taxonID	type	description	language	source
C7090247CE37E268D7CF15D8FCEE2D40.taxon	type_taxon	Type species: Lethia varia Menge, 1869 from Prussia. It is considered a junior synonym of Ciniflo humilis Blackwall, 1855 (= Lathys h., from England). The type specimens of L. varia seem to be lost. It is unclear as to whether L. varia and L. humilis are synonyms. L. varia may also be a senior synonym of L. nielseni (Schenkel, 1932), the coloration and habitat data of Menge (1869) may refer to both species. Lathys humilis is considered by several arachnologists, for example Lehtinen (1967), Thaler (1981) and Platnick (2009), to be the type species of the genus. However, Gertsch (1946) and Chamberlin and Gertsch (1958) clearly indicated that Lethia varia was the generotype, even though it is a junior synonym of L. humilis.	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE37E268D7CF15D8FCEE2D40.taxon	discussion	Lehtinen (1967) seems to have been the first to split Lathys into eight species groups. The third group was named humilis. Lehtinen (1967) included three species in this group: L. alticola (Denis, 1954); L. brevitibialis Denis, 1956 (still known from males only) and L. sexpustulata (Simon, 1878) and seems to have forgotten to include L. humilis in the list. It is not clear whether all three species belong to this group. One subspecies, L. humilis meridionalis (Simon, 1874), known from Spain, France, Corsica and North Africa (Platnick 2009) is not mentioned by Lehtinen (1967). Its status remains unclear, because it has not been studied by taxonomists in recent years. All three taxonomic entries for this species belong to Simon (Platnick 2009). Following the removal of L. nielseni and L. annulata Bösenberg & Strand, 1906 from synonymy with L. humilis (Thaler 1981; Ono 2003), and the recent synonymisation of L. alticola with L. sexpustulata (Ledoux et al. 2008) the L. humilis - group now includes five species and one subspecies. Only three of these (L. humilis, L. annulata and L. nielseni) have been properly studied and undoubtedly belong to the humilis group. The detailed morphology of the male palp in Lathys in general, and in its type species in particular, was unknown for a long time. There was no detailed written or illustrated description of the palpal tibia and bulbus. Thaler (1981) was the first to indicate and illustrate three tibial apophyses in Lathys humilis and L. nielseni. The structure of the bulbus in the Lathys stigmatisata - group was first studied by Marusik et al. (2006). They found that members of this group had a unique modification of the conductor, consisting of a very long upper part forming several coils over one another, a very long embolus, and a totally fixed terminal part of the conductor by the tibial apophyses and cymbium [cf. Marusik et al. (2006)]. The present study revealed that L. humilis and L. nielseni have the same conformation of the bulbus in general and the conductor in particular. As a result of this and previous studies it became possible to provide a new, revised diagnosis for the genus. Lathys can be easily distinguished from other dictynids by the presence of three tibial apophyses, the long and coiled upper arm of the conductor, which totally covers the tegulum, and a screw-like terminal part of the conductor arrested by the tibial apophyses and the cymbium (Figs 13, 15 - 17, 19). Females of Lathys cannot be diagnosed so easily. In all Lathys species studied by us (L. stigmatisata - and L. humilis - groups) the insemination ducts make a kind of loop or coil around the copulatory opening (cf. Figs 27, 30 and fig. 229 b in Wiehle 1953). In addition, the epigynal fovea or the pair of copulatory openings are larger than or equal to the spermatheca or in some cases about two times smaller. The related genus Scotolathys Simon, 1884 has no loops (or coils) around the copulatory duct, and its spermatheca is much larger than its fovea (cf. Marusik et al. 2009).	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE36E263D7CF1744FE9F2DCC.taxon	description	Figs 1 - 3, 7 - 9, 13 - 16, 20 - 22, 26 - 27	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE36E263D7CF1744FE9F2DCC.taxon	materials_examined	Material examined. DENMARK: 1 ♀ (ZMT: AA 11.130), Bornholm Isl., Ibsker, Paradisbakkerne Grydedal, in small sphagnum beds around a small pond, 30.06.1967 (P. T. Lehtinen). BULGARIA: 1 ♀ (ZMMU), Blagoevgrad Distr., Rila Mt. range, ESE slope of Karpatnik Mt., Bodovitsa River Valley, right riverside, ca 3.8 km WNW of Bachevo, Pinus – Fagus forest, 41 ° 56 ’ 12 ” N, 23 ° 24 ’ 09 ” E, 1230 m, 10.08.2005 (A. Gromov). UKRAINE: 35 ♁ ♀ (ZMUT, ZMMU) Crimea, Simferopol Distr., Chatyr-Dagh Mt., 23.04.2000 (D. S. Letova); Feodosya Distr., Karadag Nature Reserve, Kara-Agach Mt., Juniperus excelsa, sweeping, 14 - 16.05.2008 (A. A. Nadolny); Yalta Distr., Martyan Cape Reserve, 30.04. - 13.05.2007 (M. M. Kovblyuk); 1 ♁ (ZMMU), Ternopil’ Area, environs of Dzvynyach Village, old nest of Sylvia atricapilla, on bush 0.9 m above the ground, 5.05.2006 (M. Fedo- ryak). AZERBAIJAN: 2 ♁ 31 ♀ (ZMMU), SE Azerbaijan, Lenkoran Dist., environs of Aurora Village, 38 ° 40 ’ N 48 ° 52 ’ E, 23 - 28.04.2001 (Yu. M. Marusik); 1 ♁ (ZMMU) same locality and collector, 21 - 29.05.2003; 2 ♀ (ZMMU), SE Azerbaijan, Lenkoran Distr., Hyrcan Reserve, environs of Apo Village, 38 ° 38 ’ N 48 ° 47 ’ E, 28.05.2003 (Yu. M. Marusik); 1 ♁ (ZMMU), SE Azerbaijan, ca 10 km W of Astara Village, Isti-Su, 38 ° 27 ’ N 48 ° 47 ’ E, 25.04.2001 (Yu. M. Marusik). IRAN: 2 ♀ (ZMMU), Mazandaran Prov., Nashtarood-Khoshkadaran, 51.033 ° E 36.750 ° N, 9 - 10.06.2000 (Yu. M. Marusik).	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE36E263D7CF1744FE9F2DCC.taxon	description	Description. Measurements (Crimean specimens). Male. Total length 1.7; carapace 0.9 long, 0.7 wide, 0.4 high; chelicerae 0.5 long. Variation (n = 2): total length 1.6 - 1.7; carapace 0.8 - 0.9 long, 0.6 - 0.7 wide; 0.4 high. Length of leg segments: femur patella tibia metatarsus tarsus total I 0.8 0.3 0.7 0.6 0.4 2.8 II 0.6 0.3 0.5 0.5 0.3 2.2 III 0.5 0.2 0.4 0.4 0.2 1.7 IV 0.6 0.2 0.5 0.5 0.2 2.0 Female. Total length 1.9; carapace 0.8 long, 0.6 wide, 0.4 high; chelicerae 0.4 long. Variation (n = 3): total length 1.9; carapace 0.8 - 0.9 long; 0.6 wide, 0.4 high. Length of leg segments: femur patella tibia metatarsus tarsus total I 0.7 0.2 0.6 0.5 0.3 2.3 II 0.6 0.2 0.4 0.4 0.2 1.8 III 0.5 0.2 0.3 0.4 0.2 1.6 IV 0.6 0.2 0.4 0.5 0.2 1.9 Colouration. Carapace in both sexes without distinct pattern. Males slightly dark- er than females. Abdomen with distinct pattern consisting of white guanine dots, black pigment (cardiac mark, sides, and wide posterior band). Legs with distinct annulations. Copulatory organs. Male palp (Figs 7 - 9, 13 - 16) with patellar apophysis, tibia with three apophyses (retrolateral dorsal, retroventral and retrolateral (or intermediate) that fix (lock) terminal part of conductor. Conductor very long with two arms. Upper arm coiled, and terminal part spine-like and slightly twisted. Epigyne as in Figs 20, 26 - 27 with one shallow fovea, and copulatory openings placed in apical-lateral part of fovea. Receptacula droplet-shaped. Insemination ducts short and forming one turn.	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE36E263D7CF1744FE9F2DCC.taxon	diagnosis	Diagnosis. Lathys humilis can be easily distinguished from the closely related L. nielseni by the abdominal pattern consisting of dark stripes and white spots formed from guanine deposits (Figs 1 - 3). White guanine deposits are totally absent in L. nielseni (Figs 4 - 6). The two species can also be separated on the basis of the copulatory organs. The epigyne of L. humilis has shorter insemination ducts turned upwards in the place where they are attached to the spermathecae. The females also differ in the shape of the fovea (cf. Figs 20, 30). In L. humilis the epigynal fovea is subdivided by the septum, fovea deep with distinct margins (there is no septum and there are no distinct margins of the fovea in L. nielseni). The male palps of the two species are more similar than the epigynes. The two species can be separated by the thicker and broader patellar apophysis in L. humilis, the shape of the dorsal tibial apophysis, and the thicker and longer tip of the conductor in L. humilis. L. humilis can be distinguished from the Japanese L. annulata, treated for a long time as a junior synonym, by its droplet-shaped spermathecae and its shorter insemination duct forming one loop only (vs. round spermathecae and insemination ducts forming several coils, cf. figs 10 - 12 in Ono 2003).	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE36E263D7CF1744FE9F2DCC.taxon	distribution	Distribution. According to the Platnick’s (2009) catalogue, this species has a Palaearctic (= trans-Palaearctic) distribution with several records from China (Shandong, Anhui, Shanxi and Gansu), Taiwan and Korea. Judging from the figures of the Chinese specimens, all records of L. humilis refer to L. nielseni or another species (males of L. annulata are unknown). Judging from the figures (cf. Fig. 32) the record of this species from Shandong (Hu 1984) refers to L. nielseni. Other records of L. humilis from eastern China based on males may also refer to L. nielseni or L. annulata. The actual species belonging of “ L. humilis ” from Gansu (Schenkel 1936) remains unclear. Figures of the epigyne made by Schenkel are dissimilar to those of L. humilis or L. nielseni. The specimen stored in the Swedish Museum of Natural History, Stockholm lacks an epigyne and the abdominal pattern is indistinct. The record of this species from Korea (Paik 1978) refers to L. maculosa (Karsch, 1879), which belongs to the Lathys stigmatisata - group. According to our studies of the Palaearctic Lathys, L. humilis seems to be distributed from western Europe to Caucasus and Mazandaran, northern Iran (see “ material examined ”). The overlapping ranges of L. humilis and L. nielseni in SW Sweden may be caused partly by misidentifications. Both species were found, however, in samples from Öland in the Swedish Museum of Natural History. Habitats. According to Hänggi et al. (1995), L. humilis is found in Europe especially in coniferous forests (both spruce and pine), on the forests’ edges, in field shrubs and hedges, and less frequently in deciduous forests. It has been collected mostly on trees, both in canopies and on stems (Hänggi et al. 1995). According to Roberts (1995), in Great Britain it occurs on bushes and trees with small, hard leaves (heather, gorse, box, yew). Harvey et al. (2002) reported this species from bushes and trees in woodland and scrub, on oak, yew, pines, gorse, etc. It may also be fairly common on ornamental evergreen and privet hedges in parks and gardens; juveniles overwinter in leaf litter, brushwood, under bark and in other similar places (Harvey et al. 2002). In Sweden the species was reported from litter in dry pine forests, from Calluna - stands and from litter in woods with oaks and on limestone (Almquist 2006).	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE36E263D7CF1744FE9F2DCC.taxon	discussion	Note. Lehtinen (1967) synonymised three species with L. humilis: L. annulata (Japan), Altella nielseni Schenkel, 1932 (Sweden) and Altella lathysoides Denis, 1937 (Algeria). The first two names were removed from synonymy by Ono (2003) and Thaler (1981) respectively.	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE3DE266D7CF17C8FD672E69.taxon	description	Figs 4 - 6, 10 - 12, 17 - 19, 23 - 25, 28 - 32	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE3DE266D7CF17C8FD672E69.taxon	materials_examined	Material examined. FINLAND: Åland Isl., Geta, Getaberget: 6 ♁, 27 ♀, 33 juv. (ZMT / ARA 28251), lichenous rocks, 24.05.1975 (P. T. Lehtinen); Humppila, Rantakallio: 1 ♀ (ZMT), 28.06.1962 (P. T. Lehtinen); Parainen, Mustfinnö: 1 ♁ (MZT), forest, 4.06.1968 (S. Koponen); same locality: 1 ♁, 15 ♀ (ZMT), Vaccinium - type forest (P. T. Lehtinen); same locality: 1 ♁, 25 ♀ (ZMT), among moss in forest, 14.06.1966 (M. Saaristo); Turku, Kärsämäki, Pomponrahka: 5 ♀ (ZMT), among Cladonia, 29.03.1967 (M. Saaristo); Dragsŋärd, Purunpää: 1 ♁ (ZMT), 6.06 - 20.07.1971 (P. T. Lehtinen); Rymättylä, Ruotsalainen: 1 ♀ (ZMT), 10.07.1971 (P. T. Lehtinen); Nauvo, Seili: 1 ♀ (ZMT), lichenous rock, 1 - 30.10.1967 (P. Häkkilä); Somero, Ruunala: 1 ♁ (ZMT), 1974 - 1975 (H. Hippa and R. Mannila); Virrat, Patalankylä, Yli-Havankajärvi: 1 juv. (ZMT), 11.07.1972 (P. T. Lehtinen); Turku, Ruissalo: 1 ♀ (ZMT), 1968 (P. T. Lehtinen); Pori, Yyteri: 1 ♀ (ZMT), Elymus dyne, 14.10. 1961 (P. T. Lehtinen); Tuusula, Ruotsinkylä: 6 ♁, 15 ♀, 5 juv. (ZMH), Calluna - type forest, 1962 - 1965 (V. Huhta); Mäntyharju, Hietaniemi, Mäkelä: 2 ♁, 2 ♀ (ZMH), Vaccinium - type pine forest among Pleurozium, 29.05.1966 (P. Palmgren); Hanko, Tvärminne by: 9 ♁, 4 ♀, 5 juv. (ZMH), Calluna - type pine forest among Cladonia and Hylocomium schreberi, 1.06.1962 (P. Palmgren); same locality and habitat: 5 ♀, 16 juv. (ZMH), 8.08.1964 (P. Palmgren); 1 ♀ (ZMH), Dragsŋärd, Högholmen: among litter in Myrtillus - type forest, 5.06.2006 (I. Österblad); 1 ♀ (ZMH), Hanko, Lappohja, Högsand: 1 ♀, pitfall-trap, sandy shore, edge of dry pine forest, 19.07 - 9.08.2004 (N. R. Fritzén); Kuusamo, Rukajärvi, Rukatunturi: 4 ♀, 15 juv. (ZMT), 10.07.1961 (P. T. Lehtinen). RUSSIA: 2 ♁, 3 ♀ (ZMMU), Bashkortostan, Ilmenski Reserve, 29.05.1959 and 8.06.1959, (Stebaev). 2 ♁ 7 ♀ [ARAN. SIB 117, MZT] Novosibirsk Area, Borovoye, 16.6.1983 (H. Hippa).	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE3DE266D7CF17C8FD672E69.taxon	description	Description. Measurements (Finnish specimens). Male. Total length 1.8; carapace 0.89 long, 0.69 wide, 0.42 high; chelicerae 0.53 long. Variation (n = 3): total length 1.7 - 1.9; carapace 0.88 - 0.90 long, 0.68 - 0.71 wide, 0.39 - 0.45 high; chelicerae 0.49 - 0.63 long. Length of leg segments: femur patella tibia metatarsus tarsus total I 0.81 0.29 0.72 0.63 0.40 2.85 II 0.70 0.27 0.52 0.50 0.35 2.32 III 0.57 0.23 0.37 0.40 0.29 1.87 IV 0.65 0.25 0.50 0.52 0.28 2.20 Female. Total length 1.8; carapace 0.78 long, 0.62 wide, 0.39 high; chelicerae 0.33 long. Variation (n = 3): total length 1.6 - 2.3; carapace 0.75 - 0.86 long, 0.58 - 0.65 wide, 0.37 - 0.40 high; chelicerae 0.26 - 0.36 long. Length of leg segments: femur patella tibia metatarsus tarsus total I 0.61 0.26 0.47 0.40 0.28 2.01 II 0.53 0.25 0.34 0.34 0.26 1.71 III 0.42 0.24 0.28 0.29 0.21 1.44 IV 0.55 0.23 0.40 0.41 0.23 1.82 Colouration. Carapace in both sexes without distinct pattern, although dark stripes distinguish the cephalic area from the thoracic region. Abdomen with distinct pattern consisting of brownish pigment: long median stripe with transverse arms. Legs without annulations. Copulatory organs. Male palp (Figs 10 - 12, 17 - 19) with patellar apophysis, tibia with three apophyses (retrolateral dorsal, retroventral and retrolateral (or intermediate) that fix (lock) terminal part of conductor. Conductor very long with two arms. Upper arm coiled, lower part spine-like and slightly twisted. Epigyne as in Figs 23, 28 - 32, with indistinct epigynal fovea and distinct round copulatory openings. Spermathecae egg-shaped. Insemination ducts long with each duct having a vertical and a horizontal loop. First duct turned downwards and then upwards.	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE3DE266D7CF17C8FD672E69.taxon	diagnosis	Diagnosis. L. nielseni can be easily distinguished from L. humilis and L. annulata by lacking white guanine spots on the abdomen (Figs 4 - 6). The epigyne of L. nielseni resembles that of L. annulata. The two species can be separated by the shape of the receptacula (egg-shaped in L. nielseni and rounded in L. annulata) and the longer insemination ducts in the Japanese species. In addition to colour pattern, males of this species can be separated from the European L. humilis by the different shapes of the patellar and tibial apophyses (cf. Figs 13 - 16 and 17 - 19), the thinner tip of the conductor and the absence of leg annulations (cf. Figs 1, 4). The females of the two species can be separated by the shape of the fovea (distinct margins and septum in L. humilis, no distinct margins and septum in L. nielseni), the shape of the spermathecae and the length and the course of the insemination ducts (cf. Figs 20, 23, 26 - 30).	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE3DE266D7CF17C8FD672E69.taxon	distribution	Distribution. It seems that this species has a trans-Palaearctic range and is distributed from the UK to Shandong (China) and possibly to Taiwan. Within Europe, this species has been reported from Austria, Belorus, the Czech Republic, Great Britain, Germany, Slovakia, Sweden and Switzerland (Helsdingen 2007). In addition, L. nielseni is also known from the St. Petersburg Area and the southern Urals in Russia. The easternmost proven record of this species lies in the Novosibirsk Area (ca 85 ° E). The northernmost records are from Finland (where the species is often found up to 63 ° N) and Kuusamo, 66 ° 10 ’ N (Map 1). A comparison of figures of the epigyne (Figs 31 - 32) made from Finnish and Shandong specimens (identified by Hu 1984 as L. humilis) leaves no doubt that the Chinese specimens belong to L. nielseni. Other records from eastern China based on males may also refer either to L. nielseni or L. annulata (known exclusively from females). The identity of L. humilis from Gansu (Schenkel 1936) remains unclear. Figures of the epigyne made by Schenkel are dissimilar to those of both L. humilis and L. nielseni. The specimen stored in the Swedish Museum of Natural History, Stockholm, lacks the epigyne and the abdominal pattern is indistinct due to bleaching. Habitats. Thaler (1981) reported L. nielseni from warm pine wood steppe (as high as 1500 m a. s. l.), and Buchar and Růžička (2002) mentioned that it occurs within moss and lichens in pine forests (at 400 m). In England this species occurs in moist places at ground level on heathland, under stones or among damp, dead Molinia caerulea litter between the tussocks (Harvey et al. 2002). Almquist (2006) reported the species from dune heaths. In Finland it has been collected mainly from dry habitats, among litter, moss and lichens, also on sand dunes with Elymus. It seems that this species occurs only in litter, while the sibling L. humilis inhabits bushes and trees, and is found in litter occasionally.	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
C7090247CE3DE266D7CF17C8FD672E69.taxon	discussion	Discussion. The taxonomy of Lathys remains poorly and improperly studied in several respects. The limits of this genus are unclear (Lathys insulana Ono, 2003 seems to belong to Argenna or an undescribed genus; several Nearctic species appear to be distantly related to L. humilis). Scotolathys, which has long been considered a synonym of Lathys, was recently revalidated (Marusik et al. 2009). Many Lathys species remain unstudied since their original description, with many species known only from one sex. Many species appear to have been incorrectly synonymised with L. stigmatisata. Only a few species have been illustrated adequately. One of the reasons why the genus has been studied unsatisfactorily is a lack of developed species criteria. For example, in his revision, Lehtinen (1967) paid attention to the tip of the conductor, which is very similar in many species, or the structure of the epigynal fovea (also similar in many distantly related species) (P. T. Lehtinen pers. comm.). The species criteria were poorly defined because the conformation of the male palp was unknown until recently. The first detailed and correct figures of the Lathys male palpal tibia were published by Thaler (1981) and the structure of the bulbus was shown for the first time in 2006 (Marusik et al. 2006).	en	Marusik, Yuri, Koponen, Seppo, Fritzén, Niclas (2009): On two sibling Lathys species (Araneae, Dictynidae) from northern Europe. ZooKeys 16 (16): 181-195, DOI: 10.3897/zookeys.16.228
