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OEDOTHORAX TRILOBATUS (BANKS, 1896) (FIGS 7P, 8B, 9B, 17; SUPPORTING INFORMATION, FIG. S1B) Dicyphus trilobatus Banks, 1896: 64 (Dm). Hypomma trilobata Crosby, 1905: 310. Lophocarenum trilobatum Emerton, 1909: 191, pl. 3, fig. 1 (m). Oedothorax trilobatus Bishop & Crosby, 1935: 268, pl. 22, figs 7, 8 (m, Df). Oedothorax trilobatus Paquin & Dupérré, 2003: 115, figs 1190–1193 (mf). Oedothorax trilobatus Marusik et al., 2010: 287, figs 3–11 (mf). Type material: Bishop & Crosby (1935) stated that the type locality was Ithaca, New York, but no type designation data was provided by Banks (1896). The original and subsequent descriptions, nevertheless, make the identification of this species unequivocal. Examined material: USA: New York, McLean 2♂ 8.v.1919 (AMNH); Laborador Pond, 1♂ 14.v.1921, Tarris Sherman, C. Bishop Collection (AMNH); Alaska, Matanuska (61° N, 149° W), 1♂ 20.v.1945, coll. J. C. Chamberlin, det. Wilton Ivie (AMNH); Matanuska (61°32’ N, 149°12’ W), 1♂ v.–vi.1944, coll. J. C. Chamberlin, det. Wilton Ivie (AMNH); Wyoming, Teton Park (Two Ocean Lake), (43° N, 110° W), 1♂ 18.viii.1950, coll. D. C. Lowrie (AMNH); Ontario, Lake Temagami, Island 1024 (46°59’ N, 80°3’ W), 1♂ 8♀ 15.– 25.viii.1946, coll. Gertsch, Ivie, Kurata, det. Wilton Ivie (AMNH); Utah, Smith & Morehouse Canyon (40°47’ N, 111°6’ W), 2♂ 2♀ 3.vi.1934, coll. and det. Wilton Ivie (AMNH). Canada: British Columbia, Terrace, 1♂ 1.–10.vi.1931, leg. Hippisley, Sherman C. Bishop Collection (AMNH). Diagnosis: Males: This species can be identified by the shape of post-ocular groove and hump, different from all other species with similar prosomal modifications by the anterior side of the hump having one central and two lateral lobes. Females: Similar to females of other species in the Oedothorax s.s., but can be identified by its long and maneuvering copulatory ducts in epigyne (Fig. 17E, F); distinguished from Oe. fuscus, Oe. agrestis, Oe. meridionalis and Oe. tingitanus by having one sub- AME seta instead of two. Description: Male (Matanuska, 61° N, 149°W): Total length: 2.22. Prosoma: 0.98 long, 0.79 wide, postocular region with transverse hairy groove, posterior to the groove a hump anteriorly divided into three lobs; interocular region with thin, translucent setae directed laterally (Fig. 7P). Eyes: AME-AME: 0.03, AME width: 0.05, AME-ALE: 0.03, ALE width: 0.07, ALE-PLE: 0.01, PLE width: 0.07, PLE-PME: 0.05, PME width: 0.05, PME-PME: 0.14. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.62 long, 0.62 wide. Chelicerae: mastidia absent; stridulatory striae rows widely and evenly spaced (Fig. 8B). Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.21, 0.19, 1.19 and 1.06 times diameter of tibia, respectively; Tm I: 0.68. Pedipalp: TPA long, rod-like and scaly at tip, prolateral side laterally extended, median site with smaller rod-like projection with scaly tip; BT absent (Fig. 17C); PC short, base visible from dorsal view, distal setae at median position (Fig. 17A); T without papillae, PT with papillae; TS short, without papillae (Fig. 17D); DSA tip broad and smoothly serrated (Fig. 17A); EM short, distally oriented, cylindrical, with long papillae (Fig. 17A); TP without small protuberances; E long, prolaterally spiral, not broadened at basal part (Fig. 17B). Opisthosoma: dark brown, evenly coloured (Fig. 9B). Female (Ontario, Lake Temagami): Total length: 2.74. Prosoma: 1.17 long, 0.91 wide. Eyes: AME-AME: 0.03, AME width: 0.06, AME-ALE: 0.03, ALE width: 0.09, ALE-PLE: 0.01, PLE width: 0.09, PLE-PME: 0.04, PME width: 0.08, PME-PME: 0.07. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.69 long; 0.65 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.62, 1.64, 1.89 and 2.15 times diameter of tibia, respectively; Tm I: 0.75. Epigyne: Clade 13 characteristic morphology, borders between dorsal and ventral plates parallel, copulatory duct long, curved (Fig. 17E–G). Opisthosoma: brown, evenly coloured. Variation: The measurements are based on examined material. Males ( N = 10, means in parentheses): Total length 1.91–2.40 (2.17). Prosoma: 0.94–1.04 (0.99) long, 0.76– 0.81 (0.79) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.11–0.23 (0.18, N = 8), 0.15–0.40 (0.20, N = 7), 0.33–1.24 (0.82, N = 9) and 0.77–1.72 (1.36, N = 7) times diameter of tibia, respectively; Tm I: 0.63–0.69 (0.66, N = 9). Females ( N = 10, means in parentheses): Total length 2.57–3.04 (2.81). Prosoma: 1.07–1.25 (1.19) long, 0.83– 0.98 (0.92) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.52–1.94 (1.64), 1.52–1.94 (1.64), 1.70–2.06 (1.87) and 1.80–2.44 (2.02) times diameter of tibia, respectively; Tm I: 0.35–0.75 (0.65). Distribution: USA, Canada, Russia. Habitat: In vegetation in/next to ponds or swamps., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 450-451, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Banks N, 1896. A few new spiders. The Canadian Entomologist 28: 62 - 65.","Crosby CR. 1905. A catalogue of the Erigoneae of North America, with notes and descriptions of new species. Proceedings of the Academy of Natural Sciences of Philadelphia 57: 301 - 343.","Emerton JH. 1909. Supplement to the New England spiders. Transactions of the Connecticut Academy of Arts and Sciences 14: 171 - 236.","Crosby CR, Bishop SC. 1935. A new species of Hybocoptus from New York. Entomological News 46: 125 - 127.","Paquin P, Duperre N, 2003. Guide d'identification des araignees (Araneae) du Quebec, Fabreries, Supplement 11. Varennes: Association des entomologistes amateurs du Quebec.","Marusik YM, Ryabukhin AS, Kuzminykh GV. 2010. New data on spiders and harvestmen (Arachnida: Aranei & Opiliones) from western Koryakia, Kamchatka Peninsula. Arthropoda Selecta 19: 283 - 293."]}

‘OEDOTHORAX’ CAPORIACCOI ROEWER, 1942 INCERTAE SEDIS Oedothorax dubius di Caporiacco, 1935: 171, pl. 1, fig. 23 (Dm; NB: preoccupied by O. Pickard-Cambridge, 1898, sub- Lepthyphantes). Oedothorax caporiaccoi Roewer, 1942: 640 (replacement name). Type material: No type designated. Additional material: Mongolia: Takht-i-Suleiman, 2000 m, 1♂ 3.ix; 2♀, same locality, 5.ix (di Caporiacco, 1935) (not examined). Diagnosis: Males: No clear description of prosoma and palpal structures in di Caporiacco (1935). Females: Unknown. Distribution: Mongolia: Karakorum. Remarks: In the description of di Caporiacco (1935), only one prosoma lateral view drawing was available (fig. 23 in pl. 1), not sufficient for determining its relationship with other taxa. Status unclear., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 542-543, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Roewer CF. 1942. Katalog der Araneae von 1758 bis 1940. Bremen: Natura, Buchhandlung fur Naturkunde und exakte Wissenschaften Paul Budy.","di Caporiacco L. 1935. Aracnidi dell'Himalaia e del Karakoram, raccolti dalla Missione italiana al Karakoram (1929 - VII). Memorie della Societa Entomologica Italiana, Genova 13: 161 - 263."]}

EMERTONGONE MONTIFERA (EMERTON, 1882) COMB. NOV. (FIGS 7A, 66, 67C, 68C; SUPPORTING INFORMATION, FIG. S5B) Lophocarenum montiferum Emerton, 1882: 47, pl. 13, fig. 2 (Dmf). Gongylidium montiferum Simon, 1884: 500. Erigone montifera Marx, 1890: 535. Neriene montifera Simon, 1894: 633. Oedothorax montiferus Crosby, 1905: 312, 335. Diplocephalus montiferus Banks, 1916: 73. Oedothorax montiferus Bishop & Crosby, 1935: 266, pl. 22, figs 74–78 (mf). Oedothorax montiferus Kaston, 1948: 173, figs 486– 492 (mf). Oedothorax montifer Paquin & Dupérré, 2003: 115, figs 1186–1189 (mf). Type material: Not designated. Type locality: Brookline and Salem, Mass. (Bishop & Crosby 1935). The clear illustrations of the male prosomal and palpal structures and epigyne in the original description allow the unambiguous identification of this species. Examined material: USA: Pennsylvania, Lower Friederick Township (40°16’ N, 75°30’ W), 4♂ 3♀ iii.1954, Wilton Ivie coll. & det; Massachusetts, Middlwsex, Peperell, 1♀ 24.x.1970, det. Platnick; Massachusetts, Brookline (42°19’ N, 71°6’ W), 1♂ 1♀ 10.vi.1873, coll. J. H. Emerton; Ohio, Wayne County, Neil McCoy’s Woods, under rotting wood, 2♀ 25.iv.1959, J. A. Beatty. Description: Male (Pennsylvania): Total length: 1.70. Prosoma: 0.89 long, 0.67 wide, post-PME region largely elevated, with post-PME groove (Fig. 7A). Eyes: AME-AME: 0.02, AME width: 0.04, AME-ALE: 0.12, ALE width: 0.06, ALE-PLE: 0.01, PLE width: 0.05, PLE-PME: 0.02, PME width: 0.04, PME-PME: 0.31. Clypeus: hirsute. Sternum: 0.48 long, 0.45 wide. Chelicerae: stridulatory striae rows widely and evenly spaced (Fig. 67C). Legs: tibia chaetotaxy 2-2-1-1, dorsal proximal macroseta on tibia I, II, III and IV 0.35, 0.33, 0.33 and 0.49 times diameter of tibia, respectively; Tm I: 0.61. Metatarsi I–III with trichobothrium, IV without trichobothrium. Pedipalp: PC base visible from dorsal view, distal seta distally situated, distal clasp small; PT without papillae; TS absent; MSA round; DSA complex (Fig. 66A); EM arised from distal part of DSA; R and E fused; E prolaterally spiral, tip retrolateral to DSA; TP, ARP, LER, and VRP absent (Fig. 66C, D). Opisthosoma: evenly coloured, dark grey (Fig. 68C). Female (Pennsylvania): Total length: 1.70. Prosoma: 0.69 long, 0.65 wide. Eyes: AME-AME: 0.02, AME width: 0.04, AME-ALE: 0.04, ALE width: 0.06, ALE- PLE: 0.01, PLE width: 0.06, PLE-PME: 0.02, PME width: 0.05, PME-PME: 0.1. Clypeus: hirsute. Sternum: 47 long; 42 wide. Legs: tibia chaetotaxy 2-2-1-1, dorsal proximal macroseta on tibia I, II, III and IV 0.79, 0.77, 0.71 and 0.82 times diameter of tibia, respectively; Tm I: 0.56. Epigyne: dorsal plate extended anteriorly, bordering ventral plate by two lateral slits and an anterior slit, CO at lateral ends of anterior slit (Fig. 66E–G). Opisthosoma: evenly coloured, dark grey. Variation: The measurements are based on examined material. Males ( N = 5, means in parentheses): Total length 1.70 – 1.89 (1.79). Prosoma: 0.86 – 0.95 (0.89) long, 0.66 – 0.75 (0.68) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.29 – 0.35 (0.32), 0.25 – 0.33 (0.30, N = 3), 0.19–0.33 (0.27, N = 4) and 0.30 – 0.49 (0.42) times diameter of tibia, respectively; Tm I: 0.53 – 0.61 (0.56). Females ( N = 7, means in parentheses): Total length 1.70 – 2.37 (2.09). Prosoma: 0.69 – 0.91 (0.81) long, 0.62 – 0.77 (0.69) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.46 – 0.80 (0.69, N = 3), 0.52 – 0.78 (0.66, N = 4), 0.44 – 0.71 (0.58, N = 4) and 0.57 – 0.82 (0.72, N = 4) times diameter of tibia, respectively; Tm I: 0.53 – 0.62 (0.57). Distribution: USA. Habitat: Maple swamps (Emerton, 1882).

‘ OEDOTHORAX’ CUNUR TANASEVITCH, 2015 INCERTAE SEDIS (FIGS 7E, 67F, 68F, 75) Oedothorax cunur Tanasevitch, 2015: 383, figs 26–34 (Dmf). Type material: Holotype: India: Madras, Nilgiri, Coonoor, 1600 m, sifting in forest below town, ♂ 22.xi.1972, leg. C. Besuchet & I. Löbl (MHNG, examined). Paratype: 1♀, collected together with the holotype (MHNG, examined). Diagnosis: Males: This species can be identified by the lack of prosomal modification and the distinct shapes of palpal tibia prolateral apophysis and embolic division. Females: Epigyne simple as in Oedothorax s.s., Callitrichia and Mitrager. Description: Male (holotype): Total length: 2.10. Prosoma: 1.02 long, 0.87 wide, unmodified (Fig. 7E). Eyes: AME- AME: 0.03, AME width: 0.04, AME-ALE: 0.04, ALE width: 0.09, ALE-PLE: 0, PLE width: 0.09, PLE-PME: 0.06, PME width: 0.08, PME-PME: 0.06. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.62 long, 0.59 wide. Chelicerae: stridulatory striae rows compressed proximally (Fig. 67F). Legs: tibia chaetotaxy 2-2-1- 1, dorsal proximal macroseta on tibia I 1.69 times diameter of tibia; Tm I: 0.55. All metatarsi with trichobothrium. Pedipalp: patella prolateral proximal vertical macrosetae abesent; tibia with one prolateral, two retrolateral trichobothria; TPS absent; TPA apical side scaly, with several setae with serrated tips on the upper margin; TRA absent (Fig. 75A, D, E); PC median-sized, base not visible from dorsal view, distal setae close to distal clasp, distal clasp with inconspicuous striae, directed retrolaterally (Fig. 75A); T without papillae; PT with long papillae; TS short, with few papillae; MSA present; DSA short, irregular ar distal margin (Fig. 75F); EM flat, anterior margin without papillae, length equals ARP; ARP flat, slightly sclerotized; LER small, without striae, not extended dorsal to E; VRP long, anteriorly directed; TP tip narrow; E retrolaterally spiral, anterior margin at base slightly wavy (Fig. 75C). Opisthosoma: dorsal pattern see Fig. 68F; PMS with mAP, two AC; PLS with triad, 3+ AC (Fig. 75I, J). Female (paratype): Total length: 2.22. Prosoma: 1.02 long, 0.87 wide. Eyes: AME-AME: 0.02, AME width: 0.05, AME-ALE: 0.03, ALE width: 0.09, ALE-PLE: 0.01, PLE width: 0.09, PLE-PME: 0.06, PME width: 0.08, PME- PME: 0.05. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.62 long; 0.58 wide. Legs: tibia chaetotaxy 2-2-1-1, dorsal proximal macroseta on tibia II 2.09 times diameter of tibia; Tm I: 0.62. Epigyne: Clade 13 characteristic morphology, entrance of copulatory ducts into spermathecae ectal to exits of fertilization ducts (Fig. 75G, H). Opisthosoma: PMS with mAP, two AC, one CY; PLS with triad, two CY, 3+ AC (Fig. 75K). Distribution: India, only known from the type locality. Habitat: Lower layer of forests. Remarks: According to the results of our phylogenetic analysis, and the lack of the distinctive characteristics of Oedothorax s.s., this species is more closely related to the generus Atypena, which is also distributed in the Oriental realm. We provisionally leave the taxonomic status of this species unchanged until further data become available., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 544, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2015. Notes on the spider genus Oedothorax Bertkau, 1883 with description of eleven new species from India (Linyphiidae: Erigoninae). Revue suisse de Zoologie 122: 381 - 398."]}

HOLMELGONIA JOCQUÉ & SCHARFF, 2007 Type species: Elgonia nemoralis Holm, 1962. Diagnosis, description and taxonomic remarks: The resemblance of Holmelgonia to Callitrichia and other related genera (i.e. Oedothorax, Toschia and Ophrynia) has not been mentioned in previous publications. Regarding the shape of embolic division, the enlarged terminal part of paracymbium, the vastly erected palpal tibia prolateral apophysis and the general epigyne morphology, Holmelgonia is similar to numerous Callitrichia species. Nzigidahera & Jocqué (2014) described Holmelgonia as characterized by the absence of cheliceral stridulation ridges, the presence of a double ventral row of setae on the femora, the long tibial spines (two to three times as long as the diameter of the segment), tibial chaetotaxy 2-2-1-1, TmI between 0.32 and 0.7, the absence of prosomal modification and the apophysis on the dorsal side of the palpal tibia. However, all these features are not unique to the members of this genus. One potential diagnostic feature, the absence of cheliceral stridulatory ridges, is actually present in our studied species (Ho. basalis, Fig. 22B), and the chelicera of Ho. disconveniens Nzigidahera & Jocqué, 2014 (SEM photo, figs 2–3 in Nzigidahera & Jocqué 2014) shows a scaly lateral surface without setae, which could also be interpreted as stridulatory striae (fig. 3 shows the detail of the anterior surface, instead of the lateral surface). Despite the lack of defining features for this genus, and the sister-relationship of Ho. basalis and Callitrichia shown in the present study, this genus differs from Callitrichia in the lack of truncated protegulum and the presence of a central embolar apophysis (Fig. 36B, D, arrow). The latter structure is present in all species with known males, except Ho. discoveniens and Ho. afromontana Nzigidahera & Jocqué, 2014. Due to the lack of synapomorphies defining this genus, the monophyly of Holmelgonia remains untested (Nzigidahera & Jocqué 2014). In our phylogenetic hypothesis (Fig. 3), Ho. basalis resulted sister to Callitrichia (Clade 40), a relationship supported by the presence of a rim at the distal area of the protegulum (Ch 38, synapomorphic) and posterior lateral spinneret with more than three aciniform gland spigots (Ch 116, homoplastic). Since only one Holmelgonia species was represented in our analysis, it is yet premature to decide whether the genus is a monophyletic assemblage sister to Callitrichia, or a paraphyletic set of taxa that should in turn require taxonomic actions. Further phylogenetic analyses with a thorough species representation of Holmelgonia, including the type species Ho. nemoralis, as well as Callitrichia and other related taxa, is needed. Distribution: Ivory Coast, Cameroon, Congo, Burundi, Tanzania, Uganda, Kenya, Malawi and Mozambique. Natural history: Species of this genus were found in lower vegetation, forest litter and by pitfall traps (Jocqué 1981; Jocqué & Scharff 1986; Scharff 1990a). Most species live at high altitude and have small endemic ranges, while some have a wide distribution at midaltitude (Nzigidahera & Jocqué 2014)., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 482, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Jocque R, Scharff N, 2007. Holmelgonia, a new name for the genus Elgonia Holm, 1989 (Araneae, Linyphiidae). Journal of Afrotropical Zoology 3: 161.","Holm A. 1962. The spider fauna of the East African mountains. Part I: Fam. Erigonidae. Zoologiska Bidrag fran Uppsala 35: 19 - 204.","Nzigidahera B, Jocque R. 2014. On the Afrotropical genus Holmelgonia (Araneae, Linyphiidae), with the description of three new species from the Albertine Rift. European Journal of Taxonomy 77: 1 - 18.","Jocque R. 1981. Erigonid spiders from Malawi (Araneida, Linyphiidae). Revue Zoologique Africaine 95: 470 - 492.","Jocque R, Scharff N, 1986. Spiders (Araneae) of the family Linyphiidae from the Tanzanian mountain areas Usambara, Uluguru and Rungwe. Annalen Zoologische Wetenschappen 248: 1 - 61.","Scharff N, 1990 a. Spider of the family Linyphiidae from the Uzungwa mountains, Tanzania (Araneae). Entomologica Scandinavica, Supplement 36: 1 - 95."]}

‘OEDOTHORAX’ BIANTU ZHAO & LI, 2014 INCERTAE SEDIS Oedothorax biantu Zhao & Li, 2014: 36, figs 67A–F, 68A, B (Df). Type material: Holotype: China, Yunnan: Menglun Town: Xishuangbanna Tropical Botanical Garden (21°54’38’’ N, 101°16’518’’ E), c. 627 m, bamboo plantation, fogging, ♀ 22.xi.2009 (not examined). Diagnosis: Females: This species differs from all Oedothorax, Mitrager, Callitrichia and their related genera by the tibial chaetotaxy (1-1-1-1), the knob on each side of ventral plate of the epigyne and the long slits on the ventral plate (figs 67A and 68A in Zhao & Li, 2014). Males: Unknown. Distribution: Known only from type locality. Habitat: See collecting data of holotype. Remarks: This species has tibial chaetotaxy 1-1- 1-1, unlike the uniform pattern 2-2-1-1 among all other examined species in this study. The epigynal structure of this species is also prominently different from the simple composition in other Oedothorax species by having a knob on each side of the ventral plate and the long slits on the ventral plate of the epigyne. Therefore, this species does not belong to Oedothorax, and its relationship to other erigonines remains unknown.

MITRAGER VAN HELSDINGEN, 1985 Type species: Mitrager noordami van Helsdingen, 1985. Diagnosis: Mitrager is similar to other taxa on Clade 13 in palpal and somatic features, but is characterized and can be distinguished by the following unique combination of features: 1. Paracymbium: medium- to large-sized (small in Oedothorax); base not visible from dorsal view of male pedipalp (visible in Oedothorax); distal setae-bearing area not prominently laterally elevated (prominently laterally elevated in some Callitrichia); distal clasp with striae in many species (no striae in other taxa in Clade 13). 2. Copulatory bulb: embolus base protuberance absent (present in Oedothorax); embolus retrolaterally spiral (prolaterally spiral in Oedothorax), covered dorsally by lateral extension of radix (LER) in most species, except M. elongata and M. tholusa, in which it is retrolateral to the embolus (LER absent in most other species in Clade 13 except Ca. convector); tegular papillae absent (present in many Oedothorax and Ca. macrophthalma). 3. Palpal tibia: prolateral apophysis varies from small or absent to prominent in some species, but never elevated vertically except M. noordami (vertically elevated in many Callitrichia); scaly prolateral spike present except M. noordami, retrolaterally directed in most species (absent in other species in Clade 13); retrolateral apophysis short, retrolaterally curved in many species (absent in Oedothorax and Callitrichia except Ca. convector). 4. The general structure of the epigyne in Mitrager is extremely similar across species, and also similar to that of Callitrichia, Holmelgonia, Oedothorax and other related taxa. It can be distinguished from Callitrichia and Holmelgonia by the position of the entrance of copulatory ducts into the spermathecae, more mesal to the exits of the fertilization ducts. For a general description of epigynal conformation, see shared features defining Clade 13 above. Monophyly: This genus is supported by the following unambiguous character transformations: the retrolateral bending of palpal tibia prolateral spike (Ch 53, synapomorphic; lost in M. noordami) and the wavy prolateral margin of the embolus (Ch 14, homoplastic). Description: The genus includes medium-sized erigonines mostly with a variegated opisthosoma. Male prosoma varies in degree of modification, ranging from unnoticeable (M. unicolor and M. hirsuta) to prominent post-PME humps, post-PME grooves, PME lobe, inter- AME-PME lobe, clypeal hump, cheliceral apophysis, pre-PME groove and modified setae. Chelicerae without mastidia. Clypeus with one sub-AME seta. For palpal and epigynal feature see diagnosis. Species included: This genus comprises 25 species (re-described below), among which 24 species are transferred from Oedothorax. Distribution: Nepal, India, Indonesia (Java). Natural history: Most species have been collected from broad-leaved or coniferous forest litter., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 484, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Van Helsdingen PJ. 1985. Mitrager noordami, an erigonine novelty from Java. Bulletin of the British Arachnological Society 6: 353 - 358."]}

‘OEDOTHORAX’ BIANTU ZHAO & LI, 2014 INCERTAE SEDIS Oedothorax biantu Zhao & Li, 2014: 36, figs 67A–F, 68A, B (Df). Type material: Holotype: China, Yunnan: Menglun Town: Xishuangbanna Tropical Botanical Garden (21°54’38’’ N, 101°16’518’’ E), c. 627 m, bamboo plantation, fogging, ♀ 22.xi.2009 (not examined). Diagnosis: Females: This species differs from all Oedothorax, Mitrager, Callitrichia and their related genera by the tibial chaetotaxy (1-1-1-1), the knob on each side of ventral plate of the epigyne and the long slits on the ventral plate (figs 67A and 68A in Zhao & Li, 2014). Males: Unknown. Distribution: Known only from type locality. Habitat: See collecting data of holotype. Remarks: This species has tibial chaetotaxy 1-1- 1-1, unlike the uniform pattern 2-2-1-1 among all other examined species in this study. The epigynal structure of this species is also prominently different from the simple composition in other Oedothorax species by having a knob on each side of the ventral plate and the long slits on the ventral plate of the epigyne. Therefore, this species does not belong to Oedothorax, and its relationship to other erigonines remains unknown., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 542, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Zhao Q, Li S. 2014. A survey of linyphiid spiders from Xishuangbanna, Yunnan Province, China (Araneae, Linyphiidae). ZooKeys 460: 1 - 181."]}

EMERTONGONE GEN. NOV. Type species: Lophocarenum montiferum Emerton, 1882. Derivatio nominis: The genus name is a combination of Emerton, in honor of the American arachnologist James Henry Emerton, the original describer of the type species, and the ending of the generic name ‘ Erigone ’. Genus gender feminine. Diagnosis: Males: Small-sized, dark-brown-coloured erigonine. This genus is characterized by its male prosomal modification comprising a post-PME groove, post- PME lobe, the anteriorly protruded hirsute clypeus and the shorter distance from PLE to clypeal lower margin compared to PME. Despite its superficial partial similarity of prosomal modification with Oe. gibbosus, this genus has no separate radix and embolus connected by a membranous region, and the embolic membrane is erected from distal part of distal suprategular apophysis. These characters distinguish it from all taxa in Clade 13. Females: Can be distinguished from all other examined taxa by the shorter distance from ALE to clypeus margin compared to distance from AME to clypeus margin, the hirsute clypeus, and the transverse slit on the epigyne between the two copulatory openings (Fig. 65E, G). Species included: Emertongone montifera (Emerton, 1882) comb. nov. Phylogenetic justification: Although the prosomal modification of Em. montifera superficially resembles that of Oe. gibbosus, the palpal configuration of this species is different from that of Oedothorax as delimited in the present study. In the phylogeny (Fig. 2), this species is sister to Clade 12; i.e. more closely related to the Walkenaeria and Gonatium representatives than to all Oedothorax, Callitrichia, Mitrager and other representatives in Clade 13. The similarity in prosomal morphology is, therefore, a result of homoplasy. After a comprehensive literature research of erigonine male palpal structures, no morphologically resembling species was found that can suggest at least a preliminary (i.e. phenetic) close relatedness. Base on these findings and its resulting phylogenetic placement, this species neither belongs to Oedothorax, nor can it be transferred to any other established taxon.We, therefore, propose the erection of Emertongone gen. nov. for this unique species., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 525, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Emerton JH. 1882. New England spiders of the family Theridiidae. Transactions of the Connecticut Academy of Arts and Sciences 6: 1 - 86."]}

TMETICUS MAXIMUS EMERTON, 1882 Tmeticus maximus Emerton, 1882: 55, pl. 16, fig. 5 (Dm). Gongylidium maximum Simon, 1884: 500. Erigone maxima Marx, 1890: 534. Oedothorax maximu Crosby, 1905: 311. Unexamined material: USA: New Hampshire, Mt. Washington, half way up, in moss, 1♂ (Emerton 1882). No type was designated. Diagnosis: Males: This species is characterized by the large body size (nearly 3 mm long), the presence of cheliceral mastidia and the palpal morphology, including the short, rough-ended tibial apophysis and the bifurcated paracymbium tip. According to figure 5 in plate 16 in Emerton (1882), this species seems not to have the pointed, prolaterally spiral anterior radical process similar to Oedothorax, and the curvature of the embolus does not resemble that of Oedothorax, either. Females: Unknown. Distribution: USA: Mt. Washington. Remarks: One drawing of a male pedipalp in retrolateral view was provided in Emerton (1882), which shows a bifurcated tip of the paracymbium and the curvature of the embolus, both features significantly different from Oedothorax s.s.. We thus propose its transfer back to its original combination until further investigation is conducted. Due to the unavailability of specimens and the limited information from the original description, the relationship of this species to other erigonines remains unclear., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 537, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Emerton JH. 1882. New England spiders of the family Theridiidae. Transactions of the Connecticut Academy of Arts and Sciences 6: 1 - 86.","Simon E. 1884. Les arachnides de France. Tome cinquieme, deuxieme et troisieme partie. Roret Paris, 180 - 885.","Marx G. 1890. Catalogue of the described Araneae of temperate North America. Proceedings of the United States National Museum 12: 497 - 594.","Crosby CR. 1905. A catalogue of the Erigoneae of North America, with notes and descriptions of new species. Proceedings of the Academy of Natural Sciences of Philadelphia 57: 301 - 343."]}

MITRAGER GLOBICEPS (THALER, 1987) COMB. NOV. (FIGS 38I, 39I, 40I, 51; SUPPORTING INFORMATION, FIG. S4G) Oedothorax globiceps Thaler, 1987: 36, figs 16–21 (Dm). Type material: Holotype: India: Pahalgam, coniferous forest, 2400m, ♂ 14.v.1976, leg. J. Martens (SMF 33832, examined). Diagnosis: Males: This species shares the prosomal postocular hump and transverse groove between the hump and ocular region with M. dismodicoides, M. lineata, M. lucida, M. sexoculata, M. sexoculorum (Tanasevitch, 1998) comb. nov. and M. tholusa, but can be distinguished by the position of the PME at the lateral side of the groove (PME above the groove in M. dismodicoides, M. lineata, M. sexoculorum and M. tholusa; PME inside the pre-PME groove in M. lucida, M. sexoculata and M. sexoculorum). Description: Male (holotype, SMF 33832): Total length: 1.98. Prosoma: 0.91 long, 0.70 wide, transverse groove between PME, hump posteriorly (Fig. 38I). Eyes: AME- AME: 0.03, AME width: 0.04, AME-ALE: 0.04, ALE width: 0.07, ALE-PLE: 0, PLE width: 0.08, PLE-PME: 0.02, PME width: 0.06, PME-PME: 0.18. Clypeus: not hirsute. Sternum: 0.49 long, 0.52 wide. Chelicerae: mastidia absent; stridulatory striae rows widely and evenly spaced (Fig. 39I). Legs: Tm I: 0.85. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPS scaly, distal-retrolaterally pointed; TPA median-sized; TRA apically oriented; PC mediansized, base not visible from dorsal view, distal setae close to distal clasp, distal clasp without striae, clasp directed apically (Fig. 51A); T without papillae; PT apical part with small papillae; TS short, without papillae (Fig. 51B); MSA present; DSA tip round; EM flat, anterior margin without papillae, approximately equals ARP in length; ARP pointed; LER without striae, extended dorsal to E; VRP absent; TP tip narrow; E retrolaterally spiral, anterior margin at base smoothly curved (Fig. 51E, F). Opisthosoma: dorsal pattern see Fig. 40I; PMS with mAP, two AC; PLS with triad, 3+ AC (Fig. 51G, H)., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 503-504, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Thaler K. 1987. Uber einige Linyphiidae aus Kashmir (Arachnida: Araneae). Courier Forschungsinstitut Senckenberg 93: 33 - 42."]}

CALLITRICHIA LEGRANDI (JOCQUÉ, 1985) COMB. NOV. (FIGS 19M, 22M, 24M, 29; SUPPORTING INFORMATION, FIG. S2J) Oedothorax legrandi Jocqué, 1985: 206, figs 21–27 (Dmf). Type material: Holotype: Comoros: Mohéli, Miringoni, Chalet St. Antoine, 700m, rain forest, sieving litter, ♂ 11.xi.1983 (RMCA 161.072, examined). PARATYPES: 5♀, same collecting site as holotype, sweepnet, 7.–8. xi.1984 (RMCA, examined); 1♂ 2♀, same collecting site as holotype, sweeping by night, 8.xi.1983 (RMCA, examined); 2♂, same location, pitfall, 6.–11.xi.1983 (RMCA, examined). Diagnosis: Males: This species can be identified by the lack of prosomal modification and the particular shape of the tibial apophyses, with a vertically elevated TPS and a small TPA with one enlarged, slightly sclerotized setal base at tip. Females: Can be distinguished from other Callitrichia species by the dorsal pattern of the opisthosoma, the shape and position of the spermatheca (e.g. less oval than Ca. holmi; less anteriorly situated than Ca. longiducta; separation between spermathecae wider than in Ca. juguma) and the position of copulatory opening (e.g. more anterior than Ca. muscicola). Description: Male (paratype, RMCA 161.055): Total length: 1.67. Prosoma: 0.72 long, 0.58 wide, unmodified (Fig. 19M). Eyes: AME-AME: 0.03, AME width: 0.05, AME- ALE: 0.02, ALE width: 0.06, ALE-PLE: 0.01, PLE width: 0.05, PLE-PME: 0.04, PME width: 0.06, PME- PME: 0.04. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.42 long, 0.44 wide. Chelicerae: mastidia absent; stridulatory striae rows widely and evenly spaced (Fig. 22M). Legs: Tm I: 0.62. Pedipalp: patella prolateral proximal vertical macrosetae absent; TPS vertically oriented, tip scaly; TPA small, with one enlarged, slightly sclerotized setal base at tip (Fig. 29C, D); PC large, base not visible from dorsal view, distal setae close to distal clasp, distal-setae-bearing area slightly wide, distal clasp extended apically (Fig. 29A); T without papillae, PT without papillae, distal rim thin and smooth at margin; TS absent (Fig. 29E); MSA present; DSA tip straight; EM absent (Fig. 29A); ARP pointed; LER absent; VRP absent; TP tip slender; E retrolaterally spiral (Fig. 29E). Opisthosoma: dorsal pattern see Fig. 24M; PMS with mAP, AC absent; PLS with triad, AC absent (Fig. 29I). Female (paratype, RMCA 160.870): Total length: 1.69. Prosoma: 0.66 long, 0.52 wide. Eyes: AME-AME: 0.02, AME width: 0.04, AME-ALE: 0.02, ALE width: 0.04, ALE-PLE: 0.01, PLE width: 0.05, PLE-PME: 0.05, PME width: 0.04, PME-PME: 0.05. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.40 long; 0.41 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 2.40, 2.43, 2.89 and 2.81 times diameter of tibia, respectively; Tm I: 0.69. Epigyne: Clade 13 characteristic morphology, borders between ventral and dorsal plates converging anteriorly and posteriorly, entrances of copulatory ducts into spermathecae directly dorsal to the exits of fertilization ducts (Fig. 29G, H). Opisthosoma: dorsal pattern similar to male; PMS with mAP, two AC, CY; PLS with triad, two CY, 3+ AC (Fig. 29J). Variation: The measurements are based on examined material. Males ( N = 3, means in parentheses): Total length 1.51 – 1.67 (1.60). Prosoma: 0.63 – 0.76 (0.70) long, 0.52 – 0.62 (0.57) wide. Females ( N = 6, means in parentheses): Total length 1.50 – 1.99 (1.73). Prosoma: 0.65 – 0.75 (0.69) long, 0.50 – 0.55 (0.53) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 2.04 – 2.46 (2.30), 2.00 – 2.64 (2.38, N = 5), 2.32 – 2.89 (2.66) and 2.52 – 3.30 (2.95) times diameter of tibia, respectively; Tm I: 0.58 – 0.69 (0.63). Distribution: Comoros, only known from the type locality. Habitat: Rain forest litter. Remarks: Following the results of our phylogenetic analysis, this species is transferred to Callitrichia, a placement also consistent with the morphological traits of the species. CALLITRICHIA LONGIDUCTA (BOSMANS, 1988) COMB., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 468-469, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Jocque R. 1985. Linyphiidae (Araneae) from the Comoro Islands. Revue de Zoologie Africaine 99: 197 - 230.","Bosmans R. 1988. Scientific report of the Belgian Cameroon expeditions 1981 and 1983. No. 18. Further Erigoninae and Mynogleninae (Araneae: Linyphiidae) from Cameroonian highlands. Revue Zoologique Africaine 102: 5 - 32."]}

‘OEDOTHORAX’ KODAIKANAL TANASEVITCH, 2015 INCERTAE SEDIS (FIGS 7D, 67E, 68E, 76) Oedothorax kodaikanal Tanasevitch, 2015: 386, figs 44–50 (Dm). Type material: Holotype: India: Madras, Palni Hills, 10 km north-west of Kodaikanal, 2150 m, edge of Rhododendron forest with fern, sifting litter near river, ♂ 15.xi.1972, leg. C. Besuchet & I. Löbl (MHNG, examined). Paratypes: India: Madras, Palni Hills, 23 km west of Kodaikanal, Lake Berijam, 2150 m, Rhododendron forest, sifting litter, 1♂ (MHNG, examined) 1♂ (ZMMU) 14.xi.1972, leg. C. Besuchet & I. Löbl. Diagnosis: Males: This species can be diagnosed by the lack of external prosomal modification and the palpal features as described below. Description: Male (holotype): Total length: 2.27. Prosoma: 1.10 long, 0.85 wide, unmodified (Fig. 7D). Eyes: AME- AME: 0.02, AME width: 0.05, AME-ALE: 0.03, ALE width: 0.10, ALE-PLE: 0, PLE width: 0.10, PLE-PME: 0.05, PME width: 0.08, PME-PME: 0.07. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.66 long, 0.60 wide. Chelicerae: stridulatory striae rows compressed at proximal end (Fig. 67E). Legs: tibia chaetotaxy 2-2-1-1; Tm I: 0.55. All metatarsi with trichobothrium. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPS short, prolaterally situated at base of TPA; TPA long, dorsally elevated; TRA apically oriented (Fig. 76A, B, C); PC mediansized, base not visible from dorsal view, distal setae close to distal clasp, distal setae bearing area wide, distal clasp without striae, clasp extended slightly apically (Fig. 76A); T without papillae; PT without papillae; TS short, without papillae; MSA present; DSA tip straight (Fig. 76F); EM flat, anterior margin with small papillae, not exceeding ARP; ARP tip blunt, with a round process ventrally; LER without striae, retrolaterally oriented; VRP with sclerotized apophysis distally; TP tip round; E retrolaterally spiral, anterior margin at base smoothly curved (Fig. 76D). Opisthosoma dorsal pattern see Fig. 68E. PMS with one nubbin (vestigial mAP), one AC; PLS with triad, one AC (Fig. 76G, H). Male (paratype): Total length: 2.14. Prosoma: 1.02 long, 0.81 wide. Tm I: 0.54. Female: Unknown. Distribution: India. Habitat: Forest litter. Remarks: According to the results of our phylogenetic analysis, and the lack of characteristics of Oedothorax s.s., this species does not belong to Oedothorax, but is more closely related to Atypena and some ‘ Oedothorax ’ incertae sedis species which, as in this species, are distributed in the Oriental realm. Although this species was scored in the present study as possessing TPS (the defining feature of Mitrager), its minute size and the position at the base of TPA in this species differ significantly from the retrolaterally bent form in Mitrager. In addition, the LER in this species does not bend over the embolus like in Mitrager. Since it shares no clear synapomorphic features with other taxa, its taxonomic affinity should be scrutinized in future studies, including other oriental erigonine species like ‘Oe.’ myanmar Tanasevitch, 2017 and ‘Oe.’ khasi Tanasevitch, 2017., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 546-548, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2015. Notes on the spider genus Oedothorax Bertkau, 1883 with description of eleven new species from India (Linyphiidae: Erigoninae). Revue suisse de Zoologie 122: 381 - 398."]}

CALLITRICHIA HIRSUTA NOM. NOV. Atypena pilosa Jocqué & Scharff, 1986: 23, figs 62–66 (Dm). Callitrichia pilosa Scharff, 1990b: 124 (Tm from Atypena). Junior secondary homonym of Callitrichia pilosa (Wunderlich, 1978) Z o o b a n k r e g is t r a t i o n: urn: lsid: zoobank. org:act: E74AA2B3-AF77-4220-9439-12C04F97E2EA Derivatio nominis: The name refers to the setae on the interocular region, from Latin hirsutus, hairy. Remarks: Due to the transfer of the older name Oedothorax pilosa to Callitrichia, the combination is not available and thus, a replacement name is here provided. CALLITRICHIA HOLMI (WUNDERLICH, 1978) COMB., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 464, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Jocque R, Scharff N, 1986. Spiders (Araneae) of the family Linyphiidae from the Tanzanian mountain areas Usambara, Uluguru and Rungwe. Annalen Zoologische Wetenschappen 248: 1 - 61.","Scharff N, 1990 b. A catalogue of African Linyphiidae (Araneae). Steenstrupia 16: 117 - 152.","Wunderlich J. 1978. Zur Kentniss der Gattung Oedothorax Bertkau 1883, Callitrichia Fage 1936 und Toschia Caporiacco 1949. Senkenbergiana Biologica 58: 257 - 260."]}

CALLITRICHIA FAGE, 1936 Type species: Callitrichia hamifera Fage, 1936, by original designation. Ophrynia Jocqué, 1981 synon. nov. (type species Ophrynia superciliosa Jocqué, 1981, by original designation) Toschia Caporiacco, 1949 synon. nov. (type species Toschia picta di Caporiacco, 1949, by original designation). Taxonomic remarks: The genus was originally established by Fage in Fage & Simon (1936) for Callitrichia hamifer Fage, 1936 and two other species. Beside other somatic characheristics mentioned in their description and common to many erigonine genera, the distinctive feature of these three species is the two longitudinal dense clusters of setae with particular morphology at the interocular region of the male. Holm (1962) described a number of new species of Callitrichia and added descriptions of the palps, including the enlarged terminal part of paracymbium and the short, stout, curved embolus, which is connected to a ‘scaphium’ (i.e. radix) with two distal apophyses (i.e. anterior radical process and ventral radical process). Based on similarities in, for example, metatarsal trichobothria, epigyne structure, eye arrangement and embolic division, he suggested that Callitrichia has a close relationship with Oedothorax, although the embolus is slender and the ventral radical process is absent in the latter genus. He also elaborated on the close relationship between Toschia and Oedothorax, as previously pointed out by di Caporiacco (1949), and further proposed some differences between them, such as the distally wide paracymbium. However, the similarity between Callitrichia and Toschia in their embolic division was not mentioned. Scharff (1990a) briefly addressed the issues about the subsequently proposed synonymies, one among Oedothorax, Callitrichia and Toschia by Wunderlich (1978) and the other between Callitrichia and Atypena (Jocqué, 1983), and argued in favour of treating all taxa as separate genera until a complete revision of all genera has been done. He also suggested a close relationship of Ophrynia with the above-mentioned genera based on resemblance in the shape of the embolic division. However, he stated that the paracymbium of Ophrynia differed significantly from that of those genera, although this may only be true for species like Ca. infecta (Jocqué & Scharff, 1986) comb. nov. (fig. 128 in Jocqué & Scharff, 1986) and Ca. uncata (Jocqué & Scharff, 1986) comb. nov. (Fig. 18A, arrow), both with the terminal area of the paracymbium ventrodistally extended (pers. obs.). In our view, the distally wide paracymbium of other Ophrynia species, including the type species Ca. superciliosa (Jocqué, 1981) comb. nov., resembles considerably that of Callitrichia and Toschia, which in turn is different from the narrow terminal part of the paracymbium of Oedothorax species from the Palearctic realm. Although the type species Ca. hamifera was not included in our analysis, we consider Ca. sellafrontis a suitable representative due to its many shared features with Ca. hamifera, including two cluster of special dense setae at interocular region, posterior median eye lobe (Fig. 19H; fig. 53c, d in Holm, 1962), male palpal tibial apophysis shape (Fig. 20A–C; fig. 53a, b in Holm, 1962), paracymbium shape (Fig. 20A; fig. 54c in Holm, 1962), protegulum morphology (Fig. 20D; fig. 21a in Fage & Simon, 1936), the shape of embolic division (Fig. 20B; fig. 54a in Holm, 1962), and the shape of epigyne (Fig. 20F; fig. 53e in Holm, 1962). We also consider Ca. uncata and Ca. juguma (Scharff, 1990) comb. nov. valid representatives for Ophrynia, since they share the following common male features with the type species of Ophrynia (Ca. superciliosa): inter-AME-PME groove; palpal tibial prolateral apophysis broad, flat, with a retrolateral branch; embolic division with a moderate, poorly sclerotized ventral radical process and a round tip of radical tail piece. Furthermore, Ca. juguma resembles Ca. superciliosa especially by the dentate tips of the bifid palpal tibial apophysis (Fig. 21A, C; figs 39, 40 in Jocqué, 1981), enlarged terminal part of paracymbium without extended ventrodistal angle (Fig. 21A; fig. 39 in Jocqué, 1981, in contrast to Ca. uncata, Fig. 18A), bifid anterior radical process (Fig. 21B; fig. 41 in Jocqué, 1981), papillae on ventral radical process, and perpendicular lateral setae on metatarsi and tarsi of legs I and II of the male, which is the only proposed synapomorphy of Ophrynia (Scharff, 1990a). The results of our phylogenetic analysis placed the two original Ophrynia species as sister to Ca. sellafrontis. Features supporting this relationship are related to the reduction of spinnerets gland spigots, including the absence of the aciniform and the minor ampullate gland spigots, as well as the absence of the aggregated gland spigot (also in Ca. convector). This high degree of reduction of spigots has not been seen in other studied species so far. This close relationship between the original Ophrynia species and Ca. sellafrontis renders the remaining Callitrichia species a paraphyletic assemblage. The truncated protegulum is a newly proposed synapomorphy of the redefined Callitricha. The original generic description of Toschia by di Caporiacco (1949) included two species, and proposed several features that are not unique for the current members described under Toschia to date. Holm’s (1962) account for this genus also includes diagnostic characteristics not exclusive to Toschia (e.g. lack of carapace modification), and states that the cheliceral stridulatory organ of the type species of Toschia (namely Ca. picta (Caporiacco, 1949) comb. nov.) is absent. However, our observations refute this previous statement; the cheliceral stridulatory organ is present in Toschia (see Fig. 22D). Jocqué (1984) mentioned the need of a detailed study of the male palp, given the difficulty to place species with certainty to either Toschia, Atypena [proposed by Jocque (1983) as a senior synonym of Callitrichia] or Oedothorax. Interestingly, however, he assigned Ca. minuta (Jocqué, 1984) comb. nov. to Toschia based on the distally wide paracymbium, a common feature shared by many Callitrichia species. Additionally, Ca. minuta share with Ca. legrandi (Jocqué, 1985) comb. nov. the vertical palpal tibial apophysis, the small apophysis retrolateral to it, the absence of ventral radical process, the less enlarged terminal part of paracymbium, and the smaller body size. We provisionally transfer Toschia spinosa Holm, 1968 to Holmelgonia (but see below) based on the presence of a central embolar apophysis (fig. 28 in Holm, 1968), a feature shared with the type species Ho. nemoralis (Holm, 1962) [fig. 55 in Jocqué & Sharff (1986) and other Holmelgonia species, and absent in Callitrichia]. In addition, despite the lack of detail in the original palpal drawing of Toschia celans Gao et al., 1996, some features can be recognized, such as the embolic division with a radix not connected to the embolus via a membrane, and the paracymbium base covered by the retrolateral apophysis (absent in other species of Toschia). Given that these palpal features are significantly different from the general configuration of Callitrichia, and that all type specimens for this species are seemingly lost (therefore, no further inspection is possible), we conclude that this species is most likely misplaced in Toschia, and we provisionally transfer it to Callitrichia [Ca. celans (Gao et al., 1996) comb. nov. incertae sedis]. Remaining Toschia species can be confidently transferred to Callitrichia according to inspection of the figures in their original descriptions by Holm (1962), Jocqué (1981) and Jocqué & Scharff (1986). Jocqué (1984) questioned the generic position of Typhistes gloriosus based on a unique ‘long downpointing apophysis’ (i.e. the ventral radical process as defined in the present study) among other features. He also proposed that the genus Typhistes is closely related to Oedothorax and Atypena (i.e. as Callitrichia). The results of our phylogenetic analysis are in agreement with Jocqué (1984), placing Typhistes gloriosus within Callitrichia; we therefore transfer this species to Callitrichia as Ca. gloriosa (Jocqué, 1984) comb. nov. Monophyly: This genus is supported by the following two unambiguous character transformations: the distal part of the protegulum forming a rim (Ch 38, synapomorphic) and the number of aciniform gland spigots on posterior lateral spinneret equal to, or more than, three (Ch 116, homoplastic). Diagnosis: Callitrichia is similar to other taxa in Clade 13 in the palpal morphology and other somatic features, but is characterized and can be distinguished by the following combination of palpal and epigynal features: 1. Paracymbium: median- to large-sized; base covered by cymbial basal retrolateral extension from dorsal view (invisible from dorsal view in Oedothorax); terminal part moderately to greatly enlarged (slender in Oedothorax; moderately enlarged in Mitrager); distal setae group with distal position (with middle position in Oedothorax); distal clasp anteriorly extended, without striae (retrolaterally extended and/or with striae in many Mitrager). 2. Copulatory bulb: embolus retrolaterally curved, short, usually stout, basal protuberance absent (prolaterally curved, with basal protuberance in Oedothorax); embolic membrane flat, with few or no papillae (cylindrical in Oedothorax); embolus–radix membranous region not extended to prolateral side of radix (extended to prolateral side in Oedothorax); ventral radical process present in most species [except Ca. macrophthalma (Locket & Russell- Smith, 1980) comb. nov., Ca. legrandi and Ca. paralegrandi (Tanasevitch, 2016) comb. nov.] (absent in Oedothorax and most Mitrager species), degree of extension varies from low [e.g. Ca. holmi (Wunderlich, 1978) comb. nov., Ca. uncata] to high [Ca. gloriosa, Ca. pileata (Jocqué & Scharff, 1986) comb. nov.]; in many species, radix with papillae in area between anterior radical process and ventral radical process [radical papillae absent in Oedothorax, Atypena and Mitrager, except M. clypeellum (Tanasevitch, 1998) comb. nov.]; lateral extension of radix absent (except in Ca. macrophthalma and Ca. convector); tegular papillae absent; central emboliar apophysis absent (present in Ho. basalis); marginal suprategular apophysis present, distal suprategular apophysis straight, distally oriented, mostly round at tip. 3. Palpal tibia: prolateral apophysis ranges from vertically highly extended (e.g. Ca. sellafrontis) to low (e.g. Ca. legrandi) (not vertically extended in Oedothorax and most Mitrager, except M. noordami); retrolateral apophysis inconspicuous (conspicuous in Mitrager, Atypena and some ‘ Oedothorax ’ incertae sedis). 4. Epigyne: different from Oedothorax in that the entrance of copulatory ducts into the spermathecae are ectal to the exits of fertilization ducts from the spermathecae. Description: The genus includes medium-sized erigonines with a mostly variegated opisthosoma [except Ca. latitibialis (Bosmans, 1988) comb. nov., Ca. legrandi, Ca. longiducta (Bosmans, 1988) comb. nov., Ca. macrophthalma, Ca. monoceros (Miller, 1970) comb. nov., Ca. monticola (Tullgren, 1910), Ca. muscicola (Bosmans, 1988) comb. nov., Ca. obtusifrons Miller, 1970, Ca. paralegrandi and Ca. pilosa (Wunderlich, 1978) comb. nov.]. Male prosoma varies in degree of modification, ranging from absent to prominent PME hump, inter-AME-PME groove and inter-AME-PME lobe. One sub-AME seta. Chelicerae of normal form and size, without mastidia.This genus also shares those features defining Clade 13 (see above). For palpal and epigynal feature see diagnosis. Species included: The genus comprise 55 species and one subspecies as follows. Twenty-two species originally placed in Callitrichia: Ca. afromontana Scharff, 1990, Ca. aliena Holm, 1962, Ca. cacuminata Holm, 1962, Ca. crinigera Scharff, 1990, Ca. glabriceps Holm, 1962, Ca. hamifera, Ca. inacuminata Bosmans, 1977, Ca. incerta Miller, 1970, Ca. kenyae Fage, 1936, Ca. marakweti Fage, 1936, Ca. meruensis Holm, 1962, Ca. mira (Jocqué & Scharff, 1986), Ca. monticola, Ca. obtusifrons, Ca. paludicola Holm, 1962, Ca. pileata (Jocqué & Scharff, 1986), Ca. hirsuta nom. nov. (Jocqué & Scharff, 1986), Ca. ruwenzoriensis Holm, 1962, Ca. sellafrontis, Ca. silvatica Holm, 1962, Ca. taeniata Holm, 1968 and Ca. turrita Holm, 1962. Twelve species (plus one subspecies) here transferred from Ophrynia: Ca. galeata (Jocqué & Scharff, 1986) comb. nov., Ca. galeata lukwangulensis (Jocqué & Scharff, 1986) comb. nov., Ca. infecta comb. nov. , Ca. insulana (Scharff, 1990) comb. nov., Ca. juguma comb. nov. , Ca. perspicuua (Scharff, 1990) comb. nov., Ca. revelatrix (Jocqué & Scharff, 1986) comb. nov., Ca. rostrata (Jocqué & Scharff, 1986) comb. nov., Ca. summicola (Jocqué & Scharff, 1986) comb. nov., Ca. superciliosa, Ca. trituberculata (Bosmans, 1988) comb. nov., Ca. truncatula (Scharff, 1990) comb. nov., Ca. uncata. Eight species here transferred from Toschia: Ca. aberdarensis (Holm, 1962) comb. nov., Ca. casta (Jocqué & Scharff, 1986) comb. nov., Ca. concolor (Caporiacco, 1949) comb. nov., Ca. cypericola (Jocqué, 1981) comb. nov., Ca. minuta, Ca. picta, Ca. telekii (Holm, 1962) comb. nov. and Ca. virgo (Jocqué & Scharff, 1986) comb. nov. Eleven species here transferred from Oedothorax: Ca. convector, Ca. holmi (= Ca. simplex (Jocqué & Scharff, 1986), synon. nov.), Ca. latitibialis, Ca. legrandi, Ca. longiducta, Ca. muscicola, Ca. pilosa, Ca. paralegrandi, Ca. monoceros , Ca. macropthalma and Ca. usitata. One species here transferred from Toschia celans: Ca. gloriosa. One species misplaced in Toschia (T. celans) here transferred as Ca. celans. The following species are here transferred to other genera: Atypena formosana (Oi, 1977) from Callitrichia; Holmelgonia spinosa (Holm, 1968) comb. nov. from Toschia. Distribution: Tanzania, Malawi, Cameroon, Algeria, Kenya, Nigeria, Uganda, Angola, South Africa, Congo, Comoros, Ethiopia, India (Himalaya), Thailand. Natural history: Species of this genus have been found in litter, low vegetations and under stones in habitats like montane rain forests, evergreen forests and gallery forests., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 451-459, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Fage L, Simon E. 1936. Arachnida. III. Pedipalpi, Scorpiones, Solifuga et Araneae (1 re partie). In: Mission scientifique de l'Omo. Memoires du Museum d'Histoire Naturelle de Paris 4: 293 - 340.","Jocque R. 1981. Erigonid spiders from Malawi (Araneida, Linyphiidae). Revue Zoologique Africaine 95: 470 - 492.","di Caporiacco L. 1949. Aracnidi della colonia del Kenya raccolti da Toschi e Meneghetti negli anni 1944 - 1946. Commentationes Pontificia Academia Scientiarum 13: 309 - 492.","Holm A. 1962. The spider fauna of the East African mountains. Part I: Fam. Erigonidae. Zoologiska Bidrag fran Uppsala 35: 19 - 204.","Scharff N, 1990 a. Spider of the family Linyphiidae from the Uzungwa mountains, Tanzania (Araneae). Entomologica Scandinavica, Supplement 36: 1 - 95.","Wunderlich J. 1978. Zur Kentniss der Gattung Oedothorax Bertkau 1883, Callitrichia Fage 1936 und Toschia Caporiacco 1949. Senkenbergiana Biologica 58: 257 - 260.","Jocque R. 1983. Sur la synonymie de Callitrichia Fage et Atypena Simon avec la redescription de quelques especes paleotropicales (Araneae, Linyphiidae). Bulletin du Museum National d'Histoire Naturelle de Paris 5: 235 - 245.","Jocque R, Scharff N, 1986. Spiders (Araneae) of the family Linyphiidae from the Tanzanian mountain areas Usambara, Uluguru and Rungwe. Annalen Zoologische Wetenschappen 248: 1 - 61.","Jocque R. 1984. Linyphiidae (Araneae) from South Africa. Part I: the collection of the Plant Protection Research Institute, Pretoria. Journal of the Entomological Society of South Africa 47: 121 - 146.","Jocque R. 1985. Linyphiidae (Araneae) from the Comoro Islands. Revue de Zoologie Africaine 99: 197 - 230.","Holm A. 1968. Spiders of the families Erigonidae and Linyphiidae from East and Central Africa. Annales, Musee Royal de l'Afrique Centrale, Sciences Zoologiques 171: 1 - 49.","Tanasevitch AV. 2015. Notes on the spider genus Oedothorax Bertkau, 1883 with description of eleven new species from India (Linyphiidae: Erigoninae). Revue suisse de Zoologie 122: 381 - 398.","Locket GH, Russell-Smith A. 1980. Spiders of the family Linyphiidae from Nigeria. Bulletin of the British Arachnological Society 5: 54 - 90.","Bosmans R. 1988. Scientific report of the Belgian Cameroon expeditions 1981 and 1983. No. 18. Further Erigoninae and Mynogleninae (Araneae: Linyphiidae) from Cameroonian highlands. Revue Zoologique Africaine 102: 5 - 32.","Tanasevitch AV. 1998 a. New Oedothorax Bertkau, 1883, from Nepal (Arachnida, Araneae, Linyphiidae). Bonner Zoologische Beitraege 47: 429 - 441.","Tu LH, Li SQ. 2004. A preliminary study of erigonine spiders (Linyphiidae: Erigoninae) from Vietnam. Raffles Bulletin of Zoology 52: 419 - 433.","Tanasevitch AV. 2016. A case of disjunct montane linyphiid species (Araneae) in the Palaeotropics, with notes on synonymy and the description of a new species. Revue Suisse de Zoologie 123: 235 - 240.","Miller F. 1970. Spinnenarten der Unterfamilie Micryphantinae und der Familie Theridiidae aus Angola. Publicacoes Culturais da Companhia de Diamantes de Angola 82: 75 - 166.","Bosmans R. 1977. Spiders of the subfamily Erigoninae from Mount Kenya. Scientific report of the Belgian Mt. Kenya Bio- Expedition, n ° 3. Revue Zoologique Africaine 91: 449 - 472.","Oi R. 1977. A new erigonid spider from Formosa. Acta Arachnologica 27: 23 - 26."]}

ATYPENA CIRRIFRONS (HEIMER, 1984) (FIGS 7G, 67H, 68H, 74) Paranasoona cirrifrons Heimer, 1984: 87, figs 1–8 (Dmf). Paranasoona cirrifrons Zhu & Sha, 1992: 42, figs 1–8 (mf). Paranasoona cirrifrons Song et al., 1999: 203, fig. 114N–Q (mf). Atypena cirrifrons Tanasevitch, 2014b: 72 (Tmf from Paranasoona = Atypena). Atypena cirrifrons Komisarenko et al., 2019: 27, figs 1, 2 (m). Examined material: Laos: Champasak, Muang Bachieng, Pak Song (15°10.6’ N, 106°14.253’ E), plateau, pine forest, 1280 m, at day, sweepnet, 1♂ 27.ix.2009, coll. P. Jäger, det. A. V. Tanasevitch, 2014 (SMF 64983-124). Diagnosis: Males: According to available illustrations of Atypena species, they are all similar in regard to their prosomal and palpal morphology. According to our observation, this species can be distinguished from A. formosana by the proportionally longer distal suprategular apophysis and ventral radical process. Females: No study of Atypena comparative female morphology exists to date; such diagnosis is, therefore, beyond the scope of the current study. Description: Male (SMF): Total length: 1.60. Prosoma: 0.73 long, 0.59 wide, PME region largely elevated, interocular region with translucent setae laterally curved (Fig. 7G). Eyes: AME-AME: 0.03, AME width: 0.06, AME- ALE: 0.03, ALE width: 0.07, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.14, PME width: 0.07, PME- PME: 0.14. Clypeus: upper-half hirsute, one sub- AME seta. Sternum: 0.42 long, 0.43 wide. Chelicerae: mastidia absent; stridulatory striae imbricated, rows compressed proximally (Fig. 67H). Legs: tibia chaetotaxy 2-2-1-1, dorsal proximal macroseta on tibia I, II, III and IV 0.46, 0.64, 0.83 and 1.81 times diameter of tibia, respectively; Tm I: 0.79. All metatarsi with trichobothrium. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPA short, pointed and sclerotized at tip; PC base not visible from dorsal view, distal setae close to distal clasp, terminal part slightly enlarged, distal clasp without striae, clasp extended apically (Fig. 74A); T without papillae; PT small, without papillae; TS long and slender, without papillae (Fig. 74A, C); MSA absent; DSA wide, truncated at tip, ventral corner more extended than dorsal corner (Fig. 74A); EM broad and flat, without papillae, exceeding ARP; ARP flat, unsclerotized; LER retrolateral to E; VRP long, slender; R without papillae; TP long, slender; E short and stout, slightly retrolaterally curved (Fig. 74B). Opisthosoma: dorsal pattern see Fig. 68H; PMS with mAP, three AC; PLS with triad, 3+ AC (Fig. 74F). Distribution: India, China, Laos, Thailand, Vietnam., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 540-541, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Heimer S. 1984. A new linyphiid spider from Vietnam (Arachnida, Araneae). Reichenbachia 22: 87 - 89.","Zhu CD, Sha YH. 1992. Two species of linyphiid spiders from south China (Arachnida: Araneae). Journal of Norman","Song DX, Zhu MS, Chen J. 1999. The spiders of China. Shijiazhuang: Hebei University of Science and Techology Publishing House.","Tanasevitch AV. 2014 b. On the linyphiid spiders from Thailand and West Malaysia (Arachnida: Aranei: Linyphiidae). Arthropoda Selecta 23: 393 - 414.","Komisarenko AA, Omelko MM, Marusik YM. 2019. An annotated list of linyphiid spiders (Aranei: Linyphiidae) of Laos. Far Eastern Entomologist 377: 26 - 32."]}

MITRAGER NOORDAMI VAN HELSDINGEN, 1985 (FIGS 37, 38O, 39O, 40P; SUPPORTING INFORMATION, FIG. S3C) Mitrager noordami van Helsdingen, 1985: 353, figs 1–14 (Dmf). Examined material: Indonesia: Central Java, Dijeng Plateau, near Gunung Prahu, 2580 m, sifted from litter among mosses, ferns and Ericaceae, 1♂ 1♀ 8.viii.1977, leg. A. Noordam (paratype, Naturalis Museum, Leiden). Diagnosis: Males: This species can be recognized by its unique, prominent prosomal modification (Fig. 38O). Females: Somatic features and coloration similar to other Mitrager and some Callitrichia species, but can be distinguished by the much more convergent borders between the dorsal and ventral plates of the epigyne (Fig. 37F, G). Description: Male (paratype, Naturalis Museum): Total length: 2.27. Prosoma: 1.13 long, 0.77 wide, with extremely high, ballon-shaped post-PME hump, inter-PME region with a protrution with two round lobes, anterior margin of lobes with row of light-coloured setae, interocular region with two dense, light-coloured setal tufts (Fig. 38O). Eyes: AME-AME: 0.03, AME width: 0.05, AME-ALE: 0.06, ALE width: 0.07, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.1, PME width: 0.07, PME-PME: 0.35. Clypeus hirsute. Sternum: 0.6 long, 0.58 wide. Chelicerae: stridulatory striae imbricated, rows widely and evenly spaced (Fig. 39O). Legs: dorsal proximal macroseta on tibia IV 1.80 times diameter of tibia; Tm I: 0.82. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPS absent; TPA large, conical, distal part narrow and flat, tip curved downwards; PC distal setae close to distal clasp, distalsetae-bearing area slightly wide, distal clasp without striae, clasp extended apically; T without papillae; PT median-long, with small papillae; TS median-long, slender, without papillae; MSA present; DSA wide, round at tip, ventral corner slightly more protruded (Fig. 37A); EM broad and flat, retrolateral to LER, with parallel folds extended until anterior margin, exceeding ARP; ARP flat, blunt, short; LER large, bent over E; VRP long, thick at base; R without papillae; TP long,round at tip; E median-long, retrolaterally curved, E tip with prolateral anterior flat extension (Fig. 37A). Opisthosoma: dorsal pattern see Fig. 40P; PMS with mAP, AC absent; PLS with triad, one AC (Fig. 37H). Female (paratype, Naturalis Museum): Total length: 2.48. Prosoma: 1.22 long, 0.84 wide. Eyes: AME-AME: 0.03, AME width: 0.05, AME-ALE: 0.03, ALE width: 0.08, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.05, PME width: 0.08, PME-PME: 0.12. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.67 long, 0.62 wide. Legs: dorsal proximal macroseta on tibia III 2.39 times diameter of tibia; Tm I: 0.62. Epigyne: Clade 13 characteristic morphology, border between ventral and dorsal plates largely converging toward the middle (Fig. 37F, G). Opisthosoma dorsal pattern same as male; PMS one CY, one mAP, AC absent; PLS with triad, two CY, one AC (Fig. 37I). Distribution: Only known from the type locality in Indonesia. Remarks: The setal tufts at interocular region and inter-PME region are aggregated setae, similar to that found in some Callitrichia species., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 485, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Van Helsdingen PJ. 1985. Mitrager noordami, an erigonine novelty from Java. Bulletin of the British Arachnological Society 6: 353 - 358."]}

CALLITRICHIA CONVECTOR (TANASEVITCH, 2014) COMB. NOV. (FIGS 19G, 22G, 23, 24G) Oedothorax convector Tanasevitch, 2014a: 408, figs 64–71 (Dm). Type material: Holotype: Thailand: Chiang Mai Province, Chomthong District, Doi Inthanon, 1780 m, ♂ 3.iii.1987, leg. P. Schwendinger (MHNG, examined). Diagnosis: Males: This species can re recognized by the shape of prosomal modification with the inter-AME-PME groove and PME lobe; it can be discriminated from Oedothorax s.s., Mitrager and other Callitrichia species by its palpal features including the presence of tibial retrolateral apophysis with setae (no setae in Mitrager), the absence of tibial prolateral trichobothrium, the shape of paracymbium (distal clasp particularly long and slender), and the radix without tail piece. Description: Male (holotype, MHN): Total length: 1.68. Prosoma: 0.75 long, 0.55 wide, PME on frontal side of elevated region, short curved setae grouped at both sides of dorsal surface of prosomal elevation behind PME, inter-PME-PLE groove, three pairs of strong setae at groove lower margin, no setae inside groove (Fig. 19G). Eyes: AME-AME: 0.03, AME width: 0.05, AME-ALE: 0.01, ALE width: 0.05, ALE-PLE: 0.01, PLE width: 0.05, PLE-PME: 0.10, PME width: 0.05, PME-PME: 0.18. Clypeus: not hirsute, one sub-AME seta, particularly long. Chelicerae: stridulatory striae rows wide and evenly spaced (Fig. 22G). Sternum: 0.46 long, 0.5 wide. Legs: dorsal proximal macroseta on tibia IV 3.13 times diameter of tibia; Tm I: 0.81. Pedipalp: TPA narrow, slightly curved; TRA extended anteriorly, covering PC base from retrolateral view; PC long, distal-setae-bearing middle part enlarged, distal clasp slender and long (Fig. 23A–C); T without papillae; PT slender and long, with scale-like papillae; TS long, with small papillae; MSA absent; DSA short and wide (Fig. 23H); EM short, without papillae, not exceeding ARP (Fig. 23E); R without TP, scaly region distributed from ARP to VRP; LER present; E prolaterally spiral, between ARP and LER (Fig. 23E, F, G). Opisthosoma: dorsal pattern see Fig. 24G; PMS with mAP, two AC; PLS with one FL, 3+ AC (Fig. 23I–K). Female: Unknown Distribution: Only known from the type locality in Thailand. Habitat: No data., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 459, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2014 a. New species and records of linyphiid spiders from Laos (Araneae, Linyphiidae). Zootaxa 3841: 67 - 89."]}

HOLMELGONIA JOCQUÉ & SCHARFF, 2007 Type species: Elgonia nemoralis Holm, 1962. Diagnosis, description and taxonomic remarks: The resemblance of Holmelgonia to Callitrichia and other related genera (i.e. Oedothorax, Toschia and Ophrynia) has not been mentioned in previous publications. Regarding the shape of embolic division, the enlarged terminal part of paracymbium, the vastly erected palpal tibia prolateral apophysis and the general epigyne morphology, Holmelgonia is similar to numerous Callitrichia species. Nzigidahera & Jocqué (2014) described Holmelgonia as characterized by the absence of cheliceral stridulation ridges, the presence of a double ventral row of setae on the femora, the long tibial spines (two to three times as long as the diameter of the segment), tibial chaetotaxy 2-2-1-1, TmI between 0.32 and 0.7, the absence of prosomal modification and the apophysis on the dorsal side of the palpal tibia. However, all these features are not unique to the members of this genus. One potential diagnostic feature, the absence of cheliceral stridulatory ridges, is actually present in our studied species (Ho. basalis, Fig. 22B), and the chelicera of Ho. disconveniens Nzigidahera & Jocqué, 2014 (SEM photo, figs 2–3 in Nzigidahera & Jocqué 2014) shows a scaly lateral surface without setae, which could also be interpreted as stridulatory striae (fig. 3 shows the detail of the anterior surface, instead of the lateral surface). Despite the lack of defining features for this genus, and the sister-relationship of Ho. basalis and Callitrichia shown in the present study, this genus differs from Callitrichia in the lack of truncated protegulum and the presence of a central embolar apophysis (Fig. 36B, D, arrow). The latter structure is present in all species with known males, except Ho. discoveniens and Ho. afromontana Nzigidahera & Jocqué, 2014. Due to the lack of synapomorphies defining this genus, the monophyly of Holmelgonia remains untested (Nzigidahera & Jocqué 2014). In our phylogenetic hypothesis (Fig. 3), Ho. basalis resulted sister to Callitrichia (Clade 40), a relationship supported by the presence of a rim at the distal area of the protegulum (Ch 38, synapomorphic) and posterior lateral spinneret with more than three aciniform gland spigots (Ch 116, homoplastic). Since only one Holmelgonia species was represented in our analysis, it is yet premature to decide whether the genus is a monophyletic assemblage sister to Callitrichia, or a paraphyletic set of taxa that should in turn require taxonomic actions. Further phylogenetic analyses with a thorough species representation of Holmelgonia, including the type species Ho. nemoralis, as well as Callitrichia and other related taxa, is needed. Distribution: Ivory Coast, Cameroon, Congo, Burundi, Tanzania, Uganda, Kenya, Malawi and Mozambique. Natural history: Species of this genus were found in lower vegetation, forest litter and by pitfall traps (Jocqué 1981; Jocqué & Scharff 1986; Scharff 1990a). Most species live at high altitude and have small endemic ranges, while some have a wide distribution at midaltitude (Nzigidahera & Jocqué 2014).

OEDOTHORAX GIBBOSUS (BLACKWALL, 1841) (FIGS 6, 7O, 8A, 9A; SUPPORTING INFORMATION, FIG. S1A) Neriene gibbosa Blackwall, 1841: 653 (Dmf). Neriene tuberosa Blackwall, 1841: 654 (Dm). Argus gibbosus Walckenaer, 1847: 513. Argus tuberosus Walckenaer, 1847: 514. Neriene tuberosa Blackwall, 1864: 279, pl. 19, fig. 192 (m). Erigone tuberosa Thorell, 1873: 447. Neriene gibbosa O. Pickard-Cambridge, 1873: 455, pl. 34, fig. 20. Erigone henkingi Dahl, 1883: 49, figs 29, 30 (Dm). Gongylidium gibbosum Simon, 1884: 489, figs 270–272 (mf). Gongylidium tuberosum Simon, 1884: 490, figs 273– 275 (m, Df). Neriene henkingi Dahl, 1886: 88. Neriene gibbosa Simon, 1894: 666, fig. 720 (m). Kulczynskiellum gibbosum F.O. Pickard-Cambridge, 1895: 39. Gongylidium gibbosum Becker, 1896: 86, pl. 9, fig. 7K (mf). Kulczynskiellum tuberosum Bösenberg, 1902: 171, pl. 15, fig. 232 (mf). Oedothorax gibbosus Bösenberg, 1902: 213, pl. 19, fig. 300 (mf). Stylothorax gibbosa Reimoser, 1919: 72. Stylothorax henkingi Reimoser, 1919: 72. Stylothorax tuberosa Reimoser, 1919: 73. Oedothorax gibbosus Simon, 1926: 451, 522, fig. 785 (mf). Oedothorax tuberosus Simon, 1926: 452, 522. Oedothorax gibbosus Bishop & Crosby, 1935: 264, pl. 22, figs 70–73 (mf). Oedothorax gibbosus Denis, 1947: 138, figs 6A, 7A, 8A, 9F, 10G, H, 11F (mf). Oedothorax tuberosus Denis, 1947: 138, figs 1A, 6B, C, 7B, 8B, 9G, 10I–K, 11G (mf). Oedothorax gibbosus Locket & Millidge, 1953: 239, figs 145A, 146A, G, H (mf). Oedothorax tuberosus Locket & Millidge, 1953: 239, figs 145A, 146B, G, H (mf). Oedothorax tuberosus Wiehle, 1960a: 454, figs 835–842 (mf). Oedothorax tuberosus Tystshenko, 1971: 251, fig. 829 (f). Oedothorax tuberosus Miller, 1971: 262, pl. LIV, figs 4–6 (f). Oedothorax gibbosus Palmgren, 1976: 89, figs 8.9–12, 14, 15 (mf). Oedothorax tuberosus Palmgren, 1976: 89, figs 8.9, 13–15 (f). Oedothorax gibbosus Růžička, 1978: 195, fig. 8H, I (f). Oedothorax gibbosus Bosmans, 1985: 65, figs 11, 25, 31 (m). Oedothorax gibbosus Roberts, 1987: 57, figs 21E, 22B (mf). Oedothorax gibbosus tuberosus Roberts, 1987: 57, fig. 21F–g (downgraded to ‘form’). Oedothorax gibbosus de Keer & Maelfait, 1988: 3. Oedothorax gibbosus Heimer & Nentwig, 1991: 224, fig. 601 (mf). Oedothorax gibbosus Hormiga, 2000: 48, figs 21A–G, pl. 49A–F, 50A–F, 51A–F (mf). Oedothorax gibbosus Tanasevitch, 2015: 382, figs 1, 2 (m). Oedothorax gibbosus Russell-Smith, 2016: 22, fig. 1 (f). Type material: In Blackwall’s description (1841), 2♂ of gibbosus morph, 1♂ of tuberosus morph and 5♀ were found under stones in a moist pasture in Oakland, United Kingdom, in May 1838 [individual number according to Walckenaer (1847)], but no type designation was made, nor does any reference provide information about where the types might have been deposited. Nevertheless, the unique prosomal modification of the gibbosus morph described in the original description makes the identification of specimens in this study unequivocal. Subsequently, the identical palpal morphology confirmed the conspecificity of examined tuberosus morph with the gibbosus morph (Roberts, 1987). Examined material: Denmark: Øjesø, near Feldballe (56°17’ N, 10°36’ E), 6♂ 4♀ 19.v.1997, leg. Peter Gejdos (ZMUC 00008860); Ulvshale Skov, toward Skovbund, 3♂ 6♀ 12.x.2002, leg. J. Peterson, det. N, Scharff, 2002 (ZMUS 00007790). England: Cambridgeshire, Wicken Fen, 2♂ 18.–22.v.1957, leg. D. J. Clark (NHM). Wales: Merionethshire, Dolgelly, 1♂ 8.vi.1954, coll. D. J. Clark, det. G. H. Locket (NHM). Diagnosis: Males: This species has two male morphs. The gibbosus - morph can be identified by the shape of post-ocular groove and hump, not divided into three lobes on the anterior side as in Oe. trilobatus; the groove is equipped with dense setae as in Oe. trilobatus, absent in the Mitrager species with post-ocular groove. The tuberosus -morph does not possess a post-ocular groove, and the prosoma is elevated at the position of the fovea, distinguishing it from other Oedothorax s.s. species. The bifid palpal tibial prolateral apophysis and the absence of basal thorn distinguish this species from other Oedothorax s.s. species. Females: Can be distinguished from other Oedothorax s.s. species by the epigynal general configuration and number of sub-AME setae (one; two in Oe. fuscus, Oe. agrestis, Oe. meridionalis and Oe. tingitanus). Distinguished from Oe. agrestis, Oe. apicatus, Oe. fuscus, Oe. gibbifer, Oe. tingitanus and Oe. retusus by the more convergent ventral plate borders. Distinguished from Oe. meridionalis by not having a wide chamber at the entrances of the copulatory ducts (Fig. 6F, in comparison to Fig. 10F); from Oe. paludigena by the narrower posterior margin of the dorsal plate; from Oe. trilobatus by the much shorter copulatory ducts. Description: Male, tuberosus-morph (ZMUC 00007790): Total length: 2.37. Prosoma: 1.05 long, 0.77 wide, postocular region with a hump posteriorly, without post-ocular groove. Eyes: AME-AME: 0.03, AME width: 0.05, AME-ALE: 0.03, ALE width: 0.07, ALE-PLE: 0, PLE width: 0.07, PLE-PME: 0.04, PME width: 0.07, PME-PME: 0.07. Sternum: 0.61 long, 0.55 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.20, 1.47, 1.80 and 2.13 times diameter of tibia, respectively; Tm I: 0.62. Male, gibbosus-morph (ZMUC 00008860): Total length: 2.35. Prosoma: 1.03 long, 0.79 wide, postocular region with wide, transverse, hirsute groove and large hump posteriorly (Fig. 7O). Eyes: AME-AME: 0.04, AME width: 0.05, AME-ALE: 0.03, ALE width: 0.06, ALE-PLE: 0, PLE width: 0.07, PLE-PME: 0.06, PME width: 0.06, PME-PME: 0.08. Sternum: 0.59 long, 0.54 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.22, 0.17, 0.78 and 0.82 times diameter of tibia, respectively; Tm I: 0.65. Male, both morphs: Clypeus: not hirsute, one sub-AME seta. Chelicerae: mastidia present; stridulatory striae scaly, rows widely and evenly spaced (Fig. 8A). Pedipalp: TPA with two scaly lobes, a depressed region in between with scaly distal margin; BT absent (Fig. 6C); PC distal setae at median position (Fig. 6A); T without papillae, protegulum with long papillae; TS short, without papillae (Fig. 6D); DSA tip broad and smoothly serrated (Fig. 6A); EM EM short, cylindrical, distally oriented, with papillae; TP without small protuberances; E not broadened at basal part (Fig. 6B). Opisthosoma: brown, evenly coloured (Fig. 9A); spinnerets see Fig. 6I. Female (ZMUC): Total length: 2.53. Prosoma: 1.13 long, 0.83 wide. Eyes: AME-AME: 0.029, AME width: 0.054, AME-ALE: 0.031, ALE width: 0.074, ALE-PLE: 0.005, PLE width: 0.074, PLE-PME: 0.053, PME width: 0.064, PME-PME: 0.068. Clypeus: not hirsute, one sub- AME seta. Sternum: 0.65 long, 0.58 wide. Legs: dorsal proximal macroseta on tibia II, III and IV 1.76, 2.02 and 2.14 times diameter of tibia, respectively; Tm I: 0.69. Chelicerae: stridulatory striae scale-like. Epigyne: Clade 13 characteristic morphology, ventral plate borders converging anteriorly, copulatory duct short (Fig. 6F–H) Opisthosoma: brown, evenly coloured. Variation: The measurements are based on examined material. Males, tuberosus-morph ( N = 7, means in parentheses): Total length 2.23–2.37 (2.28). Prosoma: 1.02–1.11 (1.06) long, 0.76–0.85 (0.80) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.82–1.39 (1.12), 0.87–1.64 (1.31, N = 5), 1.48–2.00 (1.75, N = 6) and 1.56–2.16 (1.85) times diameter of tibia, respectively; Tm I: 0.60–0.68 (0.65). Males, gibbosus-morph ( N = 5, means in parentheses): Total length 1.94–2.35 (2.22). Prosoma: 0.97–1.06 (1.02) long, 0.76–0.81 (0.79) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.19 –0.60 (0.28), 0.17–0.61 (0.30), 0.41–1.16 (0.70) and 0.45–1.52 (0.83) times diameter of tibia, respectively; Tm I: 0.59–0.71 (0.64). Females ( N = 10, means in parentheses): Total length 2.33–2.83 (2.64). Prosoma: 0.93–1.20 (1.15) long, 0.71– 0.87 (0.83) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.53–1.82 (1.69, N = 9), 1.71–1.87 (1.79, N = 8), 1.89–2.27 (2.07) and 1.90–2.37 (2.20) times diameter of tibia, respectively; Tm I: 0.59–0.69 (0.65) Distribution: Europe, Turkey. Habitat: Open, humid areas., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 428-434, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Blackwall J. 1841. The difference in the number of eyes with which spiders are provided proposed as the basis of their distribution into tribes; with descriptions of newly discovered species, and the characters of a new family and three new genera of spiders. Transactions of the Linnean Society of London 18: 601 - 670.","Walckenaer CA. 1847. Dernier Supplement. In: Walckenaer CA, Gervais P, eds. Histoire naturelles des insects, Vol. 4. Paris: Apteres, 365 - 564.","Blackwall J. 1864. A history of the spiders of Great Britain and Ireland. 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Notes d'araneologie marocaine. X, Les erigonides du Maroc. Bulletin de la Societe des Sciences Naturelles du Maroc 47: 137 - 164.","Locket GH, Millidge AF. 1953. British spiders, Vol. II. London: Ray Society.","Wiehle H. 1960 a. Spinnentiere oder Arachnoidea, XI: Micryphantidae-Zwergspinnen. Tierwelt Deutschlands 47: 1 - 620.","Tystshenko VP. 1971. Opredelitel' paukov evropejskoj casti SSSR. Leningrad: Nauka.","Miller F. 1971. Pavouci-Araneida. Klic zvireny CSSR 4: 51 - 306.","Palmgren P. 1976. Die Spinnenfauna Finnlands und Ostfennoskandiens. VII. Linyphiidae 2. Fauna Fennica 29: 1 - 126.","Ruzicka V. 1978. Revision der diagnostischen Merkmale der Weibchen der tschechoslovakischen Arten der Gattung Oedothorax (Araneae: Micryphantidae). Vestnik Ceskoslovenske Zoologicke Spolecnosti v Praze 42: 195 - 208.","Bosmans R. 1985. Etudes sur les Linyphiidae nord-africains. II. Le genre Oedothorax Bertkau en Africa du Nord, avec une revision des caracteres diagnostiques des males des especes ouest-palearctiques. Biologisch Jaarboek Dodonaea 53: 58 - 75.","De Keer R, Maelfait J-P. 1988. Oedothorax gibbosus (Blackwall) and Oedothorax tuberosus (Blackwall): one species. Newsletter of the British Arachnological Society 53: 3.","Hormiga G. 2000. Higher level phylogenetics of erigonine spiders (Araneae, Linyphiidae, Erigoninae). Smithsonian Contributions to Zoology 609: 1 - 160.","Russell-Smith A. 2016. Identification of females of British Oedothorax species. Newsletter of the British Arachnological Society 137: 22 - 24."]}

NERIENE FUEGIANA SIMON, 1902 Neriene fuegiana Simon, 1902: 17 (Df). Oedothorax fuegianus Petrunkevitch, 1911: 262. Oedothorax fuegianus Miller, 2007: 244, fig. 186C (f, misplaced in this genus). Neriene fuegiana Dupérré & Harms, 2018: 4, fig. 4A–C (f). Type material: Lectotype ♀ designated by Dupérré & Harms (2018) (Zoological Museum of Hamburg, ZMH- A0000758, not examined). Non-Type material: Argentina: South Tierra del Fuego, coastline right southwest of Kap San Pio (54°15’ S, 68°0’ W), ♀ ‘holotype’ 27.xii.92, coll. Mich (Simon, 1902; Miller, 2007) (not examined). Diagnosis: Females: This species can be identified by the epigynal morphology, with a triangular protrusion of the ventral plate in the middle over the dorsal plate (fig. 186C in Miller, 2007)., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 537, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Simon E. 1902. Arachnoideen, excl. Acariden und Gonyleptiden. In: Ergebnisse der Hamburger Magalhaensischen Sammelreise 1892 / 1893. Band Arthropoden, Vol. 6. Hamburg: Naturhistorisches Museum zu Hamburg, L. Friederichsen & Co, 1 - 47.","Petrunkevitch A. 1911. A synonymic index-catalogue of spiders of North, Central and South America with all adjacent islands, Greenland, Bermuda, West Indies, Terra del Fuego, Galapagos, etc. Bulletin of the American Museum of Natural History 29: 1 - 791.","Miller JA. 2007. Review of erigonine spider genera in the neotropics (Araneae: Linyphiidae, Erigoninae). Zoological Journal of the Linnean Society 149: 1 - 263.","Duperre N, Harms D. 2018. Raising the dead: rediscovery and redescription of some lost spider types (Araneae) described by Eugene Simon. Evolutionary Systematics 2: 1."]}

ATYPENA FORMOSANA (OI, 1977) (FIGS 7H, 67I, 68I, 73) Callitrichia formosana Oi, 1977: 23, figs 1–5 (Dmf). Oedothorax formosanus Brignoli, 1983: 349. Callitrichia formosana Song, 1987: 144, fig. 104 (Tmf from Oedothorax). Atypena formosana Tazoe, 1992: 212, figs 1–10 (mf; NB: generic placement following Jocqué, 1983). Oedothorax formosanus Okuma et al., 1993: 13, fig. 11A–B (f). Atypena formosana Barrion & Litsinger, 1994: 319, figs 1673–1676 (mf). Callitrichia formosana Song et al., 1999: 160, fig. 88K–L (mf). Callitrichia formosana Ono et al., 2009: 267, figs 95–99 (mf). Examined material: Japan: Okinawa, Iromotejima Island, Urauchi, 2♂ 3♀ 02.iv.1995, leg. A. Tanikawa, det. A, Tanasevitch (SMF 56485). Diagnosis: Males: This species can be distinguished from A. cirrifrons by the proportionally shorter distal suprategular apophysis and ventral radical process. Females: No study of Atypena comparative female morphology exists to date; such diagnosis is, therefore, beyond the scope of the current study. Description: Male (SMF): Total length: 1.76. Prosoma: 0.78 long, 0.62 wide, PME region largely elevated, interocular region with translucent setae laterally curved (Fig. 7H). Eyes: AME-AME: 0.03, AME width: 0.06, AME- ALE: 0.05, ALE width: 0.08, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.14, PME width: 0.07, PME- PME: 0.17. Clypeus: upper-half hirsute, one sub- AME seta. Sternum: 0.46 long, 0.46 wide. Chelicerae: mastidia absent; stridulatory striae imbricated, rows compressed proximally (Fig. 67I). Legs: tibia chaetotaxy 2-2-1-1, dorsal proximal macroseta on tibia I, II, III and IV 0.65, 0.82, 1.45 and 1.87 times diameter of tibia, respectively; Tm I: 0.78. All metatarsi with trichobothrium. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPA short, pointed and sclerotized at tip; PC base not visible from dorsal view, distal setae close to distal clasp, terminal part slightly enlarged, distal clasp without striae, clasp extended apically (Fig. 73A); T without papillae; PT small, without papillae; TS long and slender, without papillae (Fig. 73D); MSA absent; DSA wide, truncated at tip, ventral corner more extended than dorsal corner (Fig. 73A); EM broad and flat, without papillae, exceeding ARP; ARP flat, unsclerotized; LER retrolateral to E; VRP long, slender; R without papillae; TP long, slender; E short and stout, slightly retrolaterally curved (Fig. 73B). Opisthosoma: dorsal pattern see Fig. 68I; PMS with mAP, three AC; PLS with triad, 3+ AC. Female (SMF): See also Oi (1977); epigyne without prominent external modification, copulatory ducts slightly curved, trojactory simple, entrances to spermathecae mesal to exits of fertilixation ducts. (Fig. 73). Opisthosoma: dorsal pattern same as male; PMS with CY, mAP, 2–4 AC; PLS with triad, 2 CY, 3+ AC (Fig. 73). Distribution: Bangladesh to Japan. Remarks: As in A. cirrifrons, a scaly sac-like structure is observed at the paracymbial base (Figs 73A, C, 74A, C). According to the results of our phylogenetic analysis, and the similarities in male genital organs and prosomal modifications, this species is congeneric with A. cirrifrons. ADDITIONAL MORPHOLOGICAL DATA, Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 538-540, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Oi R. 1977. A new erigonid spider from Formosa. Acta Arachnologica 27: 23 - 26.","Brignoli PM. 1983. A catalogue of the Araneae described between 1940 and 1981. Manchester: Manchester University Press.","Song DX. 1987. Spiders from agricultural regions of China (Arachnida: Araneae). Beijing: Agriculture Publishing House.","Tazoe S. 1992. A newly recorded spider from Japan, Atypena formosana (Oi, 1977), comb. nov. (Linyphiidae). Acta Arachnologica 41: 211 - 214.","Jocque R. 1983. Sur la synonymie de Callitrichia Fage et Atypena Simon avec la redescription de quelques especes paleotropicales (Araneae, Linyphiidae). Bulletin du Museum National d'Histoire Naturelle de Paris 5: 235 - 245.","Okuma C, Kamal NQ, Hirashima Y, Alam MZ, Ogata K. 1993. Illustrated Monograph of the Rice Field Spiders of Bangladesh. Institute of Postgraduate Studies in Agriculture (Salna, Gazipur, Bangladesh) (Vol. 1). Japan International Cooperation Agency Project Publication.","Barrion AT, Litsinger JA. 1994. Taxonomy of rice insect pests and their arthropod parasites and predators. In: Heinrichs EA, ed. Biology and management of rice insects. New Delhi: Wiley Eastern, 13 - 15, 283 - 359.","Song DX, Zhu MS, Chen J. 1999. The spiders of China. Shijiazhuang: Hebei University of Science and Techology Publishing House.","Ono H, Matsuda M, Saito H. 2009. Linyphiidae, Pimoidae. In: Ono H, ed. The spiders of Japan with keys to the families and genera and illustrations of the species. Kanagawa: Tokai University Press, 253 - 344."]}

‘OEDOTHORAX’ TRILINEATUS SAITO, 1934 INCERTAE SEDIS Oedothorax trilineatus Saito, 1934: 311, pl. 13, fig. 18, pl. 15, fig. 61 (Df). Oedothorax trilineatus, Saito, 1959: 79, fig. 84a–c (f). Material: Japan: Nemuro, 4♀, leg. S. Motoda, 15.vii.1931, not examined. No type material was designated. Diagnosis: Females: This species can be distinguished by the black prosoma, the black opisthosoma with three oblique white markings at the sides, and the white/ black annulated legs and palps. Males: unknown Distribution: Japan: Nemuro. Remarks: According to Saito (1934), this species has prosoma and opisthosoma uniformly black in colour, and legs are white annulated with black, different from the yellow to dark-brown coloration of Oedothorax and Mitrager species. Furthermore, according to his description, three out of four specimens have leg II as the shortest leg, in comparison to leg III as the shortest leg in Linyphiidae in general. Therefore, this species is most likely not a linyphiid., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 556, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Saito S. 1934. Spiders from Hokkaido. Journal of the Faculty of Agriculture, Hokkaido Imperial University, Sapporo, Japan 33: 267 - 362.","Saito S. 1959. The spider book illustrated in colours. Tokyo: Hokuryukan."]}

MITRAGER LUCIDA (WUNDERLICH, 1974) COMB. NOV. (FIGS 38M, 39M, 40N, 55, 56B; SUPPORTING INFORMATION, FIG. S3A) Oedothorax lucidus Wunderlich, 1974: 180, figs 28–33 (Dm). Type material: Holotype: East Nepal: Mt. Chordung at Jiri, Abies-Rhododendron- forest, 2900 m, 1♂ 1.iv.1973 (SMF 28897, examined). Paratypes: Jiri, 1800–2000 m, 6♂ i.1970 (SMF 28898, examined). Non-type material: 1♂ subadult collected with the paratypes (SMF 28898). Diagnosis: Males: This species shares with M. sexoculata the PME located within the transverse groove between the prosomal elevation and the remaining eyes, the highly sclerotized stump-like setae on both upper and lower surfaces within the groove, and one retrolateral trichobothrium on male palpal tibia instead of the common two. This species can be distinguished from M. sexoculata by the smaller TPA and the papillaebearing PT. Description: Male (holotype, SMF 28897): Total length: 1.90. Prosoma: 0.94 long, 0.67 wide, region including PME elevated, transverse groove between elevation and other eyes, with highly sclerotized stump-like setae on both upper and lower surfaces within groove (Figs 38M, 56B). Eyes: AME-AME: 0.03, AME width: 0.04, AME-ALE: 0.04, ALE width: 0.05, ALE-PLE: 0.01, PLE width: 0.04; PME inside PME-lobe. Clypeus: slightly elevated, with small setae scarcely distributed. Sternum: 0.50 long, 0.47 wide. Chelicerae: stridulatory striae imbricated, rows widely and evenly spaced (Fig. 39M). Legs: dorsal proximal macroseta on tibia III 2.93 times diameter of tibia; Tm I: 0.85. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, one retrolateral trichobothria; TPS scaly, straight, wide at base, slightly retrolaterally directed; TPA slightly elevated; TRA distally oriented; PC median-sized, base not visible from dorsal view, distal setae close to distal clasp, distal clasp without striae, distally extended (Fig. 55A); T without papillae; PT with long papillae; TS long, thick, with several small papillae at base (Fig. 55D); MSA present; DSA not pointed, not turned retrolaterally; EM flat, without papillae, not exceeding ARP (Fig. 55C); ARP rounded at tip, striated; LER without striae, extended dorsal to E; VRP absent; TP broad (Fig. 55D); E retrolaterally spiral, anterior margin at base round (Fig. 55B). Opisthosoma: dorsal pattern see Fig. 40N; PMS with mAP, AC absent; PLS with triad, AC absent (Fig. 55E). Female: Unknown. Variation: The measurements are based on examined material. Males ( N = 7, means in parentheses): Total length 1.88 – 2.13 (1.99). Prosoma: 0.94 – 0.98 (0.96) long, 0.67 – 0.74 (0. 71) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.78 – 1.29 (1.08, N = 5), 0.77 – 1.24 (0.98, N = 6), 0.44 – 0.70 (0.59) and 1.57 – 2.51 (2.01, N = 3) times diameter of tibia, respectively; Tm I: 0.76 – 0.86 (0.82). Distribution: East Nepal. Habitat: Forests., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 508-510, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Wunderlich J. 1974. Linyphiidae aus Nepal, II. Die Gattung Oedothorax Bertkau 1883 (Arachnida: Araneae). Senckenbergiana Biologica 55: 169 - 188."]}

TMETICUS BREVIPALPUS BANKS, 1901 Tmeticus brevipalpus Banks, 1901: 580, pl. 33, fig. 14 (Dm). Gongylidium brevipalpus Banks, 1910: 29. Oedothorax brevipalpus Petrunkevitch, 1911: 261. Oedothorax brevipalpatus Roewer, 1942: 645 (unnecessary replacement name; see discussion in Buckle et al., 2001: 134). Type material: No type designated by Banks, 1901. Diagnosis: Males: The only reference for discerning this species is figure 14 in plate 33 in Banks (1901), which gives a rough image of its palpal tibial apophyses and the shape of the copulatory bulb, in which the complete structure of the embolic division cannot be recognized. Nevertheless, it shows no embolic division typical of Oedothorax and related taxa (Clade 13). In the latter taxa, the embolus has a membranous connection to a radix with an anterior radical process and a tailpiece. Females: Unknown. Distribution: USA, New Hampshire. Remarks: According to the palpal structure shown in Banks (1901), this species does not belong to Oedothorax s.s.. Therefore, we propose its transfer back to Tmeticus, the genus to which this species was originally assigned. Its taxonomic position within Erigoninae remains unknown., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 536-537, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Banks N, 1901. Some Arachnida from New Mexico. Proceedings of the Academy of Natural Sciences of Philadelphia 53: 568 - 597.","Banks N, 1910. Catalogue of Nearctic spiders. Bulletin, United States National Museum 72: 1 - 80.","Petrunkevitch A. 1911. A synonymic index-catalogue of spiders of North, Central and South America with all adjacent islands, Greenland, Bermuda, West Indies, Terra del Fuego, Galapagos, etc. Bulletin of the American Museum of Natural History 29: 1 - 791.","Roewer CF. 1942. Katalog der Araneae von 1758 bis 1940. Bremen: Natura, Buchhandlung fur Naturkunde und exakte Wissenschaften Paul Budy.","Buckle D, Carroll D, Crawford R, Roth V. 2001. Linyphiidae and Pimoidae of America north of Mexico: checklist, synonymy, and literature. Fabreries, Supplement 10: 89 - 191.","Emerton JH. 1909. Supplement to the New England spiders. Transactions of the Connecticut Academy of Arts and Sciences 14: 171 - 236."]}

CALLITRICHIA CONVECTOR (TANASEVITCH, 2014) COMB. NOV. (FIGS 19G, 22G, 23, 24G) Oedothorax convector Tanasevitch, 2014a: 408, figs 64–71 (Dm). Type material: Holotype: Thailand: Chiang Mai Province, Chomthong District, Doi Inthanon, 1780 m, ♂ 3.iii.1987, leg. P. Schwendinger (MHNG, examined). Diagnosis: Males: This species can re recognized by the shape of prosomal modification with the inter-AME-PME groove and PME lobe; it can be discriminated from Oedothorax s.s., Mitrager and other Callitrichia species by its palpal features including the presence of tibial retrolateral apophysis with setae (no setae in Mitrager), the absence of tibial prolateral trichobothrium, the shape of paracymbium (distal clasp particularly long and slender), and the radix without tail piece. Description: Male (holotype, MHN): Total length: 1.68. Prosoma: 0.75 long, 0.55 wide, PME on frontal side of elevated region, short curved setae grouped at both sides of dorsal surface of prosomal elevation behind PME, inter-PME-PLE groove, three pairs of strong setae at groove lower margin, no setae inside groove (Fig. 19G). Eyes: AME-AME: 0.03, AME width: 0.05, AME-ALE: 0.01, ALE width: 0.05, ALE-PLE: 0.01, PLE width: 0.05, PLE-PME: 0.10, PME width: 0.05, PME-PME: 0.18. Clypeus: not hirsute, one sub-AME seta, particularly long. Chelicerae: stridulatory striae rows wide and evenly spaced (Fig. 22G). Sternum: 0.46 long, 0.5 wide. Legs: dorsal proximal macroseta on tibia IV 3.13 times diameter of tibia; Tm I: 0.81. Pedipalp: TPA narrow, slightly curved; TRA extended anteriorly, covering PC base from retrolateral view; PC long, distal-setae-bearing middle part enlarged, distal clasp slender and long (Fig. 23A–C); T without papillae; PT slender and long, with scale-like papillae; TS long, with small papillae; MSA absent; DSA short and wide (Fig. 23H); EM short, without papillae, not exceeding ARP (Fig. 23E); R without TP, scaly region distributed from ARP to VRP; LER present; E prolaterally spiral, between ARP and LER (Fig. 23E, F, G). Opisthosoma: dorsal pattern see Fig. 24G; PMS with mAP, two AC; PLS with one FL, 3+ AC (Fig. 23I–K). Female: Unknown Distribution: Only known from the type locality in Thailand. Habitat: No data.

OEDOTHORAX AGRESTIS (BLACKWALL, 1853) (FIGS 5, 7V, 8H, 9H; SUPPORTING INFORMATION, FIG. S1G) Neriene agrestis Blackwall, 1853: 23 (Dmf). Neriene agrestis Blackwall, 1864: 276, pl. 19, fig. 190, pl. 22, fig. D (mf). Neriene agrestis O. Pickard-Cambridge, 1882: 4, pl. 1, fig. 2B. Gongylidium agreste Simon, 1884: 494, figs 280–282 (mf). Gongylidium agrestis Becker, 1896: 89, pl. 9, fig. 9 (mf). Kulczynskiellum agreste Bösenberg, 1902: 169, pl. 15, fig. 228 (mf). Oedothorax agrestis de Lessert, 1910: 193. Stylothorax agrestis Reimoser, 1919: 72. Oedothorax agrestis Simon, 1926: 453, 523, fig. 784 (mf). Oedothorax agrestis Denis, 1947: 140, figs 2D, 6G, 7H, 8H, 9B, 10C, 11B (mf). Oedothorax agrestis Locket & Millidge, 1953: 241, figs 145C, 146D, 147C, D (mf). Oedothorax agrestis Wiehle, 1960a: 445, figs 817–826 (mf). Oedothorax agrestis Merrett, 1963: 386, fig. 46A–F (m). Oedothorax agrestis Tystshenko, 1971: 251, fig. 827 (f). Oedothorax agrestis Miller, 1971: 262, pl. LIV, figs 17–19 (mf). Oedothorax agrestis Palmgren, 1976: 87, figs 7.11, 13–17 (mf). Oedothorax agrestis Růžička, 1978: 195, fig. 8E, F (f). Oedothorax agrestis Müller, 1983: 64, fig. 2a–c (m). Oedothorax agrestis Bosmans, 1985: 65, figs 16, 21, 33 (m). Oedothorax agrestis Roberts, 1987: 57, figs 22D, 23B (mf). Oedothorax agrestis Heimer & Nentwig, 1991: 224, fig. 606 (mf). Oedothorax agrestis Aakra 2000: 81, fig. 3A–E (mf). Type material: Not examined. According to the original description of Blackwall (1853), individuals were collected in Oakland, United Kingdom, among herbage and under stones in pastures near woods. According to O. Pickard-Cambridge (1882), Blackwall lost all type material of Oe. agrestis. The original and subsequent descriptions, nevertheless, make the identification of this species unequivocal. Examined material: England: London, Beckenham, 1♂ 2.ii.1958, coll. D. J. Clark, det. G. H. Locket (NHM); Dorset, River Allen, Noritumb, c. 650 m, 3♂ 1.ix.1965, under stones by river (AMNH, No. 3080); Cumbria, Drumburgh, salt marsh, 1♀ 15.viii.1965 (AMNH). Wales: Merionethshire, Dolgellau, Cymmel Abbey, 1♀ 6.vi.56, coll. and det. D. J. Clark (NHM). Scotland: Perth, River Tay, under stone by loch, 1♂ 2.ix.1965 (AMNH No. 3083). Switzerland: Trius, 1♀ det. Schenkel (AMNH). Sweden: Öland, Stora Rör, 1♀ 15.vii.1931, leg. Nielsen, 16.ii.1932, det. Seheukel (ZMUC 00011739). Diagnosis: Males: The lack of male prosomal modifications and the presence of an embolic base protuberance distinguishes the males of this species from the ‘ gibbosus -like species group’ (Clade 22). Oedothorax agrestis is further distinguished from Oe. paludigena by the the palpal tibia prolateral apophysis basal thorn, and from Oe. fuscus and Oe. tingitanus by the lack of a bifurcated protegulum. Females: Can be distinguished from other species by the epigynal configuration and number of sub-AME setae (two; one in Oe. gibbosus, Oe. trilobatus, Oe. apicatus, Oe. retusus, Oe. gibbifer and Oe. paludigena). Distinguished from Oe. apicatus by the more anteriorly extended copulatory ducts; from Oe. fuscus by the less sclerotized epigyne, the dorsal plate bordered by thinner dark stripes and the copulatory ducts less extended anteriorly; from Oe. gibbosus, Oe. retusus, Oe. gibbifer and Oe. meridionalis by the more curved and less convergent ventral plate borders; from Oe. tingitanus by the lateral copulatory openings (Fig. 5E, G; mesal in Oe. tingitanus, Fig. 11F); from Oe. trilobatus by the more anteriorly located copulatory openings. Description: Male (London): Total length: 2.54. Prosoma: 0.95 long, 0.80 wide, postocular region slightly elevated (Fig. 7V). Eyes: AME-AME: 0.03, AME width: 0.06, AME-ALE: 0.017, ALE width: 0.08, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.05, PME width: 0.08, PME-PME: 0.06. Clypeus: not hirsute, two sub-AME setae. Sternum: 0.57 long, 0.59 wide. Chelicerae: mastidia absent; stridulatory striae scaly, rows widely and evenly spaced (Fig. 8H). Legs: dorsal proximal macroseta on tibia I, III and IV 0.96, 1.68 and 1.90 times diameter of tibia, respectively; Tm I: 0.64. Pedipalp: TPA short, rod-like, distal part scaly, with several small denticles; basal thorn short, pointed antero-ventrally; PC distal setae at median position (Fig. 5A); T papillae scale-like, PT with long papillae; TS short, without papillae (Fig. 5D); DSA tip narrow, pointed (Fig. 5A); EM median-long, cylindrical, distally oriented, with long papillae at tip; ARP prolaterally spiral; TP with several small protuberances; E not broadened at basal part (Fig. 5B). Opisthosoma: brown, evenly coloured (Fig. 9H). Female (Wales): Total length: 3.19. Prosoma: 1.20 long, 0.91 wide. Eyes: AME-AME: 0.03, AME width: 0.07, AME-ALE: 0.02, ALE width: 0.10, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.04, PME width: 0.08, PME- PME: 0.06. Sternum: 0.68 long; 0.65 wide. Legs: dorsal proximal macroseta on tibia I, II and III 1,28, 1.47 and 1.76 times diameter of tibia, respectively; Tm I: 0.63. Chelicerae: stridulatory striae scaly, rows widely and evenly spaced. Epigyne: Clade 13 characteristic morphology, borders between dorsal and ventral plates parallel, copulatory duct short (Fig. 5E–H). Variation: The measurements are based on examined material. Males ( N = 6, means in parentheses): Total length 2.21–2.54 (2.3). Prosoma: 0.95–0.99 (0.96) long, 0.76– 0.81 (0.79) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.96–1.13 (1.06), 0.94–1.51 (1.24, N = 5), 1.66–1.88 (1.75) and 1.84–2.14 (1.96) times diameter of tibia, respectively; Tm I: 0.55 –0.70 (0.64). Females ( N = 4, means in parentheses): Total length 2.89–3.65 (3.18). Prosoma: 1.02–1.30 (1.17) long, 0.79– 0.1.01 (0.90) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.28–1.55 (1.39, N = 3), 1.2–1.67 (1.44), 1.34–1.94 (1.66) and 1.50–1.85 (1.73, N = 3) times diameter of tibia, respectively; Tm I: 0.60–0.74 (0.65). Distribution: Europe. Habitat: Among herbage and under stones in pastures; in swamp litter; stony lakeshores., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 434-435, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Blackwall J. 1853. Descriptions of some newly discovered species of Araneidea. Annals and Magazine of Natural History 11: 14 - 25.","Blackwall J. 1864. A history of the spiders of Great Britain and Ireland. Ray Society 2: 175 - 384.","Pickard-Cambridge O. 1882. Notes on British spiders, with descriptions of three new species and characters of a new genus. Annals and Magazine of Natural History (5) 9: 1 - 17.","Simon E. 1884. Les arachnides de France. Tome cinquieme, deuxieme et troisieme partie. Roret Paris, 180 - 885.","Becker L. 1896. Les arachnides de Belgique, deuxieme et troisieme parties. Annales du Musee Royal d'Histoire Naturelle de Belgique 12: 1 - 127.","Bosenberg W. 1902. Die Spinnen Deutschlands. II - IV. Zoologica (Stuttgart) 14: 97 - 384.","De Lessert R. 1910. Catalogue des invertebres de la Suisse. Fasc. 3, Araignees. Geneva: Musee d'histoire naturelle de Geneve.","Reimoser E. 1919. Katolog der echten Spinnen (Araneae) des Palaarktischen Gebietes. Abhandlungen der Zoologisch- Botanischen Gesellschaft in Wien 10: 1 - 280.","Simon E. 1926. Les arachnides de France. Synopsis generale et catalogue des especes francaises de l'ordre des Araneae. Tome VI. 2 e partie. Paris: Roret.","Denis J. 1947. Notes sur les erigonides. XI. Les especes francaises du genre Oedothorax Bertkau. Bulletin de la Societe d'Histoire Naturelle de Toulouse 82: 131 - 158.","Locket GH, Millidge AF. 1953. British spiders, Vol. II. London: Ray Society.","Wiehle H. 1960 a. Spinnentiere oder Arachnoidea, XI: Micryphantidae-Zwergspinnen. Tierwelt Deutschlands 47: 1 - 620.","Merrett P. 1963. The palpus of male spiders of the family Linyphidae. Proceedings of the Zoological Society of London 140: 347 - 467.","Tystshenko VP. 1971. Opredelitel' paukov evropejskoj casti SSSR. Leningrad: Nauka.","Miller F. 1971. Pavouci-Araneida. Klic zvireny CSSR 4: 51 - 306.","Palmgren P. 1976. Die Spinnenfauna Finnlands und Ostfennoskandiens. VII. Linyphiidae 2. Fauna Fennica 29: 1 - 126.","Ruzicka V. 1978. Revision der diagnostischen Merkmale der Weibchen der tschechoslovakischen Arten der Gattung Oedothorax (Araneae: Micryphantidae). Vestnik Ceskoslovenske Zoologicke Spolecnosti v Praze 42: 195 - 208.","Muller H-G. 1983. Zur Vorkommen von Oedothorax agrestis (Blackwall) (Araneida, Linyphiidae) in Hessen. Hessische Faunistische Briefe 3: 64 - 67.","Bosmans R. 1985. Etudes sur les Linyphiidae nord-africains. II. Le genre Oedothorax Bertkau en Africa du Nord, avec une revision des caracteres diagnostiques des males des especes ouest-palearctiques. Biologisch Jaarboek Dodonaea 53: 58 - 75.","Roberts MJ. 1987. The spiders of Great Britain and Ireland, Vol. 2: Linyphiidae and check list. Colchester: Harley Books.","Heimer S, Nentwig W. 1991. Spinnen mitteleuropas: ein Bestimmungsbuch. Berlin: Paul Parey.","Aakra K. 2000. New records of spiders (Araneae) from Norway with notes on epigynal characters of Philodromus fuscomarginatus (De Geer) (Philodromidae) and Araneus sturmi (Hahn) (Araneidae). Norwegian Journal of Entomology 47: 77 - 88."]}

GONGYLIDIOIDES INSULANUS (PAIK, 1980) COMB. NOV. Oedothorax insulanus Paik, 1980: 162, figs 9–21 (Df). Oedothorax insulanus Namkung, 2002: 205, fig. 17.57A–b (f). Oedothorax insulanus Namkung, 2003: 207, fig. 17.57A–b (f). Gongylidioides kouqianensis Tu & Li, 2006: 59, fig. 5A–G (Dm), synon. nov.. Oedothorax insulanus Seo, 2011: 37, figs 11–16 (f, Dm). Type material: Oedothorax insulanus: Holotype: South Korea: Jeunlanam-Do So Heuksan-do Island, Hang-ri, ♀ 27.vii.1979, coll. S. R. Son (not examined). Paratype: South Korea: So Heuksan-do Isl, Hang-ri, 2♀ 27.vii.1979, coll. S. R. Son (not examined); same location, 1♀ 27.vii.1979, coll. T. H. Jo (not examined). Gongylidoides kouqianensis: Holotype and paratype: China: Jilin Province, Kouqian County, 2♂ 29.vi.1989, Institute of Zoology, Chinese Academy of Sciences in Beijing, China (Tu and Li, 2006; not examined). Non-type material: Gongylidoides kouqianensis: Gyeongsangbuk-do, Ulreung-gun, Hyeonpo, 10♂ 4♀ 24.viii.2006, coll. S. Y. Kim, collection of Department of Biology, Keimyung University (Seo, 2011; not examined). Diagnosis: Males: This species can be recognized among congeners by the combination of the following features: the lack of prominent prosomal modification; the broad, apically pointed palpal tibial retrolateral apophysis without a tooth on the inner surface; lamella characteristica extal tip horn-like, with two projections, inner one triangular in ventral view, dorsal one blunt (fig. 5F in Tu & Li, 2006). Females: Can be diagnosed by the half-octagonshaped dorsal plate of the epigyne (figs 16, 17 in Paik, 1980). Distribution: South Korea; China, Jilin province. Remarks: Tu & Li (2006) and Seo (2011) have provided informative drawings of three perspectives of the male palps of Gongylidioides kouqianensis and Oe. insulanus, respectively, and their resemblance strongly suggests that these two species are identical. Since Oe. insulanus was published earlier, it has priority over Gongylidioides kouqianensis. The palpal and epigynal morphology of Gongylidioides insulanus is in accordance with the generic description of Gongylidioides in Tu & Li (2006), and supports the placement of this species in Gongylidioides., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 535, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Paik KY. 1980. The spider fauna of Dae Heuksan-do Isl., So Heuksan-do Isl. and Hong-do Isl., Jeunlanam-do, Korea. Kyungpook Educational Forum Kyungpook National University 22: 153 - 173.","Namkung J. 2002. The spiders of Korea. Seoul: Kyo-Hak Publishing.","Namkung J. 2003. The spiders of Korea, 2 nd edn. Seoul: Kyo- Hak Publishing Co.","Tu LH, Li SQ. 2006. A review of Gongylidioides spiders (Araneae: Linyphiidae: Erigoninae) from China. Revue Suisse de Zoologie 113: 51 - 65.","Seo BK. 2011. Two species of the genus Oedothorax from Korea (Araneae: Linyphiidae). Quantitative Bio-Science 30: 35 - 39."]}

‘OEDOTHORAX’ UNCIGER TANASEVITCH, 2020 INCERTAE SEDIS Oedothorax unciger Tanasevitch, 2020b: 127, figs 1–3, 7–12 (Dm). Type material: Holotype: India: Meghalaya, Sohra, plateau, 1320 m a.s.l., ♂ 14–26.xii.2013, leg. K. P. Tomkovich (ZMMU). Paratype: 1♂ (ZMMU), together with holotype. Diagnosis: Males: This species is similar to many Oriental ‘ Oedothorax ’ species, but can be distinguished by its unique hook-shaped ventral radical process (see figs 8 and 12 in Tanasevitch 2020b). Females: unknown. Distribution: Only known from the type locality in India. Remarks: According to the drawings in Tanasevitch (2020), the embolic division of this species resembles those of ‘ Oe.’ kodaikanal, ‘ Oe.’ meghalaya, ‘ Oe.’ uncus, ‘ Oe.’ cunur, ‘ Oe.’ myanmar, ‘ Oe.’ sohra, ‘ Oe. ’ khasi, ‘ Oe. ’ bifoveatus and ‘ Oe.’ stylus and is, therefore, probably closely related to these species instead of Oedothorax s.s.., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 556, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2020 b. Two new Oedothorax Bertkau, 1883 from eastern India (Aranei: Linyphiidae). Arthropoda Selecta 29: 127 - 131."]}

OEDOTHORAX RETUSUS (WESTRING, 1851) (FIGS 7S, 8E, 9E, 16; SUPPORTING INFORMATION, FIG. S1E) Erigone retusa Westring, 1851: 41 (Dmf). Neriene elevate O. Pickard-Cambridge, 1862: 7966 (Dmf). Tmeticus foveolatus Menge, 1868: 186, pl. 35, fig. 86 (Dmf). Neriene retusa O. Pickard-Cambridge, 1873: 451. Gongylidium fuscum Simon, 1884: 478, figs 252–254 (mf, misidentified). Neriene retusa Chyzer & Kulczyński, 1894: 94, pl. 4, fig. 2 (mf). Kulczynskiellum retusum F.O. Pickard-Cambridge, 1895: 39. Gongylidium fuscum Becker, 1896: 82, pl. 9, fig. 5 (mf). Kulczynskiellum retusum Bösenberg, 1902: 170, pl. 15, fig. 230 (mf). Oedothorax retusus, de Lessert, 1910: 192. Kulczynskiellum retusum Fedotov, 1912: 454, pl. 8, fig. 2 (f). Stylothorax retusa, Dahl, 1912: 603. Oedothorax retusus Denis, 1947: 145, figs 6D, 7D, 8D, 9E, 10E, 11D (mf). Oedothorax retusus Vogelsanger, 1948: 53, fig. 9 (f). Oedothorax retusus Locket & Millidge, 1953: 241, figs 145D, E, 146E, 147E, F (mf). Oedothorax retusus Wiehle, 1960a: 440, figs 807–816 (mf). Oedothorax retusus Holm, 1962: 165, fig. 61C–d (m). Oedothorax retusus Tystshenko, 1971: 251, figs 820, 831 (mf). Oedothorax retusus Miller, 1971: 262, pl. LIV, figs 20–22 (f). Oedothorax retusus Palmgren, 1976: 88, figs 7.22, 8.1– 2, 8.4–6. Oedothorax retusus Růžička, 1978: 195, figs 8B, 9B (f). Oedothorax retusus Hu & Wang, 1982: 63, fig. II.1–4 (f). Oedothorax retusus Hu, 1984: 199, fig. 209.1–4 (f). Oedothorax retusus Bosmans, 1985: 65, figs 15, 20, 32 (m). Oedothorax retusus Roberts, 1987: 57, figs 22E, 23C (mf). Oedothorax retusus Heimer & Nentwig, 1991: 224, fig. 603 (mf). Oedothorax retusus Alderweireldt, 1992: 5, fig. 1c (f). Oedothorax retusus Zhao, 1993: 199, fig. 90a–c (f). Oedothorax retusus Uhl, Nessler & Schneider, 2010: 77, fig. 1E, F (f). Oedothorax retusus Uhl et al., 2014: 348, fig. 1A–F (mf). Oedothorax retusus Kunz, Witthuhn & Uhl, 2015: 279, figs 1A–h, 2A–j (mf). Oedothorax retusus Russell-Smith, 2016: 23, fig. 2 (f). Type material: No type designation nor illustration in Westring (1851). Subsequent studies do not mention the examination of types. Nevertheless, the unique palpal tibia shape and male prosomal modification illustrated in later descriptions suffice for an identification of this species. Examined material: Scotland: Tentsmuir, damp dune slack, 1♂ 2♀ 8.ix.1965 (No. 3114 AMNH). England: Yorkschire, Askham Bog, 1♀ 12.vii.53 (AMNH); New Forest, Matley Passage, sand pit, 1♀ 15.viii.1955 (AMNH); Oxford, Noke wood, 1♀ 20. x.1954. Switzerland: Basel (47° N, 8° E), 1♂ 1♀ (AMNH); Westring, 1♂ (AMNH); Trius, 1♀ det. Schenkel (received 1946), Schenkel collection (AMNH). Norway: 1♂ xiii.1962, N, Svartisen, M, J. O. 40 (AMNH); 2♀ xiii.1960, RO/RO/S J. A. L. O. (AMNH); 1♂ xiii.1962, N, Svartisen, M, J. O. 46 (AMNH). Germany: Greifswald, 1♂ 2014, coll. and det. S.-W. Lin. Data unrecognizable: 4♂ 1♀ (AMNH). Diagnosis: Males: This species is similar to Oe. apicatus and Oe. gibbifer, all three possess post-ocular hump and lateral sulci and pits, but this species can be distinguished from Oe. apicatus by not having the knob-like shape of post-ocular hump, and can be distinguished from Oe. gibbifer by the retrolaterally bent palpal tibial basal thorn, pointing prolaterally in the latter. Females: Can be distinguished from other species by the epigynal configuration and number of sub- AME setae (one; two in Oe. fuscus, Oe. agrestis, Oe. meridionalis and Oe. tingitanus). Distinguished from Oe. apicatus by the more convergent ventral plate borders; from Oe. gibbosus by the more curved ventral plate borders; from Oe. gibbifer by the wrinkles close to the posterior margin of the central area parallel to the margin; from Oe. paludigena by the narrower posterior margin of the dorsal plate; from Oe. trilobatus by the much shorter copulatory ducts. Description: Male (AMNH): Total length: 2.14. Prosoma: 0.91 long, 0.74 wide, postocular region elevated, with lateral sulci and pits (Fig. 7S). Eyes: AME-AME: 0.02, AME width: 0.05, AME-ALE: 0.03, ALE width: 0.06, ALE-PLE: 0.01, PLE width: 0.06, PLE-PME: 0.04, PME width: 0.06, PME-PME: 0.05. Clypeus: not hirsute, one sub- AME seta. Sternum: 0.57 long, 0.53 wide. Chelicerae: mastidia absent; stridulatory striae scaly, rows widely and evenly spaced (Fig. 8E). Legs: dorsal proximal macroseta on tibia I, II and IV 1.16, 1.16 and 1.87 times diameter of tibia, respectively; Tm I: 0.69. Pedipalp: TPA broad at base, triangular, distal part scaly; BT long, pointed retrolaterally; PC no recognizable distal setae group (Fig. 16A); T with scale-like papillae, PT with long papillae; TS short, with papillae (Fig. 16D); DSA tip round(Fig. 16A); EM short, cylindrical, proximally oriented; TP without small protuberances (Fig. 16B); E broad at basal part. Opisthosoma: brown, evenly coloured (Fig. 9E); spinnerets see Fig. 16H. Female (AMNH): Total length: 2.73. Prosoma: 1.07 long, 0.82 wide. Eyes: AME-AME: 0.02, AME width: 0.06, AME-ALE: 0.03, ALE width: 0.08, ALE-PLE: 0.00, PLE width: 0.08, PLE-PME: 0.04, PME width: 0.07, PME-PME: 0.07. Clypeus: not hirsute, one sub- AME seta. Sternum: 0.63 long; 0.62 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.24, 1.35, 1.72 and 1.82 times diameter of tibia, respectively; Tm I: 0.63. Chelicerae: stridulatory striae similar to male. Epigyne: Clade 13 characteristic morphology, borders between dorsal and ventral plates converging anteriorly, copulatory duct short (Fig. 16E–G). Opisthosoma: brown, evenly coloured. Variation: The measurements are based on examined material. Males (N = 10, means in parentheses): Total length 1.88–2.23 (2.06). Prosoma: 0.88–1.04 (0.93) long, 0.69– 0.81 (0.75) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.91–1.47 (1.21), 1.11–1.62 (1.25, N = 9), 1.35–2.24 (1.60, N = 9) and 1.42–2.38 (1.75, N = 8) times diameter of tibia, respectively; Tm I: 0.58– 0.69 (0.62). Females (N = 10, means in parentheses): Total length 2.29–3.02 (2.65). Prosoma: 1.01–1.20 (1.13) long, 0.80– 0.91 (0.87) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.24–1.79 (1.51), 1.35–1.74 (1.53), 1.47–2.25 (1.85) and 1.58–2.35 (1.98) times diameter of tibia, respectively; Tm I: 0.60–0.69 (0.64). Distribution: Europe, Turkey, Caucasus, Russia to Kazakhstan, China Habitat: In low vegetation or under stones close to water., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 447-448, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Westring N, 1851. Forteckning ofver de till narvarande tid Kande, i Sverige forekommande Spindlarter, utgorande ett antal af 253, deraf 132 aro nya for svenska Faunan. Goteborgs Kungliga Vetenskaps och Vitterhets Samhalles Handlingar 2: 25 - 62.","Pickard-Cambridge O. 1862. Description of ten new species of British spiders. The Zoologist 20: 7951 - 7968.","Menge A. 1868. Preussische Spinnen. II. Abtheilung. Schriften der Naturforschenden Gesellschaft in Danzig (N. F.) 2: 153 - 218.","Pickard-Cambridge O. 1873. On British spiders. Transactions of the Linnean Society of London 28: 433 - 458.","Simon E. 1884. Les arachnides de France. Tome cinquieme, deuxieme et troisieme partie. Roret Paris, 180 - 885.","Chyzer C, Kulczynski W. 1894. Araneae Hungariae. Tomus II. Budapest: Academia Scientarum Hungaricae.","Pickard-Cambridge FO. 1895. List of the Araneidea of Cumberland and Lake District. The Naturalist 29 - 48.","Becker L. 1896. Les arachnides de Belgique, deuxieme et troisieme parties. Annales du Musee Royal d'Histoire Naturelle de Belgique 12: 1 - 127.","Bosenberg W. 1902. Die Spinnen Deutschlands. II - IV. Zoologica (Stuttgart) 14: 97 - 384.","De Lessert R. 1910. Catalogue des invertebres de la Suisse. Fasc. 3, Araignees. Geneva: Musee d'histoire naturelle de Geneve.","Fedotov D. 1912. K faounie Paoukow Mourmana i Nowoi Zemli. Contribution a la faune des araignees de la cote Murmane et de Novaja Zemlja. Lejiegod Zoologicheskii Instituta Muzeya Akademii Nauk SSSR St. Petersburg 16: 449 - 474.","Dahl F. 1912. Uber die Fauna des Plagefenn-Gebietes. In: Conwentz H, ed. Das Plagefenn bei Chorin. Berlin, 339 - 638, Araneae, 575 - 622.","Denis J. 1947. Notes sur les erigonides. XI. Les especes francaises du genre Oedothorax Bertkau. Bulletin de la Societe d'Histoire Naturelle de Toulouse 82: 131 - 158.","Vogelsanger T. 1948. Beitrag zur Kenntnis der Spinnenfauna des Kantons Graubunden. Mitteilungen der Naturforschenden Gesellschaft Schaffhausen 22: 33 - 72.","Locket GH, Millidge AF. 1953. British spiders, Vol. II. London: Ray Society.","Wiehle H. 1960 a. Spinnentiere oder Arachnoidea, XI: Micryphantidae-Zwergspinnen. Tierwelt Deutschlands 47: 1 - 620.","Holm A. 1962. The spider fauna of the East African mountains. Part I: Fam. Erigonidae. Zoologiska Bidrag fran Uppsala 35: 19 - 204.","Tystshenko VP. 1971. Opredelitel' paukov evropejskoj casti SSSR. Leningrad: Nauka.","Miller F. 1971. Pavouci-Araneida. Klic zvireny CSSR 4: 51 - 306.","Palmgren P. 1976. Die Spinnenfauna Finnlands und Ostfennoskandiens. VII. Linyphiidae 2. Fauna Fennica 29: 1 - 126.","Ruzicka V. 1978. Revision der diagnostischen Merkmale der Weibchen der tschechoslovakischen Arten der Gattung Oedothorax (Araneae: Micryphantidae). Vestnik Ceskoslovenske Zoologicke Spolecnosti v Praze 42: 195 - 208.","Hu YJ, Wang HZ. 1982. Description of three species of dwarf spiders from cotton fields in Xinxiang. Journal of Hunan Teachers College (nat. sci. ed.) 1982: 63 - 66.","Hu JL. 1984. The Chinese spiders collected from the fields and the forests. Tianjin Press of Science and Techniques.","Bosmans R. 1985. Etudes sur les Linyphiidae nord-africains. II. Le genre Oedothorax Bertkau en Africa du Nord, avec une revision des caracteres diagnostiques des males des especes ouest-palearctiques. Biologisch Jaarboek Dodonaea 53: 58 - 75.","Roberts MJ. 1987. The spiders of Great Britain and Ireland, Vol. 2: Linyphiidae and check list. Colchester: Harley Books.","Heimer S, Nentwig W. 1991. Spinnen mitteleuropas: ein Bestimmungsbuch. Berlin: Paul Parey.","Alderweireldt M, 1992. Determinatieproblematiek van de zustersoorten van het genus Oedothorax (Araneae, Linyphiidae). Nieuwsbrief van de Belgische Arachnologische Vereniging 7: 4 - 8.","Zhao JZ. 1993. Spiders in the cotton fields in China. Wuhan: Wuhan Publishing House.","Uhl G, Nessler SH, Schneider JM. 2010. Securing paternity in spiders? A review on occurrence and effects of mating plugs and male genital mutilation. Genetica 138: 75.","Uhl G, Kunz K, Vocking O, Lipke E. 2014. A spider mating plug: origin and constraints of production. Biological Journal of the Linnean Society 113: 345 - 354.","Kunz K, Witthuhn M, Uhl G. 2015. Paired and complex copulatory organs: do they really impede flexible use? Journal of Zoology 297: 278 - 285.","Russell-Smith A. 2016. Identification of females of British Oedothorax species. Newsletter of the British Arachnological Society 137: 22 - 24."]}

‘ OEDOTHORAX’ SOHRA TANASEVITCH, 2020 INCERTAE SEDIS Oedothorax sohra Tanasevitch, 2020b: 129, figs 4–6, 13–18 (Dm). Type material: Holotype: India: Meghalaya, Sohra, plateau, 1320 m a.s.l., 14–26.xii.2013, leg. K. P. Tomkovich (ZMMU). Diagnosis: Males: The palpal morphology of this species is similar to many ‘ Oedothorax ’ incertae sedis species from the Oriental Region, including ‘ Oe. ’ kodaikanal, ‘ Oe. ’ meghalaya, ‘ Oe.’ uncus, ‘ Oe. ’ cunur, ‘ Oe. ’ stylus, ‘ Oe. ’ myanmar, ‘ Oe. ’ khasi, ‘ Oe. ’ bifoveatus and ‘ Oe.’ unciger. It can be distinguished from ‘ Oe.’ cunur and ‘ Oe.’ stylus by the much longer, dorsally directed palpal tibial prolateral apophysis with distal dentation; from ‘ Oe.’ kodaikanal, ‘ Oe.’ meghalaya, ‘ Oe.’ unciger, ‘ Oe. ’ khasi, ‘ Oe. ’ bifoveatus and ‘ Oe.’ myanmar by the much longer and distally curved ventral radical apophysis; from ‘ Oe.’ uncus by the shorter and less curved ventral radical apophysis. Females: Unknown. Distribution: Only known from the type locality in India. Remarks: From the original species description and figures (Tanasevitch 2020b: 130, figs 13–18) this species is overall most similar to ‘ Oe.’ uncus, both in the lack of prosomal modification and the male palpal morphology, including the extensive papillae distribution on the protegulum, the long and curved ventral radical apophysis and the long, vertically pointed palpal tibial prolateral apophysis with distal dentation, etc. Therefore, this species is probably most closely related to ‘ Oe.’ uncus instead of Oedothorax s.s.., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 553-554, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2020 b. Two new Oedothorax Bertkau, 1883 from eastern India (Aranei: Linyphiidae). Arthropoda Selecta 29: 127 - 131."]}

‘OEDOTHORAX’ COLLINUS MA & ZHU, 1991 INCERTAE SEDIS Oedothorax collinus Ma & Zhu, 1991: 27, figs 1–9 (Dmf). Oedothorax collinus Song et al., 1999: 199, fig. 113F–G, N (mf). Oedothorax collinus Yin et al., 2012: 542, fig. 258a–e (mf). Type material: Holotype: China: Hubei Province, Shennongjia Forestry District, Dayanwu, 1600 m, ♂ 23.vi.1986, leg. Jiuchun Gao (not examined); Allotype: ♀, same data (not examined). Paratype: 1♂ 4♀, same data (not examined); Hubei Province, Shennongjia Forestry District, Hongping, 1610m, 1♀ 1.viii.1986, leg. Jiuchun Gao (not examined). Deposited in the Department of Cellular Biology, Norman Bethune University of Medical Science, Changchun, China. Non-type material: China: Hunan Province: Sangzhi County, Mt. Tianping, 1♀ 1♂ 18.vi.1981, leg. Tong Xin Wang; Shimen County, Mt. Huping, 2♂ 2♀ 3.viii.2002, leg. Tang Guo. Most of the specimens are deposited in animal specimens collection of Hunan Normal University, School of Life Science (Yin et al. 2012) (not examined). Diagnosis: Males: This species can be identified by the presence of a long, slender palpal tibial retrolateral apophysis and a shorter prolateral apophysis (figs 3–6 in Ma & Zhu, 1991), and the highly elevated post-ocular area with lateral pits (fig. 2 in Ma & Zhu, 1991). Females: General epigynal morphology similar to that of Oedothorax, but the area anterior to the copulatory openings is elevated (fig. 8 in Ma & Zhu, 1991), which distinguishes it from all species examined in the current study. Distribution: China: Hunan Province, Hubei Province. Remarks: As inferred from the drawings of Ma & Zhu (1991) and Yin et al. (2012), this species has a long, slender palpal tibial retrolateral apophysis, an enlarged part of the cymbium above the paracymbial base, a bifurcated paracymbial tip and an embolic division extremely different from the newly delimited Oedothorax in the present study. We suspect that this species does not belong to Oedothorax s.s.. Since all specimens previously deposited in the Department of Cellular Biology, Norman Bethune University of Medical Science are lost (Shuqiang Li, Beijing, personal communication, 2017), and generic assignment based on the descriptions of this species alone is not feasible, the taxonomic state of this species remains dubious., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 543, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Ma XL, Zhu CD. 1991. One new species of the spider genus Oedothorax from China (Araneae: Linyphiidae: Erigoninae). Acta Zootaxonomica Sinica 16: 27 - 29.","Song DX, Zhu MS, Chen J. 1999. The spiders of China. Shijiazhuang: Hebei University of Science and Techology Publishing House."]}

MITRAGER ASSUETA (TANASEVITCH, 1998) COMB. NOV. (FIGS 38R, 39R, 40S, 42, 43B; SUPPORTING INFORMATION, FIG. S4E) Oedothorax assuetus Tanasevitch, 1998a: 431, figs 1–5 (Dm). Type material: Holotype: Nepal: Kathmandu, Godawari, foot of Phulchoki Mt., 1700 m, ♂ 19.iii.1980, leg. Martens & A. Ausobsky (SMF 38848, examined). Paratypes: Nepal: same locality, together with holotype, 1♂, leg. J. Martens & A. Ausobsky (body: SMF 38852, palp: SMF 38841, examined). Diagnosis: Males: A depression between protegulum and tegulum is unique for this species. Further distinguished from most Mitrager species by a prosoma with a hump comprising the PME, and an ocular region bearing dense setae. Distinguished from M. falcifer, M. falciferoides and ‘ Oe. ’ meghalaya incertae sedis by the smaller hump elevation; from M. malearmata by the longer spike prolateral to the palpal tibia prolateral apophysis; and from M. modesta, M. lopchu and M. rustica by the smaller body size. Description: Male (holotype): Total length: 2.40. Prosoma: 1.01 long, 0.77 wide, PME- and postocular region slightly elevated, with one strong seta at peak of elevation pointing forwards, interocular region with strong setae pointing upwards (Figs 38R, 43B). Eyes: AME- AME: 0.04, AME width: 0.04, AME-ALE: 0.03, ALE width: 0.08, ALE-PLE: 0, PLE width: 0.07, PLE-PME: 0.07, PME width: 0.07, PME-PME: 0.11. Clypeus: not hirsute. Sternum: 0.59 long, 0.57 wide. Chelicerae: stridulatory striae imbricated, rows widely and evenly spaced (Fig. 39R). Legs: Tm I: 0.56. Pedipalp: patella prolateral proximal vertical macrosetae present; tibia with one prolateral, two retrolateral trichobothria; TPS scaly, retrolaterally pointed; TPA absent; TRA bent retrolaterally; PC short, base not visible from dorsal view, distal setae close to distal clasp, distal clasp without striae, directed retrolaterally (Fig. 42A); T without papillae; PT with long papillae, a depression between T and PT creates a discontinuous appearance from lateral view (Fig. 42D); MSA present (Fig. 42D); DSA not pointed, not retrolaterally curved (Fig. 42A); EM flat, anterior margin without obvious papillae, exceeds ARP (Fig. 42D); ARP pointed, angled at tip; LER with striae on distal margin, extended dorsal to E; VRP absent; TP round at tip, ventro-posterior area with round extension (Fig. 42B); E retrolaterally spiral, anterior margin at base slightly wavy (Fig. 42B). Opisthosoma: dorsal pattern see Fig. 40S; PMS with mAP, AC absent; PLS with triad, one AC (Fig. 42E). Male (paratype): Total length: 2.39. Prosoma: 1.05 long, 0.86 wide. Legs: Tm I: 0.59., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 491, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 1998 a. New Oedothorax Bertkau, 1883, from Nepal (Arachnida, Araneae, Linyphiidae). Bonner Zoologische Beitraege 47: 429 - 441."]}

OEDOTHORAX APICATUS (BLACKWALL, 1850) (FIGS 7R, 8D, 9D, 12; SUPPORTING INFORMATION, FIG. S1D) Neriene apicata Blackwall, 1850: 339 (Dm). Erigone gibbicollis Westring, 1851: 41 (Dm). Micryphantes tuberculatus Grube, 1859: 469 (Dm). Erigone gibbicollis Westring, 1861: 223 (Df). Neriene apicata Blackwall, 1864: 269, pl. 18, f, 183 (m, Df). Micryphantes gibbus Ohlert, 1867: 65 (Dm). Phalops gibbicollis Menge, 1871: 220, pl. 43, fig. 119 (mf). Erigone gibbicollis Thorell, 1871: 112. Erigone apicata Dahl, 1883: 46. Gongylidium apicatum Simon, 1884: 487, figs 267–269 (mf). Neriene apicata Chyzer & Kulczyński, 1894: 93, pl. 4, fig. 1 (mf). Neriene apicata Simon, 1894: 666, fig. 719 (m). Kulczynskiellum apicatum F.O. Pickard-Cambridge, 1895: 39. Gongylidium apicatum Becker, 1896: 86, pl. 11, fig. 2 (mf). Kulczynskiellum apicatum Bösenberg, 1902: 169, pl. 15, fig. 227 (mf). Oedothorax apicatus de Lessert, 1910: 191. Stylothorax apicata Dahl, 1912: 603. Oedothorax apicatus Denis, 1947: 145, figs 2A, 3–5, 6F, 7C, 8C, 9C, 10D, 11C (mf). Oedothorax apicatus Locket & Millidge, 1953: 241, figs 145F, 146F, 147G, H (mf). Oedothorax apicatus Wiehle, 1960a: 437, figs 798–806 (mf). Oedothorax apicatus Wiehle, 1960b: 477, fig. 22a, b (mf). Oedothorax apicatus Tystshenko, 1971: 251, fig. 830 (f). Oedothorax apicatus Miller, 1971: 262, pl. LIV, figs 7–9 (f). Oedothorax apicatus Palmgren, 1976: figs 88, 7.23, 8.3, 8.7, 8.8 (m). Oedothorax apicatus Millidge, 1977: 11, fig. 24 (m). Oedothorax apicatus Růžička, 1978: 195, figs 8A, 9A (f). Oedothorax apicatus Hu & Wang, 1982: 63, fig. I.1–4 (m). Oedothorax apicatus Hu, 1984: 196, fig. 206.1–4 (mf). Oedothorax apicatus Bosmans, 1985: 65, figs 14, 19, 37 (m). Oedothorax apicatus Song, 1987: 153, fig. 114 (mf). Oedothorax apicatus Tanasevitch, 1987: 355 Oedothorax apicatus Roberts, 1987: 58, figs 22F, 23D (mf). Oedothorax apicatus Hu & Wu, 1989: 174, fig. 144.1–6 (mf). Oedothorax apicatus Tanasevitch, 1990: 102, figs 24.61, 28.4 (f). Oedothorax apicatus Chen & Gao, 1990: 111, fig. 139a, b (mf). Oedothorax apicatus Heimer & Nentwig, 1991: 224, fig. 602 (mf). Oedothorax apicatus Alderweireldt, 1992: 5, fig. 1B (f). Oedothorax apicatus Zhao, 1993: 191, fig. 86a–d (mf). Oedothorax apicatus Wunderlich, 1995: 473, fig. 7 (m). Oedothorax apicatus Song, Zhu & Chen, 1999: 199, fig. 113D, E, H (mf). Oedothorax apicatus Hu, 2001: 550, fig. 370.1–3 (f). Oedothorax apicatus Russell-Smith, 2016: 23, fig. 3 (f). Type material: Not examined. Blackwall (1850) collected one adult male on a rail in Oakland, England, in February 1850, but provided no deposition data. The original and subsequent descriptions, nevertheless, make the identification of this species unequivocal. Examined material: Scotland: Perth, River Tay, drift litter, c. 400 m, 1♂ 5♀ 2.ix.1965 (AMNH No. 3085). England: Essex, Crabknowe Spit, tide litter, 1♀ 18.iv.1961 (AMNH No. 3059). Sweden: Gotland, Ljugarn, 1♂ 1♀, det. Schenkel (AMNH). Germany: Greifswald, 2♂ 2014 (ZIMG); Hessen, Bad Hersfeld, Obersberg, soil avalanche, pitfall trap, 5♂ 18.ix–18. Xi.1980, leg. V. Puthz, det. S.-W. Lin, 17.xi.2015 (SMF). China: Xinjiang, 1♂ 3♀ 4.vii.1991 (Institute of Zoology, Chinese Academy of Science, IZCSA-Ar 20637). Diagnosis: Males: This species resembles Oe. retusus and Oe. gibbifer in the post-ocular hump and lateral sulci and pits, but it can be distinguished from the latter two by the knob-like shape of the post-ocular hump and its longer palpal tibia prolateral apophysis. Females: Can be distinguished from other species by the epigynal configuration and the number of sub- AME setae (one; two in Oe. fuscus, Oe. agrestis, Oe. meridionalis and Oe. tingitanus). Distinguished from Oe. agrestis by the larger spermathecae; from Oe. gibbifer, Oe. retusus and Oe. gibbosus by the more curved and the less convergent ventral plate borders; from Oe. trilobatus and Oe. meridionalis by the much more anteriorly located copulatory openings; from Oe. tingitanus by the lateral copulatory openings (Fig. 12F, in comparison to mesal in the latter species, Fig. 11F); from Oe. paludigena by the anteriorly less convergent ventral plate borders (anteriorly more convergent in Oe. paludigena, Fig. 13F). Description: Male (ZIMG): Total length: 2.11. Prosoma: 0.91 long, 0.73 wide, postocular region elevated, with knob-like hump and lateral sulci and pits (Fig. 7R). Eyes: AME- AME: 0.03, AME width: 0.05, AME-ALE: 0.04, ALE width: 0.07, ALE-PLE: 0.01, PLE width: 0.07, PLE-PME: 0.04, PME width: 0.05, PME-PME: 0.10. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.55 long, 0.55 wide. Chelicerae: mastidia absent; stridulatory striae scale-like, rows widely and evenly spaced (Fig. 8D). Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.41, 0.42, 0.80 and 0.89 times diameter of tibia, respectively; Tm I: 0.67. Pedipalp: TPA rod-like, distal part scaly; BT long, pointed retrolaterally; PC distal setae adjacent to basal setae (Fig. 12A); T with scale-like papillae, PT with long papillae; TS short, with papillae (Fig. 12D); DSA tip broad, margin smooth (Fig. 12A); EM short, cylindrical, proximally oriented; TP without small protuberances; E broad at basal part (Fig. 12B). Opisthosoma: brown, evenly coloured (Fig. 9D). Female (ZIMG): Total length: 2.95. Prosoma: 1.23 long, 1.03 wide. Eyes:AME-AME:0.22,AME width: 0.76, AME- ALE: 0.36, ALE width: 0.10, ALE-PLE: 0.01, PLE width: 0.09, PLE-PME: 0.06, PME width: 0.08, PME-PME: 0.09. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.78 long; 0.74 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.29, 1.30, 1.49 and 1.67 times diameter of tibia, respectively; Tm I: 0.67. Chelicerae: stridulatory striae scaly, widely and evenly spaced. Epigyne: Clade 13 characteristic morphology, borders between dorsal and ventral plates parallel, copulatory duct short (Fig. 12F–H). Opisthosoma: brown, evenly coloured. Variation: The measurements are based on examined material. Males ( N = 10, means in parentheses): Total length 1.30–2.32 (2.07). Prosoma: 0.87–1.04 (0.93) long, 0.71– 0.86 (0.76) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.34–0.74 (0.51), 0.39–0.59 (0.52, N = 8), 0.61–0.91 (0.74) and 0.78–1.15 (0.91, N = 8) times diameter of tibia, respectively; Tm I: 0.61–0.67 (0.63). Females ( N = 10, means in parentheses): Total length 2.51–3.37 (2.96). Prosoma: 0.98–1.35 (1.19) long, 0.84– 1.03 (0.95) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.11–1.38 (1.27), 1.09–1.68 (1.32), 0.73–1.82 (1.39) and 1.27–1.79 (1.52) times diameter of tibia, respectively; Tm I: 0.61–0.70 (0.66). Distribution: Europe, Turkey, Caucasus, Russia to Central Asia, China. Habitat: In moss, grass and sparse vegetation., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 435-440, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Blackwall J. 1850. Descriptions of some newly discovered species and characters of a new genus of Araneida. Annals and Magazine of Natural History (2) 6: 336 - 344.","Westring N, 1851. Forteckning ofver de till narvarande tid Kande, i Sverige forekommande Spindlarter, utgorande ett antal af 253, deraf 132 aro nya for svenska Faunan. Goteborgs Kungliga Vetenskaps och Vitterhets Samhalles Handlingar 2: 25 - 62.","Grube AE. 1859. Verzeichniss der Arachnoiden Liv., Kur und Ehstlands. Archiv fur die Naturkunde Liv-, Ehst- und Kurlands 1: 415 - 486.","Westring N, 1861. Araneae svecicae. Goteborgs Kungliga Vetenskaps och Vitterhets Samhalles Handlingar 7: 1 - 615.","Blackwall J. 1864. A history of the spiders of Great Britain and Ireland. Ray Society 2: 175 - 384.","Ohlert E. 1867. Die Araneiden oder echten Spinnen der Provinz Preussen. Leipzig: Engelmann.","Menge A. 1871. Preussische Spinnen. IV. Abtheilung. Schriften der Naturforschenden Gesellschaft in Danzig, (N. F.) 2: 265 - 296.","Thorell T. 1871. Remarks on synonyms of European spiders. Part II. Uppsala: C. J. Lundstrom.","Dahl F. 1883. Analytische Bearbeitung der Spinnen Norddeutschlands mit einer anatomisch-biologischen Einleitung. Schriften des Naturwissenschaftlichen Vereins fur Schleswig-Holstein 5: 13 - 88.","Simon E. 1884. Les arachnides de France. Tome cinquieme, deuxieme et troisieme partie. Roret Paris, 180 - 885.","Chyzer C, Kulczynski W. 1894. Araneae Hungariae. Tomus II. Budapest: Academia Scientarum Hungaricae.","Simon E. 1894. Histoire naturelle des araignees, Vol. 1. Paris: Roret, 489 - 760.","Pickard-Cambridge FO. 1895. List of the Araneidea of Cumberland and Lake District. The Naturalist 29 - 48.","Becker L. 1896. Les arachnides de Belgique, deuxieme et troisieme parties. Annales du Musee Royal d'Histoire Naturelle de Belgique 12: 1 - 127.","Bosenberg W. 1902. Die Spinnen Deutschlands. II - IV. Zoologica (Stuttgart) 14: 97 - 384.","De Lessert R. 1910. Catalogue des invertebres de la Suisse. Fasc. 3, Araignees. Geneva: Musee d'histoire naturelle de Geneve.","Dahl F. 1912. Uber die Fauna des Plagefenn-Gebietes. In: Conwentz H, ed. Das Plagefenn bei Chorin. Berlin, 339 - 638, Araneae, 575 - 622.","Denis J. 1947. Notes sur les erigonides. XI. Les especes francaises du genre Oedothorax Bertkau. Bulletin de la Societe d'Histoire Naturelle de Toulouse 82: 131 - 158.","Locket GH, Millidge AF. 1953. British spiders, Vol. II. London: Ray Society.","Wiehle H. 1960 a. Spinnentiere oder Arachnoidea, XI: Micryphantidae-Zwergspinnen. Tierwelt Deutschlands 47: 1 - 620.","Wiehle H. 1960 b. Der Embolus des mannlichen Spinnentasters. Verhandlungen der Deutschen Zoologischen Gesellschaft 1960: 450 - 480.","Tystshenko VP. 1971. Opredelitel' paukov evropejskoj casti SSSR. Leningrad: Nauka.","Miller F. 1971. Pavouci-Araneida. Klic zvireny CSSR 4: 51 - 306.","Palmgren P. 1976. Die Spinnenfauna Finnlands und Ostfennoskandiens. VII. Linyphiidae 2. Fauna Fennica 29: 1 - 126.","Millidge AF. 1977. The conformation of the male palpal organs of linyphiid spiders, and its application to the taxonomic and phylogenetic analysis of the family (Araneae: Linyphiidae). Bulletin of the British Arachnological Society 4: 1 - 60.","Ruzicka V. 1978. Revision der diagnostischen Merkmale der Weibchen der tschechoslovakischen Arten der Gattung Oedothorax (Araneae: Micryphantidae). Vestnik Ceskoslovenske Zoologicke Spolecnosti v Praze 42: 195 - 208.","Hu YJ, Wang HZ. 1982. Description of three species of dwarf spiders from cotton fields in Xinxiang. Journal of Hunan Teachers College (nat. sci. ed.) 1982: 63 - 66.","Hu JL. 1984. The Chinese spiders collected from the fields and the forests. Tianjin Press of Science and Techniques.","Bosmans R. 1985. Etudes sur les Linyphiidae nord-africains. II. Le genre Oedothorax Bertkau en Africa du Nord, avec une revision des caracteres diagnostiques des males des especes ouest-palearctiques. Biologisch Jaarboek Dodonaea 53: 58 - 75.","Song DX. 1987. Spiders from agricultural regions of China (Arachnida: Araneae). Beijing: Agriculture Publishing House.","Tanasevitch AV. 1987. The linyphiid spiders of the Caucasus, USSR (Arachnida: Araneae: Linyphiidae). Senckenbergiana Biologica 67: 297 - 383.","Roberts MJ. 1987. The spiders of Great Britain and Ireland, Vol. 2: Linyphiidae and check list. Colchester: Harley Books.","Hu JL, Wu WG. 1989. Spiders from agricultural regions of Xinjiang Uygur Autonomous Region, China. Jinan: Shandong University Publishing House.","Tanasevitch AV. 1990. The spider family Linyphiidae in the fauna of the Caucasus (Arachnida, Aranei). In: Striganova BR, ed. Fauna nazemnykh bespozvonochnykh Kavkaza. Moscow: Akaedemia Nauk, 5 - 114.","Chen XE, Gao JC. 1990. The Sichuan farmland spiders in China. Chengdu: Sichuan Science and Technology Publishing House.","Heimer S, Nentwig W. 1991. Spinnen mitteleuropas: ein Bestimmungsbuch. Berlin: Paul Parey.","Alderweireldt M, 1992. Determinatieproblematiek van de zustersoorten van het genus Oedothorax (Araneae, Linyphiidae). Nieuwsbrief van de Belgische Arachnologische Vereniging 7: 4 - 8.","Zhao JZ. 1993. Spiders in the cotton fields in China. Wuhan: Wuhan Publishing House.","Wunderlich J. 1995. Funf gynandromorphe Baldachin- Spinnen aus Europa (Arachnida: Araneae: Linyphiidae). Beitrage zur Araneologie 4: 471 - 477.","Song DX, Zhu MS, Chen J. 1999. The spiders of China. Shijiazhuang: Hebei University of Science and Techology Publishing House.","Hu JL. 2001. Spiders in Qinghai-Tibet Plateau of China. Henan Science and Technology Publishing House.","Russell-Smith A. 2016. Identification of females of British Oedothorax species. Newsletter of the British Arachnological Society 137: 22 - 24.","Simon E. 1926. Les arachnides de France. Synopsis generale et catalogue des especes francaises de l'ordre des Araneae. Tome VI. 2 e partie. Paris: Roret."]}

CALLITRICHIA LATITIBIALIS (BOSMANS, 1988) COMB. NOV. (FIGS 19P, 22P, 24P, 28; SUPPORTING INFORMATION, FIG. S2M) Oedothorax latitibialis Bosmans, 1988: 17, fig. 5E–g (Dm). Type material: Holotype: Cameroon: Bambouto Mountains, 2700 m, pitfall in montane grassland, 1♂ 17.i.1983 (RMCA 165.079, examined). Paratypes: 1♂, same data (RMCA 165.087, examined). Diagnosis: Males: This species can be identified by the lack of prosomal modification and the conspicuous, characteristic male palpal tibial apophyses, similar to that of Ca. longiducta, but with a different denticle morphology on tibial prolateral apophyses; the body size of this species is larger. Description: Male (holotype): Total length: 2.11. Prosoma: 1.09 long, 0.89 wide, unmodified (Fig. 19P). Eyes: AME-AME: 0.02, AME width: 0.06, AME-ALE: 0.03, ALE width: 0.09, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.05, PME width: 0.09, PME-PME: 0.05. Chelicerae: mastidia absent; stridulatory striae rows widely and evenly spaced (Fig. 22P). Clypeus: not hirsute, one sub- AME seta. Sternum: 0.62 long, 0.61 wide. Legs: dorsal proximal macroseta on tibia I 1.61 times diameter of tibia; Tm I: 0.63. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with slender, highly scletotized, pointed, dorsally situated apophysis bent retrolaterally at tip, base connected with TPA through thin plate; TPA wide, flat, apical ridge with peculiar denticles (Fig. 28C); PC large, base not visible from dorsal view, distal setae close to distal clasp, distalsetae-bearing area wide, distal clasp without striae, clasp extended apically; T without papillae, PT without papillae, distal rim thin and smooth at margin; TS short, without papillae; MSA inconspicuous; DSA tip straight; EM flat, anterior margin with small papillae, not exceeding ARP (Fig. 28A); ARP pointed; LER absent; VRP present, retrolaterally oriented; TP tip slender; E slightly retrolaterally spiral (Fig. 28B, D). Opisthosoma: single-coloured dark grey (Fig. 24P); PLS with triad, 3+ AC (Fig. 28E). Male (paratype): Prosoma: 1.10 long, 0.88 wide. Legs: dorsal proximal macroseta on tibia II and III 1.70 and 2.31 times diameter of tibia, respectively; Tm I: 0.63. Female: Unknown. Distribution: Cameroon, only known from the type locality. Habitat: Montane grasslands. Remarks: According to the results of our phylogenetic analysis this species is transferred to Callitrichia, a placement also consistent with morphological traits of the species., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 467-468, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Bosmans R. 1988. Scientific report of the Belgian Cameroon expeditions 1981 and 1983. No. 18. Further Erigoninae and Mynogleninae (Araneae: Linyphiidae) from Cameroonian highlands. Revue Zoologique Africaine 102: 5 - 32."]}

MITRAGER GLOBICEPS (THALER, 1987) COMB. NOV. (FIGS 38I, 39I, 40I, 51; SUPPORTING INFORMATION, FIG. S4G) Oedothorax globiceps Thaler, 1987: 36, figs 16–21 (Dm). Type material: Holotype: India: Pahalgam, coniferous forest, 2400m, ♂ 14.v.1976, leg. J. Martens (SMF 33832, examined). Diagnosis: Males: This species shares the prosomal postocular hump and transverse groove between the hump and ocular region with M. dismodicoides, M. lineata, M. lucida, M. sexoculata, M. sexoculorum (Tanasevitch, 1998) comb. nov. and M. tholusa, but can be distinguished by the position of the PME at the lateral side of the groove (PME above the groove in M. dismodicoides, M. lineata, M. sexoculorum and M. tholusa; PME inside the pre-PME groove in M. lucida, M. sexoculata and M. sexoculorum). Description: Male (holotype, SMF 33832): Total length: 1.98. Prosoma: 0.91 long, 0.70 wide, transverse groove between PME, hump posteriorly (Fig. 38I). Eyes: AME- AME: 0.03, AME width: 0.04, AME-ALE: 0.04, ALE width: 0.07, ALE-PLE: 0, PLE width: 0.08, PLE-PME: 0.02, PME width: 0.06, PME-PME: 0.18. Clypeus: not hirsute. Sternum: 0.49 long, 0.52 wide. Chelicerae: mastidia absent; stridulatory striae rows widely and evenly spaced (Fig. 39I). Legs: Tm I: 0.85. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPS scaly, distal-retrolaterally pointed; TPA median-sized; TRA apically oriented; PC mediansized, base not visible from dorsal view, distal setae close to distal clasp, distal clasp without striae, clasp directed apically (Fig. 51A); T without papillae; PT apical part with small papillae; TS short, without papillae (Fig. 51B); MSA present; DSA tip round; EM flat, anterior margin without papillae, approximately equals ARP in length; ARP pointed; LER without striae, extended dorsal to E; VRP absent; TP tip narrow; E retrolaterally spiral, anterior margin at base smoothly curved (Fig. 51E, F). Opisthosoma: dorsal pattern see Fig. 40I; PMS with mAP, two AC; PLS with triad, 3+ AC (Fig. 51G, H).

‘ OEDOTHORAX’ KHASI TANASEVITCH, 2017 INCERTAE SEDIS Oedothorax khasi Tanasevitch, 2017b: 331, figs 1–7 (Dm). Type material: Holotype: India: Meghalaya, Khasi Hills, 16 km south-west of Mawsynram, between Mawsynram and Balat, 1000 m, sifting in forest, in ravine, ♂ 27.x.1978, leg. C. Besuchet & I. Löbl (MNHG, not examined). Diagnosis: Males: This species is characterized by the slightly elevated interocular region, the shape of the palpal tibial prolateral apophysis, as well as by the morphology of the embolic division (see description in Tanasevitch 2017). Females: Unknown. Distribution: Only known from the type locality in India. Habitat: Forest litter. Remarks: According to our re-delimitation of Oedothorax in the present study, this species does not belong to Oedothorax, but is probably related to ‘ Oe. ’ myanmar or ‘ Oe. ’ kodaikanal. Further comparison with more species to determine the taxonomic status of these ‘ Oedothorax ’ incertae sedis species is required. Therefore, we provisionally do not change its genus., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 546, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2017 b. Another new species of Oedothorax Bertkau, 1883 from India (Araneae, Linyphiidae). Revue Suisse de Zoologie 124: 331 - 333."]}

‘OEDOTHORAX’ CRUCIFEROIDES TANASEVITCH, 2020 INCERTAE SEDIS Oedothorax cruciferoides Tanasevitch, 2020a: 285, figs 1–3, 12–17 (Dm). Type material: Holotype: Nepal: Ilam District, 5 km north of Sanishare, feet of Siwalik Mts, 270–300 m a.s.l., mixed Shorea forest, 3–5.iv.1988, leg. J. Martens & W. Schawaller (SMF, not examined). Diagnosis: Males: This species is characterized by the small, conical elevation between the post-ocular region, the dentiform tubercles on the palpal tibial prolateral apophysis, as well as by the morphology of the embolic division (see description in Tanasevitch, 2020a). Females: Unknown. Distribution: Only known from the type localities in Nepal. Habitat: Broad-leaved forest. Remarks: According to our re-delimitation of Oedothorax in the present study, this species does not belong to Oedothorax. Further comparison with more species to determine its taxonomic status is required. Therefore, we provisionally do not change its genus., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 543-544, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2020 a. The genus Oedothorax Bertkau, 1883 in the Himalayas, with descriptions of four new species from Nepal (Aranei: Linyphiidae). Arthropoda Selecta 29: 283 - 291."]}

‘OEDOTHORAX’ PARACYMBIALIS TANASEVITCH, 2015 INCERTAE SEDIS (FIGS 7C, 67D, 68D, 79) Oedothorax paracymbialis Tanasevitch, 2015: 389, figs 70–74 (Dm). Type material: Holotype: India: Madras, Nilgiri, Hulical near Coonoor, right bank of Coonoor River, 1600 m, forest in ravin, sifting, ♂ 22.xi.1972, leg. C. Besuchet & I. Löbl (MHNG, examined). Diagnosis: Males: This species can be diagnosed by the slightly elevated postocular region, the prominent TPS and the large distal part of paracymbium. Description: Male (holotype): Total length: 1.77. Prosoma: 0.77 long, 0.63 wide, postocular region slightly elevated, with curved seta directed anteriorly (Fig. 7C). Eyes: AME-AME: 0.031, AME width: 0.04, AME-ALE: 0.03, ALE width: 0.08, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.03, PME width: 0.07, PME-PME: 0.06. Clypeus: not hirsute. Sternum: 0.48 long, 0.49 wide. Chelicerae: mastidia absent; stridulatory striae rows widely and evenly spaced (Fig. 67D). Legs: Tm I: 0.62. All metatarsi with trichobothrium. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPS striped, straight, apical-retrolaterally pointed, wide at base; TPA slightly elevated; TRA absent (Fig. 79C); PC large, base not visible from dorsal view, distal setae close to distal clasp, setae-bearing region greatly enlarged, distal clasp without striae, clasp directed slightly apically (Fig. 79A); T without papillae; PT short, without papillae, distal rim thin and smooth at margin; TS absent (Fig. 79D); MSA present; DSA straight, tip round; EM absent; ARP horizontally flat, highly sclerotized; LER small, without striae, not extended dorsal to E; VRP absent; TP slender; E retrolaterally spiral, middle part wide and striped (Fig. 79E). Opisthosoma: dorsal pattern see Fig. 68D; PMS with mAP, two AC; PLS with triad, three AC (Fig. 79F). Female: Unknown. Distribution: Only known from the type locality. Habitat: Forest litter. Remarks: According to the results of our phylogenetic analysis, and the lack of the distinctive characteristics of Oedothorax s.s., this species is more closely related to species in Clade 16 that, as this species, are distributed in the Oriental realm. We provisionally leave the taxonomic status of this species unchanged until further data become available., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 552-553, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2015. Notes on the spider genus Oedothorax Bertkau, 1883 with description of eleven new species from India (Linyphiidae: Erigoninae). Revue suisse de Zoologie 122: 381 - 398."]}

CORNITIBIA GEN. NOV. Type species: Oedothorax simplicithorax Tanasevitch, 1998 Derivatio nominis: The genus name refers to the tusklike male palpal tibia apophysis of the type species. Genus gender feminine. Diagnosis: Males: This genus is distinguished from all other erigonines by the distinct male palpal tibia, bearing one tusk-like, seta-free apophysis rising from the proximal half of tibia and pointing apically, and several long thorns from enlarged setal bases both retrolateral and prolateral to the apophysis, as well as by the unique morphology of the embolic division (as shown in Fig. 69D). The lack of a membranous connection between the radix and the embolus and the presence of radical lateral tooth clearly distinguish this species from all taxa on Clade 13. Females: Unknown. Species included: Cornitibia simplicithorax (Tanasevitch, 1998) comb. nov. Phylogenetic justification: In the original description of Oedothorax simplicithorax (Tanasevitch 1998), no account was given regarding the diagnostic characters used for assigning this species to Oedothorax, and the embolic division of this species was not clearly illustrated. Later (Tanasevitch, 2015), diagnostic characteristics of Oedothorax were described, including features related to the embolic division, which match most species on Clade 13. Examination of Co. simplicithorax revealed a greatly different embolic division configuration in this species, with an pale, ventro-prolaterally situated ‘anterior radical process’, different from that of other species examined in the present study. In the phylogeny (Fig. 2), this species is sister to a clade comprising the majority of our taxon sample (Clade 10), and not closely related to Oedothorax, Callitrichia, Mitrager or other taxa in Clade 13. After a comprehensive literature research of erigonine male palpal structures, no morphologically resembling species was found that can suggest at least a preliminary (i.e. phenetic) close relatedness. Base on these findings and its relatively basal phylogenetic placement, this species neither belongs to Oedothorax, nor can it be transferred to any other established taxon. We, therefore, propose the erection of Cornitibia gen. nov. for this species., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 529, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2015. Notes on the spider genus Oedothorax Bertkau, 1883 with description of eleven new species from India (Linyphiidae: Erigoninae). Revue suisse de Zoologie 122: 381 - 398."]}

‘ OEDOTHORAX ’ BANKSI STRAND, 1906 INCERTAE SEDIS Gongylidium sp. Banks, 1900: 480. Oedothorax banksi Strand, 1906: 445 (Df). Type material: Banks’ description (1900) was based on two females collected from Muir Glacier in 1899, but no type designation was mentioned. Diagnosis: Females: From original description in Banks (1900): ‘the epigynum is an elliptical area with a nearly square cavity in posterior part; on middle of hind margin is a denticle projecting forward’. Although difficult to comprehend the structure solely based on this description, the presence of a cavity in the posterior part of the epigye discriminates this species from all Oedothorax, Mitrager, Callitrichia species and their related genera, like Gongylidium and Timeticus. Males: Unknown. Distribution: USA, Alaska. Habitat: Glacier. Remarks: Due to the unavailability of type material and any images in the original descriptions of this species, the current status of this species is unclear.

‘OEDOTHORAX’ STYLUS TANASEVITCH, 2015 INCERTAE SEDIS (FIGS 7F, 67G, 68G, 80) Oedothorax stylus Tanasevitch, 2015: 393, figs 83–85 (Dmf). Type material: Holotype: India: Kerala, NW of Nelliampathi Hills, Kaikatty, 900 m, sifting in forest, neara spring, ♂ 30.xi.1972, leg. C. Besuchet & I. Löbl (examined). Paratypes: India: 1♀, collected together with the holotype; Madras, Anaimalai Hills, 18 km north of Valparai, 1250 m, forest, sifting litter, 1♂ 18.xi.1972, leg. C. Besuchet & I. Löbl (examined). Diagnosis: Males: This species can be diagnosed by the scaly elevation representing palpal tibial prolateral apophysis, the exceptionally long and slender ventral radical process, and the lack of prosomal modification. Females: The general appearance of this species is Oedothorax -like. It can be identified by the anterior position of the copulatory ducts opening to the spermathecae. Description: Male (paratype): Total length: 1.70. Prosoma: 0.77 long, 0.67 wide, unmodified (Fig. 7F). Eyes: AME-AME: 0.02, AME width: 0.05, AME-ALE: 0.02, ALE width: 0.08, ALE-PLE: 0.01, PLE width: 0.08, PLE-PME: 0.03, PME width: 0.08, PME-PME: 0.03. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.48 long, 0.48 wide. Chelicerae: mastidia absent; stridulatory striae rows compressed proximally (Fig. 67G). Legs: tibia chaetotaxy 2-2-1-1, dorsal proximal macroseta on tibia I 1.56 times diameter of tibia; Tm I: 0.54. All metatarsi with trichobothrium. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPS absent; TPA short, stout, apical surface scaly; TRA absent (Fig. 80C); PC median-sized, base not visible from dorsal view, distal setae close to distal clasp, distal clasp without striae, clasp directed apically (Fig. 80A); T without papillae; PT long, with median-sized papillae; TS absent (Fig. 80D); MSA present; DSA tip slightly wavy, with a protuberance on dorsal side in front of MSA; EM flat, anterior margin with papillae, length equals ARP (Fig. 80A); ARP pointed, weakly sclerotized; LER small, without striae, not extended dorsal to E; VRP long, straight, apically directed; TP tip pointed, narrow; E retrolaterally spiral, anterior margin at base wavy (Fig. 80B). Opisthosoma: dorsal pattern see Fig. 68G; PMS with mAP, two AC; PLS with triad, 3+ AC (Fig. 80G). Male (holotype): Prosoma: 0.77 long, 0.63 wide. Tm I: 0.54. Female (paratype): Total length: 1.96. Prosoma: 0.85 long, 0.66 wide. Eyes: AME-AME: 0.02, AME width: 0.05, AME-ALE: 0.02, ALE width: 0.09, ALE-PLE: 0, PLE width: 0.09, PLE-PME: 0.04, PME width: 0.08, PME-PME: 0.05. Clypeus: not hirsute, one sub- AME seta. Sternum: 0.53 long; 0.51 wide. Legs: tibia chaetotaxy 2-2-1-1, dorsal proximal macroseta on tibia I, II, III and IV 2.72, 2.82, 2.60 and 3.39 times diameter of tibia, respectively; Tm I: 0.54. All metatarsi with trichobothrium. Epigyne: Clade 13 characteristic morphology, CO anteriorly oriented (Fig. 80E, F). Opisthosoma: dorsal pattern same as male; PMS with mAP, two AC, one CY; PLS with triad, two CY, 3+ AC (Fig. 80H). Distribution: At present only known from Kerala and Madras (currently Tamil Nadu), India. Habitat: Forest litter. Remarks: According to the results of our phylogenetic analysis and the lack of the distinctive characteristics of Oedothorax s. s. (see general description of Oedothorax s.s.), this species is more closely related to ‘ Oe. ’ cunur and Atypena which, as in this species, are distributed in the Oriental realm. We provisionally leave the taxonomic status of this species unchanged until further data become available., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 554, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Tanasevitch AV. 2015. Notes on the spider genus Oedothorax Bertkau, 1883 with description of eleven new species from India (Linyphiidae: Erigoninae). Revue suisse de Zoologie 122: 381 - 398."]}

‘OEDOTHORAX’ NAZARETI SCHARFF, 1989 INCERTAE SEDIS; NEW FEMALE DESCRIPTION (FIGS 7I, 67J, 68J, 78) Oedothorax nazareti Scharff, 1989: 15, figs 7–12 (Dm). Oedothorax nazareti Tanasevitch, 2015: 382, fig. 7 (m). Type material: Holotype: Ethiopia: Shoa Administrative Province, Nazaret, under stone in cultivated farmland, 2400 m, ♂ 22.vi.1985, leg. N, Scharff (not examined). Examined material: Ethiopia, c. 15 km west of Debre Siwa, under stones in overgazed woodland, 3♂ 10♀ 11.vi.1988, det. A. Russell-Smith (RMCA 224.501). Diagnosis: Males: This species can be identified by its unique conical-shaped postocular hump and setae distribution on the hump and interocular region. Females: This species has the typical simple epigyne structures as many other Oedothorax species, but can be identified by the trajectory of the copulatory ducts (Fig. 78F). Description: Male (RMCA 224.501): Total length: 2.29. Prosoma: 1.06 long, 0.74 wide, postocular region conically elevated, covered by long, thick setae; interocular region with dense setae (Fig. 7I). Eyes: AME-AME: 0.04, AME width: 0.05, AME-ALE: 0.05, ALE width: 0.08, ALE- PLE: 0, PLE width: 0.08, PLE-PME: 0.04, PME width: 0.06, PME-PME: 0.19. Clypeus: not hirsute, one sub- AME seta. Sternum: 0.63 long, 0.56 wide. Chelicerae: mastidia absent; stridulatory striae scaly, rows widely and evenly spaced (Fig. 67J). Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.12, 0.13, 0.91 and 1.46 times diameter of tibia, respectively; Tm I: 0.67. All metatarsi with trichobothrium. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPS small, ventrally situated at TPA tip; TPA long, base prolateral to palpal tibia prolateral trichobothrium, tip bent retrolaterally (Fig. 78C); PC median-sized, base not visible from dorsal view, distal setae close to distal clasp, distal clasp without striae, clasp extended apically (Fig. 78A); T without papillae; PT long, slender, without papillae; TS long, slender, without papillae; MSA absent; DSA tip straight, simple; EM absent (Fig. 78A, E); ARP with groove fitting E, round and narrow at tip; LER absent; VRP absent; TP wide, round at tip; E retrolaterally spiral at base, prolaterally spiral at tip (Fig. 78B, E). Opisthosoma: dorsal pattern see Fig. 68J. PMS with mAP, two AC; PLS with triad, 3+ AC (Fig. 78H). Female (RMCA 224.501): Total length: 2.62. Prosoma: 1.03 long, 0.73 wide. Eyes: AME-AME: 0.02, AME width: 0.05, AME-ALE: 0.05, ALE width: 0.07, ALE- PLE: 0, PLE width: 0.07, PLE-PME: 0.04, PME width: 0.07, PME-PME: 0.07. Clypeus: not hirsute, one sub- AME seta. Sternum: 0.59 long; 0.54 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.58, 1.76, 2.26 and 2.52 times diameter of tibia, respectively; Tm I: 0.60. Epigyne: CO posteriorly situated, receptacles long, wide (Fig. 78F, G). Opisthosoma: PMS with mAP, two AC, one CY; PLS with triad, two CY, 3+ AC (Fig. 78I). Variation: The measurements are based on examined material. Males ( N = 3, means in parentheses): Total length 2.22–2.46 (2.33). Prosoma: 1.06–1.14 (1.09) long, 0.72– 0.74 (0.73) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 0.12–0.13 (0.12), 0.13–0.19 (0.16), 0.91–1.68 (1.28) and 1.46–2.28 (1.88) times diameter of tibia, respectively; Tm I: 0.58–0.67 (0.64). Females ( N = 10, means in parentheses): Total length 2.42–3.07 (2.74). Prosoma: 1.03–1.18 (1.13) long, 0.73– 0.86 (0.79) wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 1.58–1.87 (1.72), 1.63–2.13 (1.90), 1.88–2.41 (2.18) and 2.15–2.68 (2.39) times diameter of tibia, respectively; Tm I: 0.60–0.69 (0.64). Distribution: Ethiopia. Habitat: Under stones in cultivated lands or woodlands. Remarks: According to our phylogenetic analysis, this species lack the synapomorphic traits of Oedothorax s.s.. Its phylogenetic position resulted in Clade 22 sister to Clade 23. However, due to the lack of shared derived characters with other taxa in this study, its taxonomic affinity is undetermined. Therefore, we leave its taxonomic state provisionally unchanged., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 550-552, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Scharff N, 1989. New species and records of Afrotropical Linyphiidae (Araneae). Bulletin of the British Arachnological Society 8: 13 - 20.","Tanasevitch AV. 2015. Notes on the spider genus Oedothorax Bertkau, 1883 with description of eleven new species from India (Linyphiidae: Erigoninae). Revue suisse de Zoologie 122: 381 - 398."]}

CALLITRICHIA PILOSA (WUNDERLICH, 1978) COMB. NOV. (FIGS 19K, 22K, 24K, 34; SUPPORTING INFORMATION, FIG. S2H) Oedothorax pilosus Wunderlich, 1978: 258, figs 1–3 (Dm). Type material: Holotype: Ethiopia: Shewa Province, det. Wunderlich 1977 (SMF 11353, examined). Diagnosis: Males: Can be identified by the shape of palpal tibial prolateral apophysis, the embolic division morphology and the lack of prosomal modification. Description: Male (holotype, SMF): Total length: 1.85. Prosoma: 0.81 long, 0.64 wide, unmodified (Fig. 19K). Eyes: AME-AME: 0.02, AME width: 0.05, AME-ALE: 0.01, ALE width: 0.09, ALE-PLE: 0, PLE width: 0.08, PLE-PME: 0.03, PME width: 0.06, PME-PME: 0.07. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.52 long, 0.46 wide. Chelicerae: mastidia absent; stridulatory striae rows widely and evenly spaced (Fig.22K).Legs:Tm I: 0.74.Pedipalp:patella prolateral proximal vertical macrosetae absent; TPA distally with a broader retrolateral lobe and a narrower prolateral lobe, both scaly, with hollow in-between (Fig. 34C); PC base not visible from dorsal view, distal setae close to distal clasp, distal-setae-bearing area wide, distal clasp extended apically (Fig. 34A); T without papillae, PT with longitudinal folds along distal rim; TS short, without papillae (Fig. 34D); MSA present; DSA wide, tip angled at ventral side; EM flat, without papillae, not exceeding ARP; ARP pointed; LER absent; VRP long; radical part close to ARP wrinkled; TP tip pointed; E retrolaterally spiral (Fig. 34E). Opisthosoma: anterior half light-grey, posterior half dark-grey, with continuous transition in the middle (Fig. 24K); PMS with mAP, two AC; PLS with triad, 3+ AC (Fig. 34F). Female: Unknown. Distribution: Ethiopia, only known from the type locality. Habitat: Unknown. Remarks: Following the results of our phylogenetic analysis, this species is transferred to Callitrichia, a placement also consistent with morphological traits of the species. The specific epithet of this species has priority over the later published Callitrichia pilosa (Jocqué & Scharff, 1986), the latter is given a new replacement name Ca. hirsuta (see above)., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 476-478, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Wunderlich J. 1978. Zur Kentniss der Gattung Oedothorax Bertkau 1883, Callitrichia Fage 1936 und Toschia Caporiacco 1949. Senkenbergiana Biologica 58: 257 - 260.","di Caporiacco L. 1949. Aracnidi della colonia del Kenya raccolti da Toschi e Meneghetti negli anni 1944 - 1946. Commentationes Pontificia Academia Scientiarum 13: 309 - 492.","Jocque R, Scharff N, 1986. Spiders (Araneae) of the family Linyphiidae from the Tanzanian mountain areas Usambara, Uluguru and Rungwe. Annalen Zoologische Wetenschappen 248: 1 - 61."]}

‘ OEDOTHORAX’ LIMATUS CROSBY, 1905 INCERTAE SEDIS Oedothorax limatus Crosby, 1905: 311, 335, pl. 29, fig. 6 (Df). Type material: Lectotype: USA: Ithaca, NY, Nov. (AMNH, not examined). Diagnosis: Females: This species differs from Oedothorax and related taxa by the absence of trichobothrium on metatarsus IV, and by the epigynal morphology (fig. 6 in plate 29 in Crosby, 1905). Males: Unknown. Distribution: North America. Remarks: The one epigyne drawing provided in Crosby (1905) is clearly different from the characteristic conformation observed in Oedothorax s.s.., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on page 548, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Crosby CR. 1905. A catalogue of the Erigoneae of North America, with notes and descriptions of new species. Proceedings of the Academy of Natural Sciences of Philadelphia 57: 301 - 343."]}

EMERTONGONE GEN. NOV. Type species: Lophocarenum montiferum Emerton, 1882. Derivatio nominis: The genus name is a combination of Emerton, in honor of the American arachnologist James Henry Emerton, the original describer of the type species, and the ending of the generic name ‘ Erigone ’. Genus gender feminine. Diagnosis: Males: Small-sized, dark-brown-coloured erigonine. This genus is characterized by its male prosomal modification comprising a post-PME groove, post- PME lobe, the anteriorly protruded hirsute clypeus and the shorter distance from PLE to clypeal lower margin compared to PME. Despite its superficial partial similarity of prosomal modification with Oe. gibbosus, this genus has no separate radix and embolus connected by a membranous region, and the embolic membrane is erected from distal part of distal suprategular apophysis. These characters distinguish it from all taxa in Clade 13. Females: Can be distinguished from all other examined taxa by the shorter distance from ALE to clypeus margin compared to distance from AME to clypeus margin, the hirsute clypeus, and the transverse slit on the epigyne between the two copulatory openings (Fig. 65E, G). Species included: Emertongone montifera (Emerton, 1882) comb. nov. Phylogenetic justification: Although the prosomal modification of Em. montifera superficially resembles that of Oe. gibbosus, the palpal configuration of this species is different from that of Oedothorax as delimited in the present study. In the phylogeny (Fig. 2), this species is sister to Clade 12; i.e. more closely related to the Walkenaeria and Gonatium representatives than to all Oedothorax, Callitrichia, Mitrager and other representatives in Clade 13. The similarity in prosomal morphology is, therefore, a result of homoplasy. After a comprehensive literature research of erigonine male palpal structures, no morphologically resembling species was found that can suggest at least a preliminary (i.e. phenetic) close relatedness. Base on these findings and its resulting phylogenetic placement, this species neither belongs to Oedothorax, nor can it be transferred to any other established taxon.We, therefore, propose the erection of Emertongone gen. nov. for this unique species.

OEDOTHORAX S.S. BERTKAU, 1883 Type species: Neriene gibbosa Blackwall, 1841. Monophyly: This group is supported by the following unambiguous character transformations: paracymbium base visible from dorsal view of male pedipalp (Ch 4, homoplastic), median position of distal setae group on paracymbium (Ch 7, synapomorphic), embolus base horn present (Ch 12, synapomorphic) embolic membrane cylindrical (Ch 34, synapomorphic) and tegular papillae present (Ch 42, homoplastic); and the following ambiguous character transformations: embolus prolaterally spiral (Ch 11, homoplastic, ambiguous transformation), radix – embolus membranous region extend to prolateral side of radix (Ch 24, synapomorphic, ambiguous transformation). Diagnosis: The newly circumscribed Oedothorax s.s. is similar to Callitrichia, Mitrager and other species in Clade 13 in their configuration of the embolic division, the tibial chaetotaxy and the epigyne morphology. Oedothorax s.s. is characterized and can be distinguished from other taxa in Clade 13 by the following features: 1. Paracymbium: small-sized; base visible from dorsal view of male pedipalp (medium- to large-sized in other species in Clade 13; base covered by cymbium from dorsal view); distal part not enlarged (greatly enlarged in most Callitrichia species); distal setae group with middle position or indistinguishable from basal setae group (with distal position in other species in Clade 13); distal clasp distally extended, without striae (retrolaterally extended and/or with striae in many Mitrager species). 2. Copulatory bulb: embolus base protuberance present (arrow in Fig. 5B) (absent in other species in Clade 13); embolus prolaterally spiral around pointed, prolaterally spiral anterior radical process (embolus retrolaterally spiral in other species in Clade 13); embolic membrane cylindrical (imperceptible in some species in unexpanded pedipalps) (either flat and broad or absent in other species in Clade 13); embolus–radix membranous region extended to prolateral side of radix (not extended to prolateral side of radix in other species in Clade 13); ventral radical process absent (present in most Callitrichia, some Mitrager, Atypena and some ‘ Oedothorax ’ incertae sedis species); lateral extension of radix absent (present in all Mitrager, Atypena, Ca. convector and some ‘ Oedothorax ’ incertae sedis species); tegular papillae present in some species, protegulum with papillae, tegular sac short; marginal suprategular apophysis present; distal suprategular apophysis straight, distally oriented, mostly narrow and round at tip (except Oe. gibbosus and Oe. trilobatus, tip broad and smoothly serrated). 3. Tibia: moderately modified; shape of prolateral apophysis varies among species, but never elevated vertically (vertically elevated in many Callitrichia, some Mitrager and some ‘ Oedothorax ’ incertae sedis species); with basal thorn in some species (absent in other species in Clade 13); retrolateral apophysis absent (present in Mitrager, Atypena, Ca. convector and some ‘ Oedothorax ’ incertae sedis species); prolateral spike absent (present in most Mitrager species). 4. Epigyne: different from Callitrichia and Holmelgonia in the mesal entrance of copulatory ducts into the spermathecae with respect to the exits of the fertilization ducts. Description: The genus includes medium-sized (male 1.2–2.5, female 2.1–3.8) erigonines with an evenly coloured opisthosoma from light brown to dark brown. Male and female posterior median spinnerets with one minor ampullate gland spigot, two aciniform gland spigots; posterior lateral spinnerets with triad, more than three aciniform gland spigots; female posterior median spinnerets and posterior lateral spinnerets with one and two cylindrical gland spigots, respectively. Male prosoma varies in the degree of prosomal modifications, ranging from unnoticeable (Oe. paludigena, Oe. agrestis, Oe. fuscus and Oe. tingitanus) to prominent post-PME humps, post-PME grooves and lateral sulci and pits. Palpal patella prolateral proximal vertical macrosetae absent.This genus also shares those features defining Clade 13 (see above). For palpal and epigynal features, see description of Clade 13 and diagnosis. New circumscription: According to our phylogenetic analysis and descriptions from the literature, only ten species are regarded here as ‘true’ Oedothorax: Oe. gibbosus and its congeners: agrestis, apicatus, fuscus, gibbifer, meridionalis, paludigena, retusus, tingitanus and trilobatus. However, 27 additional species remain here as ‘ Oedothorax ’ incertae sedis (see section below) and are deemed not congeneric with the type species, but await future taxonomic treatment. The remaining 43 species are transferred from Oedothorax to other genera (see taxonomic actions below). Natural history: Most species are found in humid environments like in litter, under bark or stones, in grasslands, marshes or at riversides. Remarks: Although no types of Oedothorax s.s. were examined, detailed descriptions and illustrations in the literature abound, allowing clear identification of the examined specimens. New distribution: Europe, Turkey, Caucasus, Iran, Russia to Central Asia, China, Azores, North Africa, North America., Published as part of Lin, Shou-Wang, Lopardo, Lara & Uhl, Gabriele, 2022, Evolution of nuptial-gift-related male prosomal structures: taxonomic revision and cladistic analysis of the genus Oedothorax (Araneae: Linyphiidae: Erigoninae), pp. 417-584 in Zoological Journal of the Linnean Society 195 on pages 426-428, DOI: 10.1093/zoolinnean/zlab033, http://zenodo.org/record/6967774, {"references":["Blackwall J. 1841. The difference in the number of eyes with which spiders are provided proposed as the basis of their distribution into tribes; with descriptions of newly discovered species, and the characters of a new family and three new genera of spiders. Transactions of the Linnean Society of London 18: 601 - 670.","Blackwall J. 1853. Descriptions of some newly discovered species of Araneidea. Annals and Magazine of Natural History 11: 14 - 25."]}

CALLITRICHIA JUGUMA (SCHARFF, 1990) COMB. NOV. (FIGS 19I, 21, 22I, 24I; SUPPORTING INFORMATION, FIG. S2F) Ophrynia juguma Scharff, 1990a: 66, figs 164–172 (Dmf). Examined material: Tanzania: Uzungwa Mts., Iringa Region, Uzungwa Scarp Forest Reserve, 2♂ 2♀, leg. N, Scharff (ZMUC). Diagnosis: Males: Similar to Ca. rostrata, Ca. perspicuua and Ca. insulana in the shapes of the flat PME lobe and inter-AME-PME lobe, forming the narrow inter-AME- PME groove (figs 155, 164 and 173 in Scharff, 1990; fig. 116 in Jocqué & Scharff, 1986), which distinguishes them from other erigonines; this species can be further distinguished from the latter three species by the shape of the palpal tibial apophysis (Fig. 21A–C, in comparison to figs 157 and 177 in Scharff, 1990a; fig. 118 in Jocqué & Scharff, 1986). Females: Epigyne morphology similar to other Callitrichia species, but can be distinguished by the particularly close distance between the spermathecae (Fig. 21E, F, in comparison to e.g. Fig. 18G, fig. 115 in Jocqué & Scharff, 1986, and fig. 163 in Scharff, 1990a). Description: Male (ZMUC): Total length: 2.08. Prosoma: 0.89 long, 0.74 wide, with a narrow and small PME lobe; posterior part with fine, long setae over PME; frontal part ends in a ridge, pointing up and forward, curved inward in the middle, with transversal slit at the peak (Fig. 19I). AME-AME: 0.04, AME width: 0.04, AME- ALE: 0.03, ALE width: 0.07, ALE-PLE: 0.07, PLE width: 0.06, PLE-PME: 0.07, PME width: 0.04, PME- PME: 0.07. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.51 long, 0.49 wide. Chelicerae: mastidia absent; stridulatory striae ridged, rows compressed and evenly spaced (Fig. 22I). Legs: metatarsi and tarsi of legs I and II with perpendicular lateral setae, long and numerous; dorsal proximal macroseta on tibia I, II, III and IV 1.44, 1.57, 2.10 and 2.96 times diameter of tibia, respectively;Tm I: 0.79. Pedipalp: patella prolateral proximal vertical macrosetae absent; tibia with one prolateral, two retrolateral trichobothria; TPA large, broadly rounded, prolateral side straight, retrolateral side with small, round, scaly distal process (Fig. 21C); PC base not visible from dorsal view, distal setae close to distal clasp, distal-setaebearing area wide, distal clasp extended apically (Fig. 21A); T without papillae, PT truncated, apical part with papillae; TS absent (Fig. 21A); MSA present; DSA wide, median part of tip concave (Fig. 21A); EM broad and flat, without papillae, exceeding ARP; ARP thick, bifid; LER absent; VRP short, with papillae extended to medan part of R; TP long, round at tip; E short and stout, slightly retrolaterally curved (Fig. 21B, C). Opisthosoma: dorsal pattern see Fig. 24I; PMS without spigots; PLS with one FL (Fig. 21G). Female (ZMUC): Total length: 1.87. Prosoma: 0.79 long, 0.66 wide. AME-AME: 0.03, AME width: 0.04, AME-ALE: 0.02, ALE width: 0.07, ALE-PLE: 0.01, PLE width: 0.06, PLE-PME: 0.04, PME width: 0.05, PME-PME: 0.05. Clypeus: not hirsute, one sub-AME seta. Sternum: 0.51 long, 0.49 wide. Legs: dorsal proximal macroseta on tibia I, II, III and IV 2.88, 2.76, 2.67 and 3.21 times diameter of tibia, respectively; Tm I: 0.88. Epigyne of Clade 13 characteristic morphology, borders between ventral and dorsal plates converging anteriorly and posteriorly, CO posterior to spermathecae (Fig. 21E, F). Opisthosoma: dorsal pattern as male; spigots as in male, PMS with one CY; PLS with two CY (Fig. 21H). Remarks: Following the results of our phylogenetic analysis, this species is transferred to Callitrichia, a placement also consistent with morphological traits of the species.

GONGYLIDIOIDES INSULANUS (PAIK, 1980) COMB. NOV. Oedothorax insulanus Paik, 1980: 162, figs 9–21 (Df). Oedothorax insulanus Namkung, 2002: 205, fig. 17.57A–b (f). Oedothorax insulanus Namkung, 2003: 207, fig. 17.57A–b (f). Gongylidioides kouqianensis Tu & Li, 2006: 59, fig. 5A–G (Dm), synon. nov.. Oedothorax insulanus Seo, 2011: 37, figs 11–16 (f, Dm). Type material: Oedothorax insulanus: Holotype: South Korea: Jeunlanam-Do So Heuksan-do Island, Hang-ri, ♀ 27.vii.1979, coll. S. R. Son (not examined). Paratype: South Korea: So Heuksan-do Isl, Hang-ri, 2♀ 27.vii.1979, coll. S. R. Son (not examined); same location, 1♀ 27.vii.1979, coll. T. H. Jo (not examined). Gongylidoides kouqianensis: Holotype and paratype: China: Jilin Province, Kouqian County, 2♂ 29.vi.1989, Institute of Zoology, Chinese Academy of Sciences in Beijing, China (Tu and Li, 2006; not examined). Non-type material: Gongylidoides kouqianensis: Gyeongsangbuk-do, Ulreung-gun, Hyeonpo, 10♂ 4♀ 24.viii.2006, coll. S. Y. Kim, collection of Department of Biology, Keimyung University (Seo, 2011; not examined). Diagnosis: Males: This species can be recognized among congeners by the combination of the following features: the lack of prominent prosomal modification; the broad, apically pointed palpal tibial retrolateral apophysis without a tooth on the inner surface; lamella characteristica extal tip horn-like, with two projections, inner one triangular in ventral view, dorsal one blunt (fig. 5F in Tu & Li, 2006). Females: Can be diagnosed by the half-octagonshaped dorsal plate of the epigyne (figs 16, 17 in Paik, 1980). Distribution: South Korea; China, Jilin province. Remarks: Tu & Li (2006) and Seo (2011) have provided informative drawings of three perspectives of the male palps of Gongylidioides kouqianensis and Oe. insulanus, respectively, and their resemblance strongly suggests that these two species are identical. Since Oe. insulanus was published earlier, it has priority over Gongylidioides kouqianensis. The palpal and epigynal morphology of Gongylidioides insulanus is in accordance with the generic description of Gongylidioides in Tu & Li (2006), and supports the placement of this species in Gongylidioides.

‘ OEDOTHORAX’ SEXMACULATUS SAITO & ONO, 2001 INCERTAE SEDIS Oedothorax sexmaculatus Saito & Ono, 2001: 5, figs 5–9 (Dmf). Oedothorax sexmaculatus Ono et al., 2009: 284, figs 343–347 (mf). Type material: Holotype: Japan: Hunshu, Fukuschima Pref., Yama-gun, Azuma-yama Mts., Jododaria, ♂, 11.vi.1986, leg. K. Kumada (NSMT-Ar 4572, not examined). Paratypes: same data as holotype, 5♂ 6♀ (NSMT-Ar 4573–4574, not examined); Gunma Pref., Tone-gun, Katashina-mura, Marunuma, 1♂ 1.vi.1983, leg. H. Saito (NSMT-Ar 4575, not examined). Diagnosis: Males: This species can be distinguished by the variegated opisthosoma, the palpal apophysis and the depiction of the bulbal structure in figures 8 and 9 in Saito & Ono (2001), in which the individual structures of the embolic division are unfortunately not clearly recognizable. Females: This species can be identified by the dorsal pattern of opisthosoma with six dark patches, and the epigynal external morphology illustrated in figure 9 in Saito & Ono (2001). Distribution: Japan: Honshu. Remarks: The description of Saito & Ono (2001) is in accordance with the general combination of somatic characters of Oedothorax, Mitrager and other related taxa. Although the configuration of the embolic division is not fully recognizable in their drawings, the embolus seems to be prolaterally spiralled (fig. 8 in Saito & Ono, 2001), but the ARP tip is not pointed at tip and not spiral as in Oedothorax; its PC has a clearly broader distal part (fig. 6 in Saito & Ono, 2001) than in Oedothorax. The six distinct spots on the opisthosoma is seen neither in Oedothorax s.s. nor in Mitrager. Due to these dissimilarities, this species most probably belongs to another genus.