♀, Mississippi, Starkville [��� Ag. Coll. Miss. ���], 10 Apr. 1915, C.C. Greer leg., DEI-GISHym 30822 (USNM); 1 ♀, Oklahoma, Pawnee County, Pawnee, 352736 East Hwy 64, 36.292�� N, 96.716�� W, Malaise trap, 9���23 Apr. 2016, C. Apgar leg., DEI-GISHym 30796 (specimen used for barcoding) (USNM); 1 ♀, 1 ♂, Texas (USNM). Supplementary description Female Synantennomere 3 3.8 mm, filament 0.8 mm, with (6���)8���9 articles. Pulvilli present on metatarsomeres 1���4, on article 1 pulvillus 100 ��m long, on article 4 170 ��m (Fig. 7D���E). See key for additional characters. Host plant Pecan (Carya illinoinensis; cited as ��� Carya pecan A. & G.��� by Ross 1936) and possibly additional species of Carya (Dyar 1898b as M. major; Yuasa 1923 as Megaxyela sp. 1). Supposedly also the photo by Ree (2014) of gregarious larvae of M. ��� major ��� feeding on pecan refers here. Remarks Smith (1978, 1979) treated Megaxyela langstoni as a synonym of M. major. With some reservation, Smith & Schiff (1998) discussed differences in behavior, color and morphology as possible intraspecific variation. Here we reinstate M. langstoni sp. rev. as a valid species. The analysis of the barcoding region of three females, all originating from a collection site in Oklahoma, resulted in two clades (DEI-GISHym 30796 and 30767 + 30797, respectively) separated by a minimum pairwise distance of 13.5%, while the two specimens included in the clade 30767 + 30797 are separated by a mimimum pairwise distance of only 0.3% (Fig. 2). This observation is paralleled by different coloration of the adults corresponding with the type material and the descriptions of M. langstoni and M. major: Specimen 30796 agrees with M. langstoni in the predominantly red brown terga and the basally black pterostigma (Fig. 7A), while the specimens 30767 + 30797 agree with M. major in the dorsally predominantly black terga and the unicolorous yellow pterostigma (Fig. 7G). Several specimens included in the type series of M. langstoni were reared from the pecan Carya illinoinensis (Ross 1936). Ross (1936) also referred to the descriptions of larvae of M. major by Dyar (1898b) and Yuasa (1923). Dyar described the larvae as ���gregarious on the young leaves of hickory��� [= Carya spec.]. Yuasa (1923) noted ���on hickory and pecan���. If the association of larvae described by Yuasa as ��� Megaxyela sp. 1��� with M. major by J.M. Langston and H.H. Ross is correct, M. major is a ���solitary feeder on pecan and some other hickories��� (Ross 1936). Citing M. major, M. langstoni has been listed as a pest of pecan in Texas, but ���in most cases sawfly damage is just ���visual discomfort��� where the larvae have damaged some of the new foliage��� (Ree 2012, 2014). Significant defoliations, which might justify a treatment, obviously are rare (Ree 2016). Smith & Schiff (1998) recorded Megaxyela major from 14 states of the eastern US. Collection data from Mississippi completely refer to M. langstoni (type material and additional specimens). We identified additional material from Oklahoma and Texas. For the distribution of M. major, we confirm Kansas (holotype of Odontophyes ferruginea), Texas (lectotype of Xyela major) and, upon additional specimens studied, also Oklahoma and Pennsylvania. Adults from Florida, Iowa, Missouri, New York and South Carolina need to be re-identified. The state records for Alabama, Arkansas, Georgia, Tennessee and Virginia are based on larvae solely. Since the larvae of four of seven Nearctic Megaxyela are unknown, and Megaxyela larvae generally display only vague specific differences according to our experience with East Asian taxa, these records require scrutiny. Ross (1936) discriminated the larvae of M. langstoni and M. major by the presence of a single large dark area vs pairs of black spots on the pronotum and the penultimate abdominal segment. But these character states might also apply to those Nearctic species of Megaxyela for which the larvae are still unknown., Published as part of Stephan M. Blank, Katja Kramp, David R. Smith, Yuri N. Sundukov, Meicai Wei & Akihiko Shinohara, 2017, Big and beautiful: the Megaxyela species (Hymenoptera, Xyelidae) of East Asia and North America, pp. 1-46 in European Journal of Taxonomy 348 on pages 22-24, DOI: 10.5852/ejt.2017.348, http://zenodo.org/record/1042663, {"references":["Ross H. H. 1936. A new pecan sawfly, Megaxyela langstoni n. sp. (Hymenoptera: Xyelidae). Entomological News 47: 131 - 132.","Shinohara A. & Smith D. R. 1979. The Sato types of sawflies (Hymenoptera: Symphyta). Bulletin of the National Science Museum, Series A, Zoology 5: 281 - 288.","Smith D. R. & Schiff N. M. 1998. The genera Macroxyela Kirby and Megaxyela Ashmead (Hymenoptera: Xyelidae) in North America. Proceedings of the Entomological Society of Washington 100: 636 - 657.","Ree B. 2012. Insects. Texas Pecan Pest Management Newsletter 12 (1): [1 - 3].","Ree B. 2014. Sawfly. Texas Pecan Pest Management Newsletter 14 (2): [1 - 3].","Ree B. 2016: Insects. Texas Pecan Pest Management Newsletter 16 (2): [1 - 3]. Available from http: //presidio. agrilife. org / files / 2011 / 08 / Texas-Pecan-Pest-Management-Newsletter-March- 31 - 2016. pdf [accessed 13 Jul. 2017].","Dyar H. G. 1898 b. Notes on some sawfly larvae, especially the Xyelidae. Canadian Entomologist 30: 173 - 176 [July 1898]. https: // doi. org / 10.4039 / Ent 30173 - 7","Cresswell S. 2014. April Unknown - Megaxyela tricolor. Available from BugGuide, http: // bugguide. net / node / view / 908657 [accessed 20 Jul. 2016]."]}

Megaxyela inversa Blank & D.R. Smith sp. nov. urn:lsid:zoobank.org:act:5BA86B4F-CFAC-42E0-A2B8-BC81E1C0B517 Figs 6A–G, 11D, 12D Diagnosis This species is unique by the very long ovipositor sheath with the dorsal outline of valvula 3 straight and the ventral outline convex, and the ovipositor curved dorsally along its longitudinal axis. Etymology The Latin adjective inversus (fem. inversa) indicates the shape of the ovipositor, that, in contrast to other Megaxyela species, is dorsally curved. Type material Holotype USA: ♀: “USA West Virginia, Randolph Co., Job, 38.8528 N, 79.5293 W, 1-20 April 2012, J. Whitaker 10067, USGS-DRO 292937 ”, [red:] “Holotype Megaxyela inversa det. S.M. Blank & D.R. Smith 2017 ”, “ DEI-GISHym 32152 ” (USNM). Paratype UNKNOWN: 1 ♀, “ 4/22/91 ” [supposed collection date: 22. Apr. 1891, no collecting locality given, see insertion in Fig. 6A for original labels], DEI-GISHym 22356 (USNM). Description Female COLOR. Black with white pattern, black parts with bronze to blue tinge (Fig. 6A, C). Head below toruli brown, clypeus and labrum white, preapical region of labrum and mandibles brown white, labial palpomeres 1–2 white (Fig. 6C). Antenna black with basal ¾ of scape in inner surface light orange. Pronotum and metanotum with bronze tinge, mesonotum and mesepisternum with blue tinge. Tegula basally brown, distally white. Abdominal terga and sterna black, terga 1–8 and sterna with white distal margins, ventral part of terga laterally with 100 µm wide white stripe, sterna 5–7 with 40–50 µm wide lateral white stripe (Fig. 6A). Valvifer 2 red brown, valvula 3 black except for narrow brown tip, membrane between valvifer 2 and valvula 3 white (Fig. 11D). Legs orange, procoxa and mesocoxa basally infuscate, distal 0.20 of metafemur, complete metatibia and metatarsus black. Wing subhyaline, venation and pterostigma black (Fig. 6F–G). MORPHOLOGY. Body 10.2 mm, fore wing 11.2 mm long. POL: OOL: OCL = 1.0: 1.3: 1.5. Interantennal distance 1.3 × as wide as distance between torulus and eye margin, malar space 0.4 × as wide as interantennal distance. Eye 1.5 × as long as wide, frons at dorsal edge of antennal toruli 1.4 × as wide as maximum diameter of eye. Synantennomere 3 3.1 mm, filament 0.9 mm, with 9 articles. Article 3 of maxillary palp 0.4 μm. Ovipositor sheath 5.1 mm, 3.8 × as long as metatarsomere 1, valvula 3 1.3 × as long as valvifer 2, valvula 3 3.3 × as long as wide. On hind leg, femur 3.1 mm, tibia 4.5 mm, tarsus 3.6 mm, tarsomere 1 5.8 × as long as wide, longer distal spur of tibia 0.3 × as long as tarsomere 1, claws cleft, subapical tooth of claw little smaller at base than apical tooth. Head dull, lateral part of frons rugose, medial part minutely areolate with some 50 µm large pits, vertex and postocellar area mostly minutely areolate to rugulose, gena rugulose (Fig. 6C). Mesoscutum and mesoscutellum homogeneously minutely areolate. Mesepisternum medially coriarious and shining, with 20 µm large pits. Metatarsomere 1 dorsally and laterally almost glabrous, ventrally densely setose, setae 70–100 µm long (Fig. 6D). Pulvilli present on metatarsomeres 1–4, on article 1 pulvillus 150 µm long, on article 4 230 µm (Fig. 6E). Basal and medial sections of valvula 3 of ovipositor sheath straight dorsally, convex ventrally, preapical section almost straight on dorsal and ventral edge, with dense pattern of up to 200 μm long setae (Figs 6A, 11D). Ovipositor 4.6 mm long, weakly curved dorsally, without bulbous areas in middle, without ctenidia (Fig. 12D). Valvula 1 in distal 0.6 with annuli, basal annuli much more widely spaced then medial and distal annuli, medial annuli very narrowly spaced, subparallel, straight, vertical, ventral edge in distal 0.2 bearing small serrulae. Valvula 2 in distal 0.3 with widely spaced annuli, annuli not extending to ventral edge of valvula, distal 0.3 of dorsal edge of valvula 1 with partly prominent, rounded denticles. Male Unknown. Remarks The ovipositor is curved dorsally in Megaxyela inversa sp. nov. (Fig. 12D), which is an unusual condition in sawflies and unique in Macroxyelinae (Fig. 12A–B, E–H; Smith & Schiff 1998: figs 16–21). The predominant number of sawflies and woodwasps have the ovipositor curved ventrally or the ovipositor is straight (see, e.g., figures in Weltz & Vilhelmsen 2014). Among Xyelidae, the sheath is curved dorsally to a various degree in species of Pleroneura Konow, 1897 (e.g., Smith et al. 1977; Shinohara 1995, 2016) and in Xyela concava Burdick, 1961 (Burdick 1961). Among Nearctic Megaxyela this species is most similar to M. bicoloripes and M. tricolor by the slender metatarsus and the red brown metafemur bearing an extensively infuscate apex. It is readily distinguished from all other Megaxyela species by the outline of the basal and medial portion of valvula 3, which in lateral view is straight dorsally and evenly curved ventrally. The remaining Megaxyela species have either both the dorsal and the ventral outlines of valvula 3 curved, or the dorsal outline is curved and the ventral straight (Fig. 11A–C, 11E–H; Smith & Schiff 1998: figs 7–11). The paratype is presumably from North America because of its discovery in a US collection and the holotype is from West Virginia. Morphometry and figures are from the paratype since the specimen used as holotype was discovered too late to include in this paper.

Megaxyela Ashmead, 1898 comprises 13 species, four of which are described as new and one is removed from synonymy: Megaxyela euchroma Blank, Shinohara & Wei sp. nov. from China (Zheijang), M. fulvago Blank, Shinohara & Wei sp. nov. from China (Hunan, Jiangsu, Zhejiang), M. inversa Blank & D.R. Smith sp. nov. from the USA (West Virginia), M. langstoni Ross, 1936 sp. rev. from the eastern USA, and M. pulchra Blank, Shinohara & Sundukov sp. nov. from China (Hubei, Jilin, Liaoning, Shaanxi, Tibet), South Korea (Kangwon-do) and Russia (Primorskiy Kray). The male of M. parki Shinohara, 1992 is described for the first time. A lectotype is designated for M. gigantea Mocsáry, 1909. A cladogram, based on COI sequences of seven species, is presented and interpreted in view of selected morphological characters. Records of M. fulvago sp. nov. from Hunan and of M. pulchra sp. nov. from Tibet extend the known distribution of Megaxyela in the Old World 600 kilometers farther south and 2500 kilometers farther west than previous records.

We made a molecular phylogenetic analysis using mitochondrial cytochrome oxidase subunit 1 (COI) gene sequences for ten unidentified Megaxyela larvae from Hokkaido, Honshu and Shikoku, Japan, and 15 identified adults of four Megaxyela , one Macroxyela and three Xyela species. It revealed that all larvae belonged to M. togashii Shinohara, 1992, which showed rather large intraspecific genetic variability even among the individuals from the same population. This is the first distribution record of M. togashii from Hokkaido. Megaxyela togashii is a univoltine species with a very short larval feeding period, only nine days in one rearing experiment from egg to larval maturation. The larva is a solitary, external leaf-feeder on Juglans ailanthifolia , resting curled around the central leaf vein at the apex of a leaflet, and may resemble the excrement of birds. The prepupa overwinters in an earthen cell whose wall is made only of soil, neither parchment-like nor containing fiber. The mature larva is described and several life traits are discussed.

The Pristiphora ruficornis group, defined here based on the structure of the penis valve and the genetic data, includes morphologically and genetically highly similar species that remain taxonomically challenging. Study of most of the relevant type material, examination of female saws and male genitalia, some rearing experiments, and genetic data enabled us to solve most of the taxonomic problems involving northern European taxa. As a result, 17 species are recognised in northern Europe. The following synonymies are proposed: Pristiphora aterrima Lindqvist, 1977, syn. n. is synonymised with P. albitibia (Costa, 1859), P. brunniapex Lindqvist, 1960, syn. n. and P. coniceps Lindqvist, 1955, syn. n. both with P. subopaca Lindqvist, 1955, Nematus vitreipennis Eversmann in Kawall, 1864, syn. n. (nomen oblitum) with P. leucopus (Hellén, 1948) (nomen protectum), and Nematus (Pristiphora) ruficornis var. integer Hellén, 1948, syn. n. with P. ruficornis (Olivier, 1811). Lectotypes are designated for the following taxa: Nematus appendiculatus Hartig, 1837, Nematus cathoraticus Förster, 1854, Nematus (Pristiphora) bifidus Hellén, 1948, Nematus frigidus Boheman, 1865, Pristiphora adelungi Konow, 1902, Nematus vitreipennis Eversmann in Kawall, 1864, Nematus melanocarpus Hartig, 1840, Nematus wuestneii Stein, 1885, Pristiphora pusilla Malaise, 1921, and Nematus fraxini Hartig, 1837. An illustrated electronic key made with Lucid and a traditional dichotomous key are provided to facilitate identification of the species. In addition we report the first occurrence of distinctly asymmetrical penis valves in Pristiphora (in P. pusilla), a condition rarely observed in Hymenoptera.

Morphological and genetic data indicate that Xyela fusca Blank, Kramp & Shinohara spec. nov. from Shikoku, Japan, plus an undescribed species from China are closely related to the Nearctic X. bakeri Konow, 1898. These species form the X. bakeri group, which is considered to be the sister clade of the Eurasian X. julii group. In addition to the X. alpigena group, the X. bakeri group is the second species group of Xyela with representatives occurring in both the Old and the New World. While X. bakeri is associated with Pinus ( Pinus ) species as the larval hosts, collection data of imagines suggest P. ( Strobus ) parviflora for X. fusca . This appears to be the second case, in which closely related Xyela species have switched hosts between species belonging to different subgenera of Pinus . Based on distribution data, migration of the common ancestor across Beringia appears likely, which would also have involved the change of the host subgenus. The six species of Xyela of Japan are keyed and new distribution data are presented. New, most northwesterly records of imagines from Yukon Territory (Canada) and field observations in Colorado and California (USA) suggest P. contorta as an additional larval host of X. bakeri .

The species formerly known as Pachyprotasis nigronotata Kriechbaumer, 1874 in Japan is described as P. kojimai Taeger & Shinohara, sp. nov. Both taxa are not very closely related, but were hitherto mixed up because of their similar coloration. The new species is endemic to the mountains of central Honshu and P. nigronotata is to be deleted from the fauna of Japan.

The phylogeography of typical alpine plant species is well understood in Europe. However, the genetic patterns of boreo-montane species are mostly unstudied. Therefore, we analysed the AFLPs of 198 individuals of Polygonatum verticillatum over a major part of its European distribution. We obtained a total of 402 reproducible fragments, of which 96.8% were polymorphic. The average Phi(ST) over all samples was high (73.0%). The highest number of private fragments was observed in the Cantabrian Mountains; the highest genetic diversities of the populations were detected in populations from the Alps. BAPS, Principal Coordinates and Cluster analyses revealed a deep split between the Cantabrian population and all other samples. The latter further distinguished two major groups in western and eastern Europe. These results suggest a complex biogeographical history of P. verticillatum. The Cantabrian population was most probably isolated for the longest time. Furthermore, putative glacial survival centres might have existed in the western group around the glaciated Alps and in the eastern group in the foothills of the Carpathian and Balkan mountain systems. The origin of the Scandinavian populations is still unresolved, but an origin from the southeastern Alps or the western Balkans appears the most likely scenario.