Zootaxa 4614 (3): 511–528 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4614.3.5 http://zoobank.org/urn:lsid:zoobank.org:pub:0D818E35-181B-4F2C-845C-3E456D45804B

The West Palaearctic and Endophytus (, )

ANDREW LISTON1,4, MARKO PROUS1,2 & HEGE VÅRDAL3 1Senckenberg Deutsches Entomologisches Institut, Eberswalder Straße 90, 15374 Müncheberg, Germany. E-mail: [email protected] 2Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia. E-mail: [email protected] 3Swedish Museum of Natural History, Box 50007, SE-10405 Stockholm, Sweden. E-mail: [email protected] 4Corresponding author

Abstract

Six valid species of Pseudodineura are now recognised as occurring in the West Palaearctic, and the only described species of the related genus Endophytus. Larvae of all species are -miners in . An identification key to adults is provided, followed by species commentaries which include summarised data on , larval host , and distribution, with particular reference to Sweden. Whereas identification of some specimens using morphological characters may not be possible, each species apparently has a distinct COI barcode sequence. Pseudodineura heringi (Enslin, 1921) is a new junior synonym of P. parvula (Klug, 1816). Pseudodineura mocsaryi Zombori, 1976 and P. scaligera Zombori, 1979 are new junior synonyms of P. clematidisrectae Hering, 1935. Lectotypes are designated for: minutus Hartig, 1837, Pelmatopus clematidis Hering, 1932, P. enslini Hering, 1923, P. heringi Enslin, 1921, and P. mentiens var. konowi Enslin, 1921.

Key words: Symphyta, , , Sweden, key, taxonomy, distribution

Introduction

Pseudodineura Konow, 1885 is a small, Holarctic genus of nematine sawflies, containing only ten valid species (Smith 1976, Smith et al. 2010, and this work). Endophytus Hering, 1934 is monotypic, and only known from the Palaearctic (Taeger et al. 2010). These genera form together with the Nearctic Kerita (3 species) a monophyletic lineage which is sometimes called the tribe Pseudodineurini Benson, 1938 (Prous et al. 2014). The larvae of Pseu- dodineura and Endophytus mine the of Ranunculaceae, whereas the only known host of a Kerita species is Mertensia virginica, Boraginaceae (Smith 2009). With the exception of fuscula, adults of the European Pseudodin- eura species are rarely collected, probably because of their small size, short flight period, and generally narrow host range (Altenhofer & Pschorn-Walcher 2006). Furthermore, some species are mono- or oligophagous on plant taxa such as Pulsatilla species, Anemone sylvestris and recta, which have suffered massive declines in large parts of their European ranges. On the other hand, where they do occur, Pseudodineura species can sometimes reach high population levels, made evident by abundance of their conspicuous leaf-mines. Two species, enslini and mentiens, are regularly found in parks and gardens, and not infrequently regarded as a nuisance, because at high population densities they render the leaves of their hosts unsightly, and can reduce the production of (Buhr 1941, Gerlach 2005).

Material and methods

The names of collections referred to in the text are abbreviated as follows:

Accepted by A. Taeger: 23 Apr. 2019; published: 11 Jun. 2019 511 FMNH Finnish Museum of Natural History, Helsinki, Finland HNHM Hungarian Natural History Museum, Budapest, Hungary MNHN Muséum national d’Histoire naturelle, Paris, France MSNG Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy MZLU Lunds universitet, Entomology Collection, Lund, Sweden NHRS Naturhistoriska riksmuseet, Stockholm, Sweden SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany VVCT Private collection of Veli Vikberg, Turenki, Finland ZMHB Naturkundemuseum, Berlin, Germany ZSM Zoologische Staatssammlung, Munich, Germany

Registration numbers of specimens allocated by the SDEI (five digits, omitting the prefix „DEI-GISHym“) are cited in the figure captions, thus enabling the corresponding collection data to be located in the list of specimens examined. An asterisk (*) preceding a country name in the list of specimens examined indicates that we know of no previous published records from that country. Morphological terminology follows Viitasaari (2002), except for the use of sawtooth instead of serrula (see Malagón-Aldana et al. 2017). Genitalia were macerated in room-temperature 10–15% KOH for four to five hours before being mounted temporarily in glycerine on glass slides for photography. After photography, the dissected parts were gummed with Berlese fluid to a card fixed to the specimen’s pin.

Results

Character assessment

Antennae. We counted the number of flagellomeres of 79 females and 25 males of the six West Palaearctic Pseudo- species in the SDEI collection. A few additional specimens lacked antennae and could not be scored. The majority have seven flagellomeres (both antennae: 7/7), except: three females of 48 fuscula, two with 7/8 flagel- lomeres, and one with 8/8; two females and one male of 24 parvula, the females with 8/8 and 7/8 flagellomeres, the male with 8/8. The “additional” tenth antennomere is variable in proportions, even between each of the same specimen. It seems that the presence of an additional flagellomere may be rather infrequent in European populations of Pseudodineura. Until the taxonomic significance, if any, of the number of flagellomeres in this genus has been better clarified, this character should not be given undue weight. Smith et al. (2010) compared the single Alaskan specimen described in that paper as P. kasatochi D. R. Smith with the otherwise apparently similar P. heringi, and attributed high diagnostic significance to the holotype of kasatochi having both antennae with eight flagellomeres. More significant are the proportions of the flagellomeres, which in some cases show clear differences between species in the length of a basal flagellomere compared to a more apical one, or the ratio of width to length in a single flagellomere. Colour pattern. The species are similarly coloured. The head and body are black, with more or less pale mouth- parts and legs, and in some species with pale markings (yellow-brown) on the thorax and . Although the range of variability in body coloration within most species is rather narrow, P. mentiens is an exception: the abdo- men may be almost completely black, to entirely pale. The colour of the metafemur is constant within most species, except clematidis: entirely pale in mentiens and clematidisrectae, and clearly partly black in all the others. Males of Pseudodineura clematidis can have entirely pale, or slightly dark-marked femora. We have used femur colour as the entry character in our identification key, but one should examine the specimen from more than one angle, or look at more than one leg, because an entirely pale femur, which is both translucent and reflective, may still look dark if it is near to black areas of the body. Genetic data. During the past few years, genetic data (mainly COI barcodes) for all of the European Pseudodin- eura species, and Endophytus anemones, have become available (Boevé et al. 2009, Schmidt et al. 2017). In BOLD Systems (boldsystems.org), all specimens of each species group in single BINs (Barcode Index Number Uniform Resource Identifier), except that there is no BIN assigned yet to P. clematidis, because of the lack of sufficiently long barcode sequences. Based on the partial COI barcode of P. clematidis, the closest is P. clematidisrectae differ- ing at least by 4.6%. Barcoding can therefore be recommended for species identification in these genera.

512 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL We retain Endophytus as a separate genus from Pseudodineura, even though most of the morphological char- acters which distinguish them could be regarded as representing species-level variability within a single genus. Although the differences in wing venation have the quality traditionally associated with genus-level classification, these character states are very unstable in Endophytus. The main reason for treating the genera as distinct is their large genetic distance from each other (see Prous et al. 2014, fig. 2). At present, the genetic distances between En- dophytus, Pseudodineura and Kerita appear to be rather similar, but their relationships could still change when more data becomes available.

FIGURES 1–8. Pseudodineura species. 1, enslini (17050). 2, fuscula leaf-mine in Ranunculus sp. (31922). 3, clematidis- rectae ♀ (mocsaryi holotype, 31821). 4, parvula ♀ (31886). 5, clematidisrectae ♀ (11441) antenna. 6, clematidis ♀ (31812) antenna. 7, mentiens ♀ (19816) leg colour. 8, clematidis ♀ (88876) leg colour. Scale bars 1mm.

West Palaearctic Pseudodineura Zootaxa 4614 (3) © 2019 Magnolia Press · 513 Genitalia. For the sawsheath, see under that term, below. The sawteeth of the lancet of Endophytus anemones are flat (Fig. 33), contrasting with the prominently pro- jecting lobes of all Palaearctic Pseudodineura (Figs 27–32). Within Pseudodineura, examination of the lancet is of limited use for identification: only that of fuscula is readily recognisable, by its relatively much larger sawteeth. The lancets of the other species are very similar, and scarcely distinguishable. The penis valves of all species are also closely similar, except for that of enslini, which differs in the proportionately higher distal part, and the ventro-api- cal lobe (Fig. 21, vl) is markedly wider than in any other species. Note that the distal filament (Fig. 22, df) is very fragile, and easily broken during preparation. Sometimes it is folded back behind the valve pseudoceps, and is then hardly visible. The apparent curvature and position of the filament relative to other parts of the valve are greatly af- fected by preparation, and are definitely not useful characters. Hering (1923) suggested that the shape of the harpes is diagnostic for some species, but we did not detect more than very slight differences between single specimens of each species, and these seem likely to lie within the range of intraspecific variability. Possibly Hering had been misled by the appearance of the harpes in situ. They are prone to distortion in dry specimens, and should be mounted on a slide for proper comparison. Length of malar space. The very narrow malar space is one of the generic characters of Pseudodineura (Prous et al. 2014). Benson (1958) stated that the malar space of fuscula is “about as long as one and a half facets of com- pound eye”, whereas for enslini he wrote that this was “about as long as two compound eye facets [CEF]”. Although we also found differences between species in the length of the malar space (e.g. nearly absent or maximally as long as one CEF in mentiens, and about as long as one and a half to four CEF in clematidis), smaller differences such as those given by Benson are very likely to be obscured by the intrinsically high measurement uncertainty. In practice, it is hard to define exactly where the edge of the eye is, and small changes in the angle of viewing also greatly alter the perceived malar space length. Furthermore, in the same specimen left and right malar spaces often appear to be of different lengths. Sawsheath. The shape of the valvulae 3 in dorsal view, together with the arrangement of their distal setae, are of great importance in distinguishing the species. However, it is often difficult to view the valvulae 3 in their entirety, when they are partly retracted into the abdomen. Furthermore, in many reared specimens which we examined the abdomen apex was more or less “dirty”, and the profile of the valvulae thus hard to discern. In such cases, the ap- pearance of the setae is often still sufficient for a reliable identification. Although a number of authors have stated that the valvulae of heringi are distally acute, and bluntly rounded or narrowly truncate in parvula, we concluded that this character is not reliable. It seems that the tip of the lancet of most dried specimens of Pseudodineura protrudes from the sawsheath. When examining specimens, it is important not to mistake the tip of the saw for the prominent ventroapical keel on the valvulae of fuscula. Only in fuscula is this structure so strongly developed. Venation. Zombori (1979) considered the angle of the posterior portion of fore wing vein M where it joins Cu, and the position of the intersection of fore wing vein cu-a on Cu to be valuable characters at the species level, but in the material which we examined, both characters showed high intraspecific variability, and we do not consider either to be of use for identification purposes. Vikberg (1967) described several more or less variable venation char- acters for Endophytus anemones. Larvae. The following characterisation is adapted mainly from Lorenz & Kraus (1957). Habitus: see Fig. 1. Length up to about 11mm. Head flat and very wide. Antennae flat, comprising three antennomeres. Body green or whitish. Prosternum with 2 anterio-lateral flecks and one median fleck. Meso- and metasternum each with a dark or pale brown median fleck. Abdominal sternum 1 rarely with a fleck. The last two abdominal terga with black flecks. Thorax not strongly flattened, without sclerotised prothoracic shield. Second annulet of abdominal segments with 2, third with 3 setae. Surpedal and subspiracular lobes with 2–3 setae. Spiracles small and brown. Thoracic legs more or less blackened, tibia with about 7 setae. Larvae of Pseudodineura, and the similar larva of Endophytus anemones, can be distinguished from Fenusini () leaf-miners by their hardly flattened thorax (strongly flattened in Fenusini), abdominal segments with 4 annulets (2 in Fenusini), the absence of the sclerotised pronotal shield (present in Fenusini), and anal prolegs separated (fused in Fenusini). Furthermore, the host plants of West Palaearctic Fenusini species are diverse taxa of Betulaceae, Fagaceae, Geraniaciae, Malvaceae, Rosaceae, Salicaceae, Sapindaceae and Ulmaceae, but not Ranun- culaceae. Lorenz & Kraus (1957) provided a key and descriptions for the larvae of five European Pseudodineura species together with E. anemones. The larva of P. parvula remains undescribed.

514 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL Life history: general

A thorough study of the biologies of the European Pseudodineura species was made by Altenhofer & Pschorn-Wal- cher (2006), and the information which we present below is mostly taken from that work. Boevé et al. (2009) added data on the chemical defences of larvae. Little is known about sex-ratios. Although the reared material that we ex- amined contained at least a few males of all species, the sex ratio seems generally to be strongly female-biased. All West Palaearctic species are univoltine, except perhaps for mentiens, in which the temporal pattern of occurrence of larvae, with two maxima during the growing season, could be interpreted as the result of a protracted, bi-modal emergence pattern, or alternatively as an indication of bivoltinism. For all species, based on the position of newly- started mines, oviposition is normally into the leaf edge. At first, the young larva makes a narrow gallery along the leaf edge. Later, the mine becomes a “blotch”, and may occupy most of the leaflet, depending on which host plant species is involved. Pseudodineura and Endophytus mines are typically full-depth, and accordingly highly transparent (Fig. 2), unlike mines of Agromyzidae (Diptera) which are frequently encountered on the same plants. Compared with the only other leaf-mining sawflies occurring in the Palaearctic (i.e. the various genera formerly mostly placed in the Heterarthrinae, but now regarded as belonging to the Blennocampinae: see Malm & Nyman 2015), Pseudodineura species are remarkable in that their larvae can move to a new leaf, or leaflet, during their development (Buhr 1941). Larvae of the blennocampine leaf-miners lack this ability. Development from oviposition to the making of a cocoon takes about 6–8 weeks. The cocoon is made in the soil, in which a fully pigmented overwinters. This is a rare trait in West Palaearctic sawflies, otherwise only recorded for Dolerus species (Oude- mans 1925): most other taxa overwinter as eo- or pronymphs, and a few as eggs (Lorenz & Kraus 1957). Prolonged diapause is not uncommon in Pseudodineura, particularly in populations in cool and humid regions. Although most individuals which enter prolonged diapause emerge after two winters, a few pass three winters in the cocoon.

Key to West Palaearctic Pseudodineura and Endophytus species (imagines)

Note: The only described species of Endophytus is included, because of its morphological similarity to Pseudodin- eura. These genera can usually be distinguished by wing venation characters (Prous et al. 2014), which are however highly variable in Endophytus (Vikberg 1967). The male of Endophytus anemones is unknown.

1 a ♀♀ ...... 2 - aa ♂♂ ...... 8 2 a Femora entirely pale, rarely with small diffusely darkened area (Figs 3, 7–8)...... 3 - aa Femora at least basally completely black on one or both surfaces (Fig. 4)...... 5 3a Valvulae 3 (Figs 15–16) in dorsal view basally subparallel-sided; apically almost truncate, or broadly rounded b Tarsomeres 2–5, and sometimes 1, fuscous above (Fig. 8)...... 4 - aa Valvulae 3 (Fig. 14) in dorsal view tapering from base; apically narrowly rounded bb All tarsomeres pale (Fig. 7)...... Pseudodineura mentiens (Thomson, 1871)♀ 4 a Valvula 3 apically more truncate, setae shorter and denser (Fig. 15) b Flagellomere 4 is 2.8–3.0 × as long as distal width (Fig. 5)...... Pseudodineura clematidisrectae Hering, 1935♀ - aa Valvula 3 apically more rounded, setae longer and sparser (Fig. 16) bb Flagellomere 4 is approx. 4.0 × as long as distal width (Fig. 6) ...... Pseudodineura clematidis (Hering, 1932)♀ 5 a Upper half of mesepisternum unsculptured and shiny between the setae (Fig. 9) b Valvulae 3 in dorsal view less than 2 × as long as basal width, subtriangular or basally subparallel-sided (Figs 18–20) c Fore wing vein 2A+3A basally straight, not curved towards 1A (Fig. 12) ...... 6 - aa Upper half of mesepisternum sculptured and dull, contrasting with shiny, unsculptured lower half (Fig. 10) bb Valvulae 3 in dorsal view more than 2 × as long as basal width, basally subparallel-sided (Fig. 17) cc Fore wing vein 2A+3A basally usually strongly curved towards 1A (Fig. 11), but can be incomplete...... Endophytus anemones (Hering, 1924)♀ 6 a Valvulae 3 in dorsal view basally subparallel-sided, apically broadly rounded or medially emarginate, about as long as basal width (Figs 19, 20)...... 7 - aa Valvulae 3 in dorsal view subtriangular, apically appearing acute or very narrowly truncate, about 1.5 × as long as basal width (Fig. 18) ...... Pseudodineura parvula (Klug, 1816)♀ 7 a Valvulae 3 (Fig. 19) in dorsal view apically rounded, with a carinate medial projection; medial setae pale, almost as long as lateral ones...... Pseudodineura fuscula (Klug, 1816)♀ - aa Valvulae 3 (Fig. 20) in dorsal view apically more or less truncate or medially emarginate; medial setae dark, much shorter than lateral ones...... Pseudodineura enslini (Hering, 1923)♀ 8 a Femora entirely pale, rarely with small diffusely darkened area (Figs 3, 7–8)...... 9

West Palaearctic Pseudodineura Zootaxa 4614 (3) © 2019 Magnolia Press · 515 - aa Femora at least basally completely black on one or both surfaces (Fig. 4)...... 10 9 a Metatarsus dorsally fuscous (darker than base of metatibia) (Fig. 8) b Upper half of mesepisternum may be pale (Fig. 13) ...... Pseudodineura clematidis (Hering, 1932)♂, Pseudodineura clematidisrectae Hering, 1935♂ - aa Metatarsus entirely pale (Fig. 7) bb Mesepisternum entirely black (Fig. 9)...... Pseudodineura mentiens (Thomson, 1871)♂ 10 a Penis valve (Fig. 21) distally very high; ventroapical lobe strongly enlarged...... enslini (Hering, 1923)♂ - aa Penis valve (Figs 22, 23) distally narrow; ventroapical lobe not enlarged...... 11 11 a Penis valve ventroapical lobe distally with a few sparse pores (Fig. 22)...... Pseudodineura parvula (Klug, 1816)♂ - aa Penis valve ventroapical lobe distally with a small group of closely spaced pores (Fig. 23) ...... Pseudodineura fuscula (Klug, 1816)♂

FIGURES 9–20. Pseudodineurini species. 9, enslini ♀ (19535) mesepisternum. 10, Endophytus anemones ♀ (15403) mesepis- ternum. 11, Endophytus anemones ♀ (15403) wings. 12, enslini ♀ (15348) wings. 13, clematidis ♀ (31812) mesepisternum. 14, mentiens ♀ (19896) valvulae 3. 15, clematidisrectae ♀ (31880) valvulae 3. 16, clematidis ♀ (31840) valvulae 3. 17, Endophytus anemones ♀ (15403) valvulae 3. 18, parvula ♀ (31886) valvulae 3. 19, fuscula ♀ (31817) valvulae 3. 20, enslini ♀ (31816) valvulae 3.

516 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL FIGURES 21–26. Pseudodineura species, penis valves; 21, enslini (15347) vl = ventroapical lobe. 22, parvula (31894) df = distal filament. 23, fuscula (31891). 24, clematidisrectae (31889). 25, clematidis (31843). 26, mentiens (31893).

Species treatments

Endophytus anemones (Hering, 1924)

Pelmatopus anemones Hering, 1924: 238–240. Holotype ♀ (GBIF-GISHym2806; images: https://doi.org/10.6084/ m9.figshare.7880252) in ZMHB, examined. Type locality: Berlin, Finkenkrug (Germany). Endophytus anemones: Hering 1934: 353. Neopelmatopus anemones: Conde 1934: 181. Pelmatopus tenuiserra Lindqvist, 1949: 66. Holotype ♀ (http://id.luomus.fi/GL.7592) in FMNH, examined. Type locality: Kar- jalohja (Finland). Synonymy with anemones by Vikberg (1967). Pseudodineura tenuiserra: Hellén 1960: 6. Pseudodineura anemones: Verzhutskii 1981: 151–152.

Additional description. Body length: ♀ 3.0–4.0mm [♂ unknown]. Entirely black, except for pale labrum, palps, more or less clypeus, tibiae, more or less tarsi, and apical parts of femora. Wing venation anomalies, such as the presence or absence of fore wing veins 2r-rs and 2m-cu (Fig. 11), are particularly common in E. anemones (see Vikberg 1967 for more details). The structure of the frontal area also varies significantly (Vikberg 1967). Lancet: Fig. 33. Total number of specimens examined: 18♀. Similar species. The very narrow valvulae 3 in dorsal view, rough upper mesepisternum, and usually nearly closed basal part of the fore wing anal cell are all characters which readily distinguish E. anemones from the Euro- pean Pseudodineura species. Life history. Host plants: Anemone nemorosa (Hering 1924, Altenhofer & Pschorn-Walcher 2006), and Anem- one ranunculoides (Hering 1935) in Europe; according to Verzhutskii (1981) Anemone reflexa in Siberia. Distribution. Central and northern Europe, north to southern Finland (Vikberg 1967), eastern Siberia (Ver- zhutskii 1981). Occurrence in Sweden: Skåne (Benander 1966), Västmanland, Ångermanland (own records: mines). Specimens examined. Austria: 2♀. Finland: 4♀ (SDEI). Including: 1♀ (DEI-GISHym15403), Helsinki, 04.05.1973, leg. M. Viitasaari (SDEI). 1♀ (PR264VV), Helsinki, 24.05.1958, leg. O. Ranin (VVCT). Germany: 7♀ (SDEI, ZMHB). Some reared from Anemone nemorosa. Sweden: Västmanland: 1 empty leaf-mine, Anemone nemorosa, Lindesberg 11km NW, 90 m, 59.67°N 15.14°E, 1.vi.2013, A. Liston (photographic record). Ångermanland: 1 leaf-mine with larva, Anemone nemorosa, Ramvik, 40 m, 62.80°N 17.87°E, 03.06.2013, A. Liston (photographic record).

West Palaearctic Pseudodineura Zootaxa 4614 (3) © 2019 Magnolia Press · 517 Pseudodineura clematidis (Hering, 1932)

Pelmatopus clematidis Hering, 1932: 157–158. Syntypes ♀♂. Lectotype ♀ (GBIF-GISHym2809; images: https://doi. org/10.6084/m9.figshare.7880402) in ZMHB, here designated. Type locality: Switzerland, Zernez. Paralectotypes: see un- der Specimens examined. Pseudodineura clematidis: Hering 1936: 161.

Notes: Hering (1932) selected one male and one female as “Typus”. Other specimens belonging to the type series are labelled “Paratypen”. The type series is therefore considered to have been comprised of syntypes. The lectotype designation agrees with Hering’s intention. Additional description. Body length: ♀ 3.5–5.5mm, ♂ 4.0–5.0mm. Body colour varies considerably. Speci- mens in the type series are rather pale, with the upper mesepisternum pale, but the reared specimens from Austria (DEIGisHym31840 to 31843) are very dark, and thus not distinguishable in coloration from clematidisrectae. Two reared males from Austria even show slight infuscation of small parts of the metafemur. Lancet: Fig. 29. Male: penis valve; Fig. 25. Total number of specimens examined: 9♀, 7♂. Similar species. Female Pseudodineura clematidis resemble clematidisrectae very closely in all but two char- acters which we examined: see key. Both sexes of clematidis usually have the upper mesepisternum clearly brown, with the lower part black, while in clematidisrectae the mesepisternum is usually completely black, or upper part at most obscurely paler, but exceptions occur in both taxa. Slight differences can be seen between the illustrated lancets of these two species (Figs 28, 29). The lamnium of clematidis is longer in proportion to its height, and the shape of the tangium is different. We cannot state whether these are constant differences, which might enable reli- able separation of the species. In the rather few male specimens available, the antenna of clematidis is as long as the fore wing costa and , whereas no longer than the fore wing costa in clematidisrectae. Their penis valves appear to be indistinguishable. Life history. Host plants: Clematis alpina (Hering 1932, Altenhofer & Pschorn-Walcher 2006). Distribution. Central Europe: the Alps, as well as the Tatra and Pieniny Mts in Poland (Taeger et al. 2006, Mi- chalska et al. 2010), and Urals (Sundukov 2017). According to Zhelochovtsev & Zinovjev (1995) possibly also in eastern Siberia. Occurrence in Sweden: published records; none. Material examined: no Swedish specimens. Specimens examined. Paralectotypes: all with the same data: Zernez (Schweiz), Mine Clematis alpina, 28.4.1930, Z. Nr. 3462, leg. W. Hopp. 1♀ 1♂ in ZMHB, 1♀ in ZSM, 1♀ 1♂ in NHRS. Other material: Austria: Salzburg: 1♀ (DEIGisHym31812), 1♂ (DEIGisHym31813), Obertauern [presumably Radstätter Tauernpaß: see Pschorn-Walcher & Altenhofer 2000], Clematis alpina, coll. 10.07.1997, emerged respec- tively 26.03.1998 / 11.03.1999, leg. E. Altenhofer (SDEI). Steiermark: 4♀ (including DEIGisHym88876, 31840, 31841), 4♂ (including DEIGisHym31842, 31843), NP Gesäuse, Hartelsgraben, approx. 1000m a.s.l., Clematis al- pina, coll. 15.09.2016, emerged 06–09.06.2018, leg. E. Altenhofer (SDEI).

Pseudodineura clematidisrectae Hering, 1935

Pseudodineura clematidis-rectae Hering, 1935: XII. Holotype ♂ (GBIF-GISHym2807; images: https://doi.org/10.6084/ m9.figshare.7880444) in ZMHB, examined. Type locality: Karlstadt (Maintal) [Germany, Bavaria]. Pseudodineura mocsaryi Zombori, 1976: 211–213. Holotype ♀ (DEI-GISHym31821; images: https://doi.org/10.6084/ m9.figshare.7880555) in HNHM, examined. Type locality: Karst [region in south-west Slovenia]. New synonym. Pseudodineura scaligera Zombori, 1979: 237–238. Holotype ♀ (DEI-GISHym31814; images: https://doi.org/10.6084/ m9.figshare.7880669) in MSNG, examined. Type locality: Turbigo (Lomb.), Ponte Ticino [Italy, Lombardy]. New syn- onym.

Taxonomy. Zombori (1976) compared his mocsaryi with a syntype female of clematidis. He did not mention clema- tidisrectae, and it seems likely that this taxon was unknown to him. Some of the differences noted by Zombori are characters which we also found to be useful in separating clematidis and clematidisrectae, but the holotype of moc- saryi is apparently somewhat faded, particularly the abdomen. In our opinion, all other characters of mocsaryi fit clematidisrectae, and we therefore consider them to be conspecific. Again, in his description of scaligera, Zombori (1979) does not mention clematidisrectae. Instead, he states that because of its similarly structured face, scaligera

518 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL “belongs to the mentiens-parvula species pair”. He supposed that the strongly bent posterior part of vein M of the scaligera holotype was an especially valuable diagnostic character, separating it “from all other congeners”. In fact, this character is more or less variable in several Pseudodineura species, and the vein is often strongly bent in clematidisrectae. Neither do other characters of scaligera differ from clematidisrectae, and we therefore treat them as synonyms. Additional description. Body length: ♀ 4.5–5.0mm, ♂ 4.5mm. Black: anterior edge of clypeus sometimes pale. Tegulae pale. Pronotum completely pale, to black with pale dorsal and posterior edges. Female: at least parts of some apical abdominal terga and sterna pale, and sometimes also distal part of valvula 3. Lancet: Fig. 28. Male: sternum 9 and more or less parts of adjacent terga and sterna pale. Penis valve: Fig. 24. Total number of specimens examined: 19♀, 7♂. Similar species. Most similar is Pseudodineura clematidis: see there. The female of enslini has a similarly shaped sawsheath, but apart from its darker femora, the lancet is probably distinguishable: see under enslini. Life history. Host plant: Clematis recta (Hering 1935, Altenhofer & Pschorn-Walcher 2006). Distribution. Central Europe (Taeger et al. 2006), south to Slovenia (Maček 1974). Occurrence in Sweden: published records; none. Material examined: no Swedish specimens; a specimen standing in the NHRS collection under this name is enslini. Specimens examined. Austria: 7♀, 6♂, reared from Clematis recta (SDEI, ZSM). Including: Lower Austria, 1♂ (DEI-GISHym31889), Zöbing, 23.05.1993, em. 29.04.1994, Clematis recta, leg. E. Altenhofer (SDEI). Ger- many: 10♀, netted from C. recta (SDEI). Including: 1♀ (DEIGisHym11441), Bavaria, Lower Bavaria, Landkreis Dingolfing-Landau, Rosenau, 13.05.2004, leg. Liston (SDEI). 1♀ (DEIGisHym31880), same locality as preceding, but coll. 12.05.2004 (SDEI). 1♀ (DEIGisHym11442), same locality as preceding, but coll. 16.05.2003 (SDEI).

Pseudodineura enslini (Hering, 1923)

Pelmatopus enslini Hering, 1923: 194–197. Syntypes ♀♂. Lectotype ♀ (GBIF-GISHym2802; images: https://doi.org/10.6084/ m9.figshare.7880810) in ZMHB, here designated. Type locality: Botanischen Garten, Berlin-Dahlem [Germany]. Paralec- totypes: see Specimens examined. Pseudodineura enslini: Hering 1937: 536.

Notes: Hering (1923) selected one male and one female each as “Typus” and referred to the remaining specimens as “Paratypen”. The type series therefore comprised syntypes. The lectotype designation agrees with Hering’s inten- tion. Additional description. Body length: ♀ 3.0–4.5mm, ♂ 4.5mm. Female: pronotum completely black, to pale- edged posteriorly. Tegulae black to brown. Abdomen black except more or less for valvula 3. Lancet: Fig. 30. Male: pronotum and tegulae black. Sternum 9 black, apically more or less pale. Penis valve: Fig. 21. Total number of specimens examined: 31♀, 16♂. Similar species. The sawsheath of enslini and its setation is closest to clematidisrectae, but the females of the latter differ in their prodominantly black femora (entirely pale in clematidisrectae). The lancet of enslini (Fig. 30) has smaller and proportionately somewhat narrower sawteeth than clematidisrectae (Fig. 28). Possibly there are differences in the shape of the tangium and the distribution of campaniform sensilla (“pores”), but this needs check- ing in a greater number of specimens. Externally, the male of enslini closely resembles the other two species with black-marked femora (fuscula and parvula), but is readily distinguished by its penis valve (see key). Life history. Host plants: Trollius species; in Europe on T. europaeus in semi-natural / natural vegetation (Al- tenhofer & Pschorn-Walcher 2006), and several other non-native species in parks and gardens (Buhr 1941), includ- ing T. hybridus hort. in Sweden (Wahlgren 1951). Distribution. Central and northern Europe, including the British mainland (Taeger et al. 2006), and eastern Siberia (Sundukov 2017). Frequently occurs on cultivated Trollius in gardens, as was the case for the type series. Occurrence in Sweden: published records; Skåne (Wahlgren 1944, 1951), Västergötland (Wahlgren 1963). Material examined: Småland, Dalarna, Torne Lappmark. Altenhofer & Pschorn-Walcher (2006) noted that enslini has been found in the Alps only at altitudes up to 1600m, i.e. within the subalpine zone. Our observations of mines at two localities above the tree-line in the Torne Träsk area, at approx. 700 and 900m above sea level, make an interesting comparison.

West Palaearctic Pseudodineura Zootaxa 4614 (3) © 2019 Magnolia Press · 519 Specimens examined. Paralectotypes: all with the same data: Berlin-Dahlem, rearing no. 1742, Trollius euro- paeus, various emergence dates in 1922. 1♀ (GBIF-GISHym2803) 1♂ (GBIF-GISHym2801) in ZMHUB, 1♀ 1♂ (GBIF-GISHym3890) in SDEI, 4♀ (GBIF-GISHym3476–3478, 3482) 2♂ in ZSM, 2♀ 1♂ in NHRS. Not exam- ined: 1♀ 1♂ in MSNG (Zombori 1985). Other material: Austria: 13♀, 10♂, reared from Trollius europaeus (SDEI). Including: 1♀ (DEI-GISHym19535), Lower Austria, Allentsteig, 27.06.1991, emerged 13.05.1992, leg. E. Altenhofer (SDEI). 1♀ (DEI-GISHym15348), same collection data as preceding (SDEI). 1♂ (DEI-GISHym15347), same locality as preceding, but coll. 17.06.1991 (SDEI). Salzburg: leaf-mine with larva (DEI-GISHym17050), Tappenkarsee, 1500–1750m, 47.18°N 13.32°E, 23.07.2010, leg. Liston & Taeger(SDEI). 1♀ (DEI-GISHym31816), locality unknown, reared Ranunculus aconiti- folius [misidentification of Trollius], leg. E. Altenhofer (SDEI). Germany: 5♀, two reared from Trollius europaeus (SDEI). Latvia: 1♀, 1♂, reared from Trollius europaeus (SDEI). Russia: 1♀, Siberia, Irkutsk, Coll. Konow (SDEI) [Pelmatopus parvulus det. Enslin, Pseud. fuscula det Morice] Scotland: 1♂, reared from Trollius europaeus (SDEI). Sweden: Småland: 1♀, Jönköping, 57.78°N 14.18°E, 25.05.1927, leg. Wieslander (NHRS-HEVA000003432), previously determined as P. clematidisrectae (NHRS). Dalarna: 1 large leaf-mine, Trollius europaeus, Stenis 1km N, 60.95°N 14.47°E, 15.06.2013, leg. Liston, Prous & Taeger (SDEI). Torne Lappmark: 5 young leaf-mines, Trollius europaeus, Björkliden, 500–600m, 68.41°N 18.64°E, 09.07.2016, leg. Liston & Prous (SDEI). 3 young leaf-mines, Trollius europaeus, Abisko, Mt Njulla, above tree-line, ca. 900m, 68.36°N 18.73°E, 29.07.2017, leg. Liston & Prous (SDEI). 5 young leaf-mines, Trollius europaeus, Abisko, Abisko River, ca. 380m, 68.36°N 18.77°E, 2.8.2017, leg. Liston & Prous (SDEI).

Pseudodineura fuscula (Klug, 1816)

Tenthredo () fuscula Klug, 1816: 70–71. Syntype(s) ♀, lost [leg. Gravenhorst: presumed to have been deposited in the Gravenhorst Collection]. Type locality: am Göttinger Walle [Germany, Göttingen, town wall]. Dineura despecta Hartig, 1837: 228. Syntype(s) ♀, should be in ZMHB (Hartig wrote “Mus. Kl[ug].”), but apparently lost; no type locality given. Listed as a synonym of fuscula by Konow (1905). Dolerus minutus Hartig, 1837: 244–245. Syntype(s) ♀, no type locality given. Lectotype ♀ (GBIF-GISHym2762; images: https://doi.org/10.6084/m9.figshare.7880864) in ZMHB, here designated. Type locality: Rügen [Germany, Mecklenburg- Vorpommern]. Listed as a synonym of fuscula by Konow (1905). Pseudodineura despecta: Konow 1885: 297. Pseudodineura fuscula: Konow 1890: 249. Phyllopais fusculus: Hering 1934: 353.

Taxonomy. Hering (1923) thought that specimens reared from Ranunculus auricomus belonged to a species sepa- rate from fuscula. Buhr (1941) noted that the phenology of Pseudodineura on R. auricomus was two or three weeks earlier than that of fuscula on other species of Ranunculus. A male and female reared by Hering from R. auricomus and identified as Pelmatopus fusculus are in the ZMHB, and a further pair reared by Hering from R. auricomus in the ZSM, labelled “Pelmatopus auricomi” [unpublished name]. The latter are missing the apex of the abdomen and genitalia. We have not seen other specimens reared from this host. No external morphological differences were detected between the specimens reared from auricomus and specimens from other Ranunculus species. Additional description. Body length: ♀ 2.5–5.0mm, ♂ 3.5–4.5mm. Pronotum completely black. Female: te- gulae usually black, sometimes brown. Abdomen black except more or less for apex of valvula 3. Lancet: Fig. 27. Male: pronotum and tegulae black. Sternum 9 black. Penis valve: Fig. 23. Total number of specimens examined: 79♀, 6♂. Similar species. Of the species with black-marked femora, only the female of parvula has a somewhat similarly shaped sawsheath. In cases where the shape of the sheath is not clearly observable, the larger sawteeth on the lancet of fuscula will distinguish them. Externally, the male closely resembles the other two species with black-marked femora (enslini and parvula), but can be separated by its penis valve (see key). Life history. Host plants: a large number of Ranunculus species. Altenhofer & Pschorn-Walcher (2006) found mines most abundantly on auricomus agg., repens, and platanifolius, but many other host species are used, such as

520 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL acris, lanuginosus, montanus (Altenhofer & Pschorn-Walcher 2006), cassubicus (Conde 1934), polyanthemos, bul- bosus, caucasicus, kerneri, sardous (Buhr 1941), reptans (Pschorn-Walcher & Altenhofer 2000), alpestris (Huber 1969), aconitifolius and carpaticus (reared specimens in Hering Collection, ZMHB). Pschorn-Walcher & Alten- hofer (2000) and Altenhofer & Pschorn-Walcher (2006) stated that aconitifolius is a frequent host, but specimens in SDEI labelled as reared from this plant, leg. Altenhofer, all belong to P. enslini.

FIGURES 27–30. Pseudodineura species, lancets; 27, fuscula (19541). 28, clematidisrectae (11442). 29, clematidis (31812). 30, enslini (15348).

Distribution. Southern, central and northern Europe, including British Isles (Taeger et al. 2006), Armenia, Ka- zakhstan, and eastern Siberia (Sundukov 2017). Introduced to North America (Smith 1976). Occurrence in Sweden: published records; Skåne (Benander 1966), Öland (Wahlgren 1944). Material examined: Skåne, Småland, Öland, Gotland, Gotska Sandön, Södermanland, Uppland, Ångermanland, Norrbotten. Specimens examined. Austria: 18♀, 2♂ (SDEI). Including: 1♂ (DEIGisHym31891), Lower Austria, Rapot- tenstein, Ritterkamp, 05.06.1988, em. April 1989, Ranunculus platanifolius, leg. E. Altenhofer (SDEI). 1♀ (DEI- GisHym19541), Lower Austria, Kohlveith, 6.6.1992, em. 6.5.1993, Ranunculus lanuginosus, leg. E. Altenhofer (SDEI).

West Palaearctic Pseudodineura Zootaxa 4614 (3) © 2019 Magnolia Press · 521 Canada: 1♀ (SDEI). Estonia: 2♀, 1♂ (SDEI). Germany: 23♀, 3♂ (SDEI). *Portugal: Viana do Castelo: 2♀, Paredes de Coura 6 km NNE, 480 m, 41.95°N -8.51°E, 13.05.2012, leg. Blank, Liston & Taeger (DEIGisHym15402, 19633) (SDEI). Sweden: Skåne: 1♀, Krankesjön, 55.70°N 13.47°E, 19.05.1968, leg. H. Andersson (MZLU). Småland: 2♀, [no further data], leg. Boheman (NHRS-HEVA000006299–6300) (NHRS). Öland: 1♀, Station Linné 1km E, 56.62°N 16.51°E, 01.04.2017, leg. Liston & Prous (DEIGisHym80263) (SDEI). Gotland: 1♀, [no further data], leg. Bohe- man, (NHRS-HEVA000006312) (NHRS). 1♀, Roma, 57.50°N 18.45°E, 8.6.2017, leg. Liston (SDEI); leaf-mine with larva, same data (DEI-GISHym31922) (SDEI). Gotska Sandön: 1♀, 58.37°N 19.25°E, 17.07.1946, leg. Jans- son (NHRS-HEVA000006322) (NHRS). Södermanland: 3♀, Sorunda, 59.01°N 17.82°E, 21.05.1933, leg. Malaise (NHRS-HEVA000006306–8) (NHRS). Uppland: 1♀, [no further data], leg. Lundblad (NHRS-HEVA000006302) (NHRS). 1♀, Harparbol, 59.87°N 18.00°E, 23.05.1952, leg. Lundblad (NHRS-HEVA000003431) (NHRS). 3♀, Holmiae (Stockholm), leg. Boheman (NHRS-HEVA000006309–311) (NHRS). 1♀, Danderyd, 59.41°N 18.04°E, 02.05.1937, leg. Roman (NHRS-HEVA000006319) (NHRS). 1♀, Uppsala, 59.87°N 17.63°E, 05.1891, leg. Ro- man (NHRS-HEVA000006301) (NHRS). 1♀, Uppsala, 59.87°N 17.63°E, 08.05.1936, leg. Hedgren (NHRS- HEVA000006321) (NHRS). 2♀, Erken, 59.84°N 18.65°E (NHRS-HEVA000006303–4) (NHRS). 1♀, Djurgården, 10.05.1937, leg. Malaise (NHRS-HEVA000006305) (NHRS). 1♀, Resarö, 19.05.1918, leg. Malaise (NHRS- HEVA000006314) (NHRS). 3♀, Resarö, [no dates], leg. Malaise (NHRS-HEVA000006315–7) (NHRS). Västman- land: 1♀, Nora, 59.56°N 15.01°E, 05.04.1935 (NHRS-HEVA000006320) (NHRS). Ångermanland: 1♀, [no further data], leg. Stål, (NHRS-HEVA000006313) (NHRS). Norrbotten: 1♀, Kamlunge, 65.99°N 22.86°E, 28.05.2014, leg. Liston & Prous (SDEI). 1♀, Kalix, Vassen, 65.85°N 23.16°E, 30.05.2014, leg. Liston & Prous (SDEI). 1♀ (DEI-GISHym31817), Råneå 10km SW, 65.79°N 22.16°E, 30.05.2014, leg. Liston & Prous (SDEI). 1♀, Björkfors, 65.92°N 23.46°E, 01.06.2014, leg. Liston & Prous (SDEI). 1♀, Haparanda 5km W, 65.82°N 24.03°E, 03.06.2014, leg. Liston & Prous (SDEI).

Pseudodineura mentiens (Thomson, 1871)

Blennocampa mentiens Thomson, 1871: 221–222. Holotype ♀ in MZLU (GBIF-GISHym17570; images: https://doi.org/10.6084/ m9.figshare.7880894). Type locality: Öland [Sweden]. Dineura hepaticae Brischke, 1883: 237. Syntypes ♀, probably destroyed (Blank & Taeger 1998). Type locality: Heiligenbrunn [Poland, near Gdansk]. Synonymy with mentiens by Konow (1899). Pseudodineura mentiens: Konow 1885: 297. Pseudodineura hepaticae: Konow 1885: 297. Pelmatopus mentiens var. konowi Enslin, 1921: 182 (key), 184. Syntypes ♀. Lectotype ♀ (GBIF–GISHym3893; images: https:// doi.org/10.6084/m9.figshare.7884782) in SDEI, here designated. Type locality: Rostock [Germany, Mecklenburg-Vorpom- mern]. Paralectotypes: see under Specimens examined.

Notes: the description of mentiens is based on a single specimen: “Anm. Jag har ett exemplar[..]” (Thomson 1871: 222). The name konowi was coined for female specimens with a pale abdomen. Similarly pale males also occur (see below). The pale male which we examined was reared from larvae collected at the same date and locality as a normal, dark female. All the very pale specimens were collected and emerged in August, whereas the few specimens collected in May were all dark-bodied. Additional description. Body length: ♀ 4.5–6.0mm, ♂ 4.5mm. Pronotum narrowly edged with pale, to nearly completely pale. Tegulae pale. Female: antennae black. Abdomen black, except for row of small pale spots along midline of dorsum, area of hypopygium, and more or less apex of valvula 3, to completely pale (yellowish) except more or less valvula 3. Lancet: Fig. 31. Male: terga black and sterna pale, to abdomen completely pale. Antennae of European males black, but in three males from the Russian Far East these are black dorsally and pale ventrally. Penis valve: Fig. 26. Total number of specimens examined: 9♀, 4♂. Similar species. Larger female specimens could be misidentified as clematidis, but the shape of the sawsheath should distinguish them. Furthermore, the distal sawteeth of the lancet of mentiens (Fig. 31) are more triangular and apically acute than those of clematidis (Fig. 29).

522 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL Life history. Host plants: Hepatica nobilis (= triloba) (Altenhofer & Pschorn-Walcher 2006), and transsil- vanica (=angulosa) (Buhr 1941, Liston & Blank 2006). Distribution. Central and northern Europe, north to Finland, introduced to the British mainland (Taeger et al. 2006, Liston & Blank 2006), Urals and Russian Far East (Sundukov 2017). Occurrence in Sweden: published records; Skåne (Wahlgren 1944, 1951), Öland (Thomson 1871, Wahlgren 1944), Västergötland, Uppland (Wahlgren 1963). Material examined: Öland. Specimens examined. Austria: Lower Austria, 1♀ (DEIGisHym31824), Neulengbach, c. 08.1988, em. 02.08.1990, Hepatica triloba, leg. E. Altenhofer (SDEI). 1♂ (DEIGisHym19538), same data, except c. 07.08.1989, em. 07.08.1991 (SDEI). 1♂ (DEIGisHym31893), same data, except c. 07.09.1999, em. 30.07.2001 (SDEI).1♂, same data, except c. 25.08.1999, em. 06.08.2002 (SDEI).1♂, same data, except c. 27.08.1999, em. 02.08.2001 (SDEI). 2♀, same data, except c. 25.09.1999, em. 03.08.2002 and 06.08.2002 (SDEI). Germany: 1♀ (DEIGisHym3892), Rostock, 4.5.[18]89, Coll. Konow (SDEI: paralectotype of mentiens var. konowi Enslin, 1921). 1♀ (DEIGisHym3891; images: https://doi.org/ 10.6084/m9.figshare.7880948), Rostock, Coll. Konow (SDEI: paralectotype of mentiens var. konowi Enslin, 1921). Russia: Khabarovskiy kray, 1♀, Boitsovo, 20km N Bikin Bolshoi Solntsepyok Hill, 300m, 47.02°N 134.21°E, 26.5.1993, leg. A. Taeger (SDEI). Sweden: Öland: 1♀ (DEIGisHym19896), Station Linné 1km E, 56.62°N 16.51°E, 30.05.2013, leg. Liston, Prous & Taeger (SDEI).

FIGURES 31–33. Pseudodineurini species, lancets. 31, mentiens (31824). 32, parvula (31804). 33, Endophytus anemones (PR264VV).

West Palaearctic Pseudodineura Zootaxa 4614 (3) © 2019 Magnolia Press · 523 Pseudodineura parvula (Klug, 1816)

Tenthredo (Allantus) parvula Klug, 1816: 71. Holotype ♀ (GBIF-GISHym2763; images: https://doi.org/10.6084/ m9.figshare.7881005) in ZMHB, examined. Type locality: Oesterreich [Austria]. Tenthredo minuta Lepeletier & Serville, in Latreille et al. 1828: 570. Syntype(s) ♀, may be in MNHN; environs de Paris [France, Paris area]. Listed in synonymy of parvula by Dalla Torre (1894). Primary homonym of Tenthredo minuta Christ, 1791 [ minuta]. Pelmatopus heringi Enslin, 1921: 183 (key), 185. Syntypes ♀♂. Lectotype ♀ (GBIF-GISHym3483; images: https://doi. org/10.6084/m9.figshare.7881065) in ZSM, here designated. Type locality: Rüdersdorf [Germany, Brandenburg]. Paralec- totypes: see under Specimens examined. New synonym. Pseudodineura parvula: Konow 1885: 297. Pseudodineura heringi: Hering 1935: 57.

Taxonomy. After a first examination of three females from Switzerland reared from Pulsatilla alpina, we thought that these might represent a species distinct from parvula. Compared to European specimens reared from lowland Pulsatilla species and Anemone sylvestris, the specimens from alpina are distinctive because of their differently pro- portioned flagellomeres: antennomere 3 1.25–1.45 × as long as antennomere 8, whereas in lowland specimens an- tennomere 3 1.53–1.80 × as long as antennomere 8. The three individuals reared from P. alpina also seemed to differ in the only slightly curved long setae on valvula 3, and the median mesoscutal lobes flatter, with groove shallow and obsolete on posterior. However, the specimens from Russia (Sajan) possess antennomeres proportioned within the range normal for lowland European specimens, but with sawsheath setation and median mesoscutal lobes more like the Swiss specimens. We conclude that all these specimens are conspecific, but some genetic data for popula- tions occurring at higher altitudes would still be desirable. Regarding the statement by Enslin (1921), that parvula can be distinguished from heringi by its deeper frontal pit and differently shaped sawsheath, we noted considerable variability in both of these characters, but found no correlation between the character state and the identity of the host plant from which they had been reared, i.e. Anemone sylvestris versus Pulsatilla. Having observed no stable differences between the lancets and male genitalia of specimens reared from these hosts, or genetic differences, we propose that heringi should be treated as a synonym of parvula. Additional description. Body length: ♀ 2.5–3.5mm, ♂ 3.0–4.0mm. Pronotum completely black. Female: tegu- lae brown. Abdomen completely black. Lancet: Fig. 32. Male: tegulae black or brown. Sternum 9 black except for extreme apex. Penis valve: Fig. 22. Total number of specimens examined: 14♀, 8♂. Similar species. Of the species with black-marked femora, only the female of fuscula has a somewhat similarly shaped sawsheath. When the shape of the sheath is not clearly visible, the smaller sawteeth on the lancet of parvula will distinguish them. Externally, the male closely resembles the other two species with black-marked femora (en- slini and fuscula), but can be separated by its penis valve (see key). Life history. Host plants: Anemone sylvestris (Enslin 1921, Altenhofer & Pschorn-Walcher 2006: as P. heringi), and Pulsatilla species (as P. parvula); P. patens (Conde 1934), P. vulgaris, P. grandis (Pschorn-Walcher & Alten- hofer 2000), P. narcissiflora (Hering 1935), P. pratensis bohemica (Kulfan 2008), and P. alpina (Hering 1932). Distribution. Central Europe, north to Estonia (Taeger et al. 2006, as heringi), and eastern Siberia (Sundukov 2017, as parvula and heringi). Occurrence in Sweden: not known, but could be present, particularly where the host plants occur as native species. Specimens examined. Paralectotypes of Pelmatopus heringi Enslin: all with the same data: 1♀ (GBIF- GisHym2805), Bln. Rüdersdorf, 13.2.1921, Nr. 1424, Mt. Hering (ZMHB). 1♂ (GBIF-GisHym2804), Bln. Rüdersdorf, 26.1.1921, Nr. 1424, Mt. Hering (ZMHB). 3♀ (GBIF-GisHym3484–3486), 1♂ (GBIF-GisHym3487), Bln. Rüdersdorf, various emergence dates in 1921, Nr. 1424, M. Hering (ZSM). Austria: Lower Austria, 4♀ (including DEIGisHym31804), 4♂ (including DEIGisHym31894), Langenlois, reared from Pulsatilla vulgaris, leg. E. Altenhofer (SDEI). Lower Austria, 2♀, 3♂, Sieghartskirchen, reared from Anemone sylvestris, leg. E. Altenhofer, previously identi- fied as heringi (SDEI). Germany: 2♀, 1♂, Brandenburg, Rüdersdorf [“Berlin Rüdersdf”], reared from Anemone sylvestris, leg. Hering (SDEI), previously identified as heringi (SDEI). Not types, because emergence dates in 1922 and 1924. Russia: 3♀, Ostsajan, Lager am Schwarz. Irkut, 1878m, 51°54’37’’N 100°45’41’’E, 02.07.2012, leg. W. H. Liebig (SDEI).

524 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL Switzerland: 1 ♀ (DEIGisHym31886), Graubünden, Vals, reared from Anemone alpina, 7.5.1930 [emergence date], Zucht Nr. 3433, leg. W. Hopp (ZMHB). 2 ♀ (DEIGisHym31887 and 31888), same data as previous except 8.5.1930 (ZMHB). Reared from mines on Pulsatilla alpina, collection dates 23.6.1929 and 26.6.1929 (Hering 1932).

Discussion

Our decision to synonymise Pseudodineura heringi with P. parvula was not taken lightly, because these taxa have been consistently treated as separate by other recent specialists. Presumably, a main reason for regarding them as distinct was their hypothetical specificity on different host plants: respectively Anemone sylvestris, and various Pulsatilla species. A host plant spectrum including more than one genus would indeed be exceptional in Pseudodi- neura. However, in this context it should be borne in mind that many recent botanical studies treat Pulsatilla as a synonym of Anemone (e.g. Jiang et al. 2017). Furthermore, a host plant range that is wider than most other Pseu- dodineura species is already well documented for fuscula. According to Paun et al. (2005), the Ranunculus species which we list as its hosts, excluding auricomus and two other species not included in that work (cassubicus, kerneri) belong to four main lineages, of which lineages I and VIII are the most distantly related within the Ranunculus core clade. Currently, the only apparent taxonomic uncertainty affecting West Palaearctic Pseudodineura is whether specimens of P. fuscula feeding on Ranunculus auricomus agg. are conspecific with those on other Ranunculus spe- cies. According to the available data, none of the West Palaearctic Pseudodineura, or Endophytus, shares the same host plant species. A re-examination of Pseudodineura specimens in Swedish collections did not reveal any species not previously recorded in the country, but because its host plants occur naturally in Sweden, P. parvula might perhaps have es- caped detection. In the past, some Swedish specimens had in fact been determined as parvula, but they all belong to other species: usually the widespread and rather abundant fuscula. The three Swedish (and Scandinavian) Pseudo- dineura species are thus the same ones as are known in Finland (Viramo 1969). Of these three, mentiens is naturally limited in Sweden to areas in the southern half of the country, north to approximately Örebro (www.artportalen.se), where its host Hepatica nobilis occurs. Pseudodineura fuscula is widespread through most of Sweden, and is by far the most frequently found species in the country, but is apparently absent from areas with a subarctic climate, e.g. the Torne Träsk Region. Considering that Trollius europaeus is a widespread and abundant plant in Sweden, it is surprising that there are so few records of P. enslini. It seems that, as in Finland (Viramo 1969), enslini may be rather a rare species in Sweden. Nevertheless, the few Swedish records are spread from the southernmost Province Skåne north to the Torne Träsk Region. Our records of mines from the latter area are remarkable for the high altitude (above the treeline) of two of the localities, and partly for the late date of occurrence: very small mines were found as late as the start of August.

Acknowledgments

This work was funded by the Swedish Taxonomy Initiative. For material examined, as well as other assistance and information, we particularly thank Ewald Altenhofer (Etzen, Austria) for his valuable reared adults, but also Rune Bygebjerg and Christer Hansson (MZLU), Frank Koch, Jenny Pohl and Viola Richter (ZMHUB), Manfred Kraus (Nürnberg, Germany), W. H. Liebig (Bad Muskau), Marko Mutanen (Oulu, Finland), Stefan Schmidt (ZSM), Maria Tavano (MSNG), Zoltán Vas (HNHM), Matti Viitasaari (Helsinki, Finland), and Veli Vikberg (Turenki, Finland). Katrin Elgner and Ute Kascinski (SDEI) kindly obtained some of the literature.

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