Zoosyst. Evol. 100 (3) 2024, 1017-1028 | DOI 10.3897/zse.100.125826 Gy huseue ror BERLIN On the identity and placement of Xysticus lendli Kulczynski, 1897 (Araneae, Thomisidae): an integrative approach Nikolett Gallé-Szpisjak, Robert Gallé'*, Krisztian Szab6°, Tamas Sziits? 1 HUN-REN, CER, IEB “Lendiilet” Landscape and Conservation Ecology Research Group, H-2163 Alkotmany road 2-4, Vacratot, Hungary 2 Department of Ecology, University of Szeged, H-6726, Kézép fasor 52., Szeged, Hungary 3 Molecular Ecology Research group, Department of Zoology, University of Veterinary Medicine Budapest, Rottenbiller u. 50., Budapest, H-1077, Hungary https://zoobank. org/B38CF 1 74-7F 50-41 FC-B520-2217BE1665D6 Corresponding author: Nikolett Gallé-Szpisjak (szpisjak.nikolett@ecolres.hu) Academic editor: Danilo Harms # Received 20 April 2024 # Accepted 9 June 2024 Published 25 July 2024 Abstract The species Xysticus lendli is known only from its original description of a single male and one doubtful record so far. Here, we illustrate and redescribe the species based on 34 specimens in total and describe its female for the first time. We illustrated the male palp via compound micrographs and scanning electron micrographs. We generated a DNA barcode and placed it into a current phylogenetic scaffold to confirm the species’ placement of Spiracme, a long-debated sister- or subgroup of Xysticus; hence, a new combination of Spiracme lendli (Kulczynski, 1897), comb. nov. is proposed. We illustrated the visually similar Xysticus mongoli- cus and the type species of Spiracme, S. striatipes, and compared them to S. /endli to aid future distinctions between those species. Key Words Central Europe, crab spiders, new combination, redescription, Spiracme Introduction Hungary has a rich arachnofauna, which has been well studied for a long time. Due to its unique location, gla- ciation history, and topological arrangement (the Car- pathian Basin), numerous species have been described in this region. The Pannonian biogeographic region lies on a unique meeting point of various other biogeograph- ic regions (Molnar et al. 2008; Fekete et al. 2016); for instance, the Eurasian steppe/forest steppe zone’s west- ernmost part is located here (Batori et al. 2018). On the eastern part of Hungary, the forest steppe biome is present in the Kiskunsag National Park, offering a unique oppor- tunity to study the forest steppe biota (Gallé et al. 2022a) (Fig. 1A). The arachnofauna of Kiskunsag National Park has been intensively studied due to the increased interest of ecologists and conservation biologists, who have con- ducted frequent studies in the last few decades (Gallé et al. 2022a, 2022b, 2022c, and references therein). During these studies, several new species have been discovered (Parasyrisca arrabonica Szinetar & Eichard, 2009; Ser- nokorba betyar Gallé-Szpisjak et al., 2023), and a few other “forgotten” taxa have been rediscovered as well. One of these taxa is Xysticus lendli Kulczynski, 1897, an enigmatic thomisid that was known by a single male specimen and did not have a verified record besides its original description more than a century ago. Xysticus has been the subject of several classification attempts in the past 50 or so years (see Lehtinen 2002; Jantscher 2002; and Breitling 2019 for details). Almost all of these papers used morphological data, whereas the last treatment by Breitling (2019) used the phylogeny of barcode sequences alone. One main result of the latter was the resurrection of the genus Spiracme Menge, 1876, with the type species Spiracme striatipes (L. Koch, 1870) and including nine other species: Spiracme baltistana (Caporiacco, 1935), Spiracme dura (Sorensen, 1898), Spiracme keyserlingi (Bryant, 1930), Spiracme lehtineni Copyright Gallé-Szpisjak, N. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 1018 Gallé-Szpisjak, N. et al.: The identity and placement of Xysticus lendli Kulczynski, 1897 Budapest 50 100 km Figure 1. A. Map of Europe and Hungary with the forest-steppe zone (green patches), the Hungarian sampling site (black cir- cle), the Slovakian sampling site (black star, Purgat et al. 2021), and the Serbian sampling site (black square, Grbi¢ et al. 2019); B. Forest-steppe patch; C. Open sand grassland. (Fomichev, Marusik & Koponen, 2014), Spiracme nigro- maculata (Keyserling, 1884), Spiracme quadrata (Tang & Song, 1988), Spiracme triangulosa (Emerton, 1894), Spiracme vachoni (Schenkel, 1963). Later, Purgat et al. (2021) added Spiracme mongolica (Schenkel, 1963) to the genus. While identifying our specimens, we noticed that the palp shows a high overall similarity to that of S. mongoli- ca, as illustrated by Purgat et al. (2021). Still, at the same time, significant differences could be observed between the illustrations of the original description of Schenkel (1963) and the specimen identified and pictured by Fo- michev (2015). The aims of this paper are to provide illustrations and a description for Xysticus lendli Kulczynski, 1897, based on freshly collected material, describe the hitherto zse.pensoft.net unknown female of the species, and reevaluate current faunistic records that may involve this species. We pro- vide a DNA barcode for the species and test its phylo- genetic position by reanalysing the matrix of Breitling (2019) with this new sequence added, and we discuss the accompanying results using a comparison with the type species of Spiracme. Materials and methods Specimens were collected in the Kiskunsag region, Hun- gary, in 2017, 2018, and 2021 by sweep net from semi-nat- ural sandy forest steppe patches (Fig. 1B, C). The spec- imens were photographed with a Nikon D300S camera attached to a Nikon S800 stereomicroscope and a Tucsen Zoosyst. Evol. 100 (3) 2024, 1017-1028 Truechrome Metrics camera attached to an Eclipse E200 compound microscope. Multifocal images were compiled using HeliconFocus. The female vulva and palps in Fig. 5 were illustrated while immersed in methyl] salicylate. Scanning electron micrographs were made in a low vac- uum on a Hitachi SV1000 FlexSEM scanning electron microscope in the Plant Protection Institute (NOVI) HUN-REN Centre for Agricultural Research. Drawings were made by the first author using a fineliner and 80 g/ m? of printing paper. The maps were made using Adobe Illustrator CS6 software. All measurements are given in millimetres. The material is deposited in the Museum of Natural History, Budapest (MNHB) and the first author’s personal collection in Szeged. Whole genomic DNA was extracted from one leg using standard extraction kits. For the barcoding, the LCO1940/HCO2198 primers (Folmer et al. 1994) were used. PCR has been carried out in the Department of Zo- ology of the University of Veterinary Medicine Budapest, and capillary electrophoresis has been done at Commer- cial Biomi Services (Géd6ll6, Hungary). Sequences were edited using Gap4 of the Staden Packge (Staden et al. 2000) and deposited in GenBank with the accession num- ber PP545308. Phylogenetic relationships were inferred via both maximum likelihood and Bayesian methods based on the dataset that has been published so far (Breit- ling 2019). MAFFT v.7.450 online (https://mafft.cbrce.jp/ alignment/server/) was used with default settings to align the sequences. A maximum likelihood phylogram was constructed using IQ-TREE (Nguyen et al. 2015) under the GTR+F+I+G model, performing 1000 ultrafast boot- strap replicates. Bayesian inference was performed using MrBayes v. 3.2.6 (Ronquist and Huelsenbeck 2003), us- ing the GTR+I+G model of sequence evolution. Analyses consisted of two independent runs with one cold chain and five hot chains from random starting trees, run for 10 million generations, and sampled every 1000 generations. Convergence was assessed through examination of the standard deviation of split frequencies, which was well below recommended thresholds (0.01). The two MrBayes runs were combined after the deletion of burn-in gener- ations (25%), and a majority-rule consensus phylogram was created. The resulting phylograms were edited and visualised with FigTree v.1.4 (Rambaut 2014). Abbreviations ALE — Anterior lateral eyes; AME — Anterior median eyes; Fe — Femur; MOA-WA — width of anterior me- dian ocular area, MOA-L — median ocular area length; MOA-WA — width of anterior median ocular area; MOA-WP — width of posterior median ocular area; Mt — Metatarsus; PLE — Posterior lateral eyes; PME — Posterior median eyes; RTA — Retrolateral tibial apoph- ysis; Ta — Tarsus; tb — tegular bump; Ti — tibia; tut — tutaculum; WTA — Ventral tibial apophysis. 1019 Results Taxonomy Spiracme Menge, 1876 Figs 2, 3A-C, G-I, 4A-C, 5A, B, 6-8 Type species. Spiracme striata Menge, 1864. Xysticus striata Menge, 1876, by monotypy. Note. Breitling (2019) revalidated the genus Spirac- me, but he did not provide any diagnosis or any diagnos- tic features. As delimited by Breitling (2019), it is impos- sible to give a precise diagnosis for the genus, including all and only those species. Hence, it should be subject to future revision or reconsideration. Diagnosis. Males can be recognised by the bulb lack- ing apophyses (Fig. 6A, D) or tegular apophysis (Gertsch 1953; Menge 1876; Lehtinen 2002). The tutaculum is tri- angle-shaped in this type of species. Tibia with two or three apophyses; VTA is simple and rounded; and RTA has a distinct pointed hook (Lehtinen 2002). Females can be recognised by the prominent septum (but can be absent; see Gertsch 1953). Both sex lateral eyes are 1.5 times larger than the median eyes and have light elevation (Menge 1876). Composition. Currently 10 species have been added to this genus (WSC 2024): S. striatipes (L. Koch, 1870), S. baltistana (Caporiacco, 1935), S. dura (Sorensen, 1898), S. keyserlingi (Bryant, 1930), S. lehtineni (Fomi- chev, Marusik & Koponen, 2014), S. nigromaculata (Key- serling, 1884), S. guadrata (Tang & Song, 1988), S. tri- angulosa (Emerton, 1894), S. vachoni (Schenkel, 1963). Description. see Menge (1876) and Lehtinen (2002). Distribution. Holarctic. Spiracme lendli (Kulczynski, 1897), comb. nov. Figs 2, 3A-C, 4A-C, 5A, B, 6A-C, 7, 8 Xysticus lendli Kulczynski, 1897: Chyzer and Kulczynski 1897: 301, Tab X, fig. 64 (3). Spiracme mongolica: Purgat et al. 2021: fig. 6 (misidentification). Xysticus cf. lendli: Ponomarev and Shmatko 2021: 217, fig. 16. Type material. Holotype male: HuNcary Orkény (thoroughly searched for in the collection of the Hun- garian Natural History Museum, without any success, likely lost). Material examined. HUNGARY * | male; Szank; forest steppe; 46.602°N, 19.571°E; 11 Sep. 2017; N. Gallé-Szpis- jak and R. Gallé leg.; sweep net; (HNHM Araneae-11039) ¢ | female; Zsana; forest steppe; 46.415°N, 19.621°E; 11 Sep. 2017; N. Gallé-Szpisjak and R. Gallé leg.; sweep net: (HNHM Araneae-10719) * 1 male; Harkakotony; forest steppe, 46.507°N, 19.568°E; 11 Sep. 2017; N. Gallé-Sz- pisjak and R. Gallé leg.; sweep net * 2 males; Kiskun- halas; forest steppe; 46.477°N, 19.432°E; 11 Sep. 2017; zse.pensoft.net 1020 Gallé-Szpisjak, N. et al.: The identity and placement of Xysticus lendli Kulczynski, 1897 B Figure 2. Habitus of Spiracme lendli: A, B. Male; C, D. Female; A, C. Specimen in alcohol, bleached out; B, D. Fresh specimens; Scale bar: 2.0 mm. N. Gallé-Szpisjak and R. Gallé leg.; sweep net * 3 males; Kiskunhalas; forest steppe; 46.483°N, 19.452°E; 09 Sep. 2018; N. Gallé-Szpisjak and R. Gallé leg.; sweep net « 2 males; Kiskunhalas; forest steppe; 46.493°N, 19.415°E; 21 Sep. 2018; N. Gallé-Szpisjak and R. Gallé leg.; sweep net « 1 male; Soltvadkert; forest steppe; zse.pensoft.net 46.536°N, 19.384°E; 11 Sep. 2017; N. Gallé-Szpisjak and R. Gallé leg.; sweep net * 1 male; Soltvadkert; forest steppe; 46.527°N, 19.374°E; 11. Sep. 2017, N. Gallé-Sz- pisjak and R. Gallé leg.; sweep net * 3 males; Soltvad- kert; forest steppe; 46.530°N, 19.401°E; 11. Sep. 2017, N. Gallé-Szpisjak and R. Gallé leg.; sweep net * 3 males; Zoosyst. Evol. 100 (3) 2024, 1017-1028 1021 Figure 3. Male palps: A—C. Spiracme lendli, D—F. Xysticus mongolicus; G-I. S. striatipes; A, D, G. Prolateral view; B, E, H. Ventral view; C, F, I. Retrolateral view. Scale bar: 0.2 mm. Abbreviations: tb — tegular bump; tut — tutaculum. Soltvadkert; forest steppe; 46.532°N, 19.385°E; 21. Sep. 2017, N. Gallé-Szpisjak and R. Gallé leg.; sweep net ¢ 2018, N. Gallé-Szpisjak and R. Gallé leg.; sweep net» 1 3 males; Zsana; forest steppe; 46.415°N, 19.621°E; 09. male; Zsana; forest steppe; 46.409°N, 19.621°E:; 11. Sep. Sep. 2018, N. Gallé-Szpisjak and R. Gallé leg.; sweep net zse.pensoft.net 1022 ¢ 3 males; Zsana; forest steppe; 46.396°N, 19.647°E; 09. Sep. 2018, N. Gallé-Szpisjak and R. Gallé leg.; sweep net ¢ 2 males; Imrehegy; forest steppe; 46.446°N, 19.318°E; 11. Sep. 2017, N. Gallé-Szpisjak and R. Gallé leg.; sweep net ¢ 1 male; Pirto; forest steppe; 46.493°N, 19.415°E; 20. Sep.—05. Oct. 2018, N. Gallé-Szpisjak and R. Gallé leg.; pitfall trap * 1 male; Pirto; forest steppe; 46.472°N, 19.436°E; 09. Sep. 2018, N. Gallé-Szpisjak and R. Gallé leg.; sweep net * 4 males; Pirt6; forest steppe; 46.496°N, 19.421°E; 05. Oct. 2018, N. Gallé-Szpisjak and R. Gallé leg.; sweep net * 2 males; Bocsa; forest steppe; 46.614°N, 19.464°E; 07. Sep. 2021, N. Gallé-Szpisjak and R. Gallé leg.; sweep net. Comparative material. S. striatipes HUNGARY * 2 males, 2 females; Cseng6d; mesic grassland; 46.722°N, 19.350°E; 15 Jun. 2018; N. Gallé-Szpisjak and R. Gallé leg.; sweep net. Diagnosis. Males of this species can be readily distin- guished from congeners by the thick and bent embolus, with transverse ridges (pine cone-like pattern in Fig. SA— B) near the tip. The thick embolus makes it similar to that of S. striatipes but differs by the bent embolus (straight in S. striatipes, see Figs 3, 4), the less pronounced inden- tations (deep indentations in S. striatipes, see Fig. 6), the absence of the large triangular-shaped tutaculum (present in S. striatipes, see Fig. 6), and the slightly pointing out- ward RTA (curved towards the cymbium in S. striatipes; see Figs 3, 4) as seen from the ventral side (compare Fig. 6A vs. Fig. 6D). The bent embolus is present in Xysti- cus mongolicus, and the bulb shows an overall similari- ty. However, the indentations (absent in XY. mongolicus), the tegular bump Fig. 3C (absent in XY. mongolicus, see Fig. 3F), and the twisted embolus as seen from the side (strongly bent in_X. mongolicus) separate S. lendli from X. mongolicus. Female epigyne of S. /endl/i with an up- side-down heart-shaped median septum (see Fig. 8B), which is seemingly a plate with a pronounced anterior border (Fig. 8A, C) with a clear indention on the posterior margin (Fig. 8B, C, F), probably to accommodate the thin tip of the RTA (Figs 3C, 7A). Description. Male. Total length 3.88. Carapace: 1.84 long, 1.79 wide. Abdomen: 2.17 long, 1.71 wide. Clypeus 0.16 high, chelicera 0.70 long. Eye sizes and inter-dis- tances: MOA-WA 0.22, MOA-WP 0.23, MOA-L 0.23, AME 0.07, ALE 0.10, PME 0.06, PLE 0.09, AME-AME 0.20, AME-ALE 0.13, PME-PME 0.23, PME-PLE 0.29. Colouration (on a freshly collected specimen, Fig. 2B): Carapace dark brown sides with median, longitudi- nal, beige-coloured, wide stripe. Ocular area white. Ab- domen dark brown with mottled black spots and median folium-shaped light stripe. Around folium’s lobe, three pairs of white and black spots present (Fig. 2A, B). These spots can be seen on bleached specimen as well, where the dark brown coloured areas became light brown. Che- licera, gnathocoxae, labium, and sternum sand-yellow with small dark brown spots. Legs: sand-yellow with mottled dark brown dots (uneven in size). Tibiae I-II, metatarsi I-II, and tarsi I-II with dark zse.pensoft.net Gallé-Szpisjak, N. et al.: The identity and placement of Xysticus lendli Kulczynski, 1897 brown/blackish rings on the distal end of the segment. Hind- legs light in colour, almost white, with black dots mottled on the appendage. The proximal end of tibia I'V and the distal end of the femur, patella, tibia, and metatarsus I'V with dark rings. Leg segments: I: 7.05 (2.04+0.96+1.6+1.6+0.85),; II: 6.67 (1.95+0.88+1.45+1.48+0.91); Il: 3.87 (1.19+0.58+0.83+0.67+0.6); IV: 4.51 (1.4+0.58+0.92+0.89+0.72). Palpal femur, patella tibia, and cymbium with sand-co- loured background mottled with dark brown dots and patches. Bulb without significant appendages, but tegular ledge or bump present (Figs 3C, 4C, 7A). Embolus origi- nates on prolateral part of the bulb at 11 o’clock position, twisted as seen from the side (Fig. 7) and bent as seen from the venter (Figs 3A—C, 4A—C, 5A, B), with indenta- tions on the distal third of embolus (Figs 5A, B, 6A, B), making it pine cone-like. Female. Total length: 5.99. Carapace: 2.66 long, 2.89 wide. Abdomen: 3.68 long, 2.92 wide. Clypeus 0.24 high, chelicera 1.01 long. Eye sizes and inter-distances: MOA-WA 0.58, MOA-WP 0.53, MOA-L 0.40, AME 0.11, ALE 0.16, PME 0.09, PLE 0.12, AME- AME0.40,AME-ALE0.21, PME-PME0.37,and PME-PLE 0.45. Leg segments: I: 8.49 (2.61+1.34+1.87+1.7+0.97); II: 8.49 (2.65+1.24+1.84+1.77+0.99); II: 5,53 (1.76+0.91+0.93+1.05+0.88); IV: 6.16 (1.94+0.88+1.44+1.07+0.83). Colouration lighter than in males, carapace and abdo- men patterns similar as in males (Fig. 2C, D), but with less contrast on the pattern. Leg colouration and pattern also similar to males, but overall lighter in colour. Distribution. Pannonian: Hungary, Serbia, Slovakia. Habitat. Specimens were collected in habitats char- acterised by sandy soil, covered with open dry grassland, and small forest patches (forest steppe, Fig. 1B, C). The grassland consists of drought-tolerant plant species (e.g., Alkanna tictoria, Festuca vaginata, and Stipa borystheni- ca). The main tree species of the forest patches are Poplus alba, and the bush layer includes Juniperus communis and Crataegus monogyna (see further details in Gallé et al. 2022a). Biology and phenology. Adult specimens were col- lected in September and October. We used a sweep net to collect S. /endli specimens very close to the ground surface in the herb layer. Life history remains mainly un- known. Noteworthy, females were very rarely found, so far only one. The overall ratio of the sexes was 33:1. Xysticus mongolicus Schenkel, 1963, comb. rev. Figs 3D-F, 4D-F, 5C, D Xysticus mongolicus Schenkel, 1963: Schenkel 1963: 227, fig. 127a-c (<). X. mongolicus Song, Yu & Yang, 1982: 210 (4Q); Song 1987: 284, fig. 24la-d (3'2); Song and Zhu 1997: 95, fig. 62a-d (4); Utoch- kin 1995: 67, fig. la-g (32); Zhao 1993: 388, fig. 193a-b (49); Marusik and Logunov 1990: 47, figs 48-49 (4); Fomichev 2015: 97, figs 13-15 (3). Zoosyst. Evol. 100 (3) 2024, 1017-1028 1023 A Figure 4. Distinguishing characters of Spiracme lendli (A—C) and Xysticus mongolicus: (D—F),; D. Prolateral view; B, E. Ventral view; C, F. Retrolateral view. Scale bar: 0.2 mm. Figure 5. Comparison of male emboluses: Spiracme lendli (A, B) and Xysticus mongolicus (C, D). Scale bars: 0.2 mm. zse.pensoft.net 1024 Gallé-Szpisjak, N. et al.: The identity and placement of Xysticus lendli Kulczynski, 1897 } f “7 4) tae sie a Pen 3 Figure 6. Spiracme spp. Male palps scanning micrographs Spiracme lendli: (A—C) Spiracme striatipes: (D-F), A. Bulb, ventral view; B, C. Closeup of the embolus, ventral view; D. Bulb, ventral view; E, F. Closeup of the embolus, ventral view. Scale bar: 0.2 mm (A—B, D—-E); 0.05 mm (C, F). Abbreviation: tut — tutaculum. Note. Because Purgat et al. 2021 (fig. 6) based their pro- posal of the new combination on a misidentified S. lendli specimen, we reinstate its original combination. Material examinded. Russia * 1 male (ISEA, 001.7306), Russia, Altai Republic, 6 km SE of Cha- gan-Uzun Vill., 26.07.2021. 50.066667°N, 88.433333°E, 1900 m asl., stony semi-desert steppe, leg. & det. A.A. Fomichev.: Altai Republic, Diagnosis. The male can be identified by the spi- ral-shaped and bare embolus with fine tip. X. mongolicus has no tegular ridge. zse.pensoft.net Description. See Fomichev, 2015. Distribution. Nentwig et al. (2024) report this species from Europe to Central Asia. So far, all Central Europe- an records have turned out to be S. /endli. We could not examine the specimens used for the Ukrainian records by Polchaninova N., Prokopenko E. (2019), or the Russian records (Ponomarev 2022). Both ML and BI phylogenetic trees (Fig. 9, Suppl. material 1, respectively) place Spiracme lendli group- ing with the type species of the genus, S. striatipes. In that group, S. triangulosus is a sister. We were unable Zoosyst. Evol. 100 (3) 2024, 1017-1028 1025 Figure 7. Spiracme lendli, male palps scanning micrographs: A. Entire palp retrolateral view. male; B. Bulb closeup, showing the embolus, the tegular bump, and the tutaculum. Scale bars: 0.2 mm. Abbreviations: tb — tegular bump; tut — tutaculum. a Figure 8. Spiracme lendli epigyne A. Fresh specimen, intact epigyne, ventral view; B. Prepared epigyne, ventral view; C. Same, in wintergreen oil; D. Vulva, dorsal view; E. Epigyne, ventral view; F. Vulva, dorsal view. Scale bar: 0.2 mm. zse.pensoft.net 1026 Gallé-Szpisjak, N. et al.: The identity and placement of Xysticus lendli Kulczynski, 1897 Spiracme triangulosus “126 7 Spiracme lendli Spiracme striatipes* 7 Coriarachne_brunneipes 51 Coriarachne_depressa 85 7 Psammitis_deichmanni 100 40 49 15 Psammitis_labradorensis Psammitis sabulosus ! = Spiracme durus Spiracme nigromaculatus 00 Bassaniana_sp Bassaniana_utahensis Xysticus_acerbus Xysticus_lineatus Xysticus_luctuosus 86 = Xysticus_elliptic Xysticus_alboniger us a Oy Xysticus_ benefactor 99 56 86 74 40 69 92 70 83 100 78 99 71 99 SE mpegs Ozyptila_simplex ysticus_montanensis Xysticus_punctatus Xysticus_bifasciatus 100 700 Xysticus_emertoni 98 Xysticus_obscurus Xysticus_ephippiatus a Xysticus_britcheri . Xysticus_pretiosus 7 Xysticus_canadensis Xysticus_chippewa Xysticus_ulmi Xysticus_elegans Xysticus_lanio Xysticus_funestus Xysticus_luctans Xysticus_ampullatus = Xysticus_auctificus 73 Xysticus_discursans Xysticus_triguttatus Xysticus_ferox Xysticus_californicus *Xysticus_cunctator Xysticus_fraternus Xysticus_gulosus Xysticus_pellax Xysticus_locuples ticus_audax ysticus_cristatus Xysticus_kochi Xysticus_erraticus Ozyptila_brevipes b/——— 0.03 Figure 9. Maximum likehood tree based on the DNA barcode data (658 nt of the mitochondrial COX1 gene), inferred with IQ-TREE. to recover a monophyletic Spiracme, as S. durus and S. nigromaculatus grouped together but not with the type species. Given the limited use of a single mitochon- drial gene in systematics and the low support values on the tree, we would not draw further conclusions regard- ing the genus limits. Discussion With the revalidation of Spiracme, perhaps Pandora’s box has been opened. As it stands, the genus contains other species groups (durus and nigromaculatus). From the original description of the genus erected for S. stri- atipes (L. Koch, 1870) by Menge (1876), it is now clear that he not only specifically listed the unique shape of the embolus of the type species (“spitze schraubenformig gewundenen eindringer” |[=embolus spirally twisted]) as diagnostic, but the etymology is also based on the spirally twisted embolus (“spira and cuspis... Von der schrauben- firmigen spitze des eindringers’) as well. We assume that the genus concept of the auctor was to include species with spirally twisted emboli on top of an anapophysate zse.pensoft.net tegulum. In the genus, as delimited currently by Breit- ling (2019), most of the species have filiform emboli and various tegular outgrowths. Considering all available in- formation, including any species with filiform embolus in the genus Spiracme, on the basis of DNA barcode data alone might have been premature. Menge erected another genus, Psammitis Menge, 1876, of which at least the type species, Psammitis sab- ulosus (Hahn, 1832), has a similar indentation on the embolus (Jantscher 2001). There is little to no consensus among the plethora of opinions regarding the limits and placement of Spiracme or Psammitis. However, placing Spiracme lendli based on morphology alone provides a challenge. It does not have a prominent tutaculum (Figs 3, 4,7) versus Spiracme striatipes, which has a large, pro- miment triangular one (Figs 3H, 4H), and the tutaculum of Psammitis species is thin and points upward. Spir- acme lendli has indentations on the bent embolus (Figs 5-7), S. striatipes has deeper indentations and a straight embolus; P. sabulosus has weaker indentations and a bent embolus; and finally, S. /endli has a tegular pocket similar to P. sabulosus, which is absent in S. striatipes (Figs 31, 41). Due to the inconsistent distribution of the Zoosyst. Evol. 100 (3) 2024, 1017-1028 morphological characters used to place species into these genera, we decided to use the grouping with the type spe- cies of Spiracme and place Xysticus lendli Kulczynski, 1897, to Spiracme Menge, 1876. Acknowledgements We are thankful to Peter Batary for his support and the free use of the equipment for identification. This work was supported by the Hungarian Research and Devel- opmental Fund (Grant ID: NKFIH FK 142926). Barna Pall-Gergely and Jené Kontschan have granted us access to their scanning electron microscope in the Plant Protec- tion Institute (NOVI) HUN-REN Centre for Agricultural Research, which is highly appreciated. We are especially grateful to Galina Azarkina and Alexander A. Fomichev in the Institute for Systematics and Ecology of Animals, Siberian Branch of the Russian Academy of Sciences, for giving us access to the Xysticus mongolicus specimen. We are indebted to Gordana Grbic, Ambros Hanggi, and Nina Polchaninova for their efforts and help to confirm their identification. Abris Toth kindly translated the Lat- in original description of Xysticus lendli, which is highly appreciated. We are grateful to Eszter Lazanyi for facili- tating several opportunities to revise and search through the whole Xysticus material in the Hungarian Natural His- tory Museum’s collection to find the type specimen. We would like to thank Suresh Benjamin and Yura Marusik for their reviews of an earlier version of the manuscript. Grant McDonald has helped with the English grammar, which is highly appreciated. References Batori Z, Erdés L, Kelemen A, Deak B, Valko O, Gallé R, Bragina TM, Kiss PJ, Kroel-Dulay G, Tolgyesi C (2018) Diversity patterns in sandy forest-steppes: A comparative study from the western and central Palaearctic. Biodiversity and Conservation 27(4): 1011-— 1030. https://doi.org/10.1007/s10531-017-1477-7 Breitling R (2019) A barcode-based phylogenetic scaffold for Xysticus and its relatives (Araneae: Thomisidae: Coriarachnini). Ecologica Montenegrina 20: 198-206. https://do1.org/10.37828/em.2019.20.16 Chyzer C, Kulczynski W (1897) Araneae Hungariae. Tomus II. Aca- demia Scientarum Hungaricae, Budapest, 147-366. [Pl. VI-X] Fekete G, Kiraly G, Molnar Z (2016) Delineation of the Pannonian vegetation region. Community Ecology 17(1): 114-124. https://doi. org/10.1556/168.2016.17.1.14 Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA prim- ers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3(5): 294-299. Fomichev AA (2015) New data on the crab spider genus Xysticus C.L. Koch, 1835 from the Altai, South Siberia (Aranei: Thomisidae). Ar- thropoda Selecta 24(1): 91-97. https://do1.org/10.15298/arthsel.24.1.05 Gallé R, Toélgyesi C, Csaszar P, Batori Z, Gallé-Szpisjak N, Kaur H, Maak I, Torma A, Batary P (2022b) Landscape structure is a major driver 1027 of plant and arthropod diversity in natural European forest fragments. Ecosphere 13(1): e3905. https://doi.org/10.1002/ecs2.3905 Gallé R, Tolgyesi C, Torma A, Batori Z, Lorinczi G, Szilassi P, Gallé-Szpisjak N, Kaur H, Makra T, Modra G, Batary P (2022a) Matrix quality and habitat type drive the diversity pattern of forest steppe fragments. Perspectives in Ecology and Conservation 20(1): 60-68. https://doi.org/10.1016/).pecon.2021.11.004 Gallé R, Koranyi D, Tolgyesi C, Lakatos T, Marcolin F, Torok E, Révész K, Szabo AR, Torma A, Gallé-Szpisjak N, Marja R, Szitar K, Deak B, Batary P (2022c) Landscape-scale connectivity and frag- ment size determine species composition of grassland fragments. Basic and Applied Ecology 65: 39-49. https://doi.org/10.1016/j. baae.2022.10.001 Gertsch WJ (1953) The spider genera Xysticus, Coriarachne, and Oxyp- tila (Thomisidae, Misumeninae) in North America. Bulletin of the AMNH, American Museum of Natural History, New York, 417-482. Grbi¢ G, Hanggi A, Gajic¢ I, Vaselek S, Ivkovic S (2019) Spiders (Ara- neae) of the Deliblato Sands (Serbia). Acta Entomologica Serbica 24: 79-93. https://doi.org/10.5281/zenodo.3249916 Jantscher E (2001) Revision der Krabbenspinnengattung Xysticus C.L. Koch, 1835 (Araneae, Thomisidae) in Zentraleuropa. Dissertation, Univ. Graz, 328 pp. [81 pls] Jantscher E (2002) The significance of male pedipalpal characters for the higher systematics of the crab spider genus Xysticus CL Koch, 1835 (Araneae: Thomisidae). In European Arachnology 2000: Pro- ceedings of the 19" European Colloquium of Arachnology. Aarhus University Press, Aarhus, 329-336. Lehtinen PT (2002) Generic revision of some thomisids related to Xysticus CL Koch, 1835 and Ozyptila Simon, 1864. In European Arachnology 2000: Proceedings of the 19" European Colloquium of Arachnology. Aarhus Univ. Press, Aarhus, 315-327. Marusik YM, Logunov DV (1990) The crab spiders of Middle Asia, USSR (Aranei, Thomisidae). 1. Descriptions and notes on distribu- tion of some species. Korean Arachnology 6: 31-62. Menge A (1876) Preussische Spinnen. VIII. Fortsetzung. Schriften der Naturforschenden Gesellschaft in Danzig (N. F.) 3: 423-454. Molnar C, Molnar Zs, Barina Z, Bauer N, Bird M, Bodonczi L, Csatho AI, Csiky J, Deak JA, Fekete G, Harmos K, Horvath A, Isépy I, Mesterhazy A, Molnar A, Nagy J, Ovari M, Purger D, Schmidt D, Sramk6 G, Szénasi V, Szmorad F (2008) Vegetation-based land- scape regions of Hungary. Acta Botanica Hungarica 50(1): 47-58. https://doi.org/10.1556/ABot.50.2008.Suppl.4 Nentwig W, Blick T, Bosmans R, Gloor D, Hanggi A, Kropf C (2024) Spiders of Europe. Version 03.2024. [https://www.araneae.nmbe.ch, accessed on 21.03.2024] https://doi.org/10.24436/1 Nguyen LT, Schmidt HA, Von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maxi- mum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268-274. https://doi.org/10.1093/molbev/msu300 Polchaninova N, Prokopenko E (2019) An updated checklist of spiders (Arachnida: Araneae) of Left-Bank Ukraine. Arachnologische Mit- teilungen. Arachnologische Mitteilungen 57(1): 60-64. https://do1. org/10.30963/aramit57 11 Ponomarev AV (2022) Spiders (Arachnida: Araneae) of the southeast of the Russian Plain: catalogue, the fauna specific features. SSC RAS Publishers, Rostov-on-Don, 640 pp. Shmatko VY (2021) New data on the fau- na and taxonomy of spiders (Aranei) in southern Russia. Ponomarev AV, zse.pensoft.net 1028 Caucasian Entomological Bulletin 17(1): 211-218. https://doi. org/10.23885/181433262021171-211218 Purgat P, GajdoS P, Purkart A, Hurajtova N, Volnar U, KrajCovitova K (2021) Walckenaeria stylifrons and Spiracme mongolica (Araneae, Linyphiidae, Thomisidae), two new species to Slovakia. Check List 17(6): 1601-1608. https://doi.org/10.15560/17.6.1601 Rambaut A (2014) FigTree vi. 4.2. released 2014-07-09. http://tree. bio. ed.ac.uk/software/figtree Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian Phylogenetic Inference under Mixed Models. Bioinformatics 19(12): 1572-1574. https://doi.org/10.1093/bioinformatics/btg 180 Schenkel E (1963) Ostasiatische Spinnen aus dem Muséum d’ Histoire naturelle de Paris. Mémoires du Muséum National d’ Histoire Na- turelle de Paris A. Zool 25: 1-481. Song DX (1987) Spiders from agricultural regions of China (Arachnida: Araneae). Agriculture Publishing House, Beijing, 376 pp. Song DX, Zhu MS (1997) Fauna Sinica: Arachnida: Araneae: Thomisi- dae, Philodromidae. Science Press, Beijing, 259 pp. Song DX, Yu SY, Yang HF (1982) A supplement note on some species of spiders from China. Acta Scientiarum Naturalium Universitatis Intramongolicae 13: 209-213. Staden R, Beal KF, Bonfield JK (2000) The Staden package, 1998. In: Misener S, Krawetz SA (Eds) Bionformatics Methods and Prtocolls. Methods in Molecular Biology 132. Humana Press, Clifton, New Jersey, 115-130. https://doi.org/10.1385/1-59259- 192-2:115 zse.pensoft.net Gallé-Szpisjak, N. et al.: The identity and placement of Xysticus lendli Kulczynski, 1897 Utotchkin AS (1995) Review of the spider genus Xysticus CL Koch, 1935 (Arachnida, Aranei, Thomisidae) in the East Kazakhstan Area. Arthropoda Selecta 4: 65-69. WSC (2024) World Spider Catalog. Version 25.0. Natural History Mu- seum Bern. [http://wsc.nmbe.ch, accessed on 21.02.2024] https:// doi.org/10.24436/2 Zhao JZ (1993) Spiders in the cotton fields in China. Wuhan Publishing House, Wuhan, China, 552 pp. Supplementary material | BI phylogram based on the barcode COI dataset (657 bp), inferred with MrBayes 3.2 Authors: Nikolett Gallé-Szpisjak, Robert Gallé, Krisztian Szabo, Tamas Sztts Data type: jpg Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow us- ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zse.100.125826.suppl1