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North-Western Palaearctic species of Pristiphora (Hymenoptera, Tenthredinidae)
expand article infoMarko Prous§, Katja Kramp, Veli Vikberg|, Andrew David Liston
‡ Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany
§ University of Tartu, Tartu, Estonia
| Unaffiliated, Turenki, Finland
Open Access

Abstract

North-Western Palaearctic species of Pristiphora Latreille, 1810 are revised. Altogether, 90 species are treated, two of which are described as new: P. caraganae Vikberg & Prous, sp. n. from Finland and P. dedeara Liston & Prous, sp. n. from Germany. Host plant of P. caraganae is Caragana arborescens Lam. Pristiphora dasiphorae (Zinovjev, 1993) (previously known from East Palaearctic) and P. cadma Wong & Ross, 1960 (previously known from North America) are recorded for the first time from Europe. Nematus nigricans Eversmann, 1847 [= Pristiphora nigricans (Eversmann, 1847), comb. n.], N. breviusculus Eversmann, 1847 [= Euura melanocephalus (Hartig, 1837)], and N. caudalis Eversmann, 1847 [= E. caudalis (Eversmann, 1847), comb. n.] are removed from synonymy with P. pallidiventris (Fallén, 1808), N. paralellus Hartig, 1840 [= P. paralella (Hartig, 1840), comb. n.] is removed from synonymy with P. bufo (Brischke, 1883), and P. mesatlantica Lacourt, 1976 is removed from synonymy with P. insularis Rohwer, 1910. The following 29 new synonymies are proposed: P. nigropuncticeps Haris, 2002, syn. n. with P. albitibia (Costa, 1859); Lygaeonematus karvoneni Lindqvist, 1952, syn. n. with P. alpestris (Konow, 1903); P. (P.) anivskiensis Haris, 2006, syn. n. with P. appendiculata (Hartig, 1837); Nematus canaliculatus Hartig, 1840, syn. n with P. carinata (Hartig, 1837); P. nigrogroenblomi Haris, 2002, syn. n. with P. cincta Newman, 1837; Tenthredo flavipes Zetterstedt, 1838, syn. n., Nematus congener W.F. Kirby, 1882, syn. n., and P. thomsoni Lindqvist, 1953, syn. n. with P. dochmocera (Thomson, 1871); P. atrata Lindqvist, 1975, syn. n. with P. friesei (Konow, 1904); P. gelida Wong, 1968, syn. n. with P. frigida (Boheman, 1865); Pachynematus nigricorpus Takagi, 1931, syn. n. with P. laricis (Hartig, 1837); Nematus (Pikonema) piceae Zhelochovtsev in Zhelochovtsev and Zinovjev, 1988, syn. n. and P. (P.) hoverlaensis Haris, 2001, syn. n. with P. leucopodia (Hartig, 1837); Mesoneura arctica Lindqvist, 1959, syn. n., Pachynematus incisus Lindqvist, 1970, syn. n., Pachynematus intermedius Verzhutskii, 1974, syn. n., and P. mongololaricis Haris, 2003, syn. n. with P. malaisei (Lindqvist, 1952); Nematus anderschi Zaddach, 1876, syn. n., P. inocreata Konow, 1902, syn. n., and P. discolor Lindqvist, 1975, syn. n. with P. nigricans (Eversmann, 1847); Lygaeonematus tenuicornis Lindqvist, 1955, syn. n. with P. paralella (Hartig, 1840); Lygaeonematus concolor Lindqvist, 1952, syn. n. with P. pseudocoactula (Lindqvist, 1952); P. flavipicta Lindqvist, 1975, syn. n., P. flavopleura Haris, 2002, syn. n., P. mongoloexigua Haris, 2002, syn. n., and P. mongolofausta Haris, 2003, syn. n. with P. punctifrons (Thomson, 1871); P. listoni Lacourt, 1998, syn. n. with P. sootryeni Lindqvist, 1955; P. gaunitzi Lindqvist, 1968, syn. n. with P. testacea (Jurine, 1807); and Nematus breviusculus Eversmann, 1847, syn. n. with Euura melanocephalus (Hartig, 1837). The valid name of Pachynematus (Pikonema) carpathiensis Haris, 2001 is Nematinus carpathiensis (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 the carinata (previously Lygaeotus), micronematica (previously Lygaeophora), and rufipes (also known as thalictri or aquilegiae) groups are not keyed to the species level, because additional research is needed to delimit the species more reliably in these groups. Phylogeny of Pristiphora is reconstructed based on one mitochondrial (COI) and two nuclear (NaK and TPI) genes. Remarkably, around 50–60% (depending on the exclusion or inclusion of the carinata, micronematica, and rufipes groups) 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).

Keywords

Sawflies, revision, new synonyms, nomenclature, taxonomy, identification key, phylogeny, DNA barcoding

Introduction

Pristiphora Latreille, 1810, as defined by Prous et al. (2014), is the second largest genus in Nematinae (Tenthredinidae), including about 240 species (Taeger et al. 2010). About half of these (120) are known in the West Palaearctic. The main keys available for the majority of West Palaearctic species are those of Benson (1958) and Zhelochovtsev and Zinovjev (1988). Benson’s (1958) key is outdated and geographically restricted (British Isles). The key by Zhelochovtsev and Zinovjev (1988) includes more species because of wider geographic scope and new species described since Benson (1958). The most comprehensive key for Pristiphora in terms of number of species and geographic scope (Palaearctic) was published by Haris (2006b), but it is mostly based on previous keys and species descriptions. Both Haris (2006b) and Zhelochovtsev and Zinovjev (1988), studied only few types, included many species based on literature, and uncritically accepted characters used in previous keys. Problems in identifying species of Pristiphora, and Nematinae in general, are exacerbated by inherent difficulties caused by the large number of species and a lack of discrete characters suitable for separating them. Here we revise the species found in the North-Western Palaearctic Region (defined here as Scandinavia and its neighbouring regions) and delimit several species groups (some of them previously treated as genera or subgenera) using molecular and morphological data. We provide photos of lancets and penis valves, an electronic key employing a large number of characters, and a dichotomous key using the most reliable characters for species identification. For convenience, the recently revised ruficornis species group (Prous et al. 2016) is also included. However, delimiting species belonging to the carinata (Lygaeotus), micronematica (Lygaeophora), and rufipes (also known as thalictri or aquilegiae group) groups requires additional research.

The most comprehensive phylogenetic analyses of Pristiphora so far published were part of broader analyses dealing mainly with higher level relationships of Nematinae (Nyman et al. 2006, Nyman et al. 2010, Prous et al. 2014). Here we estimate the phylogeny of Pristiphora based on expanded taxon sampling and three genes: one mitochondrial (COI) and two nuclear (NaK and TPI).

Mitochondrial COI barcodes are widely used for species identification. This often works rather well (Pentinsaari et al. 2014, Mutanen et al. 2016), but there are indications of significant barcoding failure in some groups of sawflies (Schmidt et al. 2017), particularly in Nematinae (table 2 in Schmidt et al. 2017). However, as there are also numerous taxonomic problems within Nematinae, the exact nature of barcoding failure is in many cases uncertain. The application of COI barcoding to Pristiphora revealed that COI cannot be reliably used for species identification in roughly half of the species, and that nuclear genes seem to work better for that purpose.

Material and methods

Specimens examined or mentioned are deposited in the following collections:

ANSP Academy of Natural Sciences of Drexel University, Philadelphia, USA;

BMNH The Natural History Museum, London, United Kingdom;

CEH Collection of Erik Heibo, Lierskogen, Norway;

CMH Collection of Mikk Heidemaa, Tartu, Estonia;

CMV Collection of Matti Viitasaari, Helsinki, Finland;

CNC Canadian National Collection of Insects, Ottawa, Canada;

COL Collection of Ole Lønnve, Oslo, Norway;

CVV Collection of Veli Vikberg, Turenki, Finland;

CTN Collection of Thierry Noblecourt, Quillan, France;

EJC Collection of Ewald Jansen, Leipzig, Germany;

ETHZ Eidgenössische Technische Hochschule-Zentrum, Zurich, Switzerland;

HMUG Hunterian Museum, University of Glasgow, United Kingdom;

HNHM Hungarian Natural History Museum, Budapest, Hungary;

INHS Illinois Natural History Survey, Champaign, USA;

INRA Institut National de la Recherche Agronomique, Centre de Biologie pour la Gestion des Populations (CBGP), Montferrier-sur-Lez Cedex, France;

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium;

LPNC Collection of Pierre-Nicolas Libert, Somal, Belgium;

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, USA;

MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain;

MNHN Muséum National d’Histoire Naturelle, Paris, France;

MZH Finnish Museum of Natural History, Helsinki, Finland;

MZLU Lunds Universitet, Lund, Sweden;

MZPW Museum of the Institute of Zoology, Polish Academy of Science, Warsaw, Poland;

NFVG Niedersächsische Forstliche Versuchsanstalt, Göttingen, Germany;

NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden;

NMPC National Museum (Natural History), Prague, Czech Republic;

NMW Naturhistorisches Museum Wien, Wien [= Vienna], Austria;

NFRC Northern Forestry Centre, Edmonton, Canada;

NSM University of Nebraska State Museum, Lincoln, USA;

NSMT National Museum of Nature and Science, Tokyo, Japan;

NUORTJ Collection of Juoko Nuorteva, Helsinki, Finland;

RMNH Nationaal Natuurhistorische Museum, Leiden, Netherlands;

RSME National Museums of Scotland, Edinburgh, United Kingdom;

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany;

SMTP Swedish Malaise Trap Project, Station Linné, Öland, Sweden;

TROM Tromsø University Museum, Tromsø, Norway;

TUZ Natural History Museum, University of Tartu, Tartu, Estonia;

UEF University of Eastern Finland, Joensuu, Finland;

USNM National Museum of Natural History, Smithsonian Institution, Washington D.C., USA;

ZIN Russian Academy of Sciences, Zoological Institute, St. Petersburg, Russia;

ZMAN Universiteit van Amsterdam, Instituut voor Taxonomische Zoologie, Zoologisch Museum, Amsterdam, Netherlands;

ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany;

ZMUC Zoological Museum of the University of Copenhagen, Denmark;

ZMUM Zoological Museum, Moscow State University, Moscow, Russia;

ZMUO Zoological Museum, University of Oulu, Finland;

ZSM Zoologische Staatssammlung, München [= Munich], Germany.

Names of the mentioned host plants follow The Plant List (http://www.theplantlist.org/). Unless otherwise stated, species distribution data at the level of zoogeographic regions is taken from Taeger et al. (2010).

Collection data of the examined specimens is included in an excel file available at figshare (http://dx.doi.org/10.6084/m9.figshare.5235835).

Morphological methods

To photograph penis valves and lancets (valvula 1 or ventral part of saw), genital capsules and ovipositors were separated from the specimen and macerated in KOH (10–15%) for 6–10 hours at room temperature or treated with proteinase during DNA extraction (see below). Temporary or permanent slide preparations were made of dissected lancets and penis valves. For temporary slides, glycerine was used. After photographing, the lancets and penis valves were glued on a piece of cardboard, which was pinned with the corresponding specimen. For permanent slides, Euparal or PVA-mounting medium (Danielsson 1985) was used (these specimens are labelled as ‘PR.XXXVV’, e.g. PR.579VV). Photos were taken with a digital camera attached to a microscope. Composite images with an extended depth of field were created from stacks of images using the software CombineZP (Alan Hadley; http://www.hadleyweb.pwp.blueyonder.co.uk/). Most of the lancets were photographed in two overlapping parts and a single image created using the program Image Composite Editor (Microsoft), or in few cases with plugin MosaicJ (Thévenaz and Unser 2007) implemented in ImageJ version 1.46r (Wayne Rasband; http://imagej.nih.gov/ij/). Morphological terminology follows Vikberg (1978, 2006) and Viitasaari (2002). We do not use the term “ctenidia” to denote the bands of setae on the annuli of female lancets (Fig. 129), as used previously by Prous et al. (2016) following Viitasaari (2002, Fig. 134). The term “ctenidia” (singular “ctenidium”) correctly refers to rows of spines (i.e. protrusions of the cuticle); superficially they resemble bands of setae, which are composed of sensilla (Smith 1968, Blank and Schönitzer 1994, fig. 131 in Viitasaari 2002). Therefore we use the more neutral term “setae”.

Rearing of larvae and adults

All the rearing data newly published here were obtained by VV. These data are given under Rearing notes in the Taxonomy section. Plants chosen for ovipositing experiments were mainly those from which females were collected in the field or reared from larvae. Our main intention was to find at least one acceptable host plant for rearing particular species. Ovipositing experiments were carried out indoors, at room temperature. Suitable parts of plants were cut, the cut end put in water or wrapped in moistened filter paper (paper towel), and the leaves offered to the sawfly in a closed container. Larvae were fed with fresh leaves that were changed every day or every second day. Cocoons were placed in glass vials and overwintered outdoors in a wooden chest.

Molecular methods

DNA was extracted and purified with an EZNA Tissue DNA Kit (Omega Bio-tek) according to the manufacturer’s protocol and stored at -20 °C for later use. Typically, the middle right leg was used for DNA extraction, but for males the whole genital capsule was often additionally used to increase DNA yield and to free penis valves from muscles for photographing. One mitochondrial and two nuclear regions were used in phylogenetic analyses. Primers used for amplification and sequencing are listed in Table 1. The mitochondrial region used is a large fragment (1078 bp) of cytochrome oxidase subunit I gene (COI). The first (from the 5’ end) 658 bp of this fragment correspond to the standard barcode region of the animal kingdom (Hebert et al. 2003). If the amplification of the 1078 bp fragment failed, or was expected to fail because of low DNA quality, the region was amplified in two overlapping fragments, or only the barcoding (658 bp) region was obtained. The nuclear markers used are fragments of triose-phosphate isomerase (TPI) and sodium/potassium-transporting ATPase subunit alpha (NaK). The TPI fragment used is the nearly complete gene region, containing 676 bp of three exons and two short introns (each around 50–100 bp) in Nematinae, altogether 795–833 bp. The NaK fragment used is a nearly complete sequence of its longest exon, 1654 bp. New NaK primers were designed mainly based on four sawfly genomes and one transcriptome available in GenBank (accessions AOFN01001568, GAWW02019159, LGIB01000323, AMWH01001469, AZGP01005167), or using sequences published by Malm and Nyman (2015). Numbers in the new NaK primer names refer to the binding position of the primer’s 3’ end in the coding region of Athalia rosae mRNA (accession XM_012414227). Seven sequences of four specimens (4b, 9t, DG, DH) newly reported here were obtained from the VoSeq database (Peña and Malm 2012) maintained by Tommi Nyman (University of Eastern Finland). For these four specimens, 810 bp of COI and 997 bp of NaK had been sequenced as described previously (Nyman et al. 2006, Leppänen et al. 2012).

PCR reactions were carried out in a total volume of 15–25 μl containing 1–2 μl of extracted DNA, 1.0–1.5 μl (5.0–7.5 pmol) of primers and 7.5–12.5 μl of 2x Multiplex PCR Plus Master mix (QIAGEN). The PCR protocol consisted of an initial DNA polymerase (HotStar Taq) activation step at 95 °C for 5 min, followed by 38–40 cycles of 30 s at 95 °C, 90 s at 47–59 °C depending on the primer set used, and 30–120 s (depending on the amplicon size) at 72 °C; the last cycle was followed by a final 30 min extension step at 68 °C. 3 μl of PCR product was visualised on a 1.4% agarose gel and then purified with FastAP and Exonuclease I (Thermo Scientific). 1.0–1.8 U of both enzymes were added to 12–22 μl of PCR solution and incubated for 15 min at 37 °C, followed by 15 min at 85 °C. Purified PCR products were sent to Macrogen (Netherlands) for sequencing. To obtain unequivocal sequences, both sense and antisense strands were sequenced, using the primers listed in Table 1. Some TPI sequences were polymorphic for intron length and in those cases heterozygous insertions/deletions (indels) were reconstructed using the program Indelligent v.1.2 (Dmitriev and Rakitov 2008), available at http://dmitriev.speciesfile.org/indel.asp. Only the longest haplotype was submitted to Genbank, because Genbank does not accept more than one sequence per specimen and marker. Ambiguous positions (i.e. double peaks in chromatograms of both strands) due to heterozygosity or heteroplasmy were coded using IUPAC symbols. The COI sequence of one sequenced specimen (Pristiphora friesei DEI-GISHym11558) had a single bp deletion, which was replaced with N before submitting to GenBank.

Primers used for PCR and sequencing, with information provided on respective gene fragment, primer name, direction (forward, F or reverse, R) and location (internal, i or external, o) according to each gene fragment, primer sequence, standard PCR annealing temperature, utilization (PCR/ sequencing), and reference. Primer annealing temperatures used for sequencing at Macrogen were 47°C for COI and 50°C for nuclear genes.

Gene Region Primer name F/R i/o Primer sequence 5'–3' PCR annealing temperature (°) PCR/ Sequencing Reference
COI SymF1 F o TTTCAACWAATCATAAARAYATTGG 47 PCR, seq Prous et al. (2016)
COI SymF2 F o TTTCAACAAATCATAAARAYATTGG 47 PCR, seq Prous et al. (2016)
COI sym- C1-J1718 F i/o GGAGGATTTGGAAAYTGAYTAGTWCC 49 PCR, seq (Nyman et al. 2006)
COI symC1-J1751 F i/o GGAGCNCCTGATATAGCWTTYCC 47 PCR, seq Prous et al. (2016)
COI C1-N1760 R i/o GGTARAAATCARAATCTTATATTAT 47 PCR, seq (Prous et al. 2011)
COI SymR1 R i/o TAAACTTCWGGRTGICCAAARAATC 47 PCR, seq Prous et al. (2016)
COI SymR2 R i/o TAAACTTCTGGRTGTCCAAARAATCA 47 PCR, seq Prous et al. (2016)
COI A2590 R o GCTCCTATTGATARWACATARTGRAAATG 49 PCR, seq (Normark et al. 1999)
TPI TPI_29Fi F o GYAAATTYTTYGTTGGNGGIAA 52 PCR, seq Prous et al. (2016)
TPI TPI hym intF F i AARGGHGCNTTYACYGGNGA 56 Seq (Malm and Nyman 2015)
TPI TPI hym intR R i TCNGARTGDCCHADRATNACCCA 52 Seq (Malm and Nyman 2015)
TPI TPI385Fi F o GTRATYGCNTGYATYGGIGARA 52 PCR, seq Prous et al. (2016)
TPI TPI 275Ri R o GCCCANACNGGYTCRTAIGC 56 PCR, seq (Malm and Nyman 2015)
TPI TPI706R R o ACNATYTGTACRAARTCWGGYTT 52 PCR, seq Prous et al. (2016)
NaK NaK_263F F o CTYAGCCAYGCRAARGCRAARGA 59 PCR, seq This study
NaK NaK_809F F i/o GCWTTYTTCTCNACSAAYGCSGTNGARGG 55 PCR, seq This study
NaK NaK_907Ri R i/o TGRATRAARTGRTGRATYTCYTTIGC 54 PCR, seq This study
NaK NaK_910R R i/o TGRATRAARTGRTGRATYTCYTT 50 PCR, seq This study
NaK NaK_1250Fi F i ATGTGGTTYGAYAAYCARATYATIGA 56 Seq This study
NaK NaKFor470 F i ATGTGGTTYGAYAAYCARATYATCGA 56 Seq Leppänen et al. (2012)
NaK NaKRev475 R i TCGATRATYTGRTTRTCRAACCACAT 56 Seq Leppänen et al. (2012)
NaK NaK_1918R R o GATTTGGCAATNGCTTTGGCAGTDAT 59 PCR, seq This study

Sequences reported here have been deposited in the GenBank (NCBI) database (accession numbers KY698031KY698412 and MF426916MF426924). Some of the sequences analysed here were originally published by Nyman et al. (2006, 2010), Prous et al. (2014, 2016), and Schmidt et al. (2017).

Sequences of COI, NaK, and TPI without introns were aligned manually, among which only one COI sequence of P. friesei (DEI-GISHym11558) had a single bp deletion.

For concatenation of separate alignments of different genes we used FASconCAT-G (Kück and Meusemann 2010) (https://www.zfmk.de/en/research/research-centres-and-groups/fasconcat-g). To minimize the missing cells in the dataset, a few composite COI sequences were created from different specimens of the same species. A nearly complete 1078 bp region of COI was created in these cases by combining the barcoding region (up to 658 bp) with the 810 bp region (423 bp of which overlaps with the barcoding region) when the overlapping part was identical between the sequences. Both ID numbers of the specimens used to create the composite COI sequences are given in the figures.

For phylogenetic analyses we used the maximum likelihood method (ML) implemented in RAxML v. 8.2.9 (Stamatakis 2014) through CIPRES Science Gateway V. 3.3 (Miller et al. 2010) at https://www.phylo.org/. Robustness of reconstructed trees was estimated with 1000 rapid bootstrap replicates. For tree search, GTRCAT model with 25 site-specific rate categories was used and the final tree evaluated with the GTRGAMMA model (Stamatakis 2006; http://sco.h-its.org/exelixis/web/software/raxml/). The concatenated dataset was partitioned by genes, and model parameters estimated separately for each gene. Additionally, MEGA7 (Kumar et al. 2016) was used to calculate p-distances (proportion of nucleotide differences) between specimens and net synonymous divergence between species. Unless stated otherwise, TPI introns were excluded from calculations. Minimal p-distances between and maximal distances within BIN (Barcode Index Number) clusters were taken from BOLD (http://www.boldsystems.org/) BIN database. Alignment files and output files from phylogenetic analysesare available at figshare (http://dx.doi.org/10.6084/m9.figshare.5235832).

Some of the COI barcode sequences used here were obtained from BOLD (http://www.boldsystems.org/). In this case, DNA extraction, PCR amplification, and sequencing were conducted at the Canadian Centre for DNA Barcoding (CCDB) in Guelph, Canada, using standardised high-throughput protocols (Ivanova et al. 2006, deWaard et al. 2008), available online under www.ccdb.ca/resources.php. DNA aliquots of SDEI vouchers are deposited in the DNA storage facility of the SDEI (including those that were originally extracted in CCDB).

Preparation of the keys

The electronic identification key for the species of Pristiphora was prepared in Lucid 3.5 Builder (http://www.lucidcentral.org/) and a zip file containing all the Lucid data files is available at figshare (http://dx.doi.org/10.6084/m9.figshare.5235805). If the licence for Lucid 3.5 is lacking, the free version of Lucid 3.3 can be used to run the key. In case of ambiguities or polymorphisms in character states, we conservatively coded these as multiple states. The key contains 90 morphological features with 244 character states, and 68 male and 71 female entities (species or groups). The first choice given in the key is between female and male, one of which has to be chosen to see all other characters. After that, characters can be chosen freely or one can use ‘Best’ and ‘Next Best’ tools in Lucid to suggest the most efficient sequence of characters for identification. A traditional dichotomous key was constructed manually to emphasise the most reliable characters (usually penis valve for males, and valvula 3 or lancet for females).

Results

Definition of Pristiphora and its separation from other Nematinae

The genus was recently delimited mainly by phylogenetic analyses of DNA sequence data by Prous et al. (2014), which is here confirmed with the addition of the nuclear gene TPI. Phylogenetic analysis of mitochondrial COI and nuclear NaK and TPI sequences (altogether 3408 bp) strongly support monophyly of Pristiphora (Fig. 1). There are no unambiguous morphological characters that define Pristiphora, but the combination of the following characters can be used to distinguish most of the species from the similar genera Nematus and Euura (see Prous et al. 2014): clypeus usually more or less truncate (Fig. 9); apex of vein C usually swollen; head length behind eyes usually small; claws often with small subapical tooth or simple; in females, valvula 3 (apical sawsheath) often with scopa (e.g. Figs 75, 86, 104–107, 127) and tangium of lancet nearly always with campaniform sensilla (Fig. 177); in males, posterior end of tergum 8 without distinct apical projection in most species (except P. armata and P. leucopus), and penis valve usually with Valvispina arising close to the ventral margin. Vikberg (1982) lists larval characters that might more reliably distinguish Pristiphora from Nematus and Euura with morphologically similar adults: 3rd abdominal segment with 6 annulets, annulets 2 and 4 with setae; no cerci; and maxillary stipes with or without seta. However, not all of these characters apply to all Pristiphora. For example, larvae of P. dasiphorae have only 3 visible annulets on abdominal segments and both species of the P. malaisei group (arctica group in Prous et al. 2014), P. malaisei and P. dasiphorae, have setae also on annulet 1 (Zinovjev 1993). More Pristiphora species should be scored for the larval characters mentioned by Vikberg (1982) to test the reliability of these characters for defining the genus.

Possible diploid males

Some of the nuclear sequences from males included polymorphic sites (double peaks in chromatograms) indicating heterozygosity or the presence of paralogous genes. Because these few polymorphic sites are restricted to synonymous positions (i.e. not affecting protein sequence) and so far there is no evidence for paralogous NaK and TPI genes at least in sawflies, these polymorphic sites might indicate heterozygosity because of diploidy. This can happen if there is no heterozygosity at the complementary sex determination (CSD) locus, preventing female development from a diploid embryo (Naito and Suzuki 1991, Heimpel and de Boer 2008, Harper et al. 2016). Survival of diploid males is known in sawflies (Naito and Suzuki 1991, Cook et al. 2013, Harper et al. 2016) and is common in nature at least in some species of Apocrita (Liebert et al. 2005, Retamal et al. 2016).

Figure 1. 

Maximum likelihood tree of Pristiphora and nematine outgroups based on three genes (3408 bp). Specimens having at least two of the three genes were included. Numbers above branches show bootstrap proportions (%). Support values for weakly supported branches (BP<70) are not shown. Part of Pristiphora is shown without support values and tip labels, which are shown in Fig. 2. The scale bar shows the number of estimated substitutions per nucleotide position.

Possible frameshift mutation in COI

COI sequence of one P. friesei specimen (DEI-GISHym11558) had a single bp deletion, which was caused by shortening of 8 bp repeat of thymines (T) to 7 thymines, compared to a different P. friesei specimen and closely related P. laricis. Because the P. friesei sequence with the deletion was of good quality (trace files at figshare http://dx.doi.org/10.6084/m9.figshare.5235832), was found in amplicons obtained with two sets of different primers, and did not show anomalous phylogenetic position, it is possible that this COI sequence is functional despite the frameshift mutation (Beckenbach et al. 2005, Rosengarten et al. 2008). Barcode sequences of two additional P. friesei specimens in BOLD that are labelled as NUMT (nuclear mitochondrial pseudogene), DEI-GISHym4993 and DEI-GISHym11557, have identical sequence to specimen DEI-GISHym11558.

Phylogeny of Pristiphora and definitions of some species groups

Phylogenetic relationships at the base of Pristiphora are not well resolved by our three-gene dataset (Fig. 1), because most of the deeper splits receive bootstrap support less than 70%. Many of the more derived clades that receive strong bootstrap support (more than 90%) are also morphologically well-supported (see group definitions below). Generally, our results strengthen the conclusions of Nyman et al. (2010) that only a minority of lineage splits correlate with host plant shifts. Species in the groups which monophyly is well supported (Figs 12) tend to share the same host plant genus or family (see group definitions below and Nyman et al. 2010 for details). We will not repeat the quantitive analysis of Nyman et al. (2010), but our broader taxon sampling yields additional support for their conclusions, e.g. we recover a Quercus-feeding clade composed of P. fausta, P. calliprina, and P. parnasia (Figs 2, 5), and an Acer-feeding depressa group (Figs 1, 3), although host plants are not known for all the species involved.

To enable easier discussion of relationships among Pristiphora species, we define the following informal species groups based on morphology and the phylogenetic analysis: abietina, alpestris, carinata, depressa, erichsonii, laricis, leucopodia, malaisei, micronematica, nigella, pallida, pallidiventris, retusa, ruficornis, and rufipes group. The names of the groups are based on the oldest valid species name in that group.

The following treated species were not assigned to any species groups: abbreviata, angulata, biscalis, bufo, cadma, cincta, condei, conjugata, dedeara sp. n., fausta, geniculata, insularis, maesta, mollis, monogyniae, paralella, pseudogeniculata, punctifrons, tenuiserra, and testacea.

Figure 2. 

Part of the maximum likelihood tree shown in Fig. 1. Numbers above branches show bootstrap proportions (%). Support values for weakly supported branches (BP<70) are not shown. Outline of the full tree is shown upper left, with the part shown here highlighted. The scale bar shows the number of estimated substitutions per nucleotide position.

Figure 3. 

Maximum likelihood tree of Pristiphora based on COI gene (1078 bp). Shortest sequence included was 462 bp (subarctica MHV00166). Numbers above branches show bootstrap proportions (%). Support values for weakly supported branches (BP<70) are not shown. BIN numbers (BOLD:XXXXXXX) referred to in the text are shown for representative specimens. Part of Pristiphora is shown without support values and tip labels, which are shown in Figs 45. The scale bar shows the number of estimated substitutions per nucleotide position.

P. abietina group (also known as the compressa complex)

Includes the following North-Western Palaearctic species: abietina, compressa, decipiens, gerula, pseudodecipiens, robusta, and saxesenii. All species feed on Picea (Beneš and Krístek 1979) and the females have a characteristically modified valvula 3 (Figs 116–118, 120–122), which is apparently correlated with oviposition in narrow conifer needles. Specifically, valvula 3 is relatively narrow in dorsal view, and in lateral view its dorsal and ventral margins are more or less parallel, with the posterior margin distinctly truncate (Figs 116–118, 120–122). In addition, the apical part of the abdomen of females is laterally compressed (Fig. 60), but this character is not reliable for unambiguous distinction of females of the abietina group from other species. Males are externally very similar to many other species outside the abietina group that are ventrally extensively pale and dorsally black, so that penis valves should be studied to recognise them. Penis valves of the abietina group have a similar structure (Figs 283–289) that can be distinguished from other Pristiphora: more or less straight ventral margin, similarly sized paravalva and pseudoceps with roughly convex dorsal margin, apically unmodified pseudoceps, and straight or slightly bent valvispina. Genetic data (no data yet for P. robusta) strongly support the monophyly of the abietina group (Figs 2, 5). We exclude the erichsonii and pallida groups from the abietina group (included in Wong 1975), because current phylogenetic analyses do not support such a clade (Figs 2, 5). Unusually, the divergence in mitochondrial sequences (maximum distance 2.2%, minimum between species distance 0.0%) in the abietina group is similar or lower than in nuclear sequences (maximum distance 2.7%, minimum between species distance 1.1%). This is a reversal of the typical condition in bilateral animals (Bilateria), in which the substitution rate in mitochondrial DNA is several-fold higher than in nuclear DNA (Ballard and Whitlock 2004, Lavrov 2007). It is unlikely that mitochondrial DNA (mtDNA) evolves more slowly than nuclear DNA in the abietina group (no reliable cases are known in Bilateria). More probable is that the reduced diversity of mtDNA is caused, for example, by infection with maternally transmitted Wolbachia (or some other Bacteria), a common occurrence in arthropods (e.g. Hurst and Jiggins 2005, Werren et al. 2008). Wolbachia infection can cause cytoplasmic incompatibility between infected males and uninfected females, meaning that mitochondria carried by infected females are preferentially spread within population or between closely related species via introgression. This could explain sharing of (near) identical COI sequences between different species in abietina group.

P. alpestris group

Includes the following North-Western Palaearctic species: alpestris, pseudocoactula, and possibly dissimilis. Penis valves and lancets are very similar in alpestris and pseudocoactula (Figs 203–204, 233–236), but females are not known for dissimilis and its penis valve is very distinctive compared to all other Pristiphora species (Fig. 237). Genetic data strongly supports monophyly of the alpestris and pseudocoactula clade (Figs 2, 5). Placement of dissimilis (no genetic data yet) in the alpestris group is tentative (Lindqvist 1971).

P. carinata group (previously Lygaeotus)

Includes the following North-Western Palaearctic species: albilabris, borea, breadalbanensis, carinata, coactula, groenblomi, lativentris, and trochanterica. Species of the carinata group have a distinctly matt mesepisternum, and completely or largely black body. Valvula 3 of females varies from simple (short, slightly tapering from base to apex, and without scopa; Figs 98–99) to somewhat square-shaped in dorsal view and with small scopa (Figs 93–97). Lancets are without setae and very similar to each other (Figs 210217). Penis valves are also hardly distinguishable (Figs 238–242). Because of the high degree of similarity and the apparently continuous variation of (nearly) all characters, it is not clear how many species should be recognised and how they should be delimited. Genetic data strongly supports monophyly of the carinata group (Figs 2, 5). More research is need to associate males and females of different species, preferably by rearing experiments and sequencing of different nuclear genes, so that morphological variation within and between the species can be assessed more reliably.

P. depressa group (previously subbifida group)

Includes the following North-Western Palaearctic species: depressa, subbifida, and tetrica. This species group was reviewed by Liston and Späth (2008) and Liston et al. (2013). As far as is known, all species feed on Acer (Liston and Späth 2008). Females have usually a relatively short and narrow valvula 3 with a small scopa (Figs 82–83), but sometimes the scopa is practically absent. Most species have bifid or subbifid (Figs 24–25) claws. Males have a long and thin valvispina that can be strongly bent (Fig. 301), or nearly straight (in P. ifranensis Lacourt, 1973). Combined analysis of mitocondrial and nuclear genes moderately supports monophyly of the depressa group (Fig. 1). Genetic data (Figs 1, 3) indicates several additional, possibly undescribed, species in Europe (Mediterranean region and Central Europe; DEI-GISHym20783, 20784) and in the Far East (DEI-GISHym86127, 80227, 80233).

P. erichsonii group (previously wesmaeli complex)

Includes the following North-Western Palaearctic species: erichsonii, glauca, and wesmaeli. All species feed on Larix (Wong 1975). Characteristically for conifer-feeders, species of the erichsonii group have a narrow valvula 3 (Figs 124, 126). P. erichsonii has a different coloration (abdomen with a red band) and valvula 3 (not apically abruptly constricted; Fig. 124) from P. glauca and P. wesmaeli (abdomen ventrally yellow, valvula 3 apically abruptly constricted Fig. 126), but penis valves (Figs 290–292) and genetic data (Figs 2, 5) support monophyly of this group. The penis valves of the erichsonii group have a somewhat rhombus- (Fig. 290) or trapez-shaped (Figs 291–292) paravalva, a rather rectangular and unmodified pseudoceps, and small and slightly bent valvispina that is asymmetrical at apex.

P. laricis group (previously Oligonematus)

Includes the following North-Western Palaearctic species: friesei and laricis. Both species feed on Larix (Adam 1973, Huflejt and Sawoniewicz 1999, Liston et al. 2006). Females have a short valvula 3 (cerci clearly extend beyond its apex), with small scopa (Figs 79–81). The absence of a velum on the anterior protibial spur (Fig. 33), and usually mostly black body, can help in recognising this group. Lancets (Figs 174–178) and penis valves (Figs 277, 279208) unambiguously distinguish the laricis group from other species (see the Key). Genetic data strongly supports monophyly of the laricis group (Figs 1, 3).

P. leucopodia group

Includes the following North-Western Palaearctic species: leucopodia and nigriceps. Both species feed on Picea (Beneš and Krístek 1979). As in many other species feeding on conifers, valvula 3 is narrow in the leucopodia group (Fig. 84). In lateral view, valvula 3 has a round outgrowth at the posterior margin (Fig. 85), which distinguishes the leucopodia group from other conifer-feeders. A relatively similar valvula 3 is found in P. insularis (Figs 86–87), but other characters distinguish this species from the leucopodia group (anterior protibial spur with velum and clearly different lancet; see Figs 33–34, 135, 181–183). A characteristic hump that is posteriorly constricted and situated at the base of the slightly bent valvispina (Figs 281–282) distinguish penis valves of the leucopodia group from other Pristiphora (see the Key). Combined mitochondrial and nuclear phylogenetic analysis strongly supports monophyly of the leucopodia group (Fig. 1), but mitochondrial COI sequences alone do not (Fig. 3).

P. malaisei group (previously arctica group, or Pristicampus)

Includes the following North-Western Palaearctic species: malaisei and dasiphorae. Both species feed on Potentilla fruticosa L. (=Dasiphora fruticosa) (Zinovjev 1993) and possibly Comarum palustre L. Quite characteristic for the group is its head shape, which is more evident in females (Fig. 58). Structure of the penis valves (apically truncate and without valvispina; Figs 297–298) is very distinct within Pristiphora, prompting Zinovjev (1993) to create even a separate tribe for the species of malaisei group. Genetic data strongly supports monophyly of the group (Fig. 1).

P. micronematica group (previously Lygaeophora)

Includes the following North-Western Palaearctic species: affinis, atripes, kontuniemii, micronematica, nordmani, reuteri, sermola, and possibly lanifica. In contrast to other group names, we use the name micronematica (Malaise 1931) for this group instead of the oldest name lanifica (Zaddach in Brischke 1883), because the identity of lanifica is uncertain and it might not belong to this group. As far as is known, species of this group feed on Salix (Lindqvist 1952, Kontuniemi 1960, 1972, Vikberg 1966, Liston 1982, Kangas 1985). Valvula 3 in females has a small scopa (Fig. 92), is more or less square-shaped in dorsal view and has two dense, lateral bundles of hairs at its apex (Fig. 91). Very similar are species in the carinata and alpestris groups, but lancets (with setae and serrulae not papilliform; Figs 205209) and penis valves (pseudoceps in most species dorsally with loose membranous region covered with hair; Figs 303–310) of the micronematica group can be readily separated from these groups (see the Key). Genetic data strongly supports monophyly of the micronematica group (Figs 2, 5). Ecologically, and in some other biological characteristics, this group is similar to the phylogenetically distant aphantoneura subgroup (Figs 2, 4; Prous et al. 2016): the majority of species (perhaps all in the micronematica group) feed on Salix, are more abundant and species rich in boreal or (sub)arctic habitats (probably because of the host plants), have low diversity in mitochondrial DNA (maximum divergence in COI barcodes is 2.5% in micronematica group, 3.3% in aphantoneura subgroup) that does not correlate with species boundaries (tentative for micronematica group), and have phenotypically exceedingly similar females, while males can in most cases be identified by clear differences in penis valves. At least ex ovo rearings or sequencing of several nuclear markers from most species is needed to associate males and females confidently in the micronematica group.

P. nigella group (previously Sharliphora)

Includes the following North-Western Palaearctic species: amphibola, nigella, and parva. All species feed on Picea (Beneš et al. 1981). The structure of valvula 3 in the nigella group is unique among Pristiphora, with the scopa positioned dorsoapically (Figs 111, 115), rather than just apically or ventroapically as in other species. This might be related to adaptation to ovipositing into closed or opening Picea buds (Nägeli 1936, Benson 1948, Grebenshchikova 1986), in contrast to other conifer-feeders which use already expanded needles. Lancets (long and narrow, with setae and flat serrulae; Figs 165–167) and penis valves (apically narrowed pseudoceps, paravalva dorsally somewhat s-shaped and with small dorsally directed valvispina; Figs 293–295) of the nigella group can also be distinguished from other species. Genetic data strongly supports monophyly of the nigella group (Figs 2, 5).

Figure 4. 

Part of the maximum likelihood tree shown in Fig. 3. Numbers above branches show bootstrap proportions (%). Support values for weakly supported branches (BP<70) are not shown. BIN numbers (BOLD:XXXXXXX) referred to in the text are shown for representative specimens. Outline of the full tree is shown upper left, with the part shown here highlighted. The scale bar shows the number of estimated substitutions per nucleotide position.

Figure 5. 

Part of the maximum likelihood tree shown in Fig. 3. Numbers above branches show bootstrap proportions (%). Support values for weakly supported branches (BP<70) are not shown. BIN numbers (BOLD:XXXXXXX) referred to in the text are shown for representative specimens. Outline of the full tree is shown upper left, with the part shown here highlighted. The scale bar shows the number of estimated substitutions per nucleotide position.

P. pallida group

Includes the following North-Western Palaearctic species: pallida and subarctica. Both species feed on Picea (Beneš and Krístek 1979). As in many other conifer-feeders, species of the pallida group have a narrow valvula 3 (as in Fig. 124) and both males and females are at least ventrally extensively pale yellow. The structure of valvula 3 cannot be distinguished from P. erichsonii, but this species has a different coloration (abdomen with a red band; as in Fig. 47). Very characteristic penis valves (Figs 276, 278) clearly distinguish this group from other similar conifer-feeders (see the Key). We cannot exclude the possible conspecifity of pallida and subarctica, because the differences between them are morphologically (Figs 171–172, 276, 278) and genetically (Fig. 5) small. Examination of more specimens and additional nuclear gene sequences are needed to decide this.

P. pallidiventris group

Includes the following North-Western Palaearctic species: nigricans and pallidiventris. Both species feed on herbaceous Rosaceae (Liston 2011, this study). Usually, in both species the abdomen is at least ventrally pale, but nearly completely black specimens of pallidiventris can occasionally be found. The structure of valvula 3 (deep scopa with long medial projection; Fig. 106) distinguishes this group from other species of Western Palaearctic Pristiphora, though the difference is small compared to many other species with similar valvula 3. When in doubt, lancets should be studied (Figs 141–144). Penis valves of both species are very similar (Fig. 223–226) and can be distinguished from other species: paravalva and pseudoceps are of similar size and shape, ventral margin of paravalva is more or less straight, and the valvispina is straight, small, and positioned in the middle or upper third of paravalva. Current genetic data (Fig. 5) does not allow separation of P. nigricans (only one COI sequence available) and P. pallidiventris.

P. retusa group

Includes the following North-Western Palaearctic species: exigua and retusa. A host plant is known only for P. retusa (Prunus padus; Kangas 1985). Characteristic for the group is small size (body length 3.0–4.5 mm), mostly black body, and distinctly asymmetrical labrum (Fig. 11). These two species appear to be rather closely related (only one COI sequence available for P. exigua) (Fig. 3).

P. ruficornis group

Includes the following North-Western Palaearctic species divided into five subgroups (Prous et al. 2016): albitibia subgroup (albitibia, astragali, sootryeni, caraganae), aphantoneura subgroup (aphantoneura, bifida, confusa, luteipes, opaca, pusilla, staudingeri, subopaca), appendiculata subgroup (appendiculata), armata subgroup (armata, leucopus), and ruficornis subgroup (melanocarpa, ruficornis, and possibly frigida). Species of the albitibia subgroup feed on Fabaceae, the aphantoneura subgroup mostly on Salix (except P. aphantoneura that feeds on Lathyrus), P. appendiculata on Ribes, the armata subgroup on Crataegus and Tilia, and the ruficornis subgroup on Betula (a host is not known for P. frigida). Morphologically, the ruficornis group is best characterised by the structure of penis valves, which have a large and usually strongly bent valvispina that almost completely (Figs 263274) or largely (Fig. 262) replaces the paravalva. Several other species have a similarly large and bent valvispina (Figs 276, 278, 299, 301), but these penis valves can nevertheless be clearly distinguished from those of the ruficornis group, in combination with differences in body coloration (see the Key). Genetic data strongly supports monophyly of the ruficornis group (Figs 2, 4).

P. rufipes group (also known as thalictri or aquilegiae group)

Includes the following North-Western Palaearctic species: brevis, dochmocera, rufipes, thalictri, and thalictrivora. The species feed on Aquilegia (P. rufipes) or Thalictrum (P. brevis, P. thalictri, P. thalictrivora, and possibly P. dochmocera). Species of this group are usually black-bodied, valvula 3 of females has a distinct scopa, and lancets are without setae. Lancets and penis valves are very similar in all species. Penis valves of the rufipes group have a somewhat rectangular and unmodified pseudoceps, a somewhat oval-, rectangular- or square-shaped paravalva that is dorsoapically abruptly narrowed before the valvispina, a small and straight valvispina that arises on the ventral part of the paravalva, and a valvar strut that is distinct along its entire length (Figs 243, 245, 247–251). Very similar penis valves (Figs 244, 246), but somewhat more elongate than in the rufipes group, are found in the closely related P. cincta (Figs 2, 5). Species of the rufipes group are also genetically closely related to each other (Figs 2, 5). Species boundaries in the rufipes group are unclear, as well as the association of males and females. At least two species seem to be involved, one feeding on Aquilegia (P. rufipes) and the other one(s) on Thalictrum. The lancet of the Aquilegia-feeding P. rufipes seems to be slightly different from the other species. The most protruding part of the serrulae of rufipes tends to be relatively acute, usually with denticles along the entire margin (Fig. 222), while the most prominent part of the serrulae of the other species tends to be relatively blunt, with denticles concentrated mostly on the apical part of the serrula. Based on mitochondrial and nuclear sequences of larvae collected from Aquilegia sp. (DEI-GISHym20983 and DEI-GISHym21482), we were able to associate three additional adults (DEI-GISHym15263, DEI-GISHym19795, and 9t from VoSeq database) with these larvae. Nuclear sequences from these adults (both genes, only NaK, or only TPI) and the larvae (both genes) form a monophyletic group to the exclusion of other species (not shown). Nuclear data suggests that there might be at least three additional lineages, which we have identified as P. brevis, P. thalictri, and P. thalictrivora (Fig. 2), but morphological separation of these forms is somewhat arbitrary. Although there is some variation in the shape of serrulae of lancets among the Thalictrum-feeders (Figs 218–221), this does not seem to correlate with external morphology, and the variation is continuous. Specimens with completely black (P. thalictri) or more or less completely pale legs (e.g. P. thalictrivora) can both have lancets of the two extreme forms (Figs 218, 221), but apparently also of intermediate form (Figs 219–220). Nevertheless, in addition to P. rufipes, we tentatively recognise four other species: brevis, dochmocera, thalictri, and thalictrivora. Pristiphora thalictri is the darkest, with completely or almost completely black body, including the legs and tegulae. Pristiphora brevis and P. thalictrivora have completely or nearly completely pale legs (at most with black hind tarsi) and tegulae. The only differences between P. brevis and P. thalictrivora may be the longer postocellar area (Fig. 7) and (usually?) rather smooth mesopostnotum (Fig. 19) in P. brevis (matt in P. thalictrivora; Fig. 20). Pristiphora dochmocera, which we tentatively keep as a separate species, might be a synonym of P. thalictri or P. thalictrivora, because its coloration is intermediate between them. Pristiphora dochmocera has black tegulae and pale (metafemur and metatibia nearly completely pale) or relatively dark legs (black metafemur and pale metatibia). Although specimens of the rufipes group usually have a completely black abdomen, we have studied numerous specimens from southern Ukraine, Armenia, and Russian Caucasus that have a mostly black to completely yellow abdomen (even the thorax is partly yellow in some specimens). Because length of postocellar area, sculpture of mesopostnotum, and coloration of tegula, pronotum, and labrum varies among these specimens, it is not clear if they can be associated with any of the species from Northern Europe (lancets are not different from those on Figs 219–220). Recently, Macek (2016) reared three species of this group: P. rufipes from Aquilegia and two species from Thalictrum. The species identified by Macek (2016) as P. sareptana Kuznetzov-Ugamskij, 1924 could actually be P. brevis or P. thalictrivora (if these are different species) based on the pictures given for adults (we have seen the types of P. sareptana and P. similis Kuznetzov-Ugamskij, 1924, which are most similar to P. brevis and could indeed be synonyms of this species). Additional studies are needed to resolve species boundaries (if there are any) of the Thalictrum-feeding complex, and to associate males with females.

Comment on species delimitation

If a well defined morphospecies corresponds to exclusive genetic clusters (not mixed with other morphospecies) based on more than one unlinked genetic marker (independently segregating gene regions that reside on different chromosomes or far from each other on the same chromosome) and individuals of that morphospecies have different relationships based on those different markers, then we are probably dealing with a single and well separated species. Unfortunately, such extensive genetic data (many markers and individuals) are not available for most species (due to lack of suitable material or/and high costs associated with sequencing). Nevertheless, it is possible to get some idea about species boundaries based on much smaller amounts of genetic data. Based on our results (see Discussion), within species divergence of mitochondrial genes seems to remain within 5% and based on nuclear genes within 1%. Although divergence higher than 2% in COI barcode sequences is commonly regarded as good evidence for different species (Ratnasingham and Hebert 2013), our data suggest that it is not unusual for within species divergence to be more than 3% or 4%. An example of this is P. albitibia, which can have COI barcode divergence of 4.6%, although neither morphology nor nuclear genes indicate that more than one species is involved. Therefore we do not consider COI divergence of less than 5% as good evidence for the existence of different species if there is no support from morphological and / or nuclear data.

On the other hand, where we have consistent morphological (based on adults or larvae) or ecological (different hosts) evidence without clear genetic evidence (even if based on all three genes we sampled) for the existence of different species within groups of closely related species, we have not synonymised these species. In our view, the genetic evidence we have so far is not sufficient (too few specimens and gene regions sampled) to decide this.

Assessment of morphological characters of the adults

For females, the shape of valvula 3 (apical sawsheath) is an important and relatively stable character for identification, but nevertheless it was necessary to key many species more than once, because of variability or intermediate character states. Still, it might be difficult to key out some species or individiuals, especially when valvula 3 is distorted (which usually happens with specimens dried from alcohol). In these cases lancets should be examined, although they can sometimes be very similar, even among distantly related species.

Males in Pristiphora usually lack good external non-colour characters, and therefore penis valves should be studied in most cases, which are stable within species and often show good differences between the species.

Coloration is the easiest character to observe, but unfortunately it often varies quite a lot within species. The abdomen can be completely pale or (almost) completely black in some species. Thorax coloration varies somewhat less, but can occasionally still vary from completely black to extensively pale. Head coloration is the most stable, in most species being black or with small pale spots. In a minority of species the head tends to be extensively pale around the eyes. Leg coloration can also vary extensively within species, but can often still be used for identification (see the Key). Coloration of the pterostigma is often a useful character for species identification, but it can be problematic in older pinned specimens. In species that normally have a dark pterostigma, the dark coloration can fade, causing the pterostigma to appear pale.

Unreliable characters. We have not used the length of the inner spur of the metatibia (in relation to the length of metatarsus for example) in species identification (e.g. Benson 1958; Zhelochovtsev and Zinovjev 1988, Haris 2006b) because the differences between the species (if there are any) are too small to be easily applied. Shape of the posterior margin of sternum 9 can be quite misleading. Several examined specimens of different species have a distinct notch, compared to other conspecific specimens with the margin entire, but because a notch is very sporadically present and does not correlate with other characters (which might indicate the existence of additional species), it seems to have no value in species identification and was therefore excluded from the key. However, in one exceptional case, the notch is large and deep and even has an outgrowth at the bottom of the notch (Fig. 52). Because it is a single specimen (based on penis valves, it is P. biscalis), it is not clear if this too is an aberration or there is an additional species involved. Vein 2r-rs of the forewing (see Fig. 1 in Prous et al. 2014) is another character that sporadically appears in some specimens, but without any phylogenetic correlation (we have observed it in P. malaisei, P. robusta [http://id.luomus.fi/GL.5198], and P. staudingeri), making it difficult to even recognise these specimens as Pristiphora.

Dichotomous key to Pristiphora adults

The key for females relies heavily on sawsheaths and ovipositors, and for males on penis valves, as these are most reliable for species identification. For females it is advisable to dissect the saw of one or more specimens of a series and for males it is in most cases necessary to pull out the genital capsule to see at least the tip of the penis valve. Fresh and clean specimens greatly help in species identification: particularly in females, where the shape of sawsheath can be of critical importance. Generally we recommend using the electronic key, which can be significantly faster (when using ‘Best’ and ‘Next Best’ options), easier to use, and more reliable because of the possibility to check more characters.

Females

Females of P. dissimilis are unknown.

1 a Head dorsally extensively pale (Fig. 8) 2
aa Head dorsally black or with small brown or pale spots (Figs 6–7) 21
2(1) a In dorsal view, valvula 3 with distinct scopa (e.g. Figs 104–107, 110–111, 115–116) 3
aa In dorsal view, valvula 3 without distinct scopa (e.g. Figs 82–83, 93–99, 108–110, 112–114, 116, 120–122, 124, 126–127) 11
3(2) a Scopa partly positioned dorsally (Figs 111, 115)
b Claws without or with small subapical tooth (Figs 21–22) 23
aa Scopa positioned posteriorly or posteroventrally (e.g. Figs 104–107, 110, 116)
bb Claws with large subapical tooth (Figs 23–24) or if as 3b, consider other characters 4
4(3) a In lateral view, posterior margin of valvula 3 truncate (Fig. 117) 5
aa In lateral view, posterior margin of valvula 3 round (Fig. 109) 7
5(4,14) a Hypopygium posteriorly excised (Fig. 63) P. decipiens
aa Hypopygium posteriorly not excised (Fig. 62) 6
6(5) a Mesopleuron (mesepisternum + mesepimeron) extensively pale (Fig. 28)
b Ventral margin of lancet not curved upwards (Fig. 163)
c Body length 6.0–9.0 mm P. pseudodecipiens
aa Mesopleuron black (Fig. 27) or slightly pale
bb Ventral margin of lancet apically strongly curved upwards (Fig. 157)
cc Body length 4.0–6.5 mm P. abietina
7(4) a Metatibia black (Fig. 29)
b Claws with large subapical tooth or bifid (Fig. 24)
c Valvula 3 with deep scopa (Fig. 107) P. fausta
aa Metatibia at least with basal 2/3 pale (Fig. 47)
bb Claws with large or small subapical tooth (Figs 22–24)
cc Valvula 3 with shallow scopa (e.g. Figs 104–106, 110) 8
8(7) a Mesepisternum completely or nearly completely pale (Fig. 336)
b Thorax dorsally extensively pale to completely black (Figs 35–36, 51)
c Claws with large subapical tooth (Fig. 335)
d Lancet with numerous setae and apical serrulae protruding (Figs 331, 333) P. cadma
aa-dd Combination of characters not as in a–d 9
9(8) a Scopa of valvula 3 shallow (Figs 108, 110)
b Claws with small subapical tooth (Fig. 22)
c Lancet with few setae and apical serrulae flat (Fig. 168) P. bufo
aa Scopa of valvula 3 not shallow (104–105)
bb Claws usually with large subapical tooth (Figs 23–24)
cc Lancet without or with numerous setae, and apical serrulae more protruding (Figs 147–148, 153–155) 10
10(9,37) a Pterostigma bicoloured (Fig. 43)
b Mesepisternum black or partly pale (Figs 27–28)
c Abdomen dorsally black or pale (Figs 49–50)
d Lancet with numerous setae (Figs 153–155) P. conjugata
aa Pterostigma unicoloured (Fig. 42)
bb Mesepisternum black (Fig. 27)
cc Abdomen nearly always completely pale (Fig. 50)
dd Lancet without setae (Figs 147–148) P. testacea
11(2) a Claws bifid or subbifid (Figs 24–25)
b Abdomen completely pale (Figs 28, 50) P. subbifida
aa Claws with small subapical tooth (Fig. 22)
bb Abdomen dorsally at least partly black (Figs 49, 51) 12
12(11) a In lateral view, posterior margin of valvula 3 distinctly truncate and about as long as ventral margin (Fig. 118), and in dorsal view, slightly tapering from base to apex (Fig. 120) 13
aa Valvula 3 with different combination of characters (e.g. Figs 94, 98, 108–110, 112–114, 116–117, 124–127) 14
13(12) a In posterior view, valvula 3 without scopa (Fig. 122)
b Pterostigma usually pale (Fig. 39) P. gerula
aa In posterior view, valvula 3 with small scopa (Fig. 121)
bb Pterostigma usually dark (Fig. 42) P. saxesenii
14(12) a In dorsal view, valvula 3 elongate, not tapering from base to apex, and with distinct scopa (Fig. 116)
b In lateral view, valvula 3 distinctly truncate (Fig. 117) 5
aa In dorsal view, valvula 3 with different combination of characters (Figs 93, 95, 97, 99, 108, 112, 124, 126)
bb In lateral view, valvula 3 round or indistinctly truncate (Figs 109, 113, 125) 15
15(14) a In dorsal view, valvula 3 abruptly constricted at apex, but not tapering completely (Fig. 126)
b In lateral view, valvula 3 indistinctly truncate (Fig. 113) 16
aa In dorsal view, valvula 3 not abruptly constricted (Figs 93, 97, 99, 108, 112, 124) at apex, or tapering completely (Fig. 95)
bb In lateral view, valvula 3 round or indistinctly truncate (Figs 109, 113, 125) 17
16(15) a Ovipositor (valvula 3 + valvifer 2, Figs 61–62) about 0.9 times as long as protibia (Benson, 1958)
b Distance between cenchri more than one and half times the breadth of one of them (Benson, 1958) P. wesmaeli
aa Ovipositor about 1.1 times as long as protibia (Benson, 1958)
bb Distance between cenchri less than one and half times the breadth of one of them (Benson, 1958) P. glauca
17(15) a In dorsal view, valvula 3 narrow (Fig. 124)
b In lateral view, valvula 3 indistinctly truncate (Fig. 125)
c Mesepisternum smooth (Fig. 44)
d Abdomen sometimes dorsally extensively pale (Fig. 50) 18
aa-dd Combination of characters not as in a–d 19
18(17) a Gap between basal serrulae small, and basal annuli with more numerous setae (Fig. 171)
b Abdomen (usually?) dorsally extensively pale (Fig. 50) P. pallida
aa Gap between basal serrulae large, and basal annuli with fewer setae (Fig. 172)
bb Abdomen (usually?) dorsally extensively black (Fig. 49) P. subarctica
19(17) a Mesepisternum matt and black (Fig. 46) few specimens of carinata group
aa Mesepisternum smooth and partly pale (Figs 28, 44) 20
20(19) a In lateral view, valvula 3 not truncate (Fig. 109)
b Anterior protibial spur with velum (Fig. 34) P. bufo
aa In lateral view, valvula 3 indistinctly truncate (Fig. 113)
bb Anterior protibial spur without velum, but with hairs (Fig. 33) P. paralella
21(1) a In dorsal view, valvula 3 with distinct scopa (Figs 75, 78, 86, 88, 100, 104, 106–107, 111, 116) 22
aa In dorsal view, valvula 3 without distinct scopa (Figs 67, 69, 72, 74–75, 77–79, 82, 84, 86, 88–89, 91, 93, 95, 97, 99, 101, 103, 116, 119, 124) 63
22(21) a Scopa partly positioned dorsally (Figs 111, 115)
b Claws without, or with small subapical tooth (Figs 21–22)
c Body length 3.0–4.5 mm 23
aa Scopa positioned posteriorly or posteroventrally (Figs 75, 78, 86, 88, 100, 104–107, 116)
bb Claws with large subapical tooth (Figs 23–25) or if as 22b consider other characters
cc Body length 3.0–11.0 mm 25
23(22,3) a Sawsheath (valvula 3 + valvifer 2; Figs 61–62) about 1.11 times longer than metatibia (Beneš et al. 1981), in dorsal view elongate
b Head dorsally black (Fig. 6)
c Posterior margin of pronotum black or narrowly pale (Fig. 35) P. amphibola
aa Sawsheath about 0.93–1.03 times longer than metatibia (Beneš et al. 1981), in dorsal view short or elongate
bb Head dorsally black to extensively pale (Figs 6–8)
cc Posterior margin of pronotum black to extensively pale (Fig. 35–36) 24
24(23) a Sawsheath about 0.93–0.97 times longer than metatibia (Beneš et al. 1981), in dorsal view short
b Tegula black or slightly pale (Fig. 35) P. nigella
aa Sawsheath about 1.00–1.03 times longer than metatibia (Beneš et al. 1981), in dorsal view elongate
bb Tegula extensively pale (Figs 36–37) P. parva
25(22) a In lateral view, posterior margin of valvula 3 truncate (Fig. 117) 26
aa In lateral view, posterior margin of valvula 3 round (Figs 85, 87, 109) 29
26(25) a Head, thorax, and abdomen black (Fig. 26)
b Hypopygium posteriorly not excised (Fig. 62)
c Body length 9–11 mm P. robusta
aa Head, thorax, and abdomen ventrally at least partly pale (Figs 27–28)
bb Hypopygium posteriorly excised (Fig. 63) or if as 26b, consider other characters
cc Body length 4–9 mm 27
27(26) a Hypopygium posteriorly excised (Fig. 63)
b Ventral margin of lancet not strongly curved upwards (Figs 158–160) 28
aa Hypopygium posteriorly not excised (Fig. 62)
bb Ventral margin of lancet apically strongly curved upwards (Fig. 157) P. abietina
28(27) a Mesopleuron usually extensively pale (Fig. 28)
b Ventral margin of lancet and its annuli more distinctly curved upwards (Fig. 160) P. decipiens
aa Mesopleuron usually black or slightly pale (Fig. 27)
bb Ventral margin of lancet and its annuli less distinctly curved upwards (Figs 158–159) P. compressa
29(25) a In lateral view, valvula 3 with round outgrowth at posterior margin (Figs 85, 87)
b In dorsal view elongate and with distinct or indistinct scopa (Figs 84, 86) 30
aa-bb Combination of characters not as in a–b 32
30(29,67) a In lateral view, valvula 3 more abruptly narrowed at apex (Fig. 87)
b In dorsal view, valvula 3 broader (Fig. 86)
c Anterior protibial spur with velum (Fig. 34) P. insularis
aa In lateral view, valvula 3 less abruptly narrowed at apex (Fig. 85)
bb In dorsal view, valvula 3 narrower (Fig. 84)
cc Anterior protibial spur without velum, but with hairs (Fig. 33) 31
31(30) a Thorax and abdomen extensively pale (Figs 28, 37, 50–51) P. nigriceps
aa Thorax and abdomen mostly black (Fig. 26), only sometimes abdomen nearly completely pale (Figs 49–50) and thorax slightly pale (Fig. 36) P. leucopodia
32(29) a Abdomen with red band (rarely in dorsal aspect entirely reduced), terga 1 and 9–10 black (Figs 47–48)
b Scopa of valvula 3 with short medial projection (Figs 100, 104) 33
aa-bb Abdomen without red band (Figs 26–28, 49–51) or scopa of valvula 3 with long medial projection (Fig. 106) 34
33(32) a Antenna ventrally partly paler than dorsally (Fig. 15)
b Supraclypeal area pale (Fig. 12)
c Postocellar area 1.5–2.5 times longer than diameter of lateral ocellus (Fig. 7) P. condei
aa Antenna usually uniformly black (Fig. 14)
bb Supraclypeal area usually black (Figs 9–10)
cc Postocellar area 1.0–1.5 times longer than diameter of lateral ocellus (Fig. 6) P. cincta
34(32) a Valvula 3 with deep scopa and without long medial projection (Fig. 107)
b Claws without, or with minute subapical tooth (Figs 21–22)
c Abdomen usually ventrally black or partly pale (Fig. 27)
d Terga 9–10 usually extensively pale (Fig. 62) P. punctifrons
aa Valvula 3 with shallow scopa or with long medial projection (Figs 75, 78, 100, 104, 106)
bb Claws with large subapical tooth (Figs 23–25) or if as 34b, consider other characters
cc Abdomen ventrally completely pale (Fig. 28) or if as 34c, consider other characters
dd Terga 9–10 black (Fig. 61) or if as 34d, consider other characters 35
35(34) a Height of eye in lateral view often about 2–3 times as long as distance from dorsal margin of eye to dorsalmost point of head (Fig. 58)
b Clypeus emarginate (Fig. 13)
c Claws with small subapical tooth (Fig. 22)
d Abdomen usually ventrally black or partly pale (Fig. 27)
e Metatibia and often metatarsus nearly completely pale (Fig. 30) P. malaisei
aa Height of eye in lateral view about 3–4 times as long as distance from dorsal margin of eye to dorsalmost point of head (Fig. 59)
bb Clypeus truncate or slightly emarginate (Figs 9–10, 12)
cc Claws without, or with small or large subapical tooth (Figs 21–25)
dd Abdomen ventrally completely pale (Fig. 28) or if as 35d, consider other characters
ee Metatibia and metatarsus black (Fig. 29) or if as 35e, consider other characters 36
36(35) a Abdomen ventrally completely or nearly completely pale (Fig. 28) 37
aa Abdomen ventrally black or partly pale (Figs 26–27) 38
37(36) a Scopa of valvula 3 with long lateral lobes and long medial projection (Fig. 106)
b Claws usually with small subapical tooth (Fig. 22) 39
aa Scopa of valvula 3 with short lateral lobes and / or short medial projection (100, 104)
bb Claws usually with large subapical tooth (Figs 23–24) 10
38(36) a Scopa of valvula 3 with long lateral lobes and long medial projection (Fig. 106)
b Postocellar area 1.0–1.5 times longer than diameter of lateral ocellus (Fig. 6)
c Claws usually with small subapical tooth (Fig. 22) 39
aa Scopa of valvula 3 with short lateral lobes and / or short medial projection (Figs 75, 78, 100, 104)
bb Postocellar area 1.0–2.5 times longer than diameter of lateral ocellus (Figs 6–7)
cc Claws without, or with small or large subapical tooth (Figs 21–25) 40
39(37,38) a Metafemur (Fig. 28) and terga 9 and 10 (Fig. 62) completely yellow or at least metafemur apically slightly black
b Pterostigma dark (Fig. 42)
c Posterovenral tip of serrulae broader and gap between serrulae smaller (Figs 142–144) P. pallidiventris
aa Metafemur completely yellow (Fig. 28) and terga 9 and 10 black (Fig. 61)
bb Pterostigma usually pale (Fig. 39)
cc Posterovenral tip of serrulae narrower and gap between serrulae larger (Fig. 141) P. nigricans
40(38) a Abdomen laterally and ventrally with pale spots (Fig. 27) 41
aa Abdomen laterally and ventrally black (Fig. 26) 42
41(40) a Clypeus at least partly pale (Figs 10, 12)
b Terga 9–10 pale (Fig. 62), often also valvula 3 pale (Fig. 65)
c Metafemur pale (Fig. 28)
d Serrulae papilliform (Fig. 203) P. alpestris
aa Clypeus black (Fig. 9)
bb Tergum 10 at least partly black (Fig. 61), valvula 3 completely black (Figs 61, 64)
cc Metafemur extensively black (Figs 26–27)
dd Serrulae flat (Fig. 130) P. maesta
42(40) a Postocellar area 2.0–2.5 times longer than diameter of lateral ocellus (Fig. 7)
b Claws with large subapical tooth (Figs 23–24) 43
aa-bb Combination of characters not as in a–b 44
43(42) a Flagellum about 2.4–2.5 times as long as width of head
b Metatarsomere 1 at least in apical 2/3 black (Fig. 29)
c Valvifer 2 pale (Fig. 62)
d Lancet with few setae and ventral margin of serrulae straight (Fig. 150) P. geniculata
aa Flagellum about 1.9–2.0 times as long as width of head
bb Metatarsomere 1 completely or nearly completely pale (Fig. 31)
cc Valvifer 2 black (Fig. 61)
dd Lancet with numerous setae and ventral margin of serrulae slightly concave (Fig. 151) P. pseudogeniculata
44(42) a Mesopostnotum smooth (Fig. 19)
b Claws without subapical tooth (Fig. 21)
c Mesepisternum smooth (Fig. 44)
d Antenna usually ventrally paler than dorsally (Fig. 15) P. appendiculata
aa-dd Combination of characters not as in a–d 45
45(44) a Postocellar area 1.5–2.5 times longer than diameter of lateral ocellus (Fig. 7)
b Mesepisternum smooth (Fig. 44)
c Lancet without setae (Figs 218–222) rufipes group
aa Postocellar area 1.0–1.5 times longer than diameter of lateral ocellus (Fig. 6)
bb Mesepisternum matt (Figs 45–46) or if as 45b, consider other characters
cc Lancet with setae (Figs 188202) or if as 45c, consider other characters 46
46(45) a Metafemur mainly pale (Figs 27–28) 47
aa Metafemur mainly black (Fig. 26) 49
47(46) a Claws with large subapical tooth (Figs 23–24)
b Antenna ventrally paler than dorsally (Fig. 15)
c Metafemur whitish (Fig. 22 in Prous et al. 2016) P. leucopus
aa Claws with small subapical tooth (Fig. 22)
bb Antenna uniformly black (Fig. 14)
cc Metafemur yellowish (Fig. 28) 48
48(47) a Lancet without setae (Figs 218–222) rufipes group
aa Lancet with numerous setae (Fig. 195) P. aphantoneura (on Lathyrus) and P. luteipes (on Salix) (see Vikberg 2006 for minor characters for separating these species)
49(46) a Claws with long subapical tooth close to apical one (bifid) (Fig. 25) 50
aa Claws with small or large subapical tooth clearly separated from apical one (Figs 22–24) 51
50(49) a Hind trochanters, trochantelli, and tibiae partly pale
b Antenna (usually?) ventrally at least slightly paler than dorsally (Fig. 15)
c Apical serrulae of lancet short and protruding, and tangium long and narrow (Fig. 199) P. bifida
aa Hind trochanters, trochantelli, and usually tibiae uniformly black or brown
bb Antenna uniformly black (Fig. 14)
cc Apical serrulae of lancet long and flat, and tangium short and broad (Fig. 200) P. frigida
51(49) a Tangium of lancet with distinct lobe (Fig. 191)
b Mesepisternum smooth (Fig. 44)
c Claws with small subapical tooth (rarely with large) (Fig. 22) 52
aa Tangium of lancet without distinct lobe (Figs 188–190, 192–194, 196–198, 201–202, 218–222)
bb Mesepisternum matt (Figs 45–46) or if as 51b, consider other characters
cc Claws with large subapical tooth (Figs 23–24) or if as 51c, consider other characters 53
52(51) a Antenna ventrally distinctly paler than dorsally (Fig. 15) P. ruficornis
aa Antenna usually uniformly black (Fig. 14), but sometimes ventrally slightly paler than dorsally P. melanocarpa
53(51) a Lancet without setae (Figs 218–222)
b Mesepisternum smooth (Fig. 44)
c Claws with small subapical tooth (Fig. 22) rufipes group
aa Lancet with numerous or few setae (Figs 188–190, 192–194, 196–198, 201–202)
bb Mesepisternum matt (Figs 45–46) or if as 53b, consider other characters
cc Claws with large subapical tooth (Figs 23–24) or if as 53c, consider other characters 54
54(53) a Inner surface of lancet with small spiny pectines (or dentes semicirculares) that reach sclerora (Figs 188–190, 194) 55
aa Inner surface of lancet without small spiny pectines (Figs 192–193, 196–198, 201–202) 58
55(54) a Pterostigma basally dark brown and apically brown (Fig. 40)
b Mesepisternum smooth (Fig. 44)
c Lancet with numerous setae (Fig. 188)
d Apical serrulae of lancet short (Figs 188, 194) 56
aa Pterostigma uniformly yellow or brown (Fig. 39)
bb Mesepisternum usually at least slightly matt (Figs 45–46)
cc Lancet with few setae (Fig. 189) or if as 55c, consider other characters
dd Apical serrulae of lancet long (Fig. 190), or if as 55d, consider other characters 57
56(55) a Antenna (usually?) ventrally paler than dorsally (Fig. 312)
b Trochanters and trochantelli (usually?) pale (Fig. 312)
c Pronotum (usually?) extensively pale (Fig. 311)
d Inner surface of lancet with (usually?) indistinct spiny pectines (or dentes semicirculares) that reach sclerora (Fig. 194) P. caraganae sp. n.
aa Antenna uniformly black (Fig. 14)
b Trochanters and trochantelli black (Fig. 26)
c Pronotum black or posterior margin narrowly pale (Fig. 35)
d Inner surface of lancet with distinct spiny pectines (or dentes semicirculares) that reach sclerora (Fig. 188) P. albitibia
57(55) a Lancet with numerous setae (Fig. 190)
b Apical serrulae of lancet long (Fig. 190) P. sootryeni
aa Lancet with few setae (Figs 189)
bb Apical serrulae of lancet short (Figs 189) P. astragali
58(54) a Lancet with few setae (Fig. 201)
b Serrulae of lancet flat (Fig. 201)
c Mesepisternum at least slightly matt (Fig. 45)
d Antenna usually ventrally slightly paler than dorsally (Fig. 15) P. pusilla
aa Lancet with numerous setae (Figs 192–193, 196–198, 202)
bb Serrulae of lancet protruding (Figs 192–193, 196, 202) or if as 58b, consider other characters
cc Mesepisternum smooth (Fig. 44) or if as 58c, consider other characters
dd Antenna uniformly black (Fig. 14) or if as 58d, consider other characters 59
59(58) a Mesepisternum (usually?) strongly matt (Fig. 46)
b Antenna uniformly black (Fig. 14)
c Pterostigma (usually?) uniformly yellow or brown (Fig. 39)
d Arctic habitats P. staudingeri
aa Mesepisternum (usually?) smooth or slightly matt (Figs 44–45)
bb Antenna ventrally paler than dorsally (Fig. 15) or if as 59b, consider other characters
cc Pterostigma uniformly yellow to dark brown (Figs 39, 42), or basally dark brown and apically brown (Fig. 40)
dd Usually non-arctic habitats 60
60(59) a Apical serrulae protruding (Figs 192–193, 196)
b Antenna often ventrally paler than dorsally (Fig. 15) 61
aa Apical serrulae flat (Figs 197–198)
bb Antenna uniformly black (Fig. 14) or if as 60b, consider other characters 62
61(60) a Pterostigma usually basally dark brown and apically brown (Fig. 40)
b Setae of lancet more distinct (Fig. 196) P. confusa
aa Pterostigma usually uniformly dark brown (Fig. 42)
bb Setae of lancet less distinct (Figs 192–193) P. armata (on Crataegus) and P. leucopus (on Tilia) (see Grearson and Liston 2012 and Prous et al. 2016 for discussion separating these species)
62(60) a Tangium of lancet without fold (Fig. 198)
b Antenna uniformly black (Fig. 14)
c Pterostigma uniformly yellow (Fig. 39) P. subopaca
aa Tangium of lancet with fold (Fig. 197)
bb Antenna ventrally slightly paler than dorsally (Fig. 15)
cc Pterostigma (usually?) basally dark brown and apically brown (Fig. 40) P. opaca
63(21) a Claws bifid or subbifid (Figs 24–25) 64
aa Claws without or with small or large subapical tooth (Figs 21–23) 66
64(63) a Pterostigma dark (Fig. 42)
b Mesepisternum smooth (Fig. 44)
c Valvula 3 in dorsal view more or less square-shaped or tapering towards apex, and longest setae not concentrated in two dense, lateroapically positioned bundles (Fig. 82) 65
aa Pterostigma pale (Fig. 39)
bb Mesepisternum often matt (45–46)
cc Valvula 3 in dorsal view more or less square-shaped, and longest setae concentrated mainly in two dense, lateroapically positioned bundles (Fig. 91) P. sermola
65(64) a Supraclypeal area black (Figs 9–10)
b Head dorsally usually black (Fig. 6)
c Legs often extensively black P. tetrica
aa Supraclypeal area pale (Fig. 12)
bb Head dorsally with pale spots (Fig. 7)
cc Legs mostly pale (Fig. 28) P. depressa
66(63) a Abdomen with red band, terga 1 and 7–10 black (Fig. 47)
b In dorsal view, valvula 3 narrow (Fig. 124)
c In lateral view, posterior margin of valvula 3 somewhat truncate, and without round outgrowth (Fig. 125) P. erichsonii
aa Abdomen without red band (Figs 26–27), or if present (few specimens of P. laricis), then valvula 3 not narrow in dorsal view (Fig. 79) and posteriorly not truncate in lateral view (Fig. 109)
bb In dorsal view, valvula 3 broad (Figs 67, 69, 72, 74–75, 77–79, 88–89, 91, 93, 95, 97, 101, 116) or if as 66b, consider other characters
cc In lateral view, posterior margin of valvula 3 with different combination of characters (Figs 73, 85, 87, 109) or if as 66c, consider other characters 67
67(66) a In lateral view, valvula 3 with round outgrowth at posterior margin (Figs 85, 87)
b In dorsal view elongate and with distinct or indistinct scopa (Figs 84, 86) 30
aa-bb Combination of characters not as in a–b 68
68(67) a In lateral view, posterior margin of valvula 3 distinctly truncate (Fig. 117)
b In dorsal view valvula 3 elongate and with scopa (Fig. 116) 69
aa-bb Combination of characters not as in a–b 71
69(68) a Head, thorax, and abdomen black (Fig. 26)
b Hypopygium posteriorly not excised (Fig. 62)
c Body length 9–11 mm P. robusta
aa Head, thorax, and abdomen ventrally at least partly pale (Fig. 27–28)
bb Hypopygium posteriorly excised (Fig. 63) or if as 69b, consider other characters
cc Body length 4–9 mm 70
70(69) a Hypopygium posteriorly excised (Fig. 63)
b Ventral margin of lancet not strongly curved upwards (Figs 158–159) P. compressa
aa Hypopygium posteriorly not excised (Fig. 62)
bb Ventral margin of lancet apically strongly curved upwards (Fig. 157) P. abietina
71(68) a Height of eye in lateral view often about 2–3 times as long as distance from dorsal margin of eye to dorsalmost point of head (Fig. 58)
b Clypeus emarginate (Fig. 13)
c In lateral view, posterior margin of valvula 3 round (Fig. 73) 72
aa Height of eye in lateral view about 3–4 times as long as distance from dorsal margin of eye to dorsalmost point of head (Fig. 59)
bb Clypeus truncate or slightly emarginate (Figs 9–10, 12)
cc In lateral view, posterior margin of valvula 3 slightly truncate (Figs 70, 123) or if as 71c, consider other characters 73
72(71) a Valvula 3 without scopa but with indistinct or distinct carina posteriorly (Figs 73, 74)
b Lancet without setae (Fig. 136)
c Pronotum extensively pale (Figs 36–37) P. dasiphorae
aa Valvula 3 with small scopa (Fig. 75)
bb Lancet with numerous setae (Fig. 137)
cc Pronotum usually darker P. malaisei
73(71) a Labrum asymmetrical with right lobe much longer than left (Fig. 11)
b Mesepisternum smooth (Fig. 44)
c Claws without subapical tooth (Fig. 21)
d Pterostigma sometimes basally paler than apically (Fig. 41)
e Body length 3.0–4.5 mm 74
aa Labrum usually symmetrical, or asymmetrical with right lobe slightly longer than left (Figs 9–10, 12)
bb Mesepisternum matt (Figs 45–46) or if as 73b, consider other characters
cc Claws with small or large subapical tooth (Figs 22–23), or if as 73c, consider other characters
dd Pterostigma usually unicoloured (Figs 39, 42)
ee Body length 3.0–8.0 mm 75
74(73) a Valvula 3 narrow (Fig. 103)
b Metafemur usually mostly black (Fig. 26) P. retusa
aa Valvula 3 broader (Figs 101–102)
bb Metafemur completely pale (Fig. 28) P. exigua
75(73) a Valvula 3 without scopa and distinct posterior carina, and slightly tapering from base to apex (Figs 95, 99), or valvula 3 long and narrow and with indistinct scopa (Fig. 119) 76
aa Valvula 3 with distinct posterior carina (Figs 66–69) or more or less square shaped in dorsal view and usually with small scopa (Figs 71–72, 76–81, 88–94, 96–97) 79
76(75) a Valvula 3 long, not tapering from base to apex, and with or without indistinct scopa (Figs 119, 123)
b Mesepisternum smooth (Fig. 44) P. tenuiserra
aa Valvula 3 short, slightly tapering from base to apex and without scopa (Figs Figs 95, 99)
bb Mesepisternum matt (Fig. 46) or if as 76b, consider other characters 77
77(76) a Mesepisternum smooth (Fig. 44)
b Pterostigma dark (Fig. 42)
c Claws without subapical tooth (Fig. 21) P. dedeara sp. n.
aa Mesepisternum matt (Fig. 46)
bb Pterostigma usually pale (Fig. 39)
cc Claws with small subapical tooth (Fig. 22) 78
78(77) a Valvifer 2 black (Fig. 61)
b Tergum 10 usually black (Fig. 61)
c Clypeus black (Fig. 9)
d Metatarsus usually mostly pale (Figs 30–31)
e Lancet with numerous setae (Fig. 156) P. mollis
aa Valvifer 2 often pale (Fig. 62)
bb Tergum 10 often pale (Fig. 62)
cc Clypeus often partly pale (Fig. 10)
dd Metatarsus mostly black (Fig. 29) or if as 78d, consider other characters
ee Lancet without setae (Figs 210217) carinata group
79(75) a In lateral view, ventral margin of valvula 3 straight along most of its length (Fig. 70)
b Valvula 3 posteriorly with distinct carina (Figs 68–69)
c Metatibia only with small dark area at apex (Fig. 30) P. angulata
aa In lateral view, ventral margin of valvula 3 bending gradually upwards (Figs 66, 71, 109)
bb Valvula 3 posteriorly without distinct carina (Figs 71–72, 76–81, 88–94, 96–97) or if as 79b, consider other characters
cc Metatibia with small dark area at apex to completely black 80
80(79) a Valvula 3 with distinct posterior carina and without scopa (Figs 66–67)
b Pronotum extensively pale (Fig. 36)
c Basal 1/3 or less of metatibia pale (Fig. 26)
d Clypeus black (Fig. 9)
e Body length 3.0–5.0 mm P. abbreviata
aa Valvula 3 without posterior carina, but usually with small scopa (Figs 71–72, 76–81, 88–94, 96–97)
bb Pronotum black to extensively pale (Figs 35–37)
cc Metatibia completely pale to completely black
dd Clypeus pale (Figs 10, 12) or if as 80d, consider other characters
ee Body length 3.0–8.0 mm 81
81(80) a Claws with large subapical tooth (Fig. 23)
b Abdomen completely black (Fig. 26)
c Metatarsomere 1 completely or nearly completely pale (Figs 30–31)
d Postocellar area 2.0–2.5 times longer than diameter of lateral ocellus (Fig. 7)
e Mesepisternum smooth (Fig. 44)
f Body length 7–8 mm P. pseudogeniculata
aa-ff Combination of characters not as in a–f 82
82(81) a Cercus clearly extends beyond apex of valvula 3 (Fig. 79)
b Anterior protibial spur without velum (Fig. 33)
c Mesepisternum smooth (Fig. 44) 83
aa-cc Combination of characters not as in a–c 84
83(82) a Labrum pale (Figs 10, 12)
b Pronotum extensively pale (Fig. 36)
c Tangium with (usually?) two campaniform sensilla and lancet with numerous setae (Figs 177–178) P. laricis
aa Labrum black or dark brown (Fig. 9)
bb Pronotum black or only posterior margin narrowly pale (Fig. 35)
cc Tangium without (usually?) campaniform sensilla and lancet with fewer setae (Figs 174–176) P. friesei
84(82) a Mesepisternum smooth (Fig. 44) 85
aa Mesepisternum matt (Fig. 46) 90
85(84) a Abdomen completely black (Fig. 26)
b Claws without subapical tooth (Fig. 21)
c Pronotum black
d Body length 3.0–4.0 mm P. monogyniae
aa-dd Combination of characters not as in a–d. 86
86(85) a Abdomen laterally and ventrally with pale patches (Fig. 27)
b Valvula 3 in dorsal view more or less square-shaped and longest setae not concentrated in two dense, lateroapically positioned bundles (Figs 77–78, 88–89) 87
aa Abdomen black (Fig. 26) or only terga 9–10 and valvifer 2 pale (Fig. 62)
bb Valvula 3 in dorsal view more or less square-shaped and longest setae concentrated mainly in two dense, lateroapically positioned bundles (Fig. 91) micronematica group
87(86) a Serrulae of lancet flat (Figs 129–130)
b Pterostigma usually dark (Fig. 42) 88
aa Serrulae of lancet distinctly papilliform (Figs 203–204)
bb Pterostigma pale (Fig. 39) 89
88(87) a Cercus black (Fig. 77)
b Clypeus at least partly pale (Fig. 10)
c Valvula 3 somewhat narrower (Figs 76–77) P. biscalis
aa Cercus usually pale (Fig. 78)
bb Clypeus black (Fig. 9)
cc Valvula 3 somewhat broader (Fig. 78) P. maesta
89(87) a Valvula 3 somewhat narrower (Fig. 89)
b Middle serrulae small compared to height of lancet (ratio around 1/10–1/7) (Fig. 204) P. pseudocoactula
aa Valvula 3 somewhat broader (Fig. 88)
bb Middle serrulae large compared to height of lancet (ratio around 1/6–1/4) (Fig. 203) P. alpestris
90(84) a Serrulae of lancet distinctly papilliform (Fig. 204)
b Lancet with setae (Fig. 204) P. pseudocoactula
aa-bb Combination of characters not as in a–b 91
91(90) a Valvula 3 in dorsal view more or less square-shaped and longest setae concentrated mainly in two dense, lateroapically positioned bundles (Fig. 91)
b Lancet with setae (Figs 205209) micronematica group
aa Valvula 3 in dorsal view tapering towards apex and / or with longest setae not concentrated in two dense, lateroapically positioned bundles (Figs 93–99)
bb Lancet without setae (Figs 210217) carinata group

Males

Males of P. aphantoneura, P. astragali, P. depressa, and P. subbifida are unknown.

1 a At least mesepisternum partly pale (Fig. 28) 2
aa At least mesepisternum black (Figs 26–27) 22
2(1) a Metatibia black (Fig. 29)
c Metafemur black (Fig. 26)
b Claws with large subapical tooth or bifid (Fig. 24) P. fausta
aa At least basal 2/3 of metatibia pale (Figs 26, 30–31)
bb Metafemur usually pale (Figs 27–28)
cc Claws usually with small subapical tooth (Fig. 22) 3
3(2) a Abdomen with red band dorsally
b Paravalva somewhat rhombus-shaped, pseudoceps somewhat rectangular and unmodified, valvispina small, slightly bent and asymmetrical at apex (Fig. 290) P. erichsonii
aa Abdomen without red band
bb Penis valve different 4
4(3) a Supraclypeal area black (Figs 9–10)
b Antenna uniformly black (Fig. 14)
c Antenna with numerous stout black setae among finer paler ones (Figs 16, 18)
d Paravalva rectangular, with straight ventral margin, and apically distinctly shorter than pseudoceps; pseudoceps somewhat rectangular and unmodified; relatively large and straight valvispina forming right or acute angle with paravalva (Fig. 254) P. insularis
aa Supraclypeal area pale (Fig. 12)
bb Antenna uniformly black to uniformly pale (Figs 14–18)
cc Antenna without or apparently without numerous stout black setae among finer paler ones (Fig. 17), or if as 4c, consider other characters
dd Penis valve different 5
5(4) a Mesepisternum completely or nearly completely pale (Fig. 336)
b Valvispina straight and positioned in middle or upper third of paravalva (Fig. 337)
c Paravalva somewhat oval shaped or rectangular (Fig. 337)
d Pseudoceps unmodified, more or less rectangular (Fig. 337) P. cadma
aa-dd Combination of characters not as in a–d 6
6(5) a Paravalva and pseudoceps more or less rectangular (Figs 224–226)
b Valvispina straight or slightly bent and positioned in middle or upper third of paravalva (Figs 224–226) P. pallidiventris
aa-bb Combination of characters not as in a–b 7
7(6) a Claws usually with large subapical tooth (Fig. 23)
b Paravalva basally much broader than apically (Figs 255, 257, 259)
c Pseudoceps more or less rectangular, not distinctly modified (Figs 255, 257, 259)
d Valvispina simple and straight or slightly bent and positioned close to but nevertheless distinctly removed from ventral margin of paravalva (Figs 255, 257, 259) P. conjugata
aa-dd Combination of characters not as in a–d 8
8(7) a Valvispina absent or indistinct (Figs 303–305)
b Pseudoceps dorsally with loose membranous region covered with hair (Figs 303–305)
c Propleuron black (Fig. 26)
d Claws sometimes bifid (Fig. 25) micronematica group
aa Valvispina distinct (Figs 275–276, 278, 283–289, 291296, 302)
bb Pseudoceps without loose membranous region (Figs 275–276, 278, 283–289, 291296, 302)
cc Propleuron often pale (Fig. 28)
dd Claws without or with small subapical tooth (Figs 21–22) 9
9(8) a Valvispina small, simple, and bent dorsally (Figs 293–295)
b Paravalva and pseudoceps similar in size and shape (Figs 293–295)
c Paravalva dorsally concave, and ventrally convex or more or less straight (Figs 293–295)
d Body length 3.0–4.5 mm 84
aa-dd Combination of characters not as in a–d 10
10(9) a Pseudoceps with spines, and narrow tip extending far from valvispina (Fig. 275)
b Valvispina small, simple, and straight (Fig. 275) P. bufo
aa Pseudoceps without spines, and with broad or narrow tip not extending far from paravalva (Figs 276, 278, 283–289, 291–292, 296, 302)
bb Valvispina variously shaped (Figs 276, 278, 283–289, 291–292, 296, 302) 11
11(10) a Valvispina large and bent, replacing half of paravalva (Figs 276, 278) 86
aa Valvispina smaller and paravalva less modified (Figs 283–289, 291–292, 296, 302) 12
12(11) a Valvispina large, gradually bent, and simple (Fig. 296) P. paralella
aa Valvispina smaller, straight, abruptly bent, or dorsally serrate (Figs 283–289, 291–292, 302) 13
13(12) a Pseudoceps with distinctly narrow tip (Fig. 302)
b Valvispina simple and more or less straight (Fig. 302) P. tenuiserra
aa Pseudoceps with broad tip (Figs 283–289, 291–292)
bb Valvispina bent, dorsally serrate (Figs 283–288, 291–292), or if as 13b, consider other characters 14
14(13) a Valvispina sharply bent, and short and broad (Figs 291–292)
b Paravalva somewhat trapez-shaped (Figs 291–292) 15
aa Valvispina not sharply bent, and narrower (Figs 283–289)
bb Paravalva not trapez-shaped, or less distinctly trapez-shaped (Figs 283–289) 16
15(14) a Ventral margins of paravalva and valvispina forming obtuse angle (Fig. 292)
b Paravalva anteriorly (usually?) slightly more concave (Fig. 292) P. wesmaeli
aa Ventral margins of paravalva and valvispina forming almost right angle (Fig. 291)
bb Paravalva anteriorly (usually?) slightly less concave (Fig. 291) P. glauca
16(14) a Valvispina small, more or less straight, and almost simple (Fig. 289)
b Paravalva broad (Fig. 289)
c Body length 8–9 mm P. robusta
aa-cc Combination of characters not as in a–c 17
17(16) a Paravalva and pseudoceps relatively narrow (Fig. 283)
b Valvispina distinctly bent, and with thin and non-serrate dorsal margin (Fig. 283)
c Body length 3.5–5.0 mm P. abietina
aa-cc Combination of characters not as in a–c 18
18(17) a Valvispina simple and angled in middle (Fig. 285) P. decipiens
aa Valvispina with at least serrate dorsal margin and angled or not angled in middle (Figs 284, 286–288) 19
19(18) a Paravalva relatively broad and abruptly narrowing at apex (Figs 284, 288) 20
aa Paravalva slightly narrower and more gradually narrowing at apex (Figs 286–287) 21
20(19) a Apical margin of paravalva more or less entirely convex (Fig. 284)
b Valvispina smaller and more or less straight (Fig. 284) P. compressa
aa Apical margin of paravalva with concave part followed by convex part (Fig. 288)
bb Valvispina larger and slightly bent (Fig. 288) P. saxesenii
21(19) a Valvispina with dorsal and ventral thinned and serrate margins (Fig. 286)
b Paravalva more elongate (Fig. 286) P. gerula
aa Valvispina only with dorsal thinned and serrate margin (Fig. 287)
bb Paravalva less elongate (Fig. 287) P. pseudodecipiens
22(1) a Tergum 8 with distinct apical projection (Fig. 57)
b Antenna ventrally distinctly paler than dorsally, or uniformly pale (Figs 16, 18)
c Claws with large subapical tooth (Figs 23–24)
d Mesepisternum smooth (Fig. 44) P. armata (on Crataegus) and P. leucopus (on Tilia) (see Grearson and Liston 2012 and Prous et al. 2016 for discussion separating these species)
aa Tergum 8 without distinct apical projection (Figs 55–56)
bb Antenna uniformly black to uniformly pale (Figs 14–18)
cc Claws without or with small subapical tooth (Figs 21–22), or if as 22c, consider other characters
dd Mesepisternum matt (Figs 45–46) or if as 22d, consider other characters 23
23(22) a Terga and sterna 2–6 at least partly pale (Fig. 27) 24
aa Terga and sterna 2–6 black (Fig. 26) 43
24(23) a Metatibia black (Fig. 29)
b Metafemur black (Fig. 26)
c Claws with large subapical tooth or bifid (Figs 23–25) 25
aa At least basal 1/3 of metatibia pale (Fig. 26)
bb Metafemur pale (Figs 27–28) or if as 24b, consider other characters
cc Claws without or with small subapical tooth (Figs 21–22), or if as 24c, consider other characters 26
25(24) a Head extensively pale around eyes (Fig. 8)
b Paravalva and valvura almost forming right angle (Fig. 299)
c Valvispina broad with curved anteroventral margin (Fig. 299) P. fausta
aa Head at most with small brown or pale spots (Figs 6–7)
bb Paravalva and valvura more or less at the same level (Fig. 301)
cc Valvispina thin and symmetrical (Fig. 301) P. tetrica
26(24) a Basal 1/3–1/2 of metatibia pale (Fig. 26)
b Supraclypeal area pale (Fig. 12)
c Abdomen with red band (Fig. 48)
d Claws usually with large subapical tooth (Fig. 23)
e Dorsoapical margin of paravalva and dorsal margin of valvispina forming distinct s-shape (Fig. 232)
f Valvispina at ventral margin of paravalva, and gradually merging with paravalva at ventral and dorsal side P. condei
aa-ff Combination of characters not as in a–f 27
27(26) a Supraclypeal area black (Figs 9–10)
b Claws usually bifid or subbifid (Figs 24–25)
c Penis valve with long, bent, and symmetrical valvispina (Fig. 301) P. tetrica
aa-cc Combination of characters not as in a–c 28
28(27) a Supraclypeal area pale (Fig. 12) 29
aa Supraclypeal area black (Figs 9–10) 32
29(28) a Abdomen with red band dorsally
b Antenna extensively pale (Figs 16, 18)
c Claws with small subapical tooth (Fig. 22)
d Paravalva somewhat rhombus-shaped, pseudoceps somewhat rectangular and unmodified, valvispina small, slightly bent, and asymmetrical at apex (Fig. 290) P. erichsonii
aa-dd Combination of characters not as in a–d 30
30(29) a Claws usually with large subapical tooth (Fig. 23)
b Abdomen often completely pale (Fig. 50)
c Pterostigma dark (Fig. 42)
d Paravalva apically distinctly narrower than pseudoceps; pseudoceps more or less rectangular, unmodified; valvispina small, simple and bent upwards (Fig. 261) P. testacea
aa-dd Combination of characters not as in a–d 31
31(30) a Dorsal margin of paravalva somewhat s-shaped (Figs 294, 296)
b Valvispina small or large, and directed dorsally (Figs 294, 296)
c Claws without, or with small subapical tooth (Figs 21–22)
d Body length 3.0–5.5 mm 76
aa Dorsal margin of paravalva not s-shaped (Figs 223–227, 231, 233–237, 243–251, 253–254, 303–309)
bb Valvispina absent, or larger, or not directed dorsally (Figs 223–227, 231, 233–237, 243–251, 253–254, 303–309)
cc Claws with large subapical tooth (Figs 23–25) or if as 31c, consider other characters
dd Body length 4.0–7.0 mm 32
32(31,28) a Anterior margin of paravalva forming acute or right angle with valvispina (Figs 253–254)
b Valvispina arising from ventral margin of paravalva (Figs 253–254) 33
aa-bb Combination of characters not as in a–b 34
33(32,53) a Valvispina larger (Fig. 254)
b Ventral margin of paravalva not protruding (Fig. 254) P. insularis
aa Valvispina smaller (Fig. 253)
bb Ventral margin of paravalva protruding (Fig. 253) P. biscalis
34(32) a Valvispina absent or indistinct (Figs 303–305)
b Pseudoceps dorsally with loose membranous region covered with setae (Figs 303–305)
c Claws sometimes bifid (Fig. 25) micronematica group
aa Valvispina distinct (Figs 223–227, 231, 233–237, 243–251)
bb Pseudoceps without loose membranous region (Figs 223–227, 231, 233–237, 243–251)
cc Claws without or with small subapical tooth (Figs 21–22) 35
35(34) a Valvispina long and ventrally bent (Fig. 237)
b Pseudoceps apically relatively narrow and ventrally bent (Fig. 237) P. dissimilis
aa Valvispina not ventrally bent (Figs 223–227, 231, 233–236, 243–251)
bb Pseudoceps not ventrally bent (Figs 223–227, 231, 233–236, 243–251) 36
36(35) a Dorsal margin of penis valve from tip of valvura to tip of pseudoceps almost forming semicircle (Figs 233–236)
b Paravalva and pseudoceps broad (Figs 233–236)
c Mesepisternum sometimes matt (Figs 45–46) 37
aa Dorsal margin of penis valve not forming semicircle (Figs 223–227, 231, 243–251)
bb Paravalva and pseudoceps narrow (Figs 223–227, 231) or if as 36b, consider other characters
cc Mesepisternum usually smooth (Fig. 44) 38
37(36) a Valvispina close to ventral margin of paravalva (Figs 234, 236) P. pseudocoactula
aa Valvispina distinctly removed from ventral margin of paravalva (Figs 233, 235) P. alpestris
38(36) a Penis valve relatively small and narrow (Fig. 231)
b Pseudoceps apically distinctly narrowed (Fig. 231)
c Valvispina starting from ventral margin of paravalva and dorsally with round hump (Fig. 231) P. punctifrons
aa-cc Combination of characters not as in a–c 39
39(38) a Paravalva and pseudoceps of similar size and shape (Figs 223–226)
b Ventral margin of paravalva more or less straight (Figs 223–226)
c Valvispina positioned in middle or upper third of paravalva (Figs 223–226) 40
aa-cc Combination of characters not as in a–c 41
40(39) a Penis valve elongate, paravalva and pseudoceps more or less rectangular (Figs 224–226)
b Pterostigma dark (Fig. 42) P. pallidiventris
aa Paravalva and pseudoceps broader, less rectangular (Fig. 223)
bb Pterostigma (usually?) pale (Fig. 39) P. nigricans
41(39) a Valvispina positioned more or less in middle of paravalva (Fig. 227)
b Dorsally, paravalva narrowing gradually at apex (Fig. 227) P. maesta
aa Valvispina positioned in lower part of paravalva (Figs 243–251)
bb Dorsally, paravalva narrowing abruptly at apex (Figs 243–251) 42
42(41) a Paravalva more elongate (Figs 244, 246)
b Valvispina longer (Figs 244, 246)
c Abdomen sometimes with red band (Fig. 47)
d Mesepisternum sometimes matt (Fig. 45) P. cincta
aa Paravalva less elongate (Figs 243, 245, 247–251)
bb Valvispina usually shorter
cc Abdomen without red band
dd Mesepisternum smooth (Fig. 44) few specimens of rufipes group
43(23) a Penis valve apically truncate (Figs 297–298)
b Valvispina absent (Figs 297–298)
c Clypeus emarginate (Fig. 13) 44
aa Penis valve apically pointed or round (Figs 224–231, 234, 236, 238254, 256, 258, 260, 262, 264, 266274, 277, 279–282, 293–296, 300, 303–310, 317, 321)
bb Valvispina usually present (Figs 224–231, 234, 236, 238254, 256, 258, 260, 262, 264, 266274, 277, 279–282, 293–296, 300, 303–310, 317, 321)
cc Clypeus truncate or slightly emarginate (Figs 9–10, 12) 45
44(43) a Penis valve apically strongly bent, L-shaped (Fig. 298)
b Apical margin of penis valve with middle notch (Fig. 298) P. malaisei
aa Penis valve apically weakly bent, not L-shaped (Fig. 297)
bb Apical margin of penis valve without middle notch (Fig. 297) P. dasiphorae
45(43) a Claws with long subapical tooth close to apical one (bifid) (Fig. 25) 46
aa Claws without or with small or large subapical tooth clearly separated from apical one (Figs 21–24) 49
46(45) a Valvispina absent or indistinct (Figs 303–305)
b Pseudoceps dorsally with loose membranous region covered with setae (Figs 303–305) micronematica group
aa Valvispina merged with paravalva forming large spine (Figs 267, 274, 317)
bb Pseudoceps with or without loose membranous region (Figs 267, 274, 317) 47
47(46) a Pseudoceps of penis valve short and broad (Figs 317)
b Antenna ventrally paler than dorsally (Fig. 319) P. caraganae sp. n.
aa Pseudoceps of penis valve longer and narrower (Figs 267, 274)
bb Antenna uniformly black (Figs 14, 17) or if as 47b, consider other characters 48
48(47) a Hind trochanters, trochantelli, and tibia partly pale
b Antenna (usually?) ventrally at least slightly paler than dorsally (Fig. 18)
c Antenna with numerous and clearly visible stout black setae among finer paler ones (Fig. 18)
d Penis valve without membranous fold near tip of valvispina and pseudoceps with distinct dorsal depression in middle or basal part (Fig. 267) P. bifida
aa Hind trochanters, trochantelli, and tibia uniformly black or brown
bb Antenna uniformly black (Figs 14, 17)
cc Antenna with only some barely visible stout black setae among finer paler ones (Fig. 17)
dd Penis valve with membranous fold near tip of valvispina and pseudoceps without dorsal depression in middle or basal part (Fig. 274) P. frigida
49(45) a Labrum asymmetrical with right lobe much longer than left (Fig. 11)
b Mesepisternum smooth (Fig. 44)
c Claws without subapical tooth (Fig. 21)
d Penis valves as in Figs 258, 260
e Body length 3.0–4.5 mm 50
aa Labrum usually symmetrical or asymmetrical with right lobe slightly longer than left (Figs 9, 10, 12)
bb Mesepisternum matt (Figs 45–46) or if as 49b, consider other characters
cc Claws with small or large subapical tooth (Figs 21–24), or if as 49c, consider other characters
dd Penis valves different (Figs 224–231, 234, 236, 238254, 256, 262, 264, 266, 268273, 277, 279–282, 293–296, 300, 303–310, 317, 321)
ee Body length 3.0–7.0 mm 51
50(49) a Anterior margin of paravalva forming acute or right angle with valvispina (Fig. 260)
b Valvispina short and bent (Fig. 260)
c Posterior margin of pronotum black or narrowly pale (Fig. 35)
d Metafemur black (Fig. 26)
e Trochanters and trochantelli from black to pale (Figs 26–27, 32) P. retusa
aa Anterior margin of paravalva forming obtuse angle with valvispina (Fig. 258)
bb Valvispina long and straight (Fig. 258)
cc Pronotum extensively pale (Fig. 35)
dd Metafemur black or pale (Figs 26–28)
ee Trochanters and trochantelli pale (Fig. 27) P. exigua
51(49) a Pseudoceps with dorsal membranous region covered with setae and / or apically bent upwards (Figs 303–309)
b Valvispina small and bent, small and straight, or absent (Figs 303–309) micronematica group
aa Pseudoceps not strongly modified (Figs 224–231, 234, 236, 238254, 256, 262, 264, 266, 268273, 277, 279–282, 293–296, 300, 310, 317, 321)
bb Valvispina distinctly present (Figs 224–231, 234, 236, 238254, 256, 262, 264, 266, 268273, 277, 279–282, 293–296, 300, 310, 317, 321) 52
52(51) a Valvispina relatively small and more or less straight (Figs 224–231, 234, 236, 238251, 253–254, 310, 321) 53
aa Valvispina large and / or bent (Figs 252, 256, 262, 264, 266, 268273, 277, 279–282, 293–296, 300, 317) 65
53(52) a Anterior margin of paravalva forms acute or right angle with valvispina (Figs 253–254)
b Valvispina relatively broad or ventral margin of paravalva distinctly protruding (Figs 253–254)
c Sternum 9 pale (Figs 52, 54) 33
aa Anterior margin of paravalva forms obtuse or rarely right angle with valvispina (Figs 224–231, 234, 236, 238251, 310, 321)
bb Valvispina thin and ventral margin of paravalva usually not distinctly protruding (Figs 224–231, 234, 236, 238251, 310, 321)
cc Sternum 9 black (Fig. 53), or if as 53c, consider other characters 54
54(53) a Pseudoceps apically distinctly narrowed (Fig. 231)
b Penis valve relatively small and narrow (Fig. 231)
c Valvispina arising from ventral margin of paravalva and dorsally with round hump (Fig. 231) P. punctifrons
aa Pseudoceps apically broad (Figs 224–230, 234, 236, 238251, 310, 321)
bb Penis valve longer or broader (Figs 224–230, 234, 236, 238251, 310, 321)
cc Valvispina arising from middle of paravalva, dorsally without round hump (Figs 224–230, 238251, 310, 321), or if as 54c, consider other characters 55
55(54) a Shape of paravalva more or less symmetrical around valvispina (Fig. 310)
b Valvispina small and positioned in middle of paravalva (Fig. 310)
c Posterior margin of pronotum and tegula extensively pale (Fig. 35)
d Metafemur black (Fig. 26)
e Mesepisternum smooth or slightly matt (Figs 44–45) P. kontuniemii (micronematica group)
aa Shape of paravalva clearly asymmetrical around valvispina (Figs 224–230, 234, 236, 238251, 321)
bb Valvispina large, positioned close to dorsal or ventral margin (Figs 224–230, 234, 236, 238251, 321), or if as 55b, consider other characters
cc Posterior margin of pronotum and tegula black (Fig. 35), or if as 55c, consider other characters
dd Metafemur pale (Figs 27–28), or if as 55d, consider other characters
ee Mesepisternum matt (Fig. 46), or if as 55e, consider other characters 56
56(55) a Paravalva and pseudoceps more or less rectangular (Figs 224–226)
b Valvispina positioned in middle or upper third of paravalva (Figs 224–226)
c Posterior margin of pronotum and tegula pale (Fig. 35)
d Supraclypeal area, clypeus, antenna ventrally, metafemur, and sternum 9 often pale (Figs 12, 18, 28, 54) P. pallidiventris
aa-dd Combination of characters not as in a–d 57
57(56) a Ventral margin of paravalva abruptly bent before valvispina (Fig. 228)
b Dorsal margin of paravalva gradually tapering towards valvispina (Fig. 228)
c Valvura slightly bent ventrally (Fig. 228) P. angulata
aa-cc Combination of characters not as in a–c 58
58(57) a Vertical distance between dorsalmost part of paravalva and valvispina relatively long (Fig. 321)
b Valvispina relatively large (Fig. 321)
c Ventral margin of paravalva slightly protruding (Fig. 321)
d Dorsobasal part of paravalva without angulate dark area (Fig. 321)
e Metatibia in basal 1/3 pale
f Mesepeisternum smooth (Fig. 44) P. dedeara sp. n.
aa-ff Combination of characters not as in a–f 59
59(58) a Vertical distance between dorsalmost part of paravalva and valvispina short (Figs 227, 229–230)
b Dorsal margin of paravalva gradually tapering towards valvispina (Figs 227, 229–230)
c Paravalva relatively narrow (Figs 227, 229–230)
d Mesepisternum smooth (Fig. 44) 60
aa Vertical distance between dorsalmost part of paravalva and valvispina long (Figs 234, 236, 238251)
bb Dorsal margin of paravalva abruptly tapering towards valvispina (Figs 242251) or if as 59b, consider other characters
cc Paravalva usually relatively broad (Figs 234, 236, 238251)
dd Mesepisternum matt (Figs 45–46) or if as 59d, consider other characters 62
60(59) a Dorsobasal part of paravalva with angulate dark area (Fig. 227) P. maesta
aa Dorsobasal part of paravalva without angulate dark area (Figs 229–230) 61
61(60) a Flagellum about 2.4–2.5 times as long as width of head
b Metatarsus mainly black (Fig. 29) P. geniculata
aa Flagellum about 2.0–2.1 times as long as width of head
bb Metatarsus mainly pale (Fig. 31) P. pseudogeniculata
62(59) a Dorsal margin of penis valve from tip of valvura to tip of pseudoceps almost forming semicircle (Figs 234, 236)
b Paravalva and pseudoceps broad (Figs 234, 236) P. pseudocoactula
aa Dorsal margin of penis valve not forming semicircle (Figs 238251)
bb Paravalva and pseudoceps narrow (e.g. Figs 244, 246) or if as 62b, consider other characters 63
63(62) a Valvar strut not clearly evident at anterior margin (Figs 238–242)
b Mesepisternum matt (Fig. 46) carinata group
aa Valvar strut distinct along its entire length (Figs 243–251)
bb Mesepisternum usually smooth (Fig. 44) 64
64(63) a Paravalva more elongate (Figs 244, 246)
b Valvispina longer (Figs 244, 246)
c Mesepisternum sometimes matt (Fig. 45) P. cincta
aa Paravalva less elongate (Figs 243, 245, 247–251)
bb Valvispina usually shorter (Figs 243, 245, 247–251)
cc Mesepisternum smooth (Fig. 44) rufipes group
65(52) a Mesopostnotum smooth (Fig. 19)
b Claws without subapical tooth (Fig. 21)
c Mesepisternum smooth (Fig. 44)
d Antenna usually ventrally paler than dorsally (Fig. 18)
e Paravalva largely present (Fig. 262)
f Valvispina large and strongly bent, about as long as apical margin of paravalva and more or less parallel to it (Fig. 262) P. appendiculata
aa Mesopostnotum (usually?) matt (Fig. 20)
bb Claws with subapical tooth (Figs 22–24) or if as 65b, consider other characters
cc Mesepisternum matt (Figs 45–46), or if as 65c, consider other characters
dd Antenna uniformly black to uniformly pale (Figs 14–18)
ee Paravalva nearly completely merged with valvispina (Figs 264, 266, 268273, 317) or largely present (Figs 252, 256, 277, 279–282, 293–296, 300)
ff Valvispina smaller, or not strongly bent, or not parallel to apical margin of paravalva, or not as long as apical margin of paravalva (Figs 252, 256, 264, 266, 268273, 277, 279–282, 293–296, 300, 317) 66
66(65) a Paravalva nearly completely merged with valvispina, forming large spine (Figs 264, 266, 268273, 317) 67
aa Paravalva largely present and valvispina usually smaller (Figs 252, 256, 277, 279–282, 293–296, 300) 75
67(66) a Penis valve with membranous fold near or covering tip of valvispina (Figs 264, 266)
b Claws with small subapical tooth (Fig. 22)
c Mesepisternum smooth (Fig. 44) 68
aa Penis valve without membranous fold (Figs 268273, 317)
bb Claws with large subapical tooth (Figs 22–23) or if as 67b, consider other characters
cc Mesepisternum matt (Figs 45–46) or if as 67c, consider other characters 69
68(67) a Valvispina of penis valve less sharply bent (forming half circle) (Fig. 266) P. ruficornis
aa Valvispina of penis valve more sharply bent (being almost L-shaped) (Fig. 264) P. melanocarpa
69(67) a Pseudoceps of penis valve short and broad (Figs 273, 317)
b Mesepisternum (usually?) smooth (Fig. 44) 87
aa Pseudoceps of penis valve longer and narrower (Figs 268–272)
bb Mesepisternum matt (Figs 44–46) or if as 69b, consider other characters 70
70(69) a Penis valve with weakly bent and broad valvispina, and with narrow pseudoceps without distinct dorsal depression in middle part (Fig. 268) P. confusa
aa Penis valve with different combination of characters (Figs 269–272) 71
71(70) a Valvispina of penis valve narrow and with blunt tip (Fig. 270)
b Antenna ventrally paler than dorsally (Fig. 18) P. opaca
aa Valvispina of penis valve broad or narrow and with sharp tip (Figs 269, 271–272)
bb Antenna uniformly black (Fig. 14) or if as 71b, consider other characters 72
72(71) a Valvispina of penis valve narrow (Fig. 272)
b Antenna uniformly black (Fig. 14) 73
aa Valvispina of penis valve broad (Figs 269, 271)
bb Antenna ventrally paler than dorsally (Fig. 18) or if as 72b, consider other characters 74
73(72) a Mesepisternum smooth to slightly matt (Figs 44–45)
b Usually non-arctic habitats P. luteipes
aa Mesepisternum usually strongly matt (Fig. 46)
bb Arctic habitats P. staudingeri
74(72) a Pseudoceps of left and right penis valve without distinct dorsal depression in middle part and with weakly bent valvispina (Fig. 269)
b Antenna uniformly black (Fig. 14) P. subopaca
aa Pseudoceps of left penis valve with distinct dorsal depression in middle part and with strongly bent valvispina (Fig. 271)
bb Antenna ventrally paler than dorsally (Fig. 18) P. pusilla
75(66) a Dorsal margin of paravalva somewhat s-shaped (Figs 293–294, 296)
b Valvispina small and narrow or large and narrow (Figs 293–294, 296)
c Supraclypeal area pale (Fig. 12)
d Body length 3.0–5.5 mm 76
aa Dorsal margin of paravalva not s-shaped (Figs 252, 256, 277, 279–282, 300)
bb Valvispina large and broad or small and narrow (Figs 252, 256, 277, 279–282, 300)
cc Supraclypeal area black (Figs 9–10)
dd Body length 3.0–7.0 mm 78
76(75) a Valvispina large (Fig. 296)
b Body length 4.0–5.5 mm P. paralella
aa Valvispina small (Figs 293, 294)
bb Body length 3.0–4.5 mm 77
77(76) a Paravalva ventrally abruptly narrowed before valvispina (Fig. 294) P. nigella
aa Paravalva ventrally gradually narrowed before valvispina (Fig. 293) P. amphibola
78(75) a Valvispina broad (Fig. 300)
b Mesepisternum matt (Fig. 46) P. mollis
aa Valvispina narrow (Figs 252, 256, 277, 279–282)
bb Mesepisternum smooth (Fig. 44) 79
79(78) a Pseudoceps covered with setae and expanded apically (Fig. 256)
b Valvispina long and abruptly bent apically (Fig. 256)
c Claws without subapical tooth (Fig. 21)
d Pronotum black
e Body length 3.0–4.0 mm P. monogyniae
aa Pseudoceps apparently without setae and not expanded apically (Figs 252, 277, 279–282)
bb Valvispina shorter or more gradually bent (Figs 252, 277, 279–282)
cc Claws with subapical tooth (Fig. 22) or if as 79c, consider other characters
dd Pronotum often pale (Fig. 35)
ee Body length 3.0–7.0 mm 80
80(79) a Valvispina long, and dorsally without posteriorly constricted hump (Fig. 252)
b Claws without subapical tooth (Fig. 21)
c Body length about 4.3 mm (based on a single male available for study) P. abbreviata
aa Valvispina short or dorsally with posteriorly constricted hump (Figs 277, 279–282)
bb Claws with small subapical tooth (Fig. 22) or if as 80b, consider other characters
cc Body length 5.0–7.0 mm 81
81(80) a Valvispina long and with posteriorly constricted hump (Figs 281–282) 82
aa Valvispina short and without posteriorly constricted hump (Figs 277, 279–280) 83
82(81) a Pseudoceps apically with narrow thickening (Fig. 282)
b Paravalva distinctly angled dorsobasally (Fig. 282) P. nigriceps
aa Pseudoceps apically with broad thickening (Fig. 281)
bb Paravalva not angled dorsobasally (Fig. 281) P. leucopodia
83(81) a Labrum pale (Figs 10, 12)
b Pronotum extensively pale or posterior margin narrowly pale (Figs 35–36)
c Flagellum about 2.4–3.2 times as long as width of head P. laricis
aa Labrum black or dark brown (Fig. 9)
bb Pronotum (usually?) black
cc Flagellum about 3.6–3.8 times as long as width of head P. friesei
84(9) a Paravalva ventrally abruptly narrowed before valvispina (Fig. 294) P. nigella
aa Paravalva ventrally gradually narrowed before valvispina (Figs 293, 295) 85
85(84) a Tegula and pronotum extensively pale (Fig. 36)
b Penis valve ventrally somewhat bent (Fig. 295) P. parva
aa Tegula and pronotum black (Fig. 35)
bb Penis valve ventrally more or less straight (Fig. 293) P. amphibola
86(11) a Depression on apical part of valvispina absent (Fig. 276) P. pallida
aa Depression on apical part of valvispina present (Fig. 278) P. subarctica
87(69) a Antenna ventrally paler than dorsally (Fig. 319)
b Ventroapical part of pseudoceps only slightly extended beyond dorsalmost margin (Fig. 317)
c Valvispina narrower and its posterior and anterior margin of similar shape (Fig. 317) P. caraganae sp. n.
aa Antenna uniformly black (Fig. 14)
bb Ventroapical part of pseudoceps distinctly extended beyond dorsalmost margin (Fig. 273; http://dx.doi.org/10.6084/m9.figshare.4690174)
cc Valvispina broader and its posterior and anterior margin of distinctly different or of similar shape (Fig. 273; http://dx.doi.org/10.6084/m9.figshare.4690174) 88
88(87) a Pterostigma (usually?) basally dark brown and apically brown (Fig. 40)
b Pseudoceps of penis valve somewhat broader (Fig. 273) P. albitibia
aa Pterostigma (usually?) uniformly yellow (Fig. 39)
bb Pseudoceps of penis valve somewhat narrower (http://dx.doi.org/10.6084/m9.figshare.4690174) P. sootryeni

Taxonomy

Pristiphora abietina group

Pristiphora abietina (Christ, 1791)

Figs 12, 157, 283

Tenthredo Pini [sic!] Retzius, 1783: 73, by indication to Degeer (1771: 1001–1002, Pl. 38. figs 5–7). Primary homonym of Tenthredo pini Linné, 1758 [= Diprion pini (Linné, 1758)]. Syntype(s) possibly in NHRS. Type locality: possibly Lövstabruk, Uppland, Sweden (place of residence of Degeer; see Taeger and Blank 1998).

Tenthredo abietina Christ, 1791: 447, by indication to Degeer (1771: 1001–1002, Pl. 38. figs 5–7). Syntype(s) possibly in NHRS. Type locality: possibly Lövstabruk, Uppland, Sweden (place of residence of Degeer; see Taeger and Blank 1998).

Tenthredo abietum Hartig in Hartig and Hartig, 1834: 984–985. Lectotype ♀ (GBIF-GISHym3183; here designated) in ZSM, examined. Type locality: not stated.

Nematus (Nematus) limbatus Dahlbom, 1835b: 9. Not available. Nomen nudum. Note. Not available through an indication, only internal reference on “Nematus abietinus No. 86”, without giving any differences.

Nematus (Nematus) abietinus Dahlbom, 1835b: 9. Note. Description by indication on “Mouche à scie du Sapin De Geer”. Syntype(s) possibly in NHRS. Type locality: possibly Lövstabruk, Uppland, Sweden (place of residence of Degeer; see Taeger and Blank 1998).

Nematus truncatus Hartig, 1837: 207. Holotype ♀ possibly lost. Type locality: Harz, Germany. Synonymised with Lygaeonematus pini by Konow (1904b).

Similar species

The most similar species is P. compressa, which is on average larger (6.0–9.0 vs. 4.0–6.5 mm) and has hypopygium posteriorly excised (not excised in P. abietina) (Figs 62–63). Males are best recognised by examining penis valves (Fig. 283). See the Key for more details.

Genetic data

Based on COI barcode sequences, P. abietina belongs to the same BIN cluster (BOLD:AAK9450) as P. compressa, P. decipiens, P. gerula, P. pseudodecipiens, P. saxesenii, and possibly P. robusta. Maximum distance within the BIN is 2.18% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9450, diverging by minimum of 3.92%, is BOLD:ACO1401 (P. euxantha Huflejt, 2006, a species not treated here). Based on nuclear data, within species divergence is 0.1% (based on two specimens and NaK) and the nearest neighbour is 1.2% different (P. saxesenii, both genes combined).

Host plants

Picea abies (L.) Karsten (Schafellner et al. 1999, Pschorn-Walcher and Altenhofer 2000), P. sitchensis (Bong.) Carriere (Austarå et al. 1984), P. pungens Engelm. (Kollár 2007, Kula et al. 2016).

Distribution and material examined

West Palaearctic. Specimens studied are from Germany, Latvia, and Sweden.

Pristiphora compressa (Hartig, 1837)

Figs 158–159, 284

Nematus compressus Hartig, 1837: 213–214. Lectotype ♀ (GBIF-GISHym3223; here designated) in ZSM, examined. Type locality: Harz, Germany.

Similar species

The most similar species are P. abietina and P. decipiens. Pristiphora abietina is on average smaller (4.0–6.5 vs 6.0–9.0 mm) and has a posteriorly sinuate hypopygium (excised in P. compressa). Pristiphora decipiens tends to be paler (with extensively pale mesepisternum, while in P. compressa it is usually black). Lancets and penis valves of these species can also be distinguished (see the Key).

Genetic data

Based on COI barcode sequences, P. compressa belongs to the same BIN cluster (BOLD:AAK9450) as P. abietina, P. decipiens, P. gerula, P. pseudodecipiens, P. saxesenii, and possibly P. robusta (Fig. 5). Maximum distance within the BIN is 2.18% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9450, diverging by a minimum of 3.92%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, within species divergence is 0.0% (based on three specimens and NaK or TPI) and the nearest neighbour is 1.3% different (P. abietina, only NaK).

Host plants

Picea sp. (Beneš and Krístek 1979), P. pungens Engelm. (Kula et al. 2016).

Distribution and material examined

Palaearctic. Specimens studied are from Germany, Slovakia, and Sweden.

Pristiphora decipiens (Enslin, 1916)

Figs 63, 116–117, 160, 285

Lygaeonematus compressus var. decipiens Enslin, 1916: 499. Lectotype ♀ (GBIF-GISHym3259; here designated) in ZSM, examined. Type locality: Mül. [abbreviation for an unidentified German locality].

Similar species

The most similar species is P. compressa, which tends to be darker (usually has a black mesepisternum, while in decipiens it is extensively pale), and has a different lancet and penis valve (see the Key).

Genetic data

Based on COI barcode sequences, P. decipiens belongs to the same BIN cluster (BOLD:AAK9450) as P. abietina, P. compressa, P. gerula, P. pseudodecipiens, P. saxesenii, and possibly P. robusta. Maximum distance within the BIN is 2.18% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9450, diverging by minimum of 3.92%, is BOLD:ACO1401 (P. euxantha). No nuclear data are available.

Host plants

Picea sp. (Beneš and Krístek 1979), P. pungens Engelm. (Kula et al. 2016).

Distribution and material examined

West Palaearctic. Specimens studied are from Finland, Germany, and Slovakia.

Pristiphora gerula (Konow, 1904)

Figs 118, 122, 161, 286

Lygaeonematus gerulus Konow, 1904a: 194, 199–200. Lectotype ♂ (GBIF-GISHym3898; here designated) in SDEI, examined. Type locality: Kalkhorst, Mecklenburg-Vorpommern, Germany.

Similar species

The most similar species is P. saxesenii, valvula 3 of which possesses a small scopa in posterior view (completely absent in P. gerula) and which usually has a dark brown pterostigma (usually pale in P. gerula). Males are best recognised by examining penis valves (see the Key).

Genetic data

Based on COI barcode sequences, P. gerula belongs to the same BIN cluster (BOLD:AAK9450) as P. abietina, P. compressa, P. decipiens, P. pseudodecipiens, P. saxesenii, and possibly P. robusta (Fig. 5). Maximum distance within the BIN is 2.18% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9450, diverging by minimum of 3.92%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data (one specimen and TPI), the nearest neighbour is 1.2% different (P. abietina).

Host plants

Picea sp. (Beneš and Krístek 1979), P. pungens Engelm. (Kula et al. 2016).

Distribution and material examined

West Palaearctic. Specimens studied are from Estonia, France, Germany, Slovakia, and Sweden.

Pristiphora pseudodecipiens Beneš & Křístek, 1976

Figs 62, 163, 287

Pristiphora pseudodecipiens Beneš & Křístek, 1976: 404–414. Holotype ♀ (Cat. No. 26 237) in NMPC, not examined. Kuničky, South Moravian Region, Czech Republic.

Similar species

The most similar species is P. decipiens, females of which can be distinguished by having a posteriorly excised hypopygium (not excised in P. pseudodecipiens). Males are best recognised by examining penis valves.

Genetic data

Based on COI barcode sequences, P. pseudodecipiens belongs to the same BIN cluster (BOLD:AAK9450) as P. abietina, P. compressa, P. decipiens, P. gerula, P. saxesenii, and possibly P. robusta (Fig. 5). Maximum distance within the BIN is 2.18% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9450, diverging by minimum of 3.92%, is BOLD:ACO1401 (P. euxantha). No nuclear data available.

Host plants

Picea abies (L.) Karsten (Beneš and Krístek 1976), P. pungens Engelm. (Kula et al. 2016).

Distribution and material examined

West Palaearctic. Specimens studied are from Czech Republic, Germany, and Ukraine.

Pristiphora robusta (Konow, 1895)

Figs 26, 164, 289

Lygaeonematus robustus Konow, 1895: 53. Holotype ♀ (GBIF-GISHym3897) in SDEI, examined. Type locality: Bohemia, Czech Republic.

Similar species

Combination of the structure of valvula 3, nearly completely black coloration, and the large size (9–11 mm) enable easy identification of the females. Males are best recognised by examining penis valves. Interestingly, a reared male specimen from Finland (http://id.luomus.fi/GL.5198) has vein 2r-rs present on both wings.

Genetic data

No data.

Host plants

Picea abies (L.) Karsten (Perkiömäki 1969, Kajmuk 1988), P. obovata Ledeb. (Kajmuk 1988), P. pungens Engelm. (Kula et al. 2016).

Distribution and material examined

Palaearctic. Specimens studied are from Czech Republic, Finland, and Sweden.

Pristiphora saxesenii (Hartig, 1837)

Figs 60, 120–121, 162, 288

Nematus (Nematus) hospes Dahlbom, 1835b: 9. Not available. Nomen nudum.

Nematus saxesenii Hartig, 1837: 212–213. Lectotype ♀ (GBIF -GISHym3409; here designated) in ZSM, examined. Type locality: Harz, Germany.

Pristiphora thalenhorsti Wong, 1975: 453–454. Holotype ♂ in CNC, not examined. Type locality: Sieber, Lower Saxony, Germany. Synonymised with P. saxesenii by Beneš and Krístek (1976).

Similar species

The most similar species is P. gerula, valvula 3 of which lacks a scopa (present in posterior view in P. saxesenii) and which usually has a pale pterostigma (usually dark brown in P. saxesenii). Males are best recognised by examining penis valves (see the Key).

Genetic data

Based on COI barcode sequences, P. saxesenii belongs to the same BIN cluster (BOLD:AAK9450) as P. abietina, P. compressa, P. decipiens, P. gerula, P. pseudodecipiens, and possibly P. robusta (Fig. 5). Maximum distance within the BIN is 2.18% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9450, diverging by minimum of 3.92%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data (one specimen), the nearest neighbour is 1.1% different (P. gerula, only TPI).

Host plants

Picea abies (L.) Karste (Pschorn-Walcher and Altenhofer 2000, ex ovo rearing experiments by VV), P. pungens Engelm. (Kula et al. 2016).

Rearing notes

Ovipositing experiment no.19/1987: Finland, South Häme, Hämeenlinna (uniform grid reference 676:336). On 5–6.VI.1987 one captured female laid eggs on new needles of Picea abies, the eggs were on outer margin of needle near its middle and entirely visible. Larvae hatched on 9.VI.1987. Four to five larval instars were observed, the development of larvae was rapid and on 21.VI.1987 prepupae were seen. No extra moult after feeding.

Distribution and material examined

West Palaearctic. Specimens studied are from Czech Republic, Estonia, Germany, Slovakia, and Sweden.

Pristiphora alpestris group

Pristiphora alpestris (Konow, 1903)

Figs 65, 88, 203, 233, 235

Pachynematus alpestris Konow, 1903: 380 (key). Lectotype ♀ (GBIF-GISHym3921; here designated) in SDEI, examined. Type locality: Switzerland.

Lygaeonematus karvoneni Lindqvist, 1952: 116–117, syn. n. Holotype ♂ (DEI-GISHym20888) in MZH, examined. Type locality: Pasila-Moor, Helsinki, Finland.

Similar species

The most similar species is P. pseudocoactula, from which it can be distinguished by usually having paler coloration (yellow clypeus, labrum, metafemur, and valvula 3) and different lancet and penis valve (see the Key). The holotype of karvoneni Lindqvist is not separable from P. alpestris (Figs 233, 235) and the figure of the penis valve given for karvoneni by Lindqvist (1952) is misleading: the valvispina is drawn close to the ventral margin of paravalva (Fig. 51 in Lindqvist 1952), while in alpestris and in the holotype of karvoneni it is distinctly removed from the ventral margin (Figs 233, 235). Furthermore, the reported host plant (Betula pendula) of karvoneni is also used by P. alpestris (Kangas, 1985).

Genetic data

Based on COI barcode sequences, P. alpestris belongs to the same BIN cluster (BOLD:AAK9467) as P. pseudocoactula (Fig. 5). Maximum distance within the BIN is 1.62% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9467, diverging by a minimum of 4.69%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data (one specimen; NaK), the nearest neighbour is 0.7% different (P. pseudocoactula).

Host plants

Betula pendula Roth (Adam 1973, Kangas 1985, ex ovo rearing experiments by VV), Betula pubescens Ehrh. (Kangas, 1985).

Rearing notes

Ovipositing experiment no. 3/1983: Finland, Janakkala, Kalpalinna. One captured female laid eggs on 7.V.1983 in pockets on undersides of the young leaves of Betula pendula, the egg pockets are attached to side veins. Larvae hatched on 11.V.1983, four larval instars were observed. They grew rapidly and prepupae were seen on 20.V.1983 and the next days. No extra moult after feeding.

Distribution and material examined

West Palaearctic. Specimens studied are from Finland, Germany, Sweden, and Switzerland.

Pristiphora dissimilis Lindqvist, 1971

Fig. 237

Pristiphora dissimilis Lindqvist, 1971: 13–14. Holotype ♂ (DEI-GISHym20906) in MZH, examined. Type locality: Helsinge, Uusimaa, Finland.

Similar species

Possibly belongs to the alpestris group. Only the holotype male is known, which can easily be distinguished from other species by its distinct penis valve (Fig. 237).

Genetic data

No data.

Host plants

Unknown.

Distribution and material examined

West Palaearctic. One male specimen studied from Finland.

Pristiphora pseudocoactula (Lindqvist, 1952)

Figs 89–90, 204, 234, 236

Lygaeonematus pseudocoactulus Lindqvist, 1952: 115–116. Holotype ♀ not found in MZH. Type locality: Utsjoki, Lapland, Finland.

Lygaeonematus concolor Lindqvist, 1952: 117–118, syn. n. Holotype ♀ (DEI-GISHym31689) in MZH, examined. Type locality: Kilpisjärvi, Lapland, Finland.

Similar species

The most similar species is P. alpestris, from which it can be distinguished by usually having darker coloration (black clypeus, labrum, metafemur, and valvula 3) and different lancet and penis valve (see the Key). The differences Lindqvist (1952) mentioned between concolor and pseudocoactula (coloration of labrum, pronotum, metafemur, and the amount of setae on the lancet) are small and not reliable, because the variation is continuous.

Genetic data

Based on COI barcode sequences, P. pseudocoactula belongs to the same BIN cluster (BOLD:AAK9467) as P. alpestris (Fig. 5). Maximum distance within the BIN is 1.62% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAK9467, diverging by minimum of 4.69%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum within species divergence is 0.4% (based on four specimens and TPI) and the nearest neighbour is 0.7% different (P. alpestris, only NaK).

Host plants

Betula pubescens Ehrh. (ex ovo rearing experiments by VV). Possibly also B. nana L. (incomplete ex ovo rearing experiments by VV) and Salix sp. (reared ex larva by Lindqvist 1952, as P. concolor).

Rearing notes

Rearing no. 56/1970 as P. concolor: Saana, Finnish Lapland. On 5.VII.1970 several larvae were found on Betula pubescens var. pumila. Next year one female emerged and buds of B. pubescens were offered her. She laid several eggs in pockets on underside of young leaves, egg pockets were near veins rather close to leaf margin.

Ovipositing experiment no. 12/1972 as P. concolor: Finland, North Karelia, Kontiolahti, Venejoki. One captured female laid two eggs in pockets on undersides of young leaves of Betula nana, young larvae died for technical reasons (food too old).

Ovipositing experiment 15/1972 as P. pseudocoactula: Finland, North Karelia, Kontiolahti, Venejoki. On 2.VI.1972 one captured female laid one egg in pocket near the midvein on underside of young leaf of Betula pubescens. She laid no eggs on Betula nana or Salix aurita. Four larval instars were observed and prepupa was seen on 16.VI.1972. No extra moult after feeding.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Finland and Sweden.

Pristiphora carinata group

Pristiphora albilabris (Boheman, 1852)

Figs 210, 238

Nematus (Nematus) albilabris Dahlbom, 1835b: 7. Not available. Nomen nudum.

Nematus albilabris Boheman, 1852: 172. Syntype(s) not found in NHRS. Type locality: Wärnaby and Anneberg, Småland, Sweden.

Nematus albilabris Thomson, 1863: 622. Primary homonym of Nematus albilabris Boheman, 1852 [= Pristiphora albilabris (Boheman, 1852)]. Holotype ♀ (NHRS-HEVA000003749) in NHRS, examined. Type locality: Bohuslän, Sweden.

Nematus collaris Stein, 1884: 305–308. 4 ♀ syntypes possibly in BMNH, not examined. Type locality: Chodov, Czech Republic. Synonymised with Lygaeonematus albilabris by Konow (1892).

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Betula sp. (Conde 1934, Benson 1958), B. pubescens Ehrh (ex ovo rearing experiments by VV).

Rearing notes

Ovipositing experiment no. 16/1972: Finland, North Karelia, Kontiolahti, Venejoki. One captured female laid eggs on 2–3.VI.1972 in pockets on underside of teeth (leaf edges) of young leaves of Betula pubescens. Larvae hatched on 8–9.VI.1972, four larval instars were observed. The final feeding instar lasted about 17 days. Rather small prepupae (male sex?) were seen on 4.VII.1972. No extra moult after feeding.

Distribution and material examined

Palaearctic. Specimens studied are from Finland and Sweden.

Pristiphora borea (Konow, 1904)

Fig. 212

Nematus astutus Cameron, 1885: 77–78. Nomen oblitum. Syntypes (♀♂) have not been found (Lindqvist 1952). Type locality: not stated. Synonymised with P. borea by Liston et al. (2006).

Lygaeonematus boreus Konow, 1904: 196–197 (key). Nomen protectum. See Liston, Taeger and Blank (2006). Syntype ♂ (GBIF-GISHym3908) in SDEI, examined (severely damaged, abdomen missing). Type locality: Kanin Peninsula, Nenets Autonomous Okrug, Russia, and Vassijaure, Torne Lappmark, Sweden.

Pachynematus lapponicus Enslin, 1916: 462–463. 2 ♀ and 1 ♂ syntypes in ZSM, examined. Type locality: Lappland, Europe. Synonymised with Lygaeonematus boreus by Lindqvist (1952).

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Betula nana L. (Bland and Liston 1999, ex ovo rearing experiments by VV).

Rearing notes

Ovipositing experiment no. 10/1970: Finland, North Karelia, Kontiolahti, Selkie. On 28–29.V. 1970 one captured female laid several eggs in pockets near leaf-margin on underside of the young leaves of Betula nana. Larvae hatched on 2.VI.1970 and 6 larval instars were observed; larvae eat margins of leaves. Last instar was some 30–40 days long. No “extra” moult after feeding, prepupae were seen on 27.VII.–12.VIII.1970.

Ovipositing experiment no. 15/1970: Finland, North Karelia, Tuupovaara. One captured female laid several eggs on leaves of Betula nana; she did not lay any eggs on Vaccinium uliginosum. Larvae developed, as in earlier experiment.

Ovipositing experiments 17/1970 and 18/1970: Finland, North Karelia, Kontiolahti, Venejoki. Two females laid many eggs on Betula nana, but respectively no and only two eggs on Vaccinium uliginosum. Otherwise very similar results as in previous two experiments.

Ovipositing experiments no. 4/1988, 5/1988, 6/1988, and 7/1988: Finland, South Häme, Janakkala, Suurisuo. Four captured females laid eggs on 26–27.V.1988 in pockets on undersurface of leaves of Betula nana, four larval instars were observed in all rearings and last instar lasted long.

Larvae observed on Betula pubescens var. pumila (L.) Govaerts (=Betula pubescens ssp. czerepanovii) in Saana, Finnish Lapland on 27.VII.1971.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Finland and Russia (Nenets Autonomous Okrug).

Pristiphora breadalbanensis (Cameron, 1882)

Fig. 214

Nematus breadalbanensis Cameron, 1882: 531–532. Syntypes possibly in BMNH, not examined. Type locality: Scotland, United Kingdom.

Lygaeonematus tromsöensis [sic!] Kiær, 1898: 48–49. 2 ♀ syntypes in TROM, not examined. Type locality: Flöifjeld and Mauken, Troms, Norway. Synonymised with Lygaeonematus breadalbanensis by Lindqvist (1952).

Lygaeonematus corpulentus Konow, 1904a: 196 (key). Syntype ♀ (GBIF-GISHym3909) in SDEI, examined. Type locality: Norway and Sweden (Konow 1904b).

Lygaeonematus arcticola Enslin, 1916: 514–515.1 ♀ and 1 ♂ syntype in ZSM, examined. Type locality: North Ural, Russia.

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Unknown.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Norway, Russia (North Ural), and United Kingdom.

Pristiphora carinata (Hartig, 1837)

Figs 98–99, 216, 239

Tenthredo pallipes Fallén, 1808: 110–111. Primary homonym of Tenthredo pallipes Spinola, 1808 [= Ametastegia (Protemphytus) pallipes (Spinola, 1808)]. Syntype ♀ in MZLU, not examined. Type locality: Västergötland, Sweden.

Nematus carinatus Hartig, 1837: 199–200. Syntype ♂ (GBIF-GISHym4689) in NFVG, examined. Type locality: Harz, Germany.

Nematus canaliculatus Hartig, 1840: 23, syn. n. Lectotype ♀ (GBIF-GISHym4691; here designated) in NFVG, examined. Type locality: Clausthal-Zellerfeld, Lower Saxony, Germany.

Nematus denudatus Hartig, 1840: 23. Lectotype ♂ (DEI-GISHym80268; here designated) in ZSM, examined. Type locality: not stated.

Pachynematus alticola Enslin, 1916: 463. Lectotype ♀ (GBIF-GISHym3192; here designated) in ZSM, examined. Type locality: Praděd (Altvater), Czech Republic.

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Vaccinium myrtillus L. (ex ovo rearing experiments by VV).

Rearing notes

Ovipositing experiment no. 3/1986: Finland, South Häme, Janakkala, Kalpalinna. On 22.V.1986 one captured female laid 5 eggs in pockets near leaf-margin on underside of the leaves of Vaccinium myrtillus. Larvae hatched on 27–28.V.1986, 5 larval instars were observed. Larvae eat margins of shoots and margins of leaves near their bases. Last instar was more than 16 days long. No “extra” moult after feeding. Larvae were put in alcohol on 27.VI.1986.

Distribution and material examined

Palaearctic. Specimens studied are from Czech Republic, Finland, and Germany.

Pristiphora coactula (Ruthe, 1859)

Fig. 217

Nematus coactulus Ruthe, 1859: 307–308. Holotype ♀ in NMW, not examined. Type locality: Iceland.

Nematus winnipeg Norton, 1867b: 198. Lectotype ♀ in ANSP (designated by Cresson 1928, spelt as “winnepeg”), not examined. Type locality: Lake Winnipeg, Manitoba, Canada. Synonymised with P. coactula by Benson (1962).

Nematus alpinus Thomson, 1871: 98. Lectotype ♀ (MZLU2014449; here designated) in MZLU, examined. Type locality: Skalstugan, Jämtland, Sweden.

Nematus winnipegensis W.F. Kirby, 1882: 143. Replacement name for Nematus winnipeg Norton, 1867.

Lygaeonematus pachyvalvis Konow, 1904a: 197 (key). Syntype ♀ (GBIF-GISHym3913) in SDEI, examined. Type locality: Mt Dore, France; Kanin Peninsula, Nenets Autonomous Okrug, Russia; and Vassijaure, Torne Lappmark, Sweden.

Pristiphora dawsoni Rohwer, 1908: 109. Holotype ♀ in NSM (Smith 1983), not examined. Type locality: Ute Creek, Costilla, Colorado, USA. Synonymised with P. coactula by Smith (1979).

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Vaccinium uliginosum L (ex ovo rearing experiments by VV).

Rearing notes

Ovipositing experiment no. 46/1969: Finland, North Karelia, Kontiolahti, Venejoki. On 14–15.VI.1969 one captured female laid several eggs in pockets near leaf-margin on underside of the young leaves of Vaccinium uliginosum. Larvae hatched on 20.VI.1969, 6 larval instars were observed. Larvae fed on the margins of leaves at night. During the day they hid near the bottom of the rearing container. The final feeding instar lasted some 30 days. No “extra” moult after feeding, prepupae were seen on 7.VIII.1969.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Finland, France, and Sweden.

Pristiphora groenblomi (Lindqvist, 1952)

Figs 10, 211

Lygaeonematus (Lygaeotus) grönblomi [sic!] Lindqvist, 1952: 101. Note. The spelling groenblomi is in predominant usage, not gronblomi, which would be correct according to Article 32.5.2 (ICZN 1999). Following Article 33.3.1, the predominantly used spelling is maintained. Holotype ♀ (DEI-GISHym31602) in MZH, examined. Type locality: Luumäki, South Karelia, Finland.

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Perhaps Betula sp. (Verzhutskii 1981), if the species was correctly identified.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Finland.

Pristiphora lativentris (Thomson, 1871)

Figs 95–96, 215

Nematus lativentris Thomson, 1871: 99–100 Lectotype ♀ (DEI-GISHym31596; here designated) in MZLU, examined. Type locality: Kälahög, Jämtland, Sweden.

Nematus scoticus Cameron, 1881: 563–564. Syntype(s) ♀ possibly in BMNH, not examined. Type locality: Braemar, Aberdeenshire, Scotland, United Kingdom. Synonymised with Lygaeonematus lativentris by Lindqvist (1952).

Nematus extremus Holmgren, 1883: 148. Syntypes ♂♀ in NHRS, not examined. Type locality: Matotschkin Scharr, Novaja Zemlya, Russia. Synonymised with Lygaeonematus lativentris by Lindqvist (1952).

Pristiphora bucoda Kincaid, 1900: 350–351. Syntype ♀ (USNMENT00778160) in USNM, not examined. Type locality: Berg Bay and Sitka, Alaska, USA. Synonymised with Pristiphora lativentris by Benson (1962).

Lygaeonematus alpicola Konow, 1904: 197 (key). Syntypes ♂♀ lost (Lindqvist 1952). Type locality: Austria and Switzerland. Tentatively synonymised with Lygaeonematus lativentris by Lindqvist (1952).

Lygaeonematus pallipes var. femoralis Zirngiebl, 1953: 32. Holotype ♂ (GBIF-GISHym3275) in ZSM, not examined. Type locality: Ammergau Alps, Bavaria, Germany. Synonymised with P. lativentris by Blank (1996).

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Unknown.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Norway and Sweden.

Pristiphora trochanterica (Lindqvist, 1952)

Fig. 213

Lygaeonematus (Lygaeotus) trochantericus Lindqvist, 1952: 101–102. Holotype ♀ (DEI-GISHym31566) in MZH, examined. Type locality: Utsjoki, Outakoski, Finland.

Similar species

Species limits in the carinata group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the carinata group are divided between four BIN clusters (BOLD:AAF4995, BOLD:ABU8603, BOLD:ACL1744, BOLD:ACL1745), which form a monophyletic group (Fig. 5). Minimal distances between these clusters are 1.2%–4.48%. Because of unresolved taxonomy, it is not yet clear how different species are divided among these BIN clusters. Based on nuclear data, maximum divergence within the group is 2.2% (based on ten specimens and TPI) and the nearest neighbour is 1.5% (P. pseudocoactula, both genes combined) or 1.0% different (P. wesmaeli, only NaK).

Host plants

Unknown.

Distribution and material examined

West Palaearctic. Specimens studied are from Finland.

Pristiphora depressa group

Pristiphora depressa (Hartig, 1840)

Figs 24, 82–83, 184

Nematus depressus Hartig, 1840: 24. Lectotype ♀ (designated by Liston and Späth 2008) in ZSM, examined. Type locality: not stated.

Pristiphora Carpentieri [sic!] Konow, 1902: 18. Lectotype ♀ (GBIF-GISHym3902; designated by Oehlke and Wudowenz 1984), examined. Type locality: Amiens, Nord-Pas-de-Calais-Picardie, France.

Similar species

The most similar species are P. subbifida and P. tetrica. Pristiphora subbifida has an extensively pale head in dorsal view (with only small pale spots in P. depressa) and at least partly pale metapostnotum (Fig. 38) (black in P. depressa). Pristiphora tetrica, on the other hand, has a darker head than P. depressa (with black supraclypeal area and usually without pale spots dorsally in P. tetrica). For more details see Liston and Späth (2008) and Liston et al. (2013). Male unknown.

Genetic data

Based on COI barcode sequences, P. depressa forms its own BIN cluster (BOLD:ABU7027) (Fig. 3). Maximum distance within the BIN is 0.61%. The nearest neighbour to BOLD:ABU7027, diverging by minimum of 7.06%, is BOLD:AAU8503 (P. subbifida).

Host plants

Acer sp. (Carpentier 1901). Adults reared from the larvae mentioned by Carpentier (1901) were described a year later as P. carpentieri by Konow (1902).

Distribution and material examined

West Palaearctic. Specimens studied are from France, Germany, Italy, and Sweden.

Pristiphora subbifida (Thomson, 1871)

Figs 28, 38, 185

Nematus subbifidus Thomson, 1871: 105. Lectotype ♀ in MZLU (designated by Lindqvist 1954), examined. Type locality: Östergöthland, Sweden.

Similar species

The most similar species is P. depressa, which has a black metapostnotum (at least partly pale in P. subbifida: Fig. 38) and head in dorsal view with small pale spots (extensively pale in P. subbifida). See Liston and Späth (2008) and Liston et al. (2013) for more details. Male unknown.

Genetic data

Based on COI barcode sequences, P. subbifida forms its own BIN cluster (BOLD:AAU8503) (Fig. 3). Maximum distance within the BIN is 2.29%. The nearest neighbour to BOLD:AAU8503, diverging by a minimum of 6.26%, is BOLD:ABU7027 (P. depressa). Based on nuclear data (one specimen and both genes combined), the nearest neighbour is 2.1% different (P. tetrica).

Host plants

Acer campestre L. (Liston 1996, Liston et al. 2013). No other Acer species has been unequivocally recorded as a host, because of previous confusion of P. subbifida with other closely related species.

Distribution and material examined

West Palaearctic, (Nearctic, see Smith 2016). Specimens studied are from Germany, Greece, and Sweden.

Pristiphora tetrica (Zaddach, 1883)

Figs 186, 301

Nematus tetricus Zaddach in Brischke, 1883b: 148–149. Holotype ♀ possibly destroyed (Blank and Taeger 1998). Type locality: Thüringen, Germany.

Nematus velatus Zaddach in Brischke, 1883b: 149. Holotype ♀ was not found in ZSM (Liston and Späth 2008). Type locality: Baiern, Germany. Synonymised with P. tetrica by Konow (1902).

Pristiphora nievesi Haris, 2004: 164–165. Holotype ♀ (DEI-GISHym20461) in MNCN, examined. Type locality: El Ventorillo, Madrid, Spain.

Similar species

The most similar species are P. depressa and P. subbifida, from which it can be distinguished by having a black supraclypeal area (pale in P. depressa and P. subbifida) and usually black head in dorsal view (at least with small pale spots in P. depressa). See Liston and Späth (2008) for more details.

Genetic data

Based on COI barcode sequences, specimens of this species are divided between five BIN clusters (BOLD:ACL2117, BOLD:ABA3515, BOLD:ACL2098, BOLD:ACL2099, BOLD:ACL2100) (Fig. 3), four of which (i.e. not BOLD:ACL2117) were previously identified as P. nievesi Haris. Minimum distances between these clusters are 1.72%–1.90%. All these clusters form a monophyletic group (Fig. 3) and we treat them as one species, because there is a continuous variation in external morphological characters used to separate P. nievesi from P. tetrica, and no clear differences in penis valves and lancets (see Liston et al. 2015). Based on nuclear data (two specimens and both genes combined), within species divergence is 0.2% and the nearest neighbour is 1.7% different (P. cretica Schedl, 1981, a species not treated here).

Host plants

Acer pseudoplatanus L. (Macek 2012b) and A. sempervirens L. (Liston et al. 2015).

Distribution and material examined

West Palaearctic. Specimens studied are from France, Germany, Greece, Italy, Morocco, Russia (Karachay-Cherkess Republic), and Spain.

Pristiphora erichsonii group

Pristiphora erichsonii (Hartig, 1837)

Figs 35, 124, 170, 290

Nematus Leachii [sic!] Dahlbom, 1835a: 27–28. Suppressed. Note. Suppressed for the purposes of the Law of Priority but not for those of the Law of Homonymy. (Opinion 906, ICZN 1970). Described from larvae, no type probably available. Type locality: Skåne (Sweden) and Zealand (Denmark). Synonymised with Nematus erichsonii by Thomson (1863).

Nematus (Nematus) leachei Dahlbom, 1835b: 10. Not available. Nomen nudum.

Nematus erichsonii Hartig, 1837: 187–188. Holotype ♀ (GBIF-GISHym3272) in ZSM, examined. Type locality: Harz, Germany.

Nematus notabilis Cresson, 1880: 7. Lectotype ♀ in ANSP (designated by Cresson 1916), not examined. Type locality: Massachusetts, USA. Synonymised with P. erichsonii by Marlatt (1896).

Similar species

Females can be distinguished from other Pristiphora species relatively easily by the shape of the valvula 3 (Fig. 124) and the colour of the abdomen (having a red band). Likewise, it should be possible to distinguish males (which are extremely rare) from other species by the dorsal red band on the abdomen and extensively pale antenna.

Genetic data

Based on COI barcode sequences (only one specimen in BOLD), P. erichsonii forms its own BIN cluster (BOLD:AAG8325). The nearest neighbour to BOLD:AAG8325, diverging by a minimum of 5.04%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data (one specimen and NaK), the nearest neighbour is 0.7% different (P. parva).

Host plants

Larix decidua Mill. (Drooz 1960, Kangas 1985, Pschorn-Walcher and Altenhofer 2000), L. kaempferi (Lamb.) Carrière (Drooz 1960, Huflejt and Sawoniewicz 1999, Pschorn-Walcher and Altenhofer 2000), L. sibirica Ledeb. (Drooz 1960, Kangas 1985), L. laricina (Du Roi) K. Koch (Drooz 1960, Jardon et al. 1994), L. occidentalis Nutt. (Drooz 1960), L. principis-rupprechtii (Mayr) Pilger (Li et al. 2013).

Distribution and material examined

Palaearctic, (Nearctic). Specimens studied are from Finland, Germany, and Norway.

Pristiphora glauca Benson, 1954

Figs 126–127, 179, 291

Pachynematus laricivorus Takagi, 1931: 28–32 (Jap.), 8–11(Engl.). Secondary homonym of Nematus laricivorus Brischke, 1883a [= Pristiphora laricis (Hartig, 1837)]. Syntypes possibly in the National Institute of Forest Science (previously Forestry Experiment Station), Seoul, South Korea (Wong 1975), not examined. Type locality: North Korea. Synonymised with P. glauca by Vikberg (1975).

Pristiphora glauca Benson, 1954a: 113–114. Holotype ♀ in BMNH, not examined. Type locality: Mortimer Forest, Hereford, England, United Kingdom.

Pristiphora takagii Wong, 1975: 459. Replacement name for Pachynematus laricivorus Takagi, 1931.

Similar species

The most similar species is P. wesmaeli. The differences in adults are small and might not be always reliable. According to Benson (1958), the ovipositor is about 1.1 times as long as the protibia in P. glauca (0.9 times in P. wesmaeli). For males, the differences in penis valves are also very slight (see the Key). The differences in larval coloration, and earlier emergence of adults and earlier larval feeding period of P. glauca distinguish the species more reliably (Benson, 1954a).

Genetic data

Based on COI barcode sequences, P. glauca belongs to the same BIN cluster (BOLD:ABY3989) as P. wesmaeli (Fig. 5). Maximum distance within the BIN is 2.17% and minimum between species distance is possibly 0.00%. The nearest neighbour to BOLD:ABY3989, diverging by minimum of 3.75%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data (one specimen), the nearest neighbour is 0.1% (only NaK) or 0.9% (only TPI) different (P. wesmaeli).

Host plants

Larix decidua Mill. (Kirkland and Styles 1955, Pschorn-Walcher and Altenhofer 2000), L. kaempferi (Lamb.) Carrière (Takagi 1931, Kirkland and Styles 1955, Pschorn-Walcher and Altenhofer 2000), L. sibirica Ledeb. (Verzhutskii 1966), Larix gmelinii (Rupr.) Kuzen. (Takagi 1931).

Distribution and material examined

Palaearctic. Specimens studied are from Germany and Russia (Primorsky Krai).

Pristiphora wesmaeli (Tischbein, 1853)

Figs 180, 292

Nematus Wesmaeli [sic!] Tischbein, 1853: 347–348. 34 ♀♀ and 32 ♂♂ syntypes probably destroyed (Horn et al. 1990). Type locality: Herrstein, Rhineland-Palatinate, Germany.

Nematus solea Snellen van Vollenhoven, 1870: 59–60. Holotype ♂ in ZMAN (Thomas 1987), not examined. Type locality: not stated. Synonymised with Lygaeonematus wesmaeli by Konow (1904a).

Similar species

The most similar species is P. glauca. The differences in adults are small and might not always be clear. According to Benson (1958), the ovipositor is about 0.9 times as long as the protibia in P. wesmaeli (1.1 times in P. glauca). For males, the differences in penis valves are also very slight (see the Key). The differences in larval coloration, later emergence of adults, and later larval feeding period distinguish P. wesmaeli from P. glauca more reliably (Benson, 1954a).

Genetic data

Based on COI barcode sequences, P. wesmaeli belongs to the same BIN cluster (BOLD:ABY3989) as P. glauca (Fig. 5). Maximum distance within the BIN is 2.17% and minimum between species distance is possibly 0.00%. The nearest neighbour to BOLD:ABY3989, diverging by a minimum of 3.75%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data (two specimens), the nearest neighbour is 0.1% (only NaK) or 0.9% (only TPI) different (P. glauca).

Host plants

Larix decidua Mill., Larix gmelinii (Rupr.) Rupr., L. kaempferi (Lamb.) Carrière, Larix × eurolepis Henry (Huflejt and Sawoniewicz 1999), and L. sibirica Ledeb. (Kangas 1985).

Distribution and material examined

Palaearctic. One male specimen studied from France. The second specimen mentioned under Genetic data is from a larva (voucher 3c, Belgium, collected on 7.VIII.2000) that we have not studied.

Pristiphora laricis group

Pristiphora friesei (Konow, 1904)

Figs 33, 79–81, 174–176, 277

Lygaeonematus Friesei [sic!] Konow, 1904a: 195, 208. Lectotype ♀ (GBIF-GISHym3914; here designated) in SDEI, examined. Type locality: Airolo, Ticino, Switzerland (not in Tirol, Austria, as written by Konow 1904a, b; see also Enslin, 1916: 509). Note. Enslin (1916: 509) and Muche (1974: 136–137) referred to the single female specimen in the SDEI as the type of P. friesei (under the name P. frisei [sic!] by Muche 1974). Neither reference constitutes a valid lectotype designation according to ICZN Article 74.5, because the specimen was not unambiguously selected from the syntype series. However, because Enslin and Muche based their concept of the species on the specimen in the SDEI, we hereby designate this specimen as the lectotype.

Pristiphora atrata Lindqvist, 1975: 13, syn. n. Holotype ♀ (DEI-GISHym20834) in MZH, examined. Type locality: Goloustnoje, Irkutsk, Russia. Note. The holotype was reared by B. N. Verzhutskii from a larva found on Vaccinium uliginosum L. on May 29, 1966 (label data; see also Verzhutskii 1981), which we interpret as an accidental find, as the host plant of P. friesei is Larix (Schedl 1976, Liston et al. 2006). Lancet and external morphology of atrata does not differ from friesei.

Similar species

The most similar species is P. laricis, which tends to be paler, but darker specimens could be mistaken for P. friesei. There are small differences in the structure of the lancets: the tangium appears to be without campaniform sensilla and there are fewer setae in P. friesei, while campaniform sensilla are present and there are more setae in P. laricis (Figs 174–178). There appear to be no consistent differences in penis valves (Figs 277, 279–280), contrary to Chevin (1974), but the antennae are longer in P. friesei (see the Key).

Genetic data

Based on COI barcode sequences, P. friesei forms its own BIN cluster (BOLD:ABV3411). When specimens that have 1 bp deletion in the barcoding region (e.g. DEI-GISHym11558) are included (excluded from calculations in BOLD), maximum distance within P. friesei is 3.5% and the nearest P. laricis specimens are only 2.3% different (based on full barcodes), but P. friesei nevertheless forms a monophyletic group (Fig. 3). The nearest neighbour to BOLD:ABV3411 (excluding the specimens with 1 bp deletion), diverging by minimum of 4.67%, is BOLD:AAQ3707 (P. laricis). No nuclear data are available.

Host plants

Larix decidua Mill. (Schedl 1976), Larix × eurolepis (Liston et al. 2006).

Distribution and material examined

Palaearctic. Specimens studied are from Germany, Italy, Russia (Irkutsk Oblast), Switzerland, and United Kingdom.

Pristiphora laricis (Hartig, 1837)

Figs 64, 177–178, 279–280

Nematus laricis Hartig, 1837: 203–204. Lectotype ♀ (GBIF-GISHym3328; here designated) in ZSM, examined. Type locality: Germany according to the title of the publication.

Nematus ruficollis Hartig, 1840: 27. Lectotype ♀ (GBIF-GISHym3405; here designated) in ZSM, examined. Type locality: not stated.

Nematus leucocnemis Förster, 1854b: 433–434. Lectotype ♀ (GBIF-GISHym3333; designated by Liston 1995) in ZSM, examined. Type locality: Aachen, North Rhine-Westphalia, Germany.

Nematus oblongus Cameron, 1882: 539. Syntype(s) possibly in BMNH, not examined. Type locality: England, United Kingdom. Synonymised with Lygaeonematus laricis by Konow (1904a).

Nematus laricivorus Brischke, 1883a: pl. I, 1e. Described from larvae, types possibly destroyed (Blank and Taeger 1998). Type locality: not stated, but probably in former East Prussia (now Kaliningrad Oblast of Russia, or Poland). Synonymised with Pristiphora laricis by Konow (1898).

Nematus rusticanus Brischke, 1884: 128–129. Holotype ♀ possibly destroyed (Blank and Taeger 1998). Type locality: not stated, but probably in former East Prussia (now Kaliningrad Oblast of Russia, or Poland). Synonymised with Lygaeonematus laricis by Konow (1904a).

Pachynematus ravidus Konow, 1903: 382 (key). Lectotype ♀ (GBIF-GISHym3855; designated by Taeger and Blank 1998) in SDEI, examined. Type locality: Zermatt, Valais, Switzerland. Synonymised with Pristiphora laricis by Koch (1989).

Lygaeonematus paedidus Konow, 1904a: 195, 205. Lectotype ♀ (GBIF-GISHym3854; designated by Koch 1989 as “holotype”) in SDEI, examined. Type locality: Ulm (Baden-Württemberg) or Erfurt (Thuringia), Germany (the specimen has two different labels, with locality names that do not match). Synonymised with Pristiphora laricis by Koch (1989).

Pachynematus nigricorpus Takagi, 1931: 32–33 (Jap.), 11–12(Engl.), syn. n. Syntypes possibly in the National Institute of Forest Science (previously Forestry Experiment Station), Seoul, South Korea, not examined. Type locality: North Korea.

Similar species

The most similar species is P. friesei, which tends to be darker (see the Key). There are small differences in the structure of the lancets: campaniform sensilla are present on the tangium and there are more setae in P. laricis, while campaniform sensilla appear to be absent and there are fewer setae in P. friesei (Figs 174–178). There appear to be no consistent differences in penis valves (Figs 277, 279–280), contrary to Chevin (1974), but the antennae are shorter in P. laricis (see the Key). One studied female from Scotland (DEI-GISHym31503) had a red band on the abdomen, like P. cincta and P. erichsonii, but all these species can be distinguished based on the shape of valvula 3 and the structure of the lancet (Figs 104–105, 145–146 for P. cincta; Figs 124–125, 170 for P. erichsonii; Figs 79–81, 177–178 for P. laricis). As already suspected by Vikberg (1975), we treat nigricorpus Takagi as synonym of laricis Hartig. The description by Takagi (1931) fits well with P. laricis (pale labrum, length of antenna in male 3–4 mm) rather than P. friesei (black or dark brown labrum and length of antenna in male about 5 mm). Haris (2006b) apparently misinterpreted P. laricis, because the figure of valvula 3 given for this species (Fig. 10 in Haris 2006b) belongs to the leucopodia group, tenuiserra or some other species where valvula 3 can extend or extends beyond cerci. However, the figure of valvula 3 (Fig. 30 in Haris 2006b) given for nigricorpus Takagi by Haris (2006b, as P. nigrocarpa) does fit with P. laricis (Fig. 79).

Genetic data

Based on COI barcode sequences, P. laricis forms its own BIN cluster (BOLD:AAQ3707). Maximum distance within the BIN is 2.09% (Fig. 3). The nearest neighbour to BOLD:AAQ3707, diverging by minimum of 4.31%, is BOLD:ABV3411 (P. friesei). Based on nuclear data (three specimens and TPI or NaK), within species divergence is 0.2% (NaK) or 1.2% (TPI) and the nearest neighbour is 2.3% different (P. nigriceps, only NaK).

Host plants

Larix decidua Mill. (Adam 1973, Huflejt and Sawoniewicz 1999), L. kaempferi (Lamb.) Carrière (Pschorn-Walcher and Altenhofer 2000), L. sibirica Ledeb. (Kangas 1985, Huflejt and Sawoniewicz 1999), L. gmelinii (Rupr.) Kuzen., L. laricina (Du Roi) K. Koch, L. occidentalis Nutt., Larix × eurolepis A. Henry (Huflejt and Sawoniewicz 1999).

Distribution and material examined

Palaearctic. Specimens studied are from Austria, Finland, Germany, United Kingdom, Italy, Japan, Slovenia, and Switzerland.

Pristiphora leucopodia group

Pristiphora leucopodia (Hartig, 1837)

Figs 56, 181–182, 281

Nematus leucopodius Hartig, 1837: 200. Lectotype ♀ (GBIF-GISHym3336; here designated) in ZSM, examined. Type locality: Harz, Germany.

Nematus nitens Borries, 1896: 232. Primary homonym of Nematus nitens Thomson, 1888 [= Euura respondens (Förster, 1854b) comb. n.]. Syntypes ♀♂ have not been located in ZMUC (Blank et al. 2009). Type locality: Dyrehaven, Region Hovedstaden (Capital Region), Denmark. Synonymised with P. leucopodia by Blank et al. (2009).

Pachynematus sagulatus Konow, 1903: 382 (key). Lectotype ♀(GBIF-GISHym3903; designated by Oehlke and Wudowenz 1984, as “Holotypus”) in SDEI, examined. Type locality: Hungaria bor. Tatra [Tatra Mountains], Poland or Slovakia.

Lygaeonematus leucopodius ab. flavipes Lindqvist, 1941: 70. Not available. Infrasubspecific name.

Nematus (Pikonema) piceae Zhelochovtsev in Zhelochovtsev and Zinovjev, 1988: 170, syn. n. Holotype ♀ in ZMUM, not examined. Type locality: Srednyaya Usva [Средняя Усьва], Perm Krai, Russia.

Pristiphora (Pristiphora) hoverlaensis Haris, 2001: 82, syn. n. Holotype ♂ (DEI-GISHym80339; http://dx.doi.org/10.6084/m9.figshare.5091895) in HNHM, examined. Type locality: Bohdan (Богдан), Zakarpattia Oblast, Ukraine.

Similar species

The most similar species is P. nigriceps, females of which have an extensively pale thorax and abdomen. In P. leucopodia, thorax and abdomen are usually black, palest specimens have a nearly completely pale abdomen and slightly pale thorax. Males are best recognised by examining penis valves. Although we did not study the holotype of piceae Zhelochovtsev, differences mentioned by Zhelochovtsev and Zinovjev (1988) in lancet, coloration and length of antennae seem to be minute. Examination of female specimens that vary in coloration and length of valvula 3 did not allow separation of two forms, as the variation seems to be continuous. Neither were reliable differences detected in the lancet. The penis valve of one male identified by Alexey Zinovjev as piceae is indistinguishable from leucopodia.

Genetic data

Based on COI barcode sequences, P. leucopodia belongs to its own BIN cluster (BOLD:AAH7553) (Fig. 3). Maximum distance within the BIN is 1.83%. The nearest neighbour to BOLD:AAH7553, diverging by minimum of 5.06%, is BOLD:AAQ3707 (P. laricis). Based on nuclear data, maximum within species divergence is 0.9% (based on three specimens and NaK) and the nearest neighbour is 2.4% different (P. nigriceps, both genes combined).

Host plants

Picea abies (L.) Karsten (Pschorn-Walcher and Altenhofer 2000), P. pungens Engelm. (Kula et al. 2016).

Distribution and material examined

West Palaearctic. Specimens studied are from Estonia, Finland, Germany, Norway, Slovakia, Sweden, Switzerland, and Ukraine.

Pristiphora nigriceps (Hartig, 1840)

Figs 37, 53, 84–85, 183, 282

Nematus nigriceps Hartig, 1840: 24. Lectotype ♀ (GBIF-GISHym3362; here designated) in ZSM, examined. Type locality: not stated.

Nematus bistriatus Thomson, 1871: 105–106. Lectotype ♀ (NHRS-HEVA000001950; here designated) in NHRS, examined. Type locality: Östergötland (Götaland), Sweden.

Similar species

The closest species is P. leucopodia, females of which usually have a black thorax and abdomen (extensively pale in P. nigriceps). In paler specimens of P. leucopodia, at least the thorax remains relatively dark compared to P. nigriceps. Males are best recognised by examining penis valves (see the Key).

Genetic data

Based on COI barcode sequences, specimens of this species are divided between two BIN clusters (BOLD:AAO1445 and BOLD:ABV9762), which form a monophyletic group (Fig. 3). Minimal distance between these two clusters is 2.87%. Based on nuclear data, within species divergence is 0.0% (based on two specimens and NaK) and the nearest neighbour is 2.3% (P. laricis, only NaK) or 2.4% different (P. leucopodia, both genes combined).

Host plants

Picea abies (L.) Karsten (Boevé 1990, Pschorn-Walcher and Altenhofer 2000, ex ovo rearing experiments by VV).

Rearing notes

Ovipositing experiment no.20/1987: Finland, South Häme, Janakkala (uniform grid reference 6757:3376). On 6.VI.1987 one captured female laid 2 eggs on new needles of Picea abies, the eggs were on outer margin of needle near its apex and mostly inside needle. Larvae hatched on 9.VI.1987. Four larval instars were observed, the development of larvae was rapid and on 22.VI.1987 one prepupa was seen. No extra moult after feeding.

Distribution and material examined

Palaearctic. Specimens studied are from Finland, Germany, Russia (Primorsky Krai), Slovakia, Sweden, Switzerland, and Ukraine.

Pristiphora malaisei group

Pristiphora dasiphorae (Zinovjev, 1993), comb. n.

Figs 73–74, 136, 297

Pristicampus dasiphorae Zinovjev, 1993: 81–84. Holotype ♀ in ZMUM, not examined. Type locality: Tibelti, Irkutsk Oblast, Russia.

Similar species

The most similar species is P. malaisei, which can be separated based on the valvula 3, the lancet, and the penis valve (see the Key).

Genetic data

Based on COI barcode sequences, P. dasiphorae belongs to its own BIN cluster (BOLD:ACO2064). Maximum distance within the BIN is 0% (based on two specimens). The nearest neighbour to BOLD:ACO2064, diverging by minimum of 8.45%, is P. malaisei (no BIN number has been assigned yet). Based on nuclear data, the nearest neighbour is 4.3% different (P. punctifrons, both genes combined).

Host plants

Potentilla fruticosa L. (=Dasiphora fruticosa) (Zinovjev 1993, our own observations on larvae in Öland).

Distribution and material examined

Palaearctic. Specimens studied are from Russia (Zabaykalsky Krai) and Sweden (Öland).

Pristiphora malaisei (Lindqvist, 1952)

Figs 13, 30, 58, 75, 137, 298

Lygaeonematus (Lygaeophora) malaisei Lindqvist, 1952: 112–113. Holotype ♀ (NHRS-HEVA000003753) in NHRS, examined. Type locality: Torneträsk, Torne Lappmark, Sweden.

Mesoneura arctica Lindqvist, 1959: 68–70, syn. n. Holotype ♂ (http://id.luomus.fi/GL.5218) in MZH, examined. Type locality: Malla, Lapland, Finland.

Pachynematus incisus Lindqvist, 1970: 103, syn. n. Holotype ♀ in MZH (http://id.luomus.fi/GL.5183), examined. Type locality: Popovo, Oljhonsk, Irkutsk, Russia.

Pachynematus intermedius Verzhutskii, 1974: 160, syn. n. Original paper not seen, cited in Zinovjev (1993): 81. Note. Described as “Pachynematus intermedius Lqv. sp. n.” from larvae feeding on Dasiphora, type material not cited. Type locality: Baikal region, Russia.

Pristiphora mongololaricis Haris, 2003: 116–117, syn. n. Holotype ♀ (DEI-GISHym80350; http://dx.doi.org/10.6084/m9.figshare.5053627) in HNHM, examined. Type locality: Nukht, Bogd Khan Mountain, Ulaanbaatar, Mongolia.

Similar species

The most similar species is P. dasiphorae, which can be separated by differences in the valvula 3, lancet, and penis valve (see the Key). Contrary to Zinovjev (1993), we treat P. incisa (Lindqvist, 1970) as a synonym, because we are unable to find consistent differences between the arctic (P. malaisei) and non-arctic forms (P. incisa).

Genetic data

Based on COI barcode sequences, P. malaisei belongs to its own cluster (no BIN number has been assigned yet). Maximum distance within P. malaisei is 0.6% (based on full barcodes). The nearest neighbour to P. malaisei, diverging by minimum of 8.45%, is BOLD:ACO2064 (P. dasiphorae). Based on nuclear data, within species divergence is 0.4% (based on two specimens and both genes combined) and the nearest neighbour is 5.1% (P. luteipes, only TPI) or 5.3% different (P. punctifrons, both genes combined).

Host plants

Potentilla fruticosa L. (=Dasiphora fruticosa) (Lindqvist 1970, Verzhutskii 1981, ex ovo rearing experiments by VV) and possibly Comarum palustre L. (incomplete ex ovo and ex larva rearing experiments by VV).

Rearing notes

Ovipositing experiment no. 1/1994: Finland, South Häme, Janakkala, Turenki, Alanko. On 11.V.1994 two females were swept from Potentilla fruticosa, one of them in copula. On the same day they laid eggs in pockets on the underside of young leaves of P. fruticosa, the eggs were near the leaf margin. On 17.V.1994 larvae hatched. Four larval instars were observed, the development of larvae was rapid. On 31.V.1994 prepupae and three cocoons were found. No extra moult after feeding.

Ovipositing experiment no. 2/1998: the same locality as above. On 2.V.1998 one female was offered leaves of Potentilla argentea agg., in which she showed no interest. On the next day small leaves and buds of Comarum palustre were offered, and the female began immediately to lay eggs in petioles of young leaves and unopened buds. Larvae hatched but did not live many days.

Larval feeding experiment on 26.VI.1998. 11 larvae from Potentilla fruticosa were put into container with Comarum palustre and Potentilla crantzii (Crantz) Beck ex Fritsch. Within an hour 6 larvae were eating Comarum but not Potentilla. Inspection on the next day: many larvae were feeding on Comarum and many leaves of Comarum had large holes. In addition, no larvae were observed to feed on Potentilla argentea L. In the evening the larvae were put again on Potentilla fruticosa.

Larval feeding experiment in July 1998. Many rather large larvae were put into a container with Potentilla anserina L., P. argentea, and P. erecta (L.) Raeusch. After 3 hours no larva was feeding on them, but they walked around, seeking better food.

Distribution and material examined

Palaearctic. Specimens studied are from Finland, France, Mongolia, Russia (Irkutsk Oblast), and Sweden.

Pristiphora micronematica group

Pristiphora affinis (Lindqvist, 1952)

Fig. 206

Lygaeonematus (Lygaeophora) affinis Lindqvist, 1952: 109–110. Holotype ♀ (DEI-GISHym20898) in MZH, examined. Type locality: Utsjoki, Outakoski, Finland.

Similar species

Species limits in the micronematica group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the micronematica group belong to the BIN cluster BOLD:ACG2488. Maximum distance within the BIN is 2.45%. The nearest neighbour to BOLD:ACG2488, diverging by minimum of 4.1%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum divergence within the group is 2.5% (based on four specimens and TPI) and the nearest neighbour is 4.7% (P. nigella, only TPI) or 0.6% different (P. siskiyouensis Marlatt, 1896, a Nearctic species not treated here; only NaK).

Host plants

Salix sp. (Lindqvist 1952, Kontuniemi 1972).

Distribution and material examined

West Palaearctic. Specimens studied are from Finland.

Pristiphora atripes (Lindqvist, 1952)

Figs 29, 205, 306

Nematus (Lygaeonematus) lanificus ab. atripes Hellén, 1948a: 46. Not available. Infrasubspecific name.

Lygaeonematus atripes Lindqvist, 1952: 111–112. Holotype ♀ (DEI-GISHym31610) in MZH, examined. Type locality: Lapponia, Europe.

Similar species

Species limits in the micronematica group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the micronematica group belong to the BIN cluster BOLD:ACG2488. Maximum distance within the BIN is 2.45%. The nearest neighbour to BOLD:ACG2488, diverging by minimum of 4.1%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum divergence within the group is 2.5% (based on four specimens and TPI) and the nearest neighbour is 4.7% (P. nigella, only TPI) or 0.6% different (P. siskiyouensis, only NaK).

Host plants

Unknown, but could be Salix herbacea L. (Kontuniemi 1972).

Distribution and material examined

West Palaearctic. Specimens studied are from Sweden.

Pristiphora kontuniemii (Lindqvist, 1952)

Fig. 310

Lygaeonematus kontuniemii Lindqvist, 1952: 113–114. Holotype ♀ (DEI-GISHym20880) in MZH, examined. Type locality: Ivalo, Finland.

Similar species

Species limits in the micronematica group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the micronematica group belong to the BIN cluster BOLD:ACG2488. Maximum distance within the BIN is 2.45%. The nearest neighbour to BOLD:ACG2488, diverging by a minimum of 4.1%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum divergence within the group is 2.5% (based on four specimens and TPI) and the nearest neighbour is 4.7% (P. nigella, only TPI) or 0.6% different (P. siskiyouensis, only NaK).

Host plants

Salix pentandra L., Salix daphnoides Vill. (ex ovo rearing experiments by VV).

Rearing notes

Rearing no. 47/1987: Finland, South Häme, Hämeenlinna, Katinen. On 14.VI.1987 larvae on Salix pentandra; head coloration characteristic.

Ovipositing experiment no. 5/2009: Finland, South Häme, Janakkala. On 8.V.2009 one female was found ovipositing in a bud of Salix pentandra. When captured she laid 12 eggs in pockets on undersides of young leaves of S. pentandra. The eggs were close to the leaf margin. On 8.V.2009 the larvae hatched, and during the next days they gnawed holes in the leaf margins. Four larval instars were observed. Development of the larvae was rapid, and prepupae were found on 22–24.V.2009. No extra moult after feeding.

Rearing no. 2/1999: Finland, Uusimaa, Espoo, Överby school. On 29.V.1999 eggs were found in large buds of Salix daphnoides subsp. acutifolia (Willd.) Ahlfv., the eggs were on underside of outer leaves near the leaf margin in the apical part of the leaf. Larvae hatched and four instars were observed. Prepupae were seen on 1–3.VI.1999. No extra moult after feeding. In next March males and females emerged.

Distribution and material examined

West Palaearctic. Specimens studied are from Finland.

Pristiphora micronematica Malaise, 1931

Figs 207, 305

Pristiphora micronematica Malaise, 1931: 58–59. Lectotype ♀ (NHRS-HEVA000003865; here designated) in NHRS, examined. Type locality: Lake Azhabachye (Ажабачье) near Nizhnekamchatsk (Нижнекамчатск), Kamchatka Krai, Russia.

Lygaeonematus (Lygaeophora) leucostoma Lindqvist, 1952: 108. Holotype ♀ (DEI-GISHym31675) in MZH, examined. Type locality: Munksnäs, Uusimaa, Finland.

Similar species

Species limits in the micronematica group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the micronematica group belong to the BIN cluster BOLD:ACG2488. Maximum distance within the BIN is 2.45%. The nearest neighbour to BOLD:ACG2488, diverging by a minimum of 4.1%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum divergence within the group is 2.5% (based on four specimens and TPI) and the nearest neighbour is 4.7% (P. nigella, only TPI) or 0.6% different (P. siskiyouensis, only NaK).

Host plants

Salix caprea L. (Verzhutskii 1981), S. phylicifolia L. (Kangas 1985, ex ovo rearing experiments by VV), S. pentandra L. (Kangas 1985).

Rearing notes

Ovipositing experiment no. 1/1973: Finland, North Karelia, Joensuu, Purola. One female was captured and buds of Salix phylicifolia and young leaves of Betula pubescens were offered her. On 18–19.V.1973 several eggs were laid in buds of Salix, the egg pockets were on undersides of leaves near the leaf margin. Four larval instars were observed, their development was rapid and on 28.V.1973 several prepupae were seen. No extra moult after feeding.

Distribution and material examined

Palaearctic, Nearctic. Specimens studied are from Finland and Russia (Kamchatka Krai).

Pristiphora nordmani (Lindqvist, 1949)

Fig. 309

Lygaeonematus nordmani Lindqvist, 1949: 85–86. Holotype ♀ (DEI-GISHym20890) in MZH, examined. Type locality: Finström, Åland, Finland.

Similar species

Species limits in the micronematica group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the micronematica group belong to the BIN cluster BOLD:ACG2488. Maximum distance within the BIN is 2.45%. The nearest neighbour to BOLD:ACG2488, diverging by minimum of 4.1%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum divergence within the group is 2.5% (based on four specimens and TPI) and the nearest neighbour is 4.7% (P. nigella, only TPI) or 0.6% different (P. siskiyouensis, only NaK).

Host plants

Unknown.

Distribution and material examined

West Palaearctic. Specimens studied are from Finland.

Pristiphora reuteri (Lindqvist, 1960)

Figs 208, 308

Lygaeonematus (Lygaeophora) reuteri Lindqvist, 1960b: 33–34. Holotype ♀ (DEI-GISHym31676) in MZH, examined. Type locality: Munksnäs, Uusimaa, Finland.

Similar species

Species limits in the micronematica group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the micronematica group belong to the BIN cluster BOLD:ACG2488. Maximum distance within the BIN is 2.45%. The nearest neighbour to BOLD:ACG2488, diverging by a minimum of 4.1%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum divergence within the group is 2.5% (based on four specimens and TPI) and the nearest neighbour is 4.7% (P. nigella, only TPI) or 0.6% different (P. siskiyouensis, only NaK).

Host plants

Salix phylicifolia L. (Vikberg 1966 and later ex ovo rearing experiments by VV). Records from Spiraea (Verzhutskii 1981) are doubtful, because of possible misidentifications.

Rearing notes

Ovipositing experiment no. 2/1973: Finland, North Karelia, Joensuu, Purola. One female was captured and offered buds of Salix phylicifolia and young leaves of Betula pubescens. On 18–19.V.1973 several eggs were laid in buds of Salix, the egg pockets were on the undersides of leaves near the leaf margin. Larvae hatched on 22.V.1973. Four larval instars were observed, their development was rapid and on 30.V.1973 some prepupae were seen. No extra moult after feeding.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Finland and Sweden.

Pristiphora sermola Liston, 1993

Figs 209, 304

Lygaeonematus (Lygaeophora) variipes Lindqvist, 1952: 106–107. Secondary homonym of Pristiphora varipes Serville, 1823. Holotype ♀ (DEI-GISHym20889) in MZH, examined. Type locality: Island Ruissalo (=Runsala), Turku, Finland.

Lygaeonematus (Lygaeophora) variipes ab. xanthopus Lindqvist, 1952: 107. Not available. Infrasubspecific name.

Lygaeonematus (Lygaeophora) variipes ab. morio Lindqvist, 1952: 107. Not available. Infrasubspecific name.

Pristiphora sermola Liston, 1993: 104. Replacement name for Lygaeonematus (Lygaeophora) variipes Lindqvist, 1952.

Similar species

Species limits in the micronematica group are still unclear.

Genetic data

Based on COI barcode sequences, specimens of the micronematica group belong to the BIN cluster BOLD:ACG2488. Maximum distance within the BIN is 2.45%. The nearest neighbour to BOLD:ACG2488, diverging by a minimum of 4.1%, is BOLD:ACO1401 (P. euxantha). Based on nuclear data, maximum divergence within the group is 2.5% (based on four specimens and TPI) and the nearest neighbour is 4.7% (P. nigella, only TPI) or 0.6% different (P. siskiyouensis, only NaK).

Host plants

Salix caprea L. [Liston 1982, as Pristiphora (Lygaeophora) lanifica], S. phylicifolia (ex larva and ex ovo rearings by J. Kangas and VV), Salix starkeana Willd. (ex ovo rearing experiments by VV). The report by Kontuniemi (1960) of P. lanifica from Salix phylicifolia L. possibly refers to P. sermola.

Rearing notes

Ovipositing experiment no. 2/1971 as P. variipes: Finland, North Karelia: Kontiolahti, Jaamankangas. On 30–31.V.1971 one captured female laid several eggs in pockets near leaf margin of Salix phylicifolia. Larvae hatched on 2–3.VI.1971. Four larval instars were observed, their development was rapid and prepupae were seen on 11–13.VI.1971. No extra moult after feeding.

Ovipositing experiment no. 23/1973: Finland, North Karelia, Kontiolahti, Jaamankangas. On 28–29.V.1973 one captured female laid several eggs in buds of Salix starkeana, in egg pockets on the undersides of leaves near the leaf margin. Larvae hatched on 1.VI.1973. Five larval instars were observed, their development was rapid and on 11.VI.1973 prepupae were seen. No extra moult after feeding.

Distribution and material examined

West Palaearctic. Specimens studied are from Finland and Sweden.

Pristiphora nigella group

Pristiphora amphibola (Förster, 1854)

Figs 165, 293

Nematus amphibolus Förster, 1854b: 329–330. Lectotype ♀ (GBIF-GISHym3195) in ZSM, examined. Type locality: Aachen, North Rhine-Westphalia, Germany.

Nematus laetus Cameron, 1883: 194–195. Syntype(s) ♀ possibly in BMNH, not examined. Type locality: Mickleham, Surrey, United Kingdom. Synonymised as fraternus with P. amphibola by Benson (1948).

Nematus fraternus Cameron, 1885: 73–74. Replacement name for Nematus laetus Cameron, 1883.

Similar species

The most similar species are P. nigella and P. parva, from which females of P. amphibola can be distinguished by the combination of dark coloration (almost completely black) and long valvula 3 (see the Key, and Beneš et al. 1981). Males are best recognised by examining penis valves (see the Key).

Genetic data

Based on COI barcode sequences, P. amphibola belongs to the same BIN cluster (BOLD:AAF5120) as P. nigella (Fig. 5). Maximum distance within the BIN is 0.71% and minimum between species distance is possibly 0.00%. The nearest neighbour to BOLD:AAF5120, diverging by minimum of 2.26%, is BOLD:ABV9415 (P. parva). Based on nuclear data (one specimen; NaK), the nearest neighbour is 0.4% different (P. nigella).

Host plants

Picea abies (L.) Karsten (Crooke 1957, Beneš et al. 1981), Picea sitchensis (Bong.) Carriere (Crooke, 1957).

Distribution and material examined

West Palaearctic. Specimens studied are from Finland and Germany.

Pristiphora nigella (Förster, 1854)

Figs 34, 111, 115, 166, 294

Tenthredo ambigua Fallén, 1808: 112–113. Primary homonym of Tenthredo ambigua O.F. Müller, 1776. Lectotype ♂ (designated by Beneš et al. 1981) in MZLU, examined. Type locality: Sweden.

Nematus occultus Förster, 1854a: 331–332. Lectotype ♂ (GBIF-GISHym3371; designated by Beneš et al. 1981) in ZSM, examined. Type locality: Aachen, North Rhine-Westphalia, Germany. Synonymised with Lygaeonematus ambiguus by Konow (1904a).

Nematus nigellus Förster, 1854a: 328–329. Lectotype ♀ (GBIF-GISHym3361; designated by Beneš et al. 1981) in ZSM, examined. Type locality: near Aachen, North Rhine-Westphalia, Germany.

Nematus furvescens Cameron, 1876: 308–311. Syntype(s) possibly in BMNH, not examined. Type locality: East Dunbartonshire or Stirling (“old road between Milngavie and Strathblane”), Scotland, United Kingdom. Synonymised with Lygaeonematus ambiguus by Konow (1904a).

Nematus obscurus Zaddach in Brischke, 1884: 126. Primary homonym of Nematus obscurus Norton, 1861 [= Euura obscura (Norton, 1861)]. Syntypes possibly destroyed (Blank and Taeger 1998). Type locality: Gdańsk [Danzig], Poland or Kaliningrad [Königsberg], Russia. Synonymised with Lygaeonematus ambiguus by Konow (1904a).

Nematus xanthomus Zaddach in Brischke, 1884: 138. Syntype ♂ and ♀ possibly destroyed (Blank and Taeger 1998). Type locality: Lüneburg, Lower Saxony, Germany. Synonymised with Sharliphora nigella by Beneš et al. (1981).

Nematus obscurior Dalla Torre, 1894: 246. Replacement name for Nematus obscurus Zaddach, 1884.

Similar species

The most similar species are P. amphibola and P. parva. In females, the shorter valvula 3 (see the Key) of P. nigella enables its separation from P. amphibola, but differences (slightly shorter valvula 3 and darker coloration) from P. parva might not always be evident. Penis valves of P. nigella, however, seem to show more reliable differences from P. amphibola and P. parva: in nigella, the paravalva is ventrally abruptly narrowed before the valvispina (Fig. 294), while in other species the narrowing is more gradual (Figs 293, 295).

Genetic data

Based on COI barcode sequences, P. nigella belongs to the same BIN cluster (BOLD:AAF5120) as P. amphibola (Fig. 5). Maximum distance within the BIN is 0.71% and minimum between species distance is possibly 0.00%. The nearest neighbour to BOLD:AAF5120, diverging by minimum of 2.26%, is BOLD:ABV9415 (P. parva). Based on nuclear data, the nearest neighbour is 0.3% different (P. parva, only TPI).

Host plants

Picea abies (L.) Karsten (Kontuniemi 1960).

Distribution and material examined

West Palaearctic. Specimens studied are from Estonia, Finland, Germany, and Sweden.

Pristiphora parva (Hartig, 1837)

Figs 167, 295

Nematus parvus Hartig, 1837: 208–210. Lectotype ♂ (GBIF-GISHym3376; designated by Beneš et al. 1981) in ZSM, examined. Type locality: Germany.

Nematus nigricornis Zaddach in Brischke, 1883b: 146–147. Primary homonym of Nematus nigricornis Serville, 1823 [= Euura nigricornis (Serville, 1823)]. Holotype ♂ possibly destroyed (Blank and Taeger 1998). Type locality: Gacko [Dammhof], West Pomeranian Voivodeship, Poland. Synonymised with Sharliphora parva by Beneš et al. (1981).

Nematus germanicus Dalla Torre, 1894: 227, 7. Replacement name for Nematus nigricornis Zaddach, 1883.

Lygaeonematus ambiguus var. flavater Enslin, 1916: 503–504. Lectotype ♂ (DEI-GISHym31699; here designated) in ZSM, examined. Type locality: Gräfenberg, Bavaria, Germany.

Similar species

Most similar species is P. nigella. Pristiphora parva tends to be paler than P. nigella, and in females the valvula 3 is slightly longer in P. parva than in P. nigella (see the Key). Penis valves seem to show more reliable differences: in P. parva, the paravalva narrows distinctly more gradually than in P. nigella (Figs 294–295).

Genetic data

Based on COI barcode sequences, P. parva belongs to its own BIN cluster (BOLD:ABV9415) (Fig. 5). Maximum distance within the BIN is 0.32%. The nearest neighbour to BOLD:ABV9415, diverging by a minimum of 2.26%, is BOLD:AAF5120 (P. amphibola and P. nigella). Based on nuclear data, within species divergence is 0.0% (based on two specimens and both genes combined) and the nearest neighbour is 0.3% different (P. nigella, only TPI).

Host plants

Picea abies (L.) Karsten (Kangas 1985).

Distribution and material examined

West Palaearctic. Specimens studied are from Estonia, Finland, Germany, and Sweden.

Pristiphora pallida group

Pristiphora pallida (Konow, 1904)

Figs 8, 125, 171, 276

Lygaeonematus pallidus Konow, 1904a: 195, 204. Lectotype ♀ (GBIF-GISHym3906; here designated) in SDEI, examined. Type locality: Thüringen, Germany.

Lygaeonematus Stecki [sic!] Nägeli, 1936: 218–219. No type specimens were found in ETHZ by Michael Greeff. Type locality: Bülach, Switzerland. Synonymised with Lygaeonematus pallidus by Forsslund (1939).

Similar species

The most similar species is P. subarctica. For females, there might be small differences in basal annuli of the lancet: in P. pallida, the gap between basal serrulae is smaller and setae are more numerous on basal three annuli than in P. subarctica. Additional differences mentioned by Forsslund (1939) are even smaller or do not seem to be reliable. For example, the lancet is supposed to be more bent in P. pallida (dorsally convex and ventrally concave) than in P. subarctica (somewhat evident also in Figs 171–172), but this difference seems to be absent according to drawings by Wong (1975: figs 17–18). Penis valves are also almost identical, the only difference being possibly the absence (P. pallida, Fig. 276) or presence (P. subarctica, Fig. 278) of a depression on the apical part of valvispina. More studies are needed to decide if P. pallida and P. subarctica are distinct species.

Genetic data

Based on COI barcode sequences, P. pallida forms its own BIN cluster (BOLD:AAX8150). Maximum distance within the BIN is 0.69%. The nearest neighbour to BOLD:AAX8150, diverging by minimum of 2.44%, is P. subarctica (no full barcode available, but possibly belonging also to BOLD:AAX8150). Based on nuclear data (one specimen and NaK), the nearest neighbour is 1.1% different (P. parva).

Host plants

Picea abies (L.) Karsten (Boevé 1990, Pschorn-Walcher and Altenhofer 2000).

Distribution and material examined

West Palaearctic. Specimens studied are from Germany.

Pristiphora subarctica (Forsslund, 1936)

Figs 172, 278

Lygaeonematus subarcticus Forsslund, 1936: 14–22. Holotype ♀ was not found in NHRS. Type locality: Tärna and Stensele, Lycksele Lappmark, Sweden.

Pristiphora pseudosaxesenii Lindqvist, 1968a: 138–139. Holotype ♀ (DEI-GISHym31526) in MZH, examined. Type locality: Forssa, Finland.

Similar species

The most similar species is P. pallida. For females, there might be small differences in basal annuli of the lancet: in P. subarctica, the gap between basal serrulae is larger and setae are less numerous on basal three annuli than in P. pallida. Additional differences mentioned by Forsslund (1939) are even smaller, or do not seem to be reliable. The penis valves are also almost identical, the only difference being possibly the presence (P. subarctica, Fig. 278) or absence (P. pallida, Fig. 276) of a depression on the apical part of valvispina. More studies are needed to decide if P. pallida and P. subarctica are distinct species.

Genetic data

No full COI barcode available, but a 462 bp fragment available in BOLD of one Finnish specimen (FISYM334-15, MHV000166) is closest to BOLD:AAX8150 (P. pallida) differing by a minimum of 2.44%. No nuclear data are available.

Host plants

Picea abies (L.) Karsten (Forsslund 1936, Rodeghiero 2006).

Distribution and material examined

West Palaearctic. Specimens studied are from Finland and Sweden.

Pristiphora pallidiventris group

Pristiphora nigricans (Eversmann, 1847), comb. n.

Figs 141, 223

Nematus nigricans Eversmann, 1847: 16. Lectotype ♀ (DEI-GISHym30030; here designated) in ZIN, examined. Type locality: prov. Casan [Republic of Tatarstan], Russia.

Nematus Anderschi [sic!] Zaddach, 1876: 62–63, syn. n. Holotype ♀ possibly destroyed (Blank and Taeger 1998). Type locality: Uncertain, but possibly Tilsit, Kaliningrad Oblast, Russia. Note. The name anderschi has apparently not been used as valid more than 25 times during the last 50 years (we found 17 publications) and therefore Article 23.9.1 (ICZN 1999) cannot be applied in this case.

Pristiphora inocreata Konow, 1902: 181, syn. n. Lectotype ♀ (GBIF-GISHym3923; designated by Oehlke and Wudowenz 1984 as “Holotypus”) in SDEI, examined. Type locality: Moravia, Czech Republic.

Pristiphora discolor Lindqvist, 1975: 13–14, syn. n. Holotype ♀ (GBIF-GISHym20870) in MZH, examined. Type locality: Popovo, Iljhonsk, Irkutsk Oblast, Russia.

Similar species

The most similar species is P. pallidiventris. Females of P. nigricans have a completely pale metafemur and black terga 9 and 10, while in P. pallidiventris metafemur and terga 9–10 are completely pale or at least the metafemur is apically slightly black. The lancet and penis valve also show consistent differences (see the Key).

Genetic data

Based on a COI barcode sequence of one confidently identified specimen (DEI-GISHym20619), P. nigricans belongs to the same BIN cluster (BOLD:AAU3334) as P. pallidiventris (Fig. 5). The nearest neighbour (BOLD:ACO0950, possibly P. pallidiventris, from China) is 2.26% different. No nuclear data available.

Host plants

Sanguisorba sp. (Verzhutskii 1981, as P. discolor). Sanguisorba officinalis L. according to the label data of the holotype of discolor Lindqvist.

Distribution and material examined

Palaearctic. Specimens studied are from Czech Republic, France, Germany, Kazakhstan, Russia (Irkutsk Oblast and Republic of Tatarstan), and Ukraine.

Pristiphora pallidiventris (Fallén, 1808)

Figs 54, 106, 142–144, 224–226

Tenthredo pallidiventris Fallén, 1808: 120–121. Lectotype ♀ (MZLU2014482; here designated) in MZLU, examined. Type locality: not stated.

Nematus (Nematus) luridus Dahlbom, 1835b: 7. Not available. Nomen nudum.

Nematus ephippiger Hartig, 1840: 24. Out of 7 ♀♀ and 2 ♂♂ syntypes, 2 ♀♀ and 2 ♂♂ were not found in ZSM. Type locality: not stated.

Nematus flavicomus Tischbein, 1846: 77. Types probably destroyed (Horn et al. 1990). Type locality: B. [Birkenfeld, Rhineland-Palatinate, Germany]. Synonymised with P. pallidiventris by Konow (1902).

Nematus gemellus Förster, 1854a: 425–427. Lectotype ♀ (GBIF-GISHym3288; here designated) in ZSM, examined. Type locality: near Aachen, North Rhine-Westphalia, Germany.

Nematus Marshalli [sic!] Cameron, 1875: 9. Syntype(s) possibly in BMNH, not examined. Type locality: Corsica, France. Synonymised with P. pallidiventris by Konow (1902).

Nematus cirrhostomus Zaddach in Brischke, 1883b: 195. Two syntypes possibly destroyed (Blank and Taeger 1998). Type locality: Finland. Synonymised with P. pallidiventris by Konow (1902).

Pristiphora pallidiventris var. denudata Konow, 1902: 165, 179. Lectotype ♀ (DEI-GISHym3905; here designated) in SDEI, examined (severely damaged, abdomen missing). Type locality: Barcelona, Spain.

Pristiphora zella Rohwer, 1909: 20. Holotype ♀ in NSM (Smith 1983), not examined. Type locality: Nebraska, USA. Synonymised with P. pallidiventris by Smith (1979).

Pristiphora pallicoxa Rohwer, 1910b: 200. Holotype ♀ (USNMENT00779341) in USNM, not examined. Type locality: Nerepis, New Brunswick, Canada. Synonymised with P. pallidiventris by Smith (1979).

Pristophora [sic!] xanthotrachela Rohwer, 1913: 281. Holotype ♀ (USNMENT00779593) in USNM, not examined. Type locality: Cabin John, Maryland, USA. Synonymised with P. pallidiventris by Smith (1979).

Pristiphora pallidiventris var. haemorrhoidalis Enslin, 1916: 526. Lectotype ♀ (GBIF-GISHym3291; here designated) in ZSM, examined. Type locality: Erlangen, Bavaria, Germany.

Pristiphora pallidiventris var. stigmatica Enslin, 1916: 526. Lectotype ♀ (GBIF-GISHym3444; here designated) in ZSM, examined. Type locality: Krefeld [Crefeld], North Rhine-Westphalia, Germany.

Pristiphora ostiaria MacGillivray, 1920: 236. Lectotype ♀ in INHS (http://inhsinsectcollection.speciesfile.org/InsectCollection.aspx) (designated by Frison 1927), not examined. Type locality: Ithaca, New York, USA. Synonymised with P. pallidiventris by Smith (1979).

Pristiphora atlantica Malaise, 1939: 1–3. Holotype ♀ in NHRS, examined. Type locality: Caramujo, Madeira, Portugal.

Nematus (Pristiphora) pallidiventris ab. flaviapex Hellén, 1948a: 45. Not available. Infrasubspecific name.

Nematus (Pristiphora) pallidiventris ab. nigrofemoratus Hellén, 1948a: 45. Not available. Infrasubspecific name.

Pristiphora pallidiventris atlantica Lacourt, 1987: 261–262. Primary homonym of Pristiphora atlantica Malaise, 1939 [= Pristiphora pallidiventris (Fallén, 1808)]. Holotype ♀ in CTN, examined. Type locality: Ifrane, Meknès-Tafilalet, Morocco.

Pristiphora pallidiventris megalpina Lacourt, 1987: 262–264. Holotype ♀ in CTN, examined. Type locality: St Véran, Provence-Alpes-Côte d’Azur, France.

Similar species

The most similar species is P. nigricans. Females of P. pallidiventris have a completely pale metafemur and terga 9–10, or at least partly black metafemur, while in P. nigricans the metafemur is completely pale and terga 9–10 are black. The lancets (Figs 141–144) and penis valves (Figs 223–226) have also consistent differences (see the Key). P. pallidiventris is rather variable (also discussed by Lacourt 1987): size of the subapical tooth of claws varies from small to large, the abdomen can be nearly completely black to completely yellow (sometimes even mesepisternum is partly pale), and the structure of the serrulae of the lancet and shape of the penis valve varies as well. Because the variation seems to be continuous and there is no unambiguous way to divide the variation into more than one taxon, we accept only one species.

Genetic data

Based on COI barcode sequences, specimens of this species are divided between at least two BIN clusters (BOLD:ACO1634 and BOLD:AAU3334) (Fig. 5). Minimal distance between these two clusters is 4.94%. Eight additional BIN numbers available in BOLD fall between these two P. pallidiventris BIN clusters. One BIN cluster from Germany (BOLD:AAK9449), five from Canada (BOLD:ACT4128, BOLD:ACO4727, BOLD:ACO4729, BOLD:ABX2590, BOLD:ACL3627) and two from China (BOLD:ACO0950 and BOLD:ACO6535) may contain P. pallidiventris rather than P. nigricans based on the pictures available in BOLD (no specimens studied), although the existence of additional species cannot be excluded. A studied male specimen of P. pallidiventris from Primorsky Krai is closest to BOLD:ACL3627 (Canada), differing by a minimum of 2.3%. Based on nuclear data (two specimens, both genes combined), within species divergence is 0.9% and the nearest neighbour is 3.1% different (P. testacea). Large genetic diversity would be expected for a widespread species (BOLD:AAU3334 contains specimens from Europe and Canada). As the specimens belonging to BIN clusters BOLD:ACO1634 and BOLD:AAU3334, and the specimen from Primorsky Krai cannot be morphologically separated, we consider pallidiventris to be a single species, that is widespread and genetically diverse.

Host plants

Filipendula ulmaria (L.) Maxim. (Loiselle 1913), Geum urbanum L. (Loth 1913), G. rivale L. (Kangas 1985), Rubus chamaemorus L. (Liston 2011), R. idaeus L. (Loth 1913), R. fruticosus agg. (Loth 1913, Chambers 1961), R. ulmifolius Schott (Liston and Zerafa 2012, as P. atlantica).

Distribution and material examined

Palaearctic, Nearctic. Specimens studied are from Austria, Estonia, Finland, France, Germany, United Kingdom, Morocco, Poland, Portugal, Russia (Primorsky Krai), Spain, Sweden, Switzerland, and Ukraine.

Pristiphora retusa group

Pristiphora exigua (Lindqvist, 1955)

Figs 101–102, 133, 258

Lygaeonematus exiguus Lindqvist, 1955a: 143. Holotype ♂ (DEI-GISHym20912) in MZH, examined. Type locality: Espoo, Uusimaa, Finland.

Similar species

The most similar species is P. retusa, females of which usually have a mostly black metafemur in (pale in P. exigua) and narrower valvula 3 (Figs 101–103). Males are best recognised by examining penis valves (see the Key).

Genetic data

Based on COI barcode sequences (only one specimen in BOLD), P. exigua belongs to its own BIN cluster (BOLD:ADD4067). The nearest neighbour to BOLD:ADD4067, diverging by a minimum of 4.3%, is BOLD:ABU7029 (P. retusa). No nuclear data are available.

Host plants

Unknown.

Distribution and material examined

West Palaearctic. Specimens studied are from Finland.

Pristiphora retusa (Thomson, 1871)

Figs 11, 41, 103, 134, 260

Nematus retusus Thomson, 1871: 109–110. Lectotype ♀ (NHRS-HEVA000003756; here designated) in NHRS, examined. Type locality: Dalarna, Sweden.

Similar species

The most similar species is P. exigua, females of which have a completely pale metafemur (usually in most part black in P. retusa) and a broader valvula 3. Males are best recognised by examining penis valves (see the Key).

Genetic data

Based on COI barcode sequences, belongs to its own BIN cluster (BOLD:ABU7029) (Fig. 3). Maximum distance within the BIN is 0.3%. The nearest neighbour to BOLD:ABU7029, diverging by a minimum of 4.3%, is BOLD:ADD4067 (P. exigua). Based on nuclear data, maximum within species divergence is 0.3% (based on three specimens and NaK or both genes combined) and the nearest neighbour is 2.4% different [P. bivittata (Norton, 1861), a Nearctic species, only NaK].

Host plants

Prunus padus L. (Benson 1954b, Kangas 1985, ex ovo rearing experiments by VV).

Rearing notes

Ovipositing experiment no. 1/1978: Finland, South Häme, Janakkala, Hangastenmäki. On 22–23.V.1978 two captured females laid eggs into pockets on undersides of young leaves of Prunus padus. Larvae hatched on 27.V.1978, and ate holes into leaves. Four larval instars were observed, the development of larvae was rapid and on 7.VI.1978 prepupae were found. No extra moult after feeding.

Distribution and material examined

Palaearctic. Specimens studied are from Germany, Russia (Primorsky Krai), and Sweden.

Pristiphora ruficornis group

Pristiphora albitibia (Costa, 1859)

Figs 6, 20, 44, 188, 273

Nematus albitibia Costa, 1859: 21. Syntype(s) ♂ possibly in MZUN, not examined. Type locality: Sila Grande, Calabria, Italy.

Nematus puncticeps Thomson, 1863: 619. Syntypes ♀♂ in MZLU, examined. Type locality: Dalarne, Stockholm, Ostergöthland, Småland, and Skåne, Sweden. Synonymised with P. albitibia by Costa (1894).

Nematus agilis Zaddach in Brischke, 1884: 142. Primary homonym of Nematus agilis Cresson, 1880 [= Euura agilis (Cresson, 1880)]. 3 ♂♀ syntypes possibly destroyed (Blank and Taeger 1998). Type locality: not specified, but probably in former East Prussia (now Kaliningrad Oblast of Russia, or Poland). Synonymised with P. staudingeri auct. by Konow (1905).

Pristiphora aterrima Lindqvist, 1977: 92. Holotype ♀ (DEI-GISHym20896) in MZH, examined. Type locality: Tolyany, Usolje, Irkutsk, Russia.

Pristiphora nigropuncticeps Haris, 2002: 75–76, syn. n. Holotype ♂ (DEI-GISHym80345; http://dx.doi.org/10.6084/m9.figshare.5057701) in HNHM, examined. Type locality: Nukht, Bogd Khan Mountain, Ulaanbaatar, Mongolia.

Similar species

Externally, the most similar species are P. armata, P. confusa, P. leucopus, P. opaca, P. sootryeni, and P. subopaca, from which it is best distinguished by the structure of the lancet (Fig. 188), the penis valve (Fig. 273), and the colour of pterostigma (Fig. 40) (see the Key).

Genetic data

Based on COI barcode sequences, specimens of this species are divided between two BIN clusters (BOLD:ACH1762 and BOLD:ADH0371) (Fig. 4). Minimum distance between the clusters is 4.6%. Based on nuclear data (four specimens and both genes combined), maximum within species divergence is 0.3% and the nearest neighbour is 0.8% different (P. astragali or P. caraganae). Two studied specimens (male and female) from Finland with a highly divergent barcode (BOLD:ADH0371; DEI-GISHym80018 and DEI-GISHym80357) are morphologically indistinguishable from other P. albitibia specimens and this is congruent with nuclear data (both genes) from the female DEI-GISHym80357 (divergence from the other three specimens is 0.1–0.3%).

Host plants

Vicia cracca L. (Stein 1885, as P. puncticeps; Vikberg 2006), V. hirsuta (L.) Gray, V. tetrasperma (L.) Schreb. (Kangas 1985, as P. puncticeps), V. baicalensis Turcz., V. unijuga A. Br. (Verzhutskii 1981, as P. puncticeps).

Rearing notes

See Vikberg (2006).

Distribution and material examined

Palaearctic. Specimens studied are from Estonia, Finland, Germany, Mongolia, Russia (Irkutsk Oblast), and Sweden.

Pristiphora aphantoneura (Förster, 1854)

Fig. 195

Tenthredo fulvipes Fallén, 1808: 113. Primary homonym of Tenthredo fulvipes Scopoli, 1763(Scopoli 1763) [= Aglaostigma (Astochus) fulvipes (Scopoli, 1763)]. Lectotype ♀ (designated by Vikberg 2006) in MZLU, examined. Type locality: Sweden.

Nematus aphantoneurus Förster, 1854b: 323–325. Lectotype ♀ (DEI-GISHym31561; designated by Vikberg 2006) in ZSM, examined. Type locality: Aachen, North Rhine-Westphalia, Germany.

Cryptocampus distinctus Costa, 1882: 198. Syntype(s) ♀ possibly in MZUN, not examined. Type locality: Oschiri, Sardinia, Italy. Synonymised with P. fulvipes by Costa (1894). Note. Identity of the type(s) is uncertain, could be P. luteipes.

Pristiphora pygmaea Lindqvist, 1964: 130. Holotype ♀ in MZH, examined. Type locality: Helsinki, Finland. Synonymised with P. aphantoneura by Vikberg (2006).

Similar species

The most similar species is P. luteipes, from which it cannot be always distinguished morphologically. Vikberg (2006) mentioned that the mesepisternum is completely smooth unlike in P. luteipes, which should show at least slightly coriaceous sculpture (Fig. 45 and Fig. 6a in Vikberg 2006). However, P. luteipes can also have a completely smooth mesepisternum, especially in southern European specimens. See Vikberg (2006) for additional minor characters for separating these species. Males are unknown.

Genetic data

Based on a COI barcode sequence of one confidently identified specimen (reared ex ovo from Lathyrus pratensis) from Finland (DEI-GISHym80037), P. aphantoneura belongs to the same BIN cluster (BOLD:AAG3568) as P. bifida, P. confusa, P. luteipes, P. opaca, P. pusilla, P. staudingeri, and P. subopaca (Fig. 1 Prous et al. 2016). Maximum distance within the BIN is 3.33% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAG3568, diverging by minimum of 2.76%, is BOLD:AAQ2302 (P. armata and P. leucopus). Based on nuclear data (if the specimen DEI-GISHym31258 is correctly identified), the nearest neighbour is 0.2% different (P. luteipes, both genes combined).

Host plants

Lathyrus pratensis L. (Vikberg 2006).

Rearing notes

See Vikberg (2006).

Distribution and material examined

Palaearctic. Specimens studied are from Estonia, Finland, and Germany.

Pristiphora appendiculata (Hartig, 1837)

Figs 19, 21, 61, 105, 187, 262

Pristiphora pallipes Serville, 1823: 75. Secondary homonym of Tenthredo pallipes Fallén, 1808 [= Pristiphora carinata (Hartig, 1837)]. Lectotype ♀ (designated by Lacourt 2000) in MNHN, not examined. Type locality: Paris, France.

Pristiphora pallipes Lepeletier, 1823: 60. Primary homonym of Pristiphora pallipes Serville, 1823 [= Pristiphora appendiculata (Hartig, 1837)]. Lectotype ♀ (designated by Lacourt 2000) in MNHN, not examined. Type locality: Paris, France.

Tenthredo (Nematus) pallicornis T.W. Harris, 1835: 583. Type(s) not available. Nomen nudum.

Tenthredo (Nematus) labrata T.W. Harris, 1835: 583. Type(s) not available. Nomen nudum.

Nematus flavipes Dahlbom, 1835a: 25–26. Nomen oblitum. Holotype ♀ in MZLU, examined. Type locality: Lund, Sweden.

Nematus appendiculatus Hartig, 1837: 202–203. Nomen protectum. See Blank et al. (2009). Lectotype ♀ (GBIF-GISHym3197; designated by Prous et al. 2016) in ZSM, examined. Type locality: Germany according to the title of the publication.

Nematus fuscicornis Hartig, 1837: 225. No syntypes were found in ZSM. Type locality: Harz, Germany. Synonymised with Nematus appendiculatus by Stein (1881).

Nematus enervis Herrich-Schäffer, 1840: 176. Replacement name for Pristiphora pallipes Lepeletier, 1823.

Nematus cathoraticus Förster, 1854: 325–326. Lectotype ♀ (GBIF-GISHym3214; designated by Prous et al. 2016) in ZSM, examined. Type locality: Aachen, North Rhine-Westphalia, Germany.

Nematus pallicornis Norton, 1861: 160. 3 ♀ syntypes in MCZ (http://140.247.119.225/mcz/Species_record.php?id=22468), although 4 specimens were mentioned in the original description, not examined. Type locality: Massachusetts, USA. Synonymised with P. pallipes by Smith (1966).

Nematus pallicornis var. labratus Norton, 1861: 160. Holotype ♀ possibly in ANSP or MCZ. Type locality: Massachusetts, USA. Synonymised with P. rufipes auct. by Smith (1979).

Pristiphora grossulariae Walsh, 1866: 123. Neotype ♀ (designated by Zinovjev and Smith 2000) in ANSP, not examined. Type locality: possibly (if the neotype belongs to the syntype series) Davenport, Iowa, USA. Synonymised with Nematus appendiculatus by Dalla Torre (1894).

Nematus Peletieri [sic!] André, 1880: 111. Replacement name for Pristiphora pallipes Lepeletier, 1823.

Nematus hypobalius Zaddach in Brischke, 1884: 154. Holotype ♀ possibly destroyed (Blank and Taeger 1998). Type locality: Hungary. Synonymised with P. pallipes by Konow (1905).

Nematus pumilus Zaddach in Brischke, 1884: 172. 2 ♂ syntypes possibly destroyed (Blank and Taeger 1998). Type locality: Chernyakhovsk [Insterburg], Kaliningrad Oblast, Russia. Synonymised with P. pallipes by Konow (1905).

Nematus Ghilianii [sic!] Costa, 1894: 73. Syntype(s) ♂ possibly in MZUN, not examined. Type locality: Alps [Alpi boreali], Europe. Synonymised with P. pallipes by Konow (1905).

Pristiphora (Pristiphora) anivskiensis Haris, 2006a: 194–195, syn. n. Holotype ♂ (DEI-GISHym80340; http://dx.doi.org/10.6084/m9.figshare.5100763) in HNHM, examined. Type locality: Novo-Aleksandrovsk, Sakhalin Oblast, Russia.

Similar species

Smooth mesopostnotum (Fig. 19), claws without subapical tooth (Fig. 21), and the structure of serrula (Fig. 187) or penis valves (Fig. 262) allow separation of this species from other similar species in the ruficornis and rufipes groups.

Genetic data

Based on COI barcode sequences, specimens of this species are divided between two BIN clusters (BOLD:AAG7866 and BOLD:AAU8684) (Fig. 1 in Prous et al. 2016). Minimum distance between the clusters is 3.26%. However, BOLD:AAU8684 might represent a cluster of nuclear mitochondrial pseudogenes (Prous et al. 2016). Based on nuclear data (two specimens and both genes combined), maximum within species divergence is 0.3% and the nearest neighbour is 1.8% different (P. pusilla).

Host plants

Ribes spp. Ribes alpinum L. (Kangas 1985, as P. rufipes), R. rubrum L. (Adam 1973, as P. pallipes), R. uva-crispa L. emend. Lam. (Adam 1973, Kangas 1985), R. aureum Pursh (Adam 1973), R. sanguineum Pursh (Adam 1973), R. nigrum L. (Adam 1973), R. spicatum Robs. (Kontuniemi 1975, as P. pallipes).

Distribution and material examined

Palaearctic, Nearctic. Specimens studied are from Austria, Canada, Estonia, Finland, Germany, Russia (Republic of Bashkortostan, Sakhalin Oblast), and Sweden.

Pristiphora armata (Thomson, 1863)

Figs 16, 57, 193, 263

Nematus crassicornis Hartig, 1837: 204–205. Primary homonym of Nematus crassicornis Stephens, 1835 [= Cladius (Cladius) pectinicornis (Geoffroy, 1785)]. 3 ♀♀ and 13 ♂♂ possible syntypes belonging to P. armata and P. leucopus in ZSM, examined. Type locality: Germany according to the title of the publication.

Nematus armatus Thomson, 1863: 619. Seven possible female syntypes belonging to P. armata and P. leucopus in MZLU, examined. Type locality: Bohus Län (Bohuslän), Stockholm, and Skåne, Sweden. Note. Because of difficulties separating P. armata from P. leucopus (see Prous et al. 2016), we refrain from selecting a lectotype at this stage.

Nematus crataegi Brischke, 1883b: pl. I(7), 6. Syntype(s) possibly destroyed (Blank and Taeger 1998). Type locality: not stated, but probably in former East Prussia (now Kaliningrad Oblast of Russia, or Poland). Synonymised with P. crassicornis by Konow (1905).

Nematus Fletcheri [sic!] Cameron, 1884: 26. Syntype(s) possibly in BMNH, not examined. Type locality: Worcester and Clydesdale, United Kingdom. Synonymised with P. crassicornis by Konow (1905).

Nematus melanostomus Zaddach in Brischke, 1884: 140–141. Holotype ♀ possibly destroyed (Blank and Taeger 1998). Type locality: Bautzen, Saxony, Germany. Synonymised with P. crassicornis by Konow (1905).

Nematus ensicornis Jacobs, 1884: XXIII. Syntype(s) ♀ possibly in IRSNB, not examined. Type locality: near Brussels, Belgium. Synonymised with P. crassicornis by Konow (1905).

Nematus nigricollis Cameron, 1885: 66. Syntype(s) possibly in BMNH, not examined. Type locality: Worcester, United Kingdom. Synonymised with P. crassicornis by Konow (1905).

Similar species

The most similar species is P. leucopus. Differences between these two species, which unfortunately are not very strong, were discussed by Grearson and Liston (2012) and Prous et al. (2016).

Genetic data

Based on COI barcode sequences, P. armata belongs to the same BIN cluster (BOLD:AAQ2302) as P. leucopus (Fig. 4). Maximum distance within the BIN is 2.41% and minimum between species distance is possibly 0.00%. The nearest neighbour to BOLD:AAQ2302, diverging by minimum of 2.76%, is BOLD:AAG3568 (P. aphantoneura, P. bifida, P. confusa, P. luteipes, P. opaca, P. pusilla, P. staudingeri, and P. subopaca). Based on nuclear TPI sequences, P. armata (one specimen) and P. leucopus (two specimens) cannot be separated either. The P. armata specimen is identical to one of the P. leucopus specimens when ambiguous positions due to heterozygosity are excluded.

Host plants

Crataegus species (Brischke 1883, Grearson and Liston 2012).

Distribution and material examined

West Palaearctic. Specimens studied are from Belgium, Finland, France, Germany, Italy, Luxembourg, and Sweden.

Pristiphora astragali Vikberg, 1978

Figs 46, 189

Pristiphora astragali Vikberg, 1978: 133–137. Holotype ♀ (PR.354VV) in MZH, examined. Type locality: Kilpisjärvi, Finland.

Similar species

Based on the external morphology, the most similar species are P. confusa, P. opaca, P. pusilla, P. sootryeni, P. staudingeri, and P. subopaca, from which it is best distinguished by the structure of the lancet (Figs 189). The lancet has weak setae (weak or well-developed in the others) and on the inner surface of the lancet there are small spiny pectines (or dentes semicirculares) that reach the sclerora (present also in P. sootryeni). However, differences from P. sootryeni (Fig. 190) are rather small. Morphologically, the subapical tooth of the claws tends be smaller, the apical serrulae of the lancet are shorter, and the number of setae on the lancet is smaller than in P. sootryeni (Vikberg 2006). Male unknown.

Genetic data

Based on a COI barcode sequences, specimens of P. astragali are divided between two BIN clusters (BOLD:AAL8292, BOLD:AAL8277), one of which (BOLD:AAL8292) includes also P. sootryeni (Fig. 4). Minimum distance between the two BINs is 2.4%. Based on nuclear data (one specimen and both genes combined), the nearest neighbour is 0.6% different (P. caraganae).

Host plants

Astragalus alpinus L. (Vikberg 1978, 2006).

Rearing notes

See Vikberg (1978, 2006).

Distribution and material examined

West Palaearctic. Specimens studied are from Finland and Sweden.

Pristiphora bifida (Hellén, 1948)

Figs 9, 25, 199, 267

Nematus (Pristiphora) bifidus Hellén, 1948b: 116–117. Lectotype ♀ (http://id.luomus.fi/GL.5214; designated by Prous et al. 2016) in MZH, examined. Type locality: Malla, Kilpisjärvi, Enontekiö, Finland.

Similar species

Externally, perhaps the most similar species is P. frigida, from which it can be distinguished by having pale hind trochanters, trochantelli, and tibiae (black or brown in P. frigida). In addition, antennae of males have numerous and clearly visible stout black setae among finer paler ones (Fig. 18), while in P. frigida only a few black setae are barely visible (Fig. 17). The lancets (Figs 199–200) and penis valves (Figs 267, 274) are also different. The apical serrulae are somewhat shorter and more protruding, and the tangium of the lancet tends to be longer and narrower (Fig. 199) than in P. frigida (Fig. 200). The penis valve lacks (Fig. 267) a membranous fold near the tip of the valvispina (present in P. frigida; Fig. 274) and the pseudoceps has a distinct dorsal depression in the middle or basal part (absent in P. frigida).

Genetic data

Based on COI barcode sequences, P. bifida belongs to the same BIN cluster (BOLD:AAG3568) as P. aphantoneura, P. confusa, P. luteipes, P. opaca, P. pusilla, P. staudingeri, and P. subopaca (Fig. 4). Maximum distance within the BIN is 3.33% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAG3568, diverging by minimum of 2.76%, is BOLD:AAQ2302 (P. armata and P. leucopus). Based on nuclear data, maximum within species divergence is 0.1% (based on two specimens and both genes combined) and the nearest neighbour is 0.0% different (P. confusa, P. luteipes, or P. beaumonti, only TPI). When including TPI introns, the nearest neighbour is still 0.0% different (P. confusa or P. luteipes).

Host plants

Salix viminalis L. (Liston and Burger 2009). In Kilpisjärvi (Finland) some other species must be the host, as S. viminalis does not occur there.

Distribution and material examined

West Palaearctic. Specimens studied are from Finland, Germany, Norway, and Sweden.

Pristiphora caraganae Vikberg & Prous, sp. n.

Figs 194, 311–319

Description

Female (holotype DEI-GISHym80209 and one paratype), Figs 194, 311–313, 315. Body 6.1–6.3 mm. Colour mostly black. Labrum brown; labial and maxillary palps pale; mandibles reddish-brown at apex; flagellum ventrally slightly paler than dorsally or extensively pale; tibia and tarsi of fore and middle legs more or less entirely pale; apex of profemur pale; metatibia and metatarsomere 1 pale with small black area at apex; trochanters and trochantelli mainly pale; pterostigma bicoloured, basally darker than apically; pronotum and tegula extensively pale. Labrum more or less symmetrical; clypeus truncate; postocellar area about 1.4 times as long as diameter of lateral ocellus; flagellum length 3.9–4.0 mm, 2.3–2.4 times as long as width of head. Mesopostnotum matt; mesepisternum smooth; claws with large subapical tooth clearly separated from apical one; anterior protibial spur with velum. Hypopygium posteriorly not excised. Sawsheath (valvula 3 + valvifer 2) with distinct scopa directed posteriorly and with short medial projection. Lancet with numerous setae on annuli; serrulae protruding, more or less triangular, and with numerous denticles on ventro-apical surface; small spiny pectines on the inner surface (dentes semicirculares) that reach sclerora weak; tangium without lobe, campaniform sensilla present.

Male (31 paratypes), Figs 314, 316–319. Body 5.0–5.6 mm (n=6). Colour mostly black, similar to female. Flagellum ventrally distinctly paler than dorsally; labial and maxillary palps pale; mandibles reddish-brown at apex; tibia and tarsi of fore and middle legs more or less entirely pale; apex of pro- and mesofemur pale; metatibia pale with small black area at apex, trochanters and trochantelli slightly pale; pterostigma bicoloured, basally darker than apically; pronotum black or apically slightly brownish, tegula black to extensively pale. Labrum more or less symmetrical; clypeus truncate; postocellar area about 1.5 times as long as diameter of lateral ocellus; flagellomeres with numerous stout black setae among finer paler ones; flagellum length 4.0–4.3 mm, 2.6–3.1 times as long as width of head. Mesopostnotum matt; mesepisternum smooth; claws with long subapical tooth close to or clearly separated from the apical one; anterior protibial spur with velum. Tergum 8 without apical projection; sternum 9 without notch. Penis valve with large valvispina that nearly entirely replaces paravalva. Valvispina strongly bent in dorsal direction, narrow and with posterior and anterior margin of similar shape, apex sharp; pseudoceps relatively short and broad, apex distinctly narrowed and ventroapical part only slightly extending beyond dorsalmost margin, dorsally without depression, distinct spines, hair, and membranous regions or folds.

Etymology

The species name refers to the host plant Caragana.

Similar species

Externally, females of this species (only the holotype and one paratype known so far) are not distinguishable from many specimens of P. confusa, as well as from P. armata and P. leucopus that have completely black metafemora. However, the lancet has small spiny pectines on its inner surface (Fig. 194) as in other species of albitibia subgroup. Unfortunately, these spiny pectines might easily be overlooked, because they are rather weakly developed (at least in the holotype) compared to other species in the group. In the albitibia subgroup, the female as well as the male of P. caraganae differ from the other species by having the antenna ventrally pale. The female differs additionally from astragali and sootryeni by having a smooth mesepisternum and different lancet (see the Key). Males seem externally most similar to P. bifida, but the penis valves of P. caraganae (Fig. 317) clearly indicate a close relationship with the albitibia subgroup. Compared to other species in the group (although males of P. astragali are unknown), the penis valves seem to be sufficiently distinct (see the Key). There appears to be one more species belonging to the albitibia subgroup that has ventrally pale antennae. Pristiphora nigromongolica Haris, 2002 (http://dx.doi.org/10.6084/m9.figshare.5057791), which is so far known only from the holotype male, differs from P. caraganae by having a smaller subapical tooth of the claw, possibly darker pterostigma (uniformly dark) and a different penis valve that is most similar to P. sootryeni (http://dx.doi.org/10.6084/m9.figshare.4690174).

Genetic data

Based on two identical COI barcode sequences, P. caraganae possibly forms its own cluster (no BIN number has been assigned yet) (Fig. 4). Based on nuclear data (two specimens and both genes combined), within species divergence is 0.0% and the nearest neighbour is 0.6% different (P. astragali).

Host plants

Caragana arborescens Lam.

Rearing notes

The type specimens were collected from Caragana arborescens that grew intermixed with Populus tremula. One larva swept on 18.VI.2017 from the same spot as the adults was offered leaves of Caragana, Populus, and Vicia cracca L., but fed only on Caragana. The larvae made a cocoon on 22.VI.2017 and a female emerged on 2.VII.2017. The reared female laid 44 eggs in leaflets of Caragana and one egg in a leaflet of Vicia. The first larvae emerged on 7.VII.2017. The larva emerged from Vicia was also feeding on Vicia, but died the following day (8.VII.2017). The larvae feeding on Caragana had four instars and possibly a few (3?) had five instars and their development was rapid. On 17–19.VII.2017, 41 prepupae were counted (three larvae had died earlier as rather young) and they made brown cocoons in filter paper in petri dishes, except four prepupae which were stored in alcohol. On 31.VII.–5.VIII.2017, 30 males emerged.

Distribution and material examined

West Palaearctic. Finland.

Holotype. Female. Finland, Hame, Janakkala, Kuumola [ETRS-TM35FIN 67558:[8]3721], 60.91707°N 24.64196°E, 28.VIII.2016, leg. V. Vikberg (MZH). Paratypes. 1♂ (DEI-GISHym80356), same data as the holotype, except 30.V.2017 (MZH), 1♀, same data as the holotype, except reared ex larva from Caragana arborescens found on 18.VI.2017, emerged 2.VII.2017 (CVV), 30 ♂♂, same data as the holotype, except reared ex ovo from the reared female, emerged 31.VII.–5.VIII.2017 (15 in CVV, 5 in MZH, 8 in SDEI, 2 in UEF), 4 prepupae (♂), same data as the holotype, except reared ex ovo from the reared female (SDEI).

Pristiphora confusa Lindqvist, 1955

Figs 196, 268

Pristiphora confusa Lindqvist, 1955: 40–41. Holotype ♀ (http://id.luomus.fi/GL.5209) in MZH, examined. Type locality: Sipoo [Sibbo], Uusimaa, Finland.

Similar species

Based on the external morphology, the most similar species are P. albitibia, P. armata, P. leucopus, P. opaca, P. pusilla, P. sootryeni, and P. subopaca. Pristiphora confusa is best distinguished by the structure of the male penis valve (Fig. 268). Unfortunately, it is rather difficult to distinguish females from P. armata, P. leucopus, P. opaca, and P. subopaca, as the differences in lancets are small (Figs 192–193, 196–198). The apical serrulae are more protruding and shorter than in P. opaca and P. subopaca (Figs 196–198). Pristiphora opaca also has a fold at the base of the tangium of the lancet (Fig. 197) that is lacking in other species in the ruficornis group. Pristiphora opaca tends also to have a smaller subapical tooth than P. confusa (Figs 22–23). The pterostigma of P. confusa is apically brown and basally dark brown, like in P. opaca (Fig. 40), but unlike in P. subopaca, in which it is uniformly yellow (Fig. 39). In P. armata and P. leucopus, the pterostigma is usually dark brown (Fig. 42), but sometimes the pterostigma can have more or less the same colour as in P. confusa. In this case, small differences in the lancet can help distinguish P. confusa from P. armata and P. leucopus, as setae tend to be more distinct in P. confusa (Figs 192–193, 196). Among the males, the most similar penis valves are of P. subopaca. The valvispina in P. confusa is barely bent and the pseudoceps is narrower compared to P. subopaca (Figs 268–269).

Genetic data

Based on COI barcode sequences, belongs to the same BIN cluster (BOLD:AAG3568) as P. aphantoneura, P. bifida, P. luteipes, P. opaca, P. pusilla, P. staudingeri, and P. subopaca (Fig. 4). Maximum distance within the BIN is 3.33% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAG3568, diverging by minimum of 2.76%, is BOLD:AAQ2302 (P. armata and P. leucopus). Based on nuclear data, maximum within species divergence is 0.7% (based on two specimens and both genes combined) and the nearest neighbour is 0.0% different (P. bifida or P. luteipes, only TPI). If TPI introns are included, the nearest neighbour is still 0.0% different (P. bifida).

Host plants

Salix caprea L. (Kangas 1985), Salix fragilis L. (Benson 1958), Salix phylicifolia L. (Benson 1958).

Distribution and material examined

Palaearctic. Specimens studied are from Estonia, Finland, France, Germany, Russia (Primorsky Krai), Sweden, and Switzerland.

Pristiphora frigida (Boheman, 1865)

Figs 17, 200, 274

Nematus frigidus Boheman, 1865: 568–569. Lectotype ♂ (NHRS-HEVA000005005; designated by Prous et al. 2016) in NHRS, examined. Type locality: “Middel Hook in Belsund” (Spitsbergen Island), Svalbard, Norway.

Pristiphora Adelungi [sic!] Konow, 1902: 162, 167–168. Lectotype ♀ (DEI-GISHym30151; designated by Prous et al. 2016) in ZIN, examined. Type locality: Hornsund (Spitsbergen Island), Svalbard, Norway.

Pristiphora gelida Wong, 1968: 185, syn. n. Holotype ♂ (USNMENT00778416) in USNM, not examined. Type locality: Point Barrow, Alaska, USA.

Similar species

Externally, perhaps the most similar species is P. bifida, from which it can be distinguished by having black or brown hind trochanters and trochantelli, and usually black tibiae (pale in P. bifida). In addition, antennae of males have only some barely visible stout black setae among finer paler ones (Fig. 17), while these are numerous and clearly visible in P. bifida (Fig. 18). On the other hand, the penis valve (Fig. 274) might indicate a closer relationship to P. melanocarpa and P. ruficornis (Figs 264, 266), because of a membranous fold near the tip of the valvispina that is missing in other species of the ruficornis group. The tangium of the lancet (Fig. 200) also resembles more closely the Betula feeding P. melanocarpa and P. ruficornis (Fig. 191) rather than P. bifida (Fig. 199): the dark sclerotized area is rather broader than long instead of longer than broad. Although we did not study the holotype of gelida Wong, six specimens from Canada identified by Wong as P. gelida could not be distinguished from P. frigida. The minute differences figured by Wong (1968: 186) between gelida and frigida penis valves and lancets are not reliable. Penis valves of one of the specimens that we studied from Canada are more similar to the figure given by Wong (1968) for frigida.

Genetic data

No data.

Host plants

Unknown.

Distribution and material examined

West Palaearctic, Nearctic. Specimens studied are from Canada and Norway (Svalbard).

Pristiphora leucopus (Hellén, 1948)

Figs 192, 265

Nematus vitreipennis Eversmann in Kawall, 1864: 295. Nomen oblitum (Prous et al. 2016). Lectotype ♀ (DEI-GISHym30027; designated by Prous et al. 2016) in ZIN, examined. Type locality: In promontor. Uralensibus [foothills of Ural mountains], Russia.

Nematus (Pristiphora) ruficornis var. leucopus Hellén, 1948b: 116. Nomen protectum (Prous et al. 2016). No syntypes were found in MZH. Type locality: Joutseno, South-Eastern Finland, Finland and Pionerskoye [Kuolemajärvi], Leningrad Oblast, Russia.

Similar species

The most similar species is P. armata. Differences between these two species, which unfortunately are not very strong, were discussed by Grearson and Liston (2012) and Prous et al. (2016).

Genetic data

Based on COI barcode sequences, P. leucopus belongs to the same BIN cluster (BOLD:AAQ2302) as P. armata (Fig. 4). Maximum distance within the BIN is 2.41% and minimum between species distance is possibly 0.00%. The nearest neighbour to BOLD:AAQ2302, diverging by minimum of 2.76%, is BOLD:AAG3568 (P. aphantoneura, P. bifida, P. confusa, P. luteipes, P. opaca, P. pusilla, P. staudingeri, and P. subopaca). Neither does nuclear sequence data allow separation of P. leucopus from P. armata. Based on nuclear data, maximum within species divergence is 0.4% (based on three specimens and TPI or NaK) and the nearest neighbour is 0.0% different (P. armata, based on TPI including introns).

Host plants

Tilia cordata Mill. (Kangas 1985, Grearson 2006, Grearson and Liston 2012), Tilia × vulgaris Hayne (Grearson 2006).

Distribution and material examined

West Palaearctic. Specimens studied are from Austria, Finland, Germany, United Kingdom, Russia (Ural mountains), and Sweden.

Pristiphora luteipes Lindqvist, 1955

Figs 14, 45

Pristiphora luteipes Lindqvist, 1955b: 47–48. Holotype ♀ (DEI-GISHym20897) in MZH, examined. Type locality: Degerby, Uusimaa, Finland.

Similar species

The most similar species is P. aphantoneura, from which it cannot be always distinguished morphologically. Sometimes the separation of P. luteipes from P. staudingeri can also be problematic, when the colour of metafemur is intermediate between completely black and completely pale (e.g. specimen DEI-GISHym80238 from an arctic habitat in Sweden has a nearly completely pale metafemur). See Vikberg (2006) and Prous et al. (2016) for more detailed discussion.

Genetic data

Based on COI barcode sequences, P. luteipes belongs to the same BIN cluster (BOLD:AAG3568) as P. aphantoneura, P. bifida, P. confusa, P. opaca, P. pusilla, P. staudingeri, and P. subopaca (Fig. 4). Maximum distance within the BIN is 3.33% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAG3568, diverging by minimum of 2.76%, is BOLD:AAQ2302 (P. armata and P. leucopus). Based on nuclear data (four specimens and TPI or both genes combined), maximum within species divergence is 0.3% (excluding DEI-GISHym80238, which could be P. staudingeri) or 0.4% (including DEI-GISHym80238) and the nearest neighbour is 0.0% different (P. bifida, P. confusa, or P. subopaca, only TPI). When including TPI introns, the nearest neighbour is still 0.0% different (P. bifida).

Host plants

Salix alba L., S. aurita L., S. babylonica L., S. repens L. S. rosmarinifolia L., S. phylicifolia L., S. viminalis L., S. purpurea L. (see Vikberg 2006); S. cinerea L. and S. fragilis L. (Loiselle 1909, as P. fulvipes).

Rearing notes

See Vikberg (2006).

Distribution and material examined

West Palaearctic. Specimens studied are from Finland, France, Germany, United Kingdom, Italy, Norway, Portugal, Spain, and Sweden.

Pristiphora melanocarpa (Hartig, 1840)

Figs 191, 264

Nematus melanocarpus Hartig, 1840: 27. Lectotype ♀ (GBIF-GISHym3349; designated by Prous et al. 2016) in ZSM, examined. Type locality: North Germany (according to introduction).

Nematus funerulus Costa, 1859: 20–21. Syntypes ♂♀ possibly in MZUN, not examined. Type locality: vicinity of Naples, Campania, Italy. Synonymised with P. wustneii [sic!] by Ghigi (1905).

Nematus wuestneii Stein, 1885 [mandatory correction of incorrect original spelling N. Wüstneii]: 304. Lectotype ♀ (designated by Prous et al. 2016) in BMNH, examined. Type locality: Chodov [Chodau], Czech Republic.

Pristiphora ortinga Kincaid, 1900: 349–350. Holotype ♀ (USNMENT00778199) in USNM, not examined. Type locality: Kukak Bay, Alaska, USA. Synonymised with P. melanocarpa by by Smith (1979).

Similar species

The most similar species is P. ruficornis, which has paler antennae compared to P. melanocarpa. Females have the ventral side of antennae uniformly black (Fig. 14) or only slightly paler, while P. ruficornis has a distinctly paler ventral side (Fig. 15). Males of P. melanocarpa also tend to have darker antennae than in P. ruficornis, but penis valves should be studied in specimens that have conspicuously pale antennae. The valvispina of the penis valve bends distinctly more sharply (being almost L-shaped) and is usually narrower (Fig. 264) than in P. ruficornis (Fig. 266).

Genetic data

Based on COI barcode sequences, specimens are divided between three BIN clusters (BOLD:AAG3540, BOLD:ACZ4465, BOLD:ACZ4466), two of them (BOLD:ACZ4465 and BOLD:ACZ4466) including also P. ruficornis (Fig. 1 in Prous et al. 2016). These BIN clusters form a monophyletic group (Fig. 4) and minimum distances between them are only 1.13–1.50%. Based on nuclear data, maximum within species divergence is 0.9% (based on seven specimens and NaK, TPI, or both genes) and the nearest neighbour is 0.0% different (P. ruficornis, only TPI including introns).

Host plants

Betula pendula Roth (Kangas 1985), B. pubescens Ehrh., B. nana L. (Prous et al. 2016). The records from Salix are doubtful (Prous et al. 2016).

Distribution and material examined

Holarctic. Specimens studied are from Canada, Estonia, Finland, France, Germany, Norway, Russia (Primorsky Krai), and Sweden.

Pristiphora opaca Lindqvist, 1955

Figs 22, 40, 197, 270

Pristiphora opaca Lindqvist, 1955b: 42–43. Holotype ♀ (http://id.luomus.fi/GL.5204) in MZH, examined. Type locality: Pihtipudas, Central Finland.

Similar species

Based on the external morphology, the most similar species are P. albitibia, P. confusa, P. pusilla, P. sootryeni, and P. subopaca. The species is best distinguished through the structure of male penis valve (Fig. 270). Unfortunately, it is rather difficult to distinguish females from P. subopaca as the differences in the lancets are small (Figs 197–198). The best character might be the structure of the tangium: on its basal part, P. opaca appears to have a fold (Fig. 197) that is absent in other species of the ruficornis group. There are also slight differences in external morphology between P. opaca and P. subopaca. In P. opaca (Fig. 40), the pterostigma is apically brown and basally dark brown (uniformly yellow in P. subopaca; Fig. 39), antennae are slightly paler ventrally (uniformly black in P. subopaca), and claws seem to have a somewhat smaller subapical tooth (Fig. 22) than in P. subopaca (Fig. 23).

Genetic data

Based on COI barcode sequences, P. opaca belongs to the same BIN cluster (BOLD:AAG3568) as P. aphantoneura, P. bifida, P. confusa, P. pusilla, P. staudingeri, and P. subopaca (Fig. 4). Maximum distance within the BIN is 3.33% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAG3568, diverging by a minimum of 2.76%, is BOLD:AAQ2302 (P. armata and P. leucopus). Based on nuclear data (one specimen and both genes combined), the nearest neighbour is 0.4% different (P. luteipes or P. pusilla).

Host plants

Unknown.

Distribution and material examined

Western Palaearctic. Specimens studied are from Finland and Sweden.

Pristiphora pusilla Malaise, 1921

Figs 18, 32, 201, 271

Pristiphora pusilla Malaise, 1921: 11–12. Lectotype ♂ (NHRS-HEVA000004942; designated by Prous et al. 2016) in NHRS, examined. Type locality: Torne Träsk, Torne Lappmark, Sweden.

Pristiphora amaura Lindqvist, 1955b: 43–45. Holotype ♀ (http://id.luomus.fi/GL.5205) in MZH, examined. Type locality: Kangasala, South Häme, Finland.

Similar species

Based on the external morphology, the most similar species are P. albitibia, P. astragali, P. confusa, P. opaca, P. sootryeni, P. staudingeri, and P. subopaca. The species is best distinguished through the structure of male penis valve (Figs 271) and female lancet (Fig. 201). In females, the lack of small spiny pectines (or dentes semicirculares) on the inner surface of the lancet, and weakly developed setae, distinguish it from other similar species. Male penis valves are asymmetric, the left one (Fig. 271) having a noticeably stronger dorsal depression in the middle of pseudoceps and a more strongly bent valvispina than the right one (Fig. 94 in Prous et al. 2016). The most similar penis valves are those of P. subopaca (Fig. 269), which have a less distinct dorsal depression in the middle of pseudoceps and a less strongly bent valvispina, but this difference is clear only when compared to the left penis valve of P. pusilla. Externally, P. pusilla can usually be distinguished from P. subopaca by having ventrally paler antennae (uniformly black in P. subopaca; Fig. 15), which is more evident in males (Fig. 18).

Genetic data

Based on COI barcode sequences, P. pusilla belongs to the same BIN cluster (BOLD:AAG3568) as P. aphantoneura, P. bifida, P. confusa, P. opaca, P. staudingeri, and P. subopaca (Fig. 4). Maximum distance within the BIN is 3.33% and minimum between species distance is 0.00%. The nearest neighbour to BOLD:AAG3568, diverging by a minimum of 2.76%, is BOLD:AAQ2302 (P. armata and P. leucopus). Based on nuclear data, maximum within species divergence is 0.2% (based on seven specimens and TPI or both genes combined) and the nearest neighbour is 0.1% different (P. confusa or P. subopaca, only TPI).

Host plants

Unknown.

Distribution and material examined

Western Palaearctic. Specimens studied are from Finland, Norway, and Sweden.

Pristiphora ruficornis (Olivier, 1811)

Figs 15, 42, 266

Nematus ruficornis Olivier in Olivier and Manuel, 1811: 167. No unambiguous type specimens were found in MNHN. Type locality: near Paris, France. Note. Two male specimens in Lepeletier’s type collection in MNHN (which we identified as P. armata, but could also be P. leucopus) were under the name N. ruficornis, but only one of them had a label “N. ruficornis 216 ♂”. The number “216” apparently refers to Olivier’s Nematus ruficornis in Lepeletier de Saint-Fargeau (1823: 71) (“216. NEMATUS RUFICORNIS. Oliv. Enc., n°9.”). However, Lepeletier de Saint-Fargeau (1823: 71) mentions females, not males. In an old box of uncertain origin in MNHN there is also a female (we identified it as P. armata?) labelled “”N. ruficornis” Lp. 216 N. crassicornis H. 37.”, but this seems to be of later origin than the publication of Lepeletier de Saint-Fargeau (1823), because the specimen is identified as Hartig’s crassicornis (Hartig 1837). In order to reserve the name ruficornis Olivier for the species concept used in the widely known identification key by Benson (1958) and recently reviewed by Prous et al. (2016), i.e. for the species feeding on Betula not on Crataegus, designation of a neotype would be desirable. However, we refrain doing so at this point, as no fresh specimens of ruficornis are available from or near the type locality (near Paris). The neotype specimen should be fresh enough to enable extraction of good quality DNA, that could greatly facilitate solving the problem of separating ruficornis from melanocarpa (if they indeed are different species).

Pristiphora testaceicornis Serville, 1823: 75. Syntype(s) ♂ not found in MNHN (Lacourt 2000). Type locality: Paris, France. Pristiphora testaceicornis Lepeletier, 1823: 60. Primary homonym of Pristiphora testaceicornis Serville, 1823 [= Pristiphora ruficornis (Olivier, 1811)]. Syntype(s) ♂ not found in MNHN (Lacourt 2000). Type locality: Paris, France. Synonymised with P. ruficornis by André (1882).

Nematus (Nematus) robustellus Dahlbom, 1835b: 9. Type(s) not available. Nomen nudum.

Nematus fraxini Hartig, 1837: 204. Lectotype ♀ (GBIF-GISHym3285; designated by Prous et al. 2016) in ZSM, examined. Type locality: Harz, Germany.

Nematus testaceicornis Jacobs, 1884: XXIII-XXIV. Syntype(s) ♀ possibly in IRSNB, not examined. Type locality: near Brussels, Belgium. Synonymised with P. ruficornis by Konow (1902).

Nematus (Pristiphora) ruficornis var. integer Hellén, 1948b: 116. Primary homonym of Nematus integer Say, 1836. Holotype ♀ (http://id.luomus.fi/GL.5212) in MZH, examined. Type locality: Hammaslahti, North Karelia, Finland.

Similar species

The most similar species is P. melanocarpa, which has darker antennae compared to P. ruficornis. Females usually have a distinctly paler ventral side of antennae (Fig. 15), while the antennae in P. melanocarpa are uniformly black (Fig. 14) or have only a slightly paler ventral side. Males of