ACTA ENTOMOLOGICA 2021 61(1): 73–81 MUSEI NATIONALIS PRAGAE doi: 10.37520/aemnp.2021.004 ISSN 1804-6487 (online) – 0374-1036 (print) www.aemnp.eu R E S E A R C H PA P E R Stenocephus janseni sp. nov., a new species of stem-sawfly from Germany (Hymenoptera: Cephidae) Andrew LISTON1,*) & Marko PROUS2,3) 1) Senckenberg Deutsches Entomologisches Institut, Eberswalder Str. 90, 15374 Müncheberg, Germany; e-mail: aliston@senckenberg.de 2) Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia; e-mail: mprous@ut.ee 3) Ecology and Genetics Research Unit, PO. Box 3000, FI-90014 University of Oulu, Finland *) corresponding author Accepted: Abstract. Stenocephus janseni sp. nov. (Hymenoptera: Cephidae) is described from Bran- 23rd February 2021 denburg, eastern Germany, known only from the female holotype. It possesses an unusual Published online: combination of “generic” morphological characters, which makes its placement in Stenoce- 1st March 2021 phus Shinohara, 1999 provisional. Compared to other Stenocephus species, differences in the morphology of the lancet are particularly striking. Genetic data for S. janseni sp. nov. place it unequivocally in the Hartigiini, but rather distantly from other genera of this tribe which have so far been sequenced. The three previously described Stenocephus species are recorded from the East Palaearctic. No genetic data are currently available for these. Pachycephus nigratus Dovnar-Zapolskij, 1931, comb. restit., is no longer treated as belonging to Phylloecus Newman, 1838, but as a member of the genus in which it was originally described. Key words. Hymenoptera, Cephidae, Hartigiini, Stenocephus, Pachycephus nigratus, new species, DNA barcoding, Germany, Palaearctic Region Zoobank: http://zoobank.org/urn:lsid:zoobank.org:pub:5B17AF86-56BB-4D13-AF8E-0AFE5DBE803E © 2021 The Authors. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Licence. Introduction and perhaps also on Lamiaceae (G 1935: see under Janus nigratus) and Asteraceae (Z The larvae of Cephidae tunnel in the stems or twigs 1968: see under Characopygus Konow, 1899). of their hosts. A few species are of major significance as Our main purpose here is to describe a new species of pests of grain crops (S H 2004). S cephid, belonging to the Hartigiini, recently collected in S (2009) provided a key to the three extant sub- eastern Germany. The specimen is morphologically highly families of Cephidae: the Cephinae (Holarctic and Orien- distinctive, with an unusual combination of characters tal), Athetocephinae (Afrotropical and Australasian), and which does not fit current circumscriptions of the genera. Australcephinae (Australasian). The latter two subfamilies Accordingly, before describing the new species, provisi- contain only three and one species respectively, whereas the onally placed in the genus Stenocephus Shinohara, 1999, Cephinae contains about 160 described species, mostly Pa- we discuss these characters, as far as they are relevant to laearctic (T et al. 2010). Suprageneric classification its placement. of the Cephinae currently remains as proposed by B (1946), i.e. divided into the three tribes Cephini, Hartigiini, and Pachycephini. The tribes, as far as the hosts of species Material and methods are known, have different groups of larval host plants: Morphology and systematics. The extensive adult ma- Cephini feed on Poaceae (T et al. 1998), Hartigiini terial in the Senckenberg Deutsches Entomologisches on Rosaceae and various other families of woody dicots Institut (DEI), Müncheberg, Germany, was used in com- (L 2020, M et al. 2020), and Pachycephini on parative morphological studies. Most observations, and poppy (Papaver, Papaveraceae) (S 1978), images, were made with an Olympus SZX12 microscope 74 LISTON & PROUS: Stenocephus janseni sp. nov. from Germany (Hymenoptera: Cephidae) and Leica DFC295 camera. The images presented here, POL2_104Fv2 and reverse primer POL2_2569R). The except that of the antenna (a single image), are compo- NaK fragment does not include any introns, but POL2 sites, derived from stacks of images taken sequentially has one short intron (109–113 bp) that was excluded (from top to bottom) at different focal planes. Helicon from phylogenetic analyses. When excluding the intron Focus was used to collate the stacks. The lancets of Ste- in POL2, algorithm-based alignment was not necessary nocephus janseni sp. nov., now gummed with all other due to the lack of insertions or deletions in the studied detached parts to cards pinned with the specimen, were specimens (length differences were only due to the extent photographed with a Leica DFC450 through an Olympus the gene regions were amplified and sequenced, and the BX51. alignment was manually adjusted accordingly). Some of We have not included in the references the works in the analyzed sequences were published previously by which taxa which we mention were first described, unless M N (2015) and S et al. (2017). COI these publications are cited for other reasons. Such infor- sequences of some cephids were extracted from published mation can be found in T et al. (2010). mitochondrial genomes (see references and GenBank ac- Phylogenetic analyses. To assess the phylogenetic place- cessions in A K 2020). Nuclear sequen- ment of Stenocephus janseni sp. nov. within the Cephidae, ces of Cephus cinctus Norton, 1872 were extracted from published as well as newly obtained mitochondrial and whole genome shotgun contigs in GenBank (accessions nuclear DNA sequences were used. DNA extraction, AMWH01001469 and AMWH01002735). Additionally, primers used, PCR protocols, and Sanger sequencing are a few of the COI sequences were obtained from BOLD described in P et al. (2019). Additionally, some of the (http://www.boldsystems.org/). The newly obtained amplicons were pooled and sequenced with the MinION DNA sequences have been submitted to NCBI GenBank R10.3 flow cell using a Ligation Sequencing Kit (SQK (accessions MW353980–MW353997). To concatenate -LSK109) (Oxford Nanopore). Each amplicon sequenced separate gene alignments, we used R (R C T with MinION was amplified using different combinations 2019) package apex (J et al. 2017). Phylogenetic of tailed forward and reverse primers (variable 4–12 bp analyses using maximum likelihood (ML) were done with added to the 5’-end) to confirm the identity of the final IQ-TREE 1.6.1 (http://www.iqtree.org/) (N et al. consensus sequences. The raw sequencing signal from 2015). By default, IQ-TREE runs ModelFinder (K - MinION was basecalled (translated into a DNA sequen- et al. 2017) to find the best-fit substitution ce) with Guppy v4.0.11 or 4.2.3 in high accuracy mode. model and then reconstructs the tree using the model Using available cephid sequences as query, corresponding selected according to Bayesian information criterion single molecule Nanopore reads were identified with (BIC). The alignments were treated as a single partition BLAST 2.9.0+ (https://www.ncbi.nlm.nih.gov/books/ to avoid over-parametrization of the small alignments NBK279690/). A maximum of 3000 single reads were and the rather similar sequences analyzed here. We com- aligned with MAFFT v7.427 (K &S 2013) plemented the default option of IQ-TREE with a SH-like and the trees were built with FastTree 2.1.11 (P et approximate likelihood ratio (SH-aLRT) test (G et al. 2010). Based on the resulting trees, separate clusters al. 2010) and ultrafast bootstrap (H et al. 2018) with of reads were identified and subsequently used to create 1000 replicates to estimate robustness of reconstructed consensus sequences. All sequences, except nuclear splits. Ultrafast bootstrap support values above 95% are sequences of two specimens (DEI-GISHym89964, DEI- generally considered reliable (H et al. 2018). GISHym86341), separated clearly at single read level. Based on 200 reads of each amplicon, MAFFT v7.427 Results + EMBOSS cons v6.6.0.0 and abPOA 1.0.4 (https:// Identity of Pachycephus nigratus github.com/yangao07/abPOA) were used to create Dovnar-Zapolskij, 1931, comb. restit. initial consensus sequences that were further polished with Medaka 1.0.1 (https://github.com/nanoporetech/ B (1946) proposed that Pachycephus nigratus medaka). Medaka variant calling was used to separate Dovnar-Zapolskij, 1931 belongs to Hartigia Schiødte, 1839 the very similar nuclear sequences of DEI-GISHym89964 (= Phylloecus Newman, 1838: L P 2014). Pre- and DEI-GISHym86341. A more detailed protocol and viously, the species had been transferred to Janus Stephens, data analysis workflow will be published separately. 1829 by G (1935). Benson’s opinion was fol- For most specimens, one mitochondrial and two nuclear lowed by T et al. (2010). However, the latter authors genes were sequenced. The mitochondrial gene used is did not give due regard to Z (1976), who had partial (1078–1087 bp) cytochrome c oxidase subunit I refuted Benson’s placement and treated the species once (COI), amplified with forward primer SymF1 or SymF4 more as a Pachycephus. We now accept Zhelochovtsev’s and reverse primer A2590 (see P et al. 2019). The decision, because his association of the previously unknown amplified COI fragment covers the entire barcode region male is convincing, and its prolonged and in lateral view (658 bp). The two nuclear markers are fragments of sodi- pre-apically constricted subgenital plate unequivocally um/potassium-transporting ATPase subunit alpha (NaK, places it in the Pachycephini. The placement of this species, 1654 bp; primers NaK_263F and NaK_1918R) and DNA which has two pre-apical metatibial spurs, is relevant to our dependent RNA polymerase II subunit RPB1 (POL2, generic placement of Stenocephus janseni sp. nov., because 1771 or 2573–2577 bp; forward primer POL2_797F or all known Phylloecus species have only one.  Acta Entomologica Musei Nationalis Pragae, volume 61, number 1, 2021 75 Figs 1–6. Stenocephus janseni sp. nov., holotype, female. 1 – lateral, scale bar 5 mm; 2 – head, dorsal; 3 – head, frontal; 4 – head, lateral; 5 – thorax, dorsal; 6 – pronotum, dorsal. Morphological character states in Hartigiini the base. In the East Palaearctic and Oriental Regions, five Shape of claw. Much use has been made of the form of other described genera have an acute basal lobe resembling the claw as a character for the separation of cephid genera, that of Janus: Jungicephus Maa, 1949, Magnitarsijanus and even of the tribes Cephiini and Hartigiini (e.g. B Wei, 2007, Megajanus Wei, 1999, Stigmatijanus Wei, 2007, 1946, G 1992). In the West Palaearctic Hartigiini the and Tibetajanus Wei, 1996. Additional East Palaearctic and strongly-developed, acutely-apexed basal lobe of Janus Oriental genera lacking a basal lobe are Miscocephus Wei, distinguishes it from Phylloecus, Syrista Konow, 1896 1999, Sinicephus Maa, 1949, Stenocephus Shinohara, 1999 and Caenocephus Konow, 1896, in which the claw lacks and Urosyrista Maa, 1944. Stenocephus janseni sp. nov. a basal lobe, although it may be gently expanded towards lacks a basal lobe, and the two teeth are not very close to-  76 LISTON & PROUS: Stenocephus janseni sp. nov. from Germany (Hymenoptera: Cephidae) Figs 7–12. Stenocephus janseni sp. nov., holotype, female. 7 – anterior thorax, lateral; 8 – mesoscutellum, dorsal; 9 – metatibial claw; 10 – metatibial spurs; 11 – abdomen base, lateral; 12 – abdomen apex, dorsal. gether, with the inner tooth markedly longer and wider than inner tooth, of about the same basal width as the outer, the outer one (Fig. 9). As regards the teeth, this is similar has a simple outline (not conspicuously “shouldered” on to Phylloecus, Sinicephus, and Syrista, whereas Urosyrista anterior edge) (Fig. 15). In these characters, S. janseni sp. has approximately equally sized teeth (S 1999), and nov. resembles previously known species of Stenocephus in previously known Stenocephus the inner tooth is shorter more closely than species in other genera of Hartigiini. than the outer (W et al. 2015). The taxonomic review of Maxillary palps. According to B (1946), and our Syrista, by W S (2010), in which the claws of both own observations, Phylloecus differs from other cephids in sexes of two species are illustrated, reveals, however, that typically with palpomere 4 about as long as 6, as opposed sexual dimorphism in claw shape, particularly the orienta- to palpomere 4 about 1.5× length of 6. Stenocephus janseni tion, proportions and size of the outer tooth relative to the sp. nov. has palpomere 4 about 1.5× length of 6 (Fig. 14). inner, can be greater than differences sometimes stated to Lancet. The lancet of Stenocephus janseni most closely be diagnostic of genera. resembles those of Phylloecus species (S 1986 [Nearc- Shape of the left mandible. Nearly all authors since tic species]; Palaearctic species examined by us), in having B (1946) have made use of this character. B rather small, simple serrulae, and distinct annuli (Fig. 17). (1946) and S (1999) provided illustrations of However, whereas Phylloecus have two ctenidial teeth its form in various genera, and a figure for Urosyrista was per annulus, S. janseni sp. nov. has only one. The lancet given by S (1999). In Stenocephus janseni sp. nov. of Caenocephus lunulatus (Strobl, 1895) is also similar, the outer (anterior) tooth is longer than the inner, and the but the serrulae are much more prominent, and ctenidial  Acta Entomologica Musei Nationalis Pragae, volume 61, number 1, 2021 77 Figs 13–17. Stenocephus janseni sp. nov., holotype, female. 13 – valvula 3 and cerci, lateral; 14 – maxillary palp; 15 – left mandible [broken tip of external tooth outlined]; 16 – antenna; 17 – lancet. teeth are absent (based on examination of the single known including Stenocephus janseni sp. nov., the abdomen gra- German specimen: L 2006). Other Stenocephus have dually widens distally from tergum 2 to about two thirds numerous ctenidial teeth on the basal annuli, and some of of the abdomen length (Fig. 1). the serrulae with a more complex outline (almost bifid) Cenchri and tegulae. W (1999) stated that the pre- (W et al. 2015). Syrista have no ctenidial teeth, indistinct sence of cenchri and absence of [fore wing] tegulae distin- annuli, and at least the apical serrulae with a more complex guish Miscocephus from all other cephids. The presence outline (W S 2010). Miscocephus has apparently of cenchri would be remarkable in the Cephidae, in which no (or indistinct) ctenidial teeth, distinct annuli, and simple their absence is generally regarded as an autapomorphy of serrulae (W 1999). Janus have no ctenidial teeth, indi- the family (e.g. V 2000). Examinations of the stinct annuli, and serrulae sub-rectangular, almost bifid type specimen of M. cyaneus Wei, 1999 by S. M. Blank (S S 1989, L et al. 2017). (personal communication) and an unidentified species Number of pre-apical metatibial spurs. This has been (one male specimen) in the DEI from Nepal, which based used by several taxonomists as the prime distinction on other characters seems to belong to this lineage, reveal between some genera of Cephidae. R (1937) first pointed that the “cenchri” are actually small pale areas, contrasting out that it is not always a stable character, although his with the surrounding black integument, but are not raised findings relate mainly to the Cephiini. In the Hartigiini, structures (i.e. they are not cenchri), and that tegulae are S (1999) remarked on variability (1–2 spurs) present, albeit inconspicuous (small and strongly down- in Stenocephus. On the other hand, the absence of a pre- wardly deflected towards the anterior). apical spine in Caenocephus, and presence of only one in Genetic data. Three gene fragments were obtained for Phylloecus, has so far been observed to be constant. B - Stenocephus janseni sp. nov. to examine its phylogenetic (1946) stated that Phylloecus (as Hartigia) sometimes position within Cephidae. Most of the new sequences could have two, but this was based on his wrong placement were obtained either with the Sanger or Nanopore method, of Pachycephus nigratus (see above). The holotype of except COI of S. janseni sp. nov. holotype that was Stenocephus janseni sp. nov. has two pre-apical spurs (Fig. sequenced with both methods. The Sanger and Nanopore 10), as in all other hartigiine genera, perhaps excepting consensus sequences of COI of S. janseni sp. nov. holo- only Magnitarsijanus (1) (but character state not known type were identical, indicating reliability of the relatively to us for Tibetajanus). new Nanopore sequencing technology. Nevertheless, a Body shape. Phylloecus, Caenocephus and Janus Nanopore consensus sequence of COI of one specimen species are stockier than most other Hartigiini, this being (DEI-GISHym89964) probably contained a deletion most apparent in the shape of the abdomen, which is nearly error in a homopolymer region (6 G instead of probable cylindrical distal to tergum 2 in the former. In the others, 7) causing a frame shift mutation that is not expected in  78 LISTON & PROUS: Stenocephus janseni sp. nov. from Germany (Hymenoptera: Cephidae) the protein coding regions. This one-nucleotide gap was expanding from apex of flagellomere 3. Flagellomere 1 replaced with an undetermined nucleotide (“N”) to pre- slightly longer than 2 (1.2 : 1.0). All flagellomeres longer serve the protein translation frame. For both alignments, than broad. Pubescence on upper head about 0.25× as long COI only and combined COI and nuclear (Figs 18–19), the as diameter of anterior ocellus, and sparser than on thorax. best-fit model chosen according to Bayesian information Most of thorax (Figs 5–8) deeply punctured, with criterion (ModelFinder implemented in IQ-TREE) was narrow interspaces, and dull. Dorsal part of propleuron GTR+F+I+G4. The maximum likelihood analyses of the glabrous and without punctures (Fig. 7). Longest setae on data (Figs 18–19) clearly place S. janseni sp. nov. within the thorax about 0.5× as long as diameter of anterior ocellus. Hartigiini (excluding Syrista), but its exact phylogenetic Pronotum in dorsal view (Fig. 6) slightly wider than long position remains uncertain. Unfortunately, genetic data are (ca. 1.1 : 1.0); anteriorly and posteriorly strongly carinate lacking for any other Stenocephus and many other, possibly on dorsal margins; a longitudinal furrow present, deeper relevant, East-Asian genera. Based on the current taxon and wider posteriorly, ending posteriorly in a shallow mar- sampling, S. janseni sp. nov. is weakly supported as sister ginal notch and anteriorly in a small triangular area without group of Phylloecus based on mitochondrial COI (Fig. 18) punctures or sculpture. Mesoscutellum almost circular in or as sister of Janus based on combined COI and nuclear outline in dorsal view (Fig. 8). Protibia without preapical data (Fig. 19). spurs; mesotibia with one preapical spur; metatibia with two preapical spurs (Fig. 10). Metatibial preapical spurs Stenocephus janseni sp. nov. longer than apical ones (Fig. 10). Metatarsomere 1 slightly Type material examined. H :  (DEI-GISHym84482), ‘Ger- longer than combined lengths of 2–4 (1.03 : 1.00). Claw many: Brandenburg, Müncheberg 4 km NW, 52.521°N 14.064°E [58 m with inner tooth close to outer tooth and clearly larger; a.s.l. in woodland beside railway track “Ostbahn”; Berlin to Kostryn], no basal lobe (Fig. 9). Fore wing stigma 8.6× as long as 20.V.2020, leg. A. Liston, M. Prous’ [white label, printed], ‘Holotype  broad. Fore wing anal cell with cross-vein. Cross-vein 3r-m Stenocephus janseni n. sp. det. A. Liston’ [red label, handwritten], ‘DEI- present in hind wing; 7 hamuli on hind wing. GISHym84482’ [white label, printed]. Deposited in the Senckenberg Deutsches Entomologisches Institut, Müncheberg. Abdominal terga shallowly and indistinctly punctate; shiny between the punctures. Tergum 1 with a complete Description. Female (Figs 1–17). Length: 10 mm, without longitudinal median incision extending anteriorly. Ter- ovipositor. gum 2 in lateral view (Fig. 11) strongly widened towards Colour (Fig. 1). Black. Parapterum whitish. Yellow posterior (distal height approx. 1.25× length). Terga 3–7 -brown are: extreme dorsal apex of profemur, all tibiae, progressively widened distally (Fig. 1). Cerci short (Figs more or less all tarsi except for distally progressively so- 12, 13): appearing about 0.15× as long as valvula 3 in dorsal mewhat darker tarsomeres (from apex of basitarsomere), view (Fig. 12). Valvifer 2 about as long as valvula 3. In abdominal terga 3 and 4 and corresponding sterna, and lateral view upper edge of valvula 3 with straight profile; tergum 2 with corresponding sternum except for their longitudinally with a ridge running approximately parallel more or less fuscous dorsal / ventral parts. Wing membrane to lower edge (Fig. 13). In dorsal view (Fig. 12) slightly subhyaline. Venation basally yellowish, apically (including expanded from base to middle, then gently narrowing pterostigma) becoming brown. towards apex. Lancet (Fig. 17): 19 annulets; a single, Head in dorsal view (Fig. 2) slightly contracted behind robust, ventrally-placed ctenidial tooth on each annulus; eyes; lateral length (from anterior of eye to most posterior serrulae narrow, and small in comparison to length of an- point) approximately 0.5× width. Temple much shorter nulet, not clearly developed on annulus 1 [numbered from than length of eye (Fig. 4). Genal carina developed from base, excluding the radix]. Lance of similar proportions malar space to about 0.33 height of eye. Ratio of distance to lancet, with about 12 proximal dorsal serrulae; annular between inner edges of toruli : distance between lower edge sutures not visible. of torulus and centre of anterior tentorial pit approx. 1.2 Male. Unknown. : 1.0 (Fig. 3). Postocellar area rather densely punctured, Differential diagnosis. As already indicated in our brief with shiny interspaces (Fig. 2). Frontal area more densely comparative review of morphological character states in and finely punctured; dull (Fig. 3). Vertex, temple and Hartigiini, Stenocephus janseni sp. nov. is readily distin- supraclypeal area shiny, with weak, scattered punctures. guished from other known taxa by an unusual combination Left mandible (Fig. 15) outer (anterior) tooth longer than of characters: claw without angled basal lobe and inner the inner; inner and outer teeth of about equal basal width; tooth longer and wider than the outer; mandible outer inner tooth anterior edge slightly convex; posterior edge tooth longer than inner and teeth of about same basal concave. Labial palp with four palpomeres, much shorter width, inner tooth with simple outline (not conspicuously than maxillary palp. Apical labial palpomere much wider “shouldered”); maxillary palpomere 4 about 1.5× length than apical maxillary palpomeres. Maxillary palp (Fig. of 6; lancet with distinct annuli, simple serrulae, and one 14): palpomere 6 arising at approximately 0.6 of length ctenidial tooth per annulus. of 5 (from base); palpomeres 5 and 6 combined length Body colour pattern alone, if stable, might distinguish slightly less than 4 (or palpomere 4 about 1.5× as long as female Stenocephus janseni sp. nov. from most other West 6). Antenna (Fig. 16) long and slender; about as long as Palaearctic cephid species. Whereas S. janseni sp. nov. has combined length of fore wing costa and stigma; 28 anten- some of the basal terga and sterna entirely pale, most others nomeres. Flagellum widest at about 0.75 from base, gently have only parts of the terga or sterna banded or flecked  Acta Entomologica Musei Nationalis Pragae, volume 61, number 1, 2021 79 Fig. 18. Maximum likelihood tree of Cephidae based only on mitochondrial COI gene. Numbers at branches show SH-aLRT support (%) / ultrafast bootstrap support (%) values. Support values for weakly supported branches (<90) are not shown. Letters “f” and “m” stand for “female” and “male” if known. Numbers at the end of the tip labels refer to the length of the sequence. The tree was rooted according to the results of M N (2015) and A K (2020). The scale bar shows the number of estimated substitutions per nucleotide position. with pale. In the Hartigiini, Syrista parreyssii (Spinola, entirely pale, and the metatibia is largely black, with the 1843) has a similarly coloured body and legs, but differs in base clear-white. its greater body length (female 15–18 mm), temple much Characters which separate Stenocephus janseni sp. nov. longer than length of eye, and a strongly downcurved val- from the four European Phylloecus species (Ph. etorofensis vula 3. Janus compressus (Fabricius, 1793) also has some (Takeuchi, 1955), Ph. faunus Newman, 1838, Ph. niger (M. entirely pale terga, but in the female at least terga 3–7 are Harris, 1779), and Ph. xanthostoma (Eversmann, 1847))  80 LISTON & PROUS: Stenocephus janseni sp. nov. from Germany (Hymenoptera: Cephidae) Fig. 19. Maximum likelihood tree of Cephidae based on combined COI and nuclear (POL2 + NaK) genes. Numbers at branches show SH-aLRT support (%) / ultrafast bootstrap support (%) values. Support values for weakly supported branches (<90) are not shown. Letters “f” and “m” stand for “female” and “male” if known. Numbers at the end of the tip labels refer to the length of the sequence. The tree was rooted according to the results of M N (2015) and A K (2020). The scale bar shows the number of estimated substitutions per nucleotide position. are the absence of any pale markings on the head capsule racters seem to fit Stenocephus better. The differences in (in the others, there is at least a small pale spot near the the lancet of S. janseni sp. nov. to those of the other three eye on the upper inner orbit), maxillary palpomere 4 about described Stenocephus species are the most disturbing. On 1.5× as long as 6 (others: about equal in length), and its the other hand, in the absence of convincing phylogenetic long and thin antennae. Its two metatibial preapical spurs hypotheses, one can question whether some of the nominal separate S. janseni sp. nov. from Phylloecus species, which genera of Hartigiini currently treated as valid are really have one, and from Caenocephus, which have none. worth maintaining as separate. Etymology. Named after Dipl.-Biol. Ewald Jansen, for We do not wish to speculate on the status of Stenocephus his contributions to the study of European Hymenoptera, janseni sp. nov. in the central European fauna. Whether it particularly the sawfly fauna of Germany. is “native”, but has previously escaped detection, or has Host plant. Unknown. recently extended its range, or has been introduced from Habitat. Woodland dominated by Pinus sylvestris, with elsewhere, may only become clearer when further speci- much Betula pendula, and some Robinia pseudoacacia and mens are collected. Quercus robur. Diverse woody plants in understorey, such as Populus tremula, Crataegus sp., Prunus spinosa, and P. Acknowledgements serotina. Field layer dominated by grasses, with patches We thank Stephan Blank (Müncheberg, Germany) for of Rubus fruticosus agg. sharing valuable insights gained from his unpublished Distribution. Germany: Brandenburg. studies on the morphological characters of Hartigiini, particularly Miscocephus. Eva Kleibusch (Müncheberg) Discussion and conclusions supported sequencing work. Marko Mutanen (Oulu, Fin- Genetic data for many cephid taxa, both genera and land) kindly allowed the use of unpublished COI sequences species, are still unavailable, particularly for most of the in BOLD. Dominik Vondráček (Prague, Czech Republic) monotypic or species-poor genera of Hartigiini proposed suggested some helpful improvements to the manuscript. in the last two decades by taxonomists in China and Japan. Partly because no robust phylogenetic analysis of the Harti- References giini is currently available, we refrain from describing a AYDEMIR M. N. & KORKMAZ E. M. 2020: Comparative mitogenomics new genus for Stenocephus janseni sp. nov. Its placement of Hymenoptera reveals evolutionary differences in structure and in Stenocephus is necessarily provisional, and was decided composition. International Journal of Biological Macromolecules upon after consideration of the extent to which existing 144: 460–472. BENSON R. B. 1946: Classification of the Cephidae (Hymenoptera characterisations of individual genera would have to be Symphyta). Transactions of the Royal Entomological Society of modified to accommodate it. Some grounds could have London 96 (6): 89–108. been found for placing it in Phylloecus, but, as outlined GOULET H. 1992: The genera and subgenera of the sawflies of Canada above (see: Morphological character states), most cha- and Alaska: Hymenoptera: Symphyta. In: The Insects and Arachnids  Acta Entomologica Musei Nationalis Pragae, volume 61, number 1, 2021 81 of Canada. Part 20. Agriculture Canada, Publication 1876, Ottawa, SCHMIDT S., TAEGER A., MORINIÈRE J., LISTON A., BLANK S. 235 pp. M., KRAMP K., KRAUS M., SCHMIDT O., HEIBO E., PROUS GUINDON S., DUFAYARD J., LEFORT V., ANISIMOVA M., HORDIJK M., NYMAN T., MALM T. & STAHLHUT J. 2017: Identification W. & GASCUEL O. 2010: New algorithms and methods to estimate of sawflies and horntails (Hymenoptera, ‘Symphyta’) through DNA maximum-likelihood phylogenies: assessing the performance of barcodes: successes and caveats. Molecular Ecology Resources 17: PhyML 3.0. Systematic Biology 59: 307–321. 670–685. GUSSAKOVSKIJ V. V. 1935: Insectes Hyménoptères, Chalastogastra SHANOWER T. G. & HOELMER K. A. 2004: Biological control of 1. Fauna SSSR, Vol. 2(1). Moskva, Leningrad, 453 pp. (in Russian). wheat stem sawflies: past and future. Journal of Agricultural and HOANG D. T., CHERNOMOR O., VON HAESELER A., MINH B. Urban Entomology 21 (4): 197–221. Q. & VINH L. S. 2018: UFBoot2: Improving the ultrafast bootstrap SHINOHARA A. 1999: A study on stem boring sawflies (Hymenoptera, approximation. Molecular Biology and Evolution 35 (2): 518–522. Cephidae) of the tribe Hartigiini from Japan and Korea. Japanese JOMBART T., ARCHER F., SCHLIEP K., KAMVAR Z., HARRIS R., Journal of Systematic Entomology 5 (1): 61–77. PARADIS E., GOUDET J. & LAPP H. 2017: apex: phylogenetics SMITH D. R. 1986: The berry and rose stem-borers of the genus Harti- with multiple genes. Molecular Ecology Resources 17: 19–26. gia in North America (Hymenoptera: Cephidae). Transactions of the KALYAANAMOORTHY S., MINH B. Q., WONG T. K. F., VON HAE- American Entomological Society 112: 129–145. SELER A. & JERMIIN L. S. 2017: ModelFinder: Fast Model Selection SMITH D. R. 1999: Identity of Syrista speciosa Mocsáry and notes on for Accurate Phylogenetic Estimates. Nature Methods 14: 587–589. the Genus Urosyrista Maa (Hymenoptera: Cephidae). Proceedings of KATOH K. & STANDLEY D. M. 2013: MAFFT Multiple Sequence the Entomological Society of Washington 101 (2): 285–289. Alignment Software Version 7: Improvements in Performance and SMITH D. R. & SCHMIDT S. 2009: A new subfamily, genus, and spe- Usability. Molecular Biology and Evolution 30: 772–780. cies of Cephidae (Hymenoptera) from Australia. Zootaxa 2034: 56–60. LACOURT J. 2020: Sawflies of Europe. Hymenoptera of Europe 2. SMITH D. R. & SOLOMON J. D. 1989: A new Janus (Hymenoptera: [translated from the original French by Anna Hallewell]. N. A. P. Cephidae) from Quercus, and a key to North American species. Editions, Verrières-le-Buisson, 876 pp. Entomological News 100 (1): 1–5. LISTON A. D. 2006: Beitrag zur Pflanzenwespenfauna von Brandenburg TAEGER A., ALTENHOFER E., BLANK S. M., JANSEN E., KRAUS und Berlin (Hymenoptera, Symphyta). Nachrichtenblatt der Bayeri- M., PSCHORN-WALCHER H. & RITZAU C. 1998: Kommenta- schen Entomologen 55 (3/4): 65–76. re zur Biologie, Verbreitung und Gefährdung der Pflanzenwespen LISTON A. D. & PROUS M. 2014: Sawfly taxa (Hymenoptera, Sym- Deutschlands (Hymenoptera, Symphyta). Pp. 49–135. In: TAEGER phyta) described by Edward Newman and Charles Healy. ZooKeys A. & BLANK S. M. (eds): Pflanzenwespen Deutschlands (Hyme- 398: 83–98. noptera, Symphyta). Kommentierte Bestandsaufnahme. Goecke & LIU L., CHU B. & WEI M. 2017: Two new species of Janus Stephens Evers, Keltern, 367 pp. (Hymenoptera: Cephidae) from China with a key to Chinese species. TAEGER A., BLANK S. M. & LISTON A. D. 2010: World Catalog of Entomotaxonomia 39 (3): 238–246. Symphyta (Hymenoptera). Zootaxa 2580: 1–1064. MACEK J., ROLLER L., BENEŠ K., HOLÝ K. & HOLUSA J. 2020: VILHELMSEN L. 2000: Before the wasp-waist: comparative anatomy Blanokřídlí České a Slovenské republiky II. Širopasí. [Hymenoptera and phylogenetic implications of the skeleto-musculature of the of the Czech and Slovak Republics II. Symphyta.] Academia, Praha, thoraco-abdominal boundary region in basal Hymenoptera (Insecta). 669 pp. (in Czech). Zoomorphology 119: 185–221. MALM T. & NYMAN T. 2015: Phylogeny of the symphytan grade of WEI M. 1999: Two new genera and three new species of Hartigiini (Hy- Hymenoptera: new pieces into the old jigsaw(fly) puzzle. Cladistics menoptera: Cephidae). Acta Zootaxonomica Sinica 24 (2): 201–205. 31: 1–17. WEI M. & NIE H. 2007: Two new genera of Cephidae (Hymenoptera) NGUYEN L., SCHMIDT H. A., VON HAESELER A. MINH B. Q. from Eastern China. Acta Zootaxonomica Sinica 32 (1): 109–113. 2015: IQ-TREE: A fast and effective stochastic algorithm for esti- WEI M., NIU G. & YAN J. 2015: Review of Stenocephus Shinohara mating maximum-likelihood phylogenies. Molecular Biology and (Hymenoptera: Cephidae), with description of a new species from Evolution 32: 268–274. China. Proceedings of the Entomological Society of Washington 117 PRICE M. N., DEHAL P. S. & ARKIN A. P. 2010: FastTree 2 – Appro- (4): 508–518. ximately Maximum-Likelihood Trees for Large Alignments. PLoS WEI M. & SMITH D. R. 2010: Review of Syrista Konow (Hymenoptera: ONE 5 (3) (e9494): 1–10. Cephidae). Proceedings of the Entomological Society of Washington PROUS M., LISTON A., KRAMP K., SAVINA H., VÅRDAL H. & 112 (2): 302–316. TAEGER A. 2019: The West Palaearctic genera of Nematinae (Hy- ZHELOCHOVTSEV A. N. 1968: Novye vidy Symphyta (Hymenoptera) menoptera, Tenthredinidae). ZooKeys 875: 63–127. fauny SSSR. [New species of Symphyta (Hymenoptera) of the fauna R CORE TEAM 2019: R: A language and environment for statistical of the USSR.]. Sbornik Trudov Zoologicheskogo Muzeya MGU 11: computing. R Foundation for Statistical Computing, Vienna, Austria. 47–56 (in Russian). URL https://www.R-project.org/. ZHELOCHOVTSEV A. N. 1976: Materialy po faune pilil’shchikov RIES D. T. 1937: A revision of the Nearctic Cephidae (Hymenoptera: i rogokhvostov sredney Azii, I. [Materials on the fauna of sawflies Tenthredinoidea). Transactions of the American Entomological and woodwasps of Central Asia, I.]. Sbornik Trudov Zoologicheskogo Society 63: 259–324. Muzeya MGU 15: 3–73 (in Russian). SCHEIBELREITER G. K. 1978: The poppy-cephid, Pachycephus smyr- nensis Stein (Hymenoptera: Cephidae). Zeitschrift für Angewandte Entomologie 86 (1): 19–25. 82 LISTON & PROUS: Stenocephus janseni sp. nov. from Germany (Hymenoptera: Cephidae)