Dictynidae O. Pickard-Cambridge, 1871

Montana, Katherine O., Cala-Riquelme, Franklyn, Crews, Sarah C., Gorneau, Jacob A., Al-Jamal, Amin M., Alequín, Luigie D., Spagna, Joseph C., Ballarin, Francesco & Esposito, Lauren A., 2025, Tailor’s drawer no more: a reappraisal of the spider family Dictynidae O. Pickard-Cambridge, 1871 sensu lato, Zoological Journal of the Linnean Society 204 (2) : -

Family Dictynidae O. Pickard-Cambridge, 1871 s.s.

Type genus: Dictyna Sundevall, 1833 (type species Aranea benigna Walckenaer, 1802 ).

Material examined: Aebutina binotata : Brazil, Amazonas, São Paulo de Olivença, Fonte Boa , Coll. Mathan, M. de (Syntypes, 3 females, MNHN-AR-AR159). Adenodictyna kudoae : Japan, Kagoshima Pref., Amami-Ōshima Is., Naze city, Kinsakubaru , 20.IV.2008, Coll. Y. Kudo (Holotype, 1 male, NMST7953 ) . Ajmonia gratiosa : Algeria, Constantine (7 males, 34 females, MNHN-AR-AR606). Ajmonia marakata : India, Ootacamund, Nilgiris , 19–20.vii.1960, Coll. Pres. Dr. W. Rae-Sherriffs (Hototype, 1 male, 1 female, NHMUK1635133 ) . Ajmonia numidica : Algeria, Mila Prov. (prev. Dept. Constantine), Zouagha Forest , 15.v.1935 (2 males, 3 females, MNHN-AR-AR657). Ajmonia smaragdula : Sri Lanka, Central Province, Nuwara Eliya to Kandy , 1.i–31.xii.1892, Coll. E. Simon (Syntypes, 1 male, 1 female, MNHN-AR-AR714). Ajmonia velifera : India, Sikkim (Syntypes, 1 male, 1 female, MNHN-AR-AR713). Anaxibia difficilis : São Tomé-et-Príncipe, São Tomé, Bombaim, Traz-os-Montes , 7.vi.1956, Coll. P. Viette (Holotype, 1 male, MNHN-AR-AR477) . São Tomé and Príncipe, São Tomé, Lobata, Muquinqui Peak, near Guadalupe , 0°22'49.2"N 6°38'47.1"E, Elev. 190 m, beating foliage, 22.xi.2016, Coll. L.A. Esposito (8 males, 6 females, 3 imm., CASENT9118598 ; 1female, 1male, 1imm., CASENT9118599 ) . São Tomé and Príncipe, São Tomé, Mé-Zóchi, São Nicolau Waterfall , 0°17'08.8"N 6°37'31.9"E, Elev. 927 m, 15.xi.2016, Coll. L.A. Esposito (1 male, CASENT9118601 ) . Archaeodictyna anguiniceps : Egypt, Siwa , Bahrein, east lake shore, 26.iv.1939, Coll. M. Cameron, Dr. G. Denis (Holotype, 1 male, 1 female, NHMUK1635121 ) . Archaeodictyna consecuta : Morocco / Algeria, Saida (7 males, 4 females, MNHN-AR-AR604). Archaeodictyna ammophila : France, Île-de-France, Seine-et- Marne Prov., Fontainebleau , Coll. E. Simon (1 male, 1 female, MNHN-AR-AR530). Ar. ulova South Africa, Natal Spioenkop Dam , south shore, 30 km SW Ladysmith, el. 900m, 28°40'60"S, 25°27'59.00"′E, mixed grasslands and dry bushveld, 1.vii.1986, Coll. T. Meikle-Griswold, C.E. Griswold (Holotype, 1 male, 1 female, NHMUK 1635102 ) . Brigittea innocens (O. Pickard-Cambridge, 1872) : Myanmar, Magway Division, Shwesettaw Wildlife Reservation , guest house, 20°3′37.3′′N 94°35′39.8′′E, at night, deciduous forest, 28.ix.2003, Coll. D. Ubick, (1 female, CASENT9017108 ) . Israel, Har Horshan, North of Ma’agan Mikh’el , 32°32'60"N 34°54'59.99"E, at night, deciduous forest, 26.iv.1987, Coll. V. D. Roth (1 male, CASENT9118698 ) . Tanzania, Tanganyika, 6 mi W of Ngare Nanyuki, Elev. 1500 m, 19.xi.1957, Coll. E.S. Ross, R. E. Leech (1 female, CASENT9118704 ) . Callevophthalmus albus : Australia, Western Australia, 1.5 km E Cranbrook , low bushes in bottom sandy land, open scrub, 11.ix.2003, Coll. E.I. Schlinger, M.E. Irwin (1 female, CASENT9118676 ) . Australia, Sydney , 1854–58 (Holotype, 1 female, NHMUK 1635107 ) . Callevophthalmus maculatus : Australia, Sydney , 1.i–31.xii.1887 (Holotype, 1 female, NHMUK 1635108 ) . Mallos dugesi (Becker, 1886) : USA, Arizona, Pima Co., Tucson , vii.1930, Coll. Gertsch (1 female, MNHN-AR-AR493). Mallos dugesi : USA, Arizona, Huachuca Mountains, Miller Canyon , 17.xi.1910, Coll. William Morton Wheeler (Holotype, 2 females, MCZ-IZ29976 ) . Mallos hesperius (Chamberlin, 1916) : Peru, San Miguel , vii.1911, Coll. H.W. Foote, Yale Peruvian Expedition (Holotype, 1 male, MCZ-IZ15690 ) . Mallos mians (Chamberlin, 1919) : USA, California (no specific locality data), 1909 and 1913 (from publication), Coll. R. V. Chamberlin (Holotype, 1 female, MCZ-IZ15096 ) . Mallos niveus O. Pickard-Cambridge, 1902 : Mexico, San Luis Potosi, Rio Frio , 29.xi.1940, Coll. Gertsch (1 male, 1 female, MNHN-AR-AR481) . USA, Arizona, Cochise Co., Chiricahua Mtns. , SWRS, 5 mi W Portal, 14.viii.1972, Coll. P. R. Craig (1 male, CASENT9118619 ) . USA, Arizona, Mohave Co., 2 mi S Davis Dam, Elev. 500 ft, salt brush, 14.vi.1968, Coll. L.D. Nelson (1 female, CASENT9118639 ) . USA, New Mexico, Otero Co., Oliver Lee State Park, Dog Canyon , 32°44'57.6"N 105°54'46.6"W, 7.x.2021, Coll. M. Pleisher (1 male, 1 female, 2 inm., CASENT9087236 ) . Mallos pallidus (Banks, 1904) : USA, California, Mt. Shasta , pre-1904 publication date, Coll. Nathan Banks, (Syntype, 1 male, 2 females, MCZ-IZ22473 ) . USA, California, Santa Barbara Co., Santa Cruz Island, Stanton Beach , 17.iv.1973, Coll. D. Haralson (1 female, CASENT9118623 ) . USA, California, Mendocino Co., Angelo Reserve, S Fork Eel R. near Wilderness Lodge, 9.7 km N Branscomb, 39°44.450′N 123°37.944′W, Elev. 420 m, redwood/ Douglas fir forest, extraction from concentrated forest, 28.ix.2013, Coll. B. Mathat (1 female, CASENT9056350 ) . USA, California, Kern Co., Indian Wells Canyon , 35°41'9"N 117°55'38.99"W, Elev. 1125 m, malaise, damp creek bed, 8–14.v.2014, Coll. M.E. Irwin (1 male, CASENT9118662 ) . Mexitlia trivittata : USA, New Mexico, Albuquerque , pre-1901 publication date, Coll. Hugo Soltau, Nathan Banks (Syntype, 1 male, 1 female, MCZ-IZ23355 ) . USA, New Mexico, Sandoval Co., Jemez Mountains, E fork Jemez River, near Battleship Rock , 27.iv.2021, Coll. M. Pleisher (2 females, CASENT9087231 ) . USA, Arizona, 17 miles NE, 8.vii.1940, Coll. Gertsch (1 male, 1 female, MNHN-AR-AR482). Nigma flavescens (Walckenaer, 1830) : Israel, Jerusalem, The Hebrew University Fac. Agric., Dept. Ent. , 2.vi.2000, Coll. Mendel (1 male, 2 females, 1 imm., CASENT9118690 ) . Nigma gertschi ( Berland and Millot 1940) : Kenya, Rift Valley Province, West Pokot District, Marich Pass Field Studies Centre , 1°32.2'S 35°27.4'E, Elev. 3000 ft, 26–29.vii.1999, Coll. W.J. Pulawski, J.S. Schweikert (1 male, CASENT9118714 ) . Nigma longipes : Kenya, Mau Forest du Kenya, 22.i.1912, Coll. C. Alluaud, R. Jeannel (Type, 1 male, MNHN-AR-AR5302) . Kenya, Nakuru Co., Naivasha , 1–31.xii.1911, Coll. C. Alluaud, R. Jeannel (Syntypes, 1 female, MNHN-AR-AR5291). Paradictyna rufoflava (Chamberlain, 1946) : New Zealand, North Island, L. Tikitapu (Blue Lake), Elev. 350 m, malaise, dry hillside, 29.x.–4.xi.1977, Coll. E.I. Schlinger (1 male, CASENT9118610 ) . New Zealand, South Island, S. Auckland, Waitomo Co., Pio Pio Dist. , 2 km NW Mangakowhai Rd., sweeping, native bush, 29.x.–8.ii.1996, Coll. B. Zuparko (1 female, CASENT9118688 ) . Phantyna bicornis Emerton, 1915 : USA, Massachusetts, Ipswich , sand near seashore, 24.vi.1905, Coll. James H. Emerton (Syntype, 1 female, MCZ-IZ43954 ) . USA, Massachusetts, Ipswich , sand near seashore, 24.vi.1905, Coll. James H. Emerton (Syntype, 1 male, MCZ-IZ20499 ) . Phantyna remota (Banks, 1924) : Ecuador, Galapagos, Daphne , 23.iv.1923, Coll. William Beebe, Williams Galapagos Expedition 1923 (Syntype, 1 male, 2 females, MCZ-IZ22879 ) . Phantyna terranea (Ivie, 1947) : USA, Washington, Vantage , 6.v.1936, Coll. M.H. Hatch (Syntype, 1 female, MCZ-IZ22451 ) . USA, Washington, Vantage , 1.v.1936, Coll. M.H. Hatch (Syntype, 1 female, MCZ-IZ24763 ) . Rhion pallidum : Sri Lanka, Kandy , Coll. E. Simon (MNHN-AR-AR378). Tahuantina zapfeae : Chile, Curicó Province, Loma Hueca-Huecan , 15 km E Curicó, 35°03'59.4"S 71°07'21.4"W, Elev. 400 m, malaise, dry hillside, 29.x.–10.ix.1997, Coll. J.E. Barriga, M.E. Irwin (1 male, CASENT9062312 ) . Chile, Prov. Coquimbo, Hacienda Illapel , 31°36'S 71°07'W, Elev. 900 m., malaise, dry hillside, 19.x.1966, Coll. E.I. Schlinger, M.E. Irwin, Peña (3 males, 6 females, 3 imm., CASENT9118813 ) . Thallumetus acanthochirus Simon, 1904 : Chile, Valparaiso Prov., Marga Marga, Quilpué , Coll. C.E. Porter (MNHN-AR-AR383) . Chile, Diguillín Prov., Ñuble Region , Sierra de Chillán, Coll. E. Simon (1 male, MNHN-AR-AR377) . Venezuela, Carabobo, Parque Nacional San Esteban , Coll. E. Simon (1 male, MNHN-AR-AR384). Thallumetus parvulus Bryant, 1942 : US Virgin Islands, St. Croix , pre-1942, Coll. Harry A. Beatty (Holotype, 1 male, MCZ-IZ22541 ) . Thallumetus pullus Chickering, 1952 : Panama, Canal Zone, Biol. Area , viii.1950, Coll. Arthur M. Chickering (Allotype, 1 female, MCZ-IZ44022 ) . Panama, Canal Zone, Biol. Area , 1–31.vii.1950, Coll. Arthur M. Chickering (Holotype, 1 male, MCZ-IZ22788 ) . Thallumetus pusillus Chickering, 1948 : Panama, Canal Zone, Biol. Area , 1-30.viii.1939, Coll. Arthur M. Chickering (Allotype, 1 female, MCZ-IZ44021 ) . Panama, Canal Zone, Biol. Area , 1–30. viii.1939, Coll. Arthur M. Chickering (Holotype, 1 male, MCZ-IZ22815 ) . Tivyna moaba : USA, California, Imperial Co., East Mesa Geothermal Site, 8.0 mi ESE Holtville , 32°42'N 115°15'W, Elev. 17 m, pitfalls, 23.iii.1978, Coll. R. Aalbu (1 female, CASENT9062468 ) . Tivyna spatula : Cuba, Cienfuegos, Soledad , 22.viii.1933, Coll. Weber (1 male, MCZ-IZ20964 ) . Peru, Cajamarca Prov., Sector La Pampa, Fundo La Paloma , 3 km W Chilete, 07°13.42'S 78°51.84'W, Elev. 865 m, malaise, dry gulch, 31.iii–7.iv.2008, Coll. M.E. Irwin, G. Antón Amaya (1 female, CASENT9061652 ) . Peru, Cajamarca Prov., Sector La Pampa, Fundo La Paloma , 3 km W Chilete, 07°13.42'S 78°51.84'W, Elev. 865 m, malaise, 17–24.i.2008, Coll. M.E. Irwin, G. Antón Amaya (1 female, CASENT9061673 ) . Mexico, Colima, Revillagigedo Archipelago , Isla Socorro , ~ 2 km NW Bahia Braithwaite, in orchard, 31.i.1966, Coll. L. Baptista (1 female, CASENT9118791 ) .

Diagnosis: Small cribellate spiders ( Figs 9A–F, 10E, F, 12A–F, 43E–F, 47A–B, 52A–C, E–F, 53A–C, 54A, 63D, 66B) with the carapace longer than broad, quite high and convex ( Figs 6E, F, 7B, D–F) (two times higher in Archaeodictyna , in part: Fig. 45A–B, E) ( Chamberlin and Gertsch 1958); clypeus typically high ( Figs 6E, F, 7D, F, 47A, 50A, 51F, 53A) (clypeus low in Lathyidae fam. n. and Argyronetidae stat. reinst.); chelicerae concave medially, gradually narrowed apically, subparallel, with a conspicuous lateral condyle sometimes developed into a horn ( Figs 6E, F, 7A–F, 43A–B, 45B); tarsi lacking trichobothria, present at the apical end of the metatarsi; male palpal tibia usually with a dictynid process with typically two ctenidia ( Figs 30E, F, 31F, 44E, 51D, E, 53E–F, 55C, D, 56B, D–F, 57A, C, 59E, F, 62A–D); embolus of variable thickness, acuminate to apex or variously modified apically ( Figs 28A, B, D–F, 29A–E, 31E, 50E, 52D, 53D, 56D–F, 57A–D, 58A–F, 59E, F); conductor upper arm variable in size but never coiled more than 225° (conductor upper arm shorter than lower arm in Argyronetidae ); conductor lower arm widens to a conductor-tegulum lock mechanism ( Fig. 27C), coiled to produce a spiral of variable form ( Figs 25B, D, F, 26C, 27C, 29F, 30B, C, 31C, 53D–F) ( Chamberlin and Gertsch 1958) that usually ends in a scaly tip ( Figs 27C, D, 30B, 31C).

Remarks: Dictynidae s.l. has long been a receptacle for a mixture of cribellate and ecribellate RTA clade spiders. Wheeler et al. (2017) argued that the cribellate genera could be a natural group considering that they all have a characteristic male palp, and many males have medially concave chelicerae. With our restructuring of the family and the transfer of species into Lathyidae fam. n. and Argyronetidae stat. reinst., Dictynidae s.l. becomes morphologically structured with clear synapomorphies. The results of Spagna and Gillespie (2008), Miller et al. (2010), Spagna et al. (2010), and Wheeler et al. (2017) have explored the molecular phylogenetics of the RTA clade using Sanger loci but with few terminals, and all of their results initially suggested Dictynidae s.l. to be monophyletic; however, the relationship between the genera has changed in each of these studies, suggesting a paraphyletic or polyphyletic composition. Most recently, Crews et al. (2020), also using Sanger data, and Kulkarni et al. (2023), using UCE + Sanger loci, recovered Dictynidae s.s. as paraphyletic, and Gorneau et al. (2023a) using UCEs + Sanger loci recovered Dictynidae s.l. as monophyletic. Our results corroborate the results obtained by Gorneau et al. (2023a). We find support for including cribellate genera that present: (i) a dictynid process (with the exception of Tivyna , Mallos , Ajmonia , Dictynomorpha Spassky, 1939 , Nigma , and Califorenigma gen. n.) and (ii) males and females with the chelicerae medially concave.

The monotypic genus Clitistes Simon, 1902 described from only a female specimen was declared a nomen dubium by Miller (2007: 259). Dupérré and Harms (2018: 8) revised the type of the species and transferred it from the Linyphiidae to Dictynidae . After examining several specimens of males and females of Cybaeolus pusillus Simon, 1884 at CAS [ CASENT9118799 (1 male), 9118750 (2 males, 1 female), 9118752 (2 males), 9118758 (1 male), 9118764 (1 male, 1 female), 9118783 (1 male, 1 juv.), 9118833 (1 female), 9118839 (1 male), 9118843 (1 male), 9118845 (1 female)], based on the morphology of Clitistes velutinus Simon, 1902 (see: Dupérré and Harms 2018: 8), we consider Clitistes velutinus Simon, 1902 a junior synonym of Cybaeolus pusillus Simon, 1884 ( Clitistes velutinus Simon, 1902 syn. n.) .

The genus Aebutina Simon, 1892 , by having reduced AME, the retromargin of the cheliceral fang furrow with one to several differential denticles ( Griswold et al. 2005: fig. 130E), and a tegular median apophysis ( Fig. 19D; Griswold et al. 2005: fig. 169C, D) is transferred to Macrobunidae . Hoplolathys Caporiacco, 1947 was originally described from a currently lost juvenile, and here we consider the genus a nomen dubium . Lathys changtunesis Hu, 2001 was previously considered a member of the genus Ajmonia by Zhang et al. (2012: 1); however, this transfer was not accepted due to lack of evidence. We conclude that L. changtunesis is not a representative of the Lathyidae fam. n. (see diagnosis of Lathyidae fam. n.). The presence of a conductor with the upper arm shorter than the lower arm, the tegular path of the spermaphor without a loop, the cymbial dorsal process developed, and the patellar dorsal apophysis present indicate that this species should be placed in Ajmonia . Here, we transfer L. changtunesis to Ajmonia [ Ajmonia changtunesis (Hu, 2001) comb. n.].

The morphology of the palp of Dictyna aguasverdes Wunderlich, 2022 , D. bispinosa Simon, 1906 , D. fuerteventurensis Schmidt, 1976 , and D. lanzarotensis Wunderlich, 2022 places these species outside the delimitation of Dictyna (see diagnosis of Dictyna below) but within Archaeodictyna ( Figs 45A–F, 46A– D) by having the embolus thick, coiled approximately 250°, originating prolaterally from the bulb ( Fig. 46B); the conductor lower arm coiled no more than 45°, with the tip directed distally in most species, and with the conductor locking mechanism longer than the conductor tip; dictynid process without an evident tubercle ( Fig. 46C, D).

Nigma albida (O. Pickard-Cambridge, 1885) is represented by a single illustration of the epigyne ( Dyal 1935: pl. 14, fig. 79) that does not resemble the epigyne of N. flavescens , the type of the genus. It likely does not belong in Nigma , but only examination of the type of N. albida will allow that determination. Nigma gertschi is also not similar to the type of N. flavescens . There is only a single illustration of the palp and the epigyne ( Berland and Millot 1940), and the authors indicate that it is similar to Dictyna viridissima Roewer, 1951 , which has been synonymized with N. walckenaeri ; however, we do not agree that they are similar to N. walckenaeri or Nigma s.s.. Lehtinen (1967) indicated that the types are located in Paris; however, we were unable to locate this specimen and did not find evidence that it was deposited there previously. The scarce morphological evidence for the monotypic genera Qiyunia Song and Xu 1989 and Tandil Mello-Leitão, 1940 suggest placing these species outside Argyronetidae stat. reinst., Lathyidae fam. n., and Dictynidae s.s.; however, despite our efforts we were unable to view the material. Based on this argument, we consider the taxa mentioned in this paragraph as incertae sedis .

Transferred to other families: Aebutina Simon, 1892 ( Macrobunidae ).

In synonymy: Clitistes velutinus Simon, 1902 = Cybaeolus pusillus Simon, 1884 .

Nomina dubium . Hoplolathys Caporiacco, 1947 . List of included genera: Adenodictyna Ono, 2008 , Ajmonia Caporiacco, 1934 [ Aj. aurita Song and Lu, 1985, Aj. bedeshai (Tikader, 1966) , Aj. capucina (Schenkel, 1936) , Aj. changtunesis (Hu, 2001) comb. n., Aj. gratiosa (Simon, 1881) , Aj. lehtineni Marusik and Koponen 1998 , Aj. marakata (Sherriffs, 1927) , Aj. numidica (Denis, 1937) , Aj. patellaris (Simon, 1911) , Aj. procera (Kulczyński, 1901) , Aj. psittacea (Schenkel, 1936) , Aj. rajaeii Zamani and Marusik 2017 , Aj. smaragdula (Simon, 1905) , and Aj. velifera (Simon, 1906) ], Anaxibia Thorell, 1898 , Arangina Lehtinen, 1967 , Archaeodictyna Caporiacco, 1928 [ Archaeodictyna aguasverdes (Wunderlich, 2022) comb. n., Archaeodictyna ammophila ( Menge, 1871) , Archaeodictyna anguiniceps (Simon, 1899) , Archaeodictyna bispinosa (Simon, 1906) comb. n., Archaeodictyna condocta (O. Pickard-Cambridge, 1876) , Archaeodictyna consecuta (O. Pickard-Cambridge, 1872) , Archaeodictyna fuerteventurensis (Schmidt, 1976) comb. n., Archaeodictyna lanzarotensis (Wunderlich, 2022) comb. n., Archaeodictyna minutissima (Miller, 1958) , Archaeodictyna sexnotata (Simon, 1890) , Archaeodictyna suedicola (Simon, 1890) , Archaeodictyna tazzeiti (Denis, 1954) , and Archaeodictyna ulova Griswold and Meikle-Griswold, 1987 ], Arethyna gen. n. [ Arethyna coloradensis (Chamberlin, 1919) comb. n., Arethyna idahoana (Chamberlin and Ivie, 1933) comb. n., Arethyna osceola ( Chamberlin and Gertsch, 1958) comb. n., Arethyna peon ( Chamberlin and Gertsch, 1958) comb. n., Arethyna personata (Gertsch and Mulaik, 1936) comb. n., Arethyna saltona ( Chamberlin and Gertsch, 1958) comb. n., Arethyna secuta (Chamberlin, 1924) comb. n., Arethyna sierra (Chamberlin, 1948) comb. n., Arethyna ubsunurica ( Marusik and Koponen, 1998) comb. n., Arethyna volucripes (Keyserling, 1881) comb. n., and Arethyna volucripes volucripoides (Ivie, 1947) comb. n.], Argennina Gertsch and Mulaik, 1936 , Atelolathys Simon, 1892 , Banaidja Lehtinen, 1967 , Brigittea Lehtinen, 1967 [ B. avicenna Zamani and Marusik, 2021 , B. civica (Lucas, 1848) , B. colona (Simon, 1906) comb. n., B. innocens (O.Pickard-Cambridge, 1872) , B. latens (Fabricius, 1775) , B. vicina (Simon, 1873) ], Califorenigma gen. n. [ C. linsdalei ( Chamberlin and Gertsch, 1958) comb. n.], Callevophthalmus Simon, 1906 , Dictyna Sundevall, 1833 [ Dictyna abundans Chamberlin and Ivie, 1941 , D. alaskae Chamberlin and Ivie, 1947 , D. albicoma Simon, 1893 , D. albovittata Keyserling, 1881 , D. alyceae Chickering, 1950 , D. apacheca Chamberlin and Ivie, 1935 , D. arundinacea , D. brevitarsus Emerton, 1915 , D. cafayate Mello-Leitão, 1941 , D. chandrai Tikader, 1966 , D. cofete Wunderlich, 2022 , D. columbiana Becker, 1886 , D. cronebergi Simon, 1889 , D. crosbyi Gertsch and Mulaik, 1940 , D. dauna Chamberlin and Gertsch, 1958 , D. ectrapela (Keyserling, 1886) , D. fluminensis Mello-Leitão, 1924 , D. guineensis Denis, 1955 , D. hamifera Thorell, 1872 , D. kosiorowiczi Simon, 1873 , D. laeviceps Simon, 1911 , D. linzhiensis Hu, 2001 , D. livida (Mello-Leitão, 1941) , D. marilina Chamberlin, 1948 , D. moctezuma Gertsch and Davis, 1942 , D. namulinensis Hu, 2001 , D. navajoa Gertsch and Davis, 1942 , D. pictella Chamberlin and Gertsch, 1958 , D. pusilla Thorell, 1856 , D. quadrispinosa Emerton, 1919 , D. ranchograndei Caporiacco, 1955 , D. saepei Chamberlin and Ivie, 1941 , D. similis Keyserling, 1878 , D. simoni Petrunkevitch, 1911 , D. sinaloa Gertsch and Davis, 1942 , D. siniloanensis Barrion and Litsinger, 1995 , D. tarda Schmidt, 1971 , D. togata Simon, 1904 , D. tristis Spassky, 1952 , D. trivirgata Mello-Leitão, 1943 , D. tullgreni Caporiacco, 1949 , D. turbida Simon, 1905 , D. uncinata Thorell, 1856 , D. uvs Marusik and Koponen, 1998 , D. vittata Keyserling, 1883 , D. vultuosa Keyserling, 1881 , and D.yongshun Yin, Bao and Kim, 2001 ], Dictynomorpha Spassky, 1939 , Emblyna Chamberlin, 1948 [ Emblyna acoreensis Wunderlich, 1992 , E. aiko ( Chamberlin and Gertsch, 1958) , E. altamira (Gertsch and Davis, 1942) , E. ampla Chamberlin, 1948 , E. angulata (Emerton, 1915) , E. annulipes (Blackwall, 1846) , E. ardea ( Chamberlin and Gertsch, 1958) , E. artemisia (Ivie, 1947) , E. borealis (O.Pickard-Cambridge, 1877) , E. borealis cavernosa (Jones, 1947) , E. branchi ( Chamberlin and Gertsch, 1958) , E. brevidens (Kulczyński, 1897) , E. budarini Marusik, 1988 , E. burjatica (Danilov, 1994) , E. callida (Gertsch and Ivie, 1936) , E. capens Chamberlin, 1948 , E. cavata (Jones, 1947) comb. n., E. chitina ( Chamberlin and Gertsch, 1958) , E. completa (Chamberlin and Gertsch, 1929) , E. completoides (Ivie, 1947) , E. consulta (Gertsch and Ivie, 1936) , E. cornupeta (Bishop and Ruderman, 1946) , E. coweta ( Chamberlin and Gertsch, 1958) , E. crocana Chamberlin, 1948 , E. decaprini (Kaston, 1945) , E. evicta (Gertsch and Mulaik, 1940) , E. florens (Ivie and Barrows, 1935) , E. formicaria Baert, 1987 , E. hentzi (Kaston, 1945) , E. horta (Gertsch and Ivie, 1936) , E. hoya (Chamberlin and Ivie, 1941) , E. joaquina ( Chamberlin and Gertsch, 1958) , E. lina (Gertsch, 1946) , E. linda ( Chamberlin and Gertsch, 1958) , E. manitoba (Ivie, 1947) , E. marissa ( Chamberlin and Gertsch, 1958) , E. melva ( Chamberlin and Gertsch, 1958) , E. nanda ( Chamberlin and Gertsch, 1958) , E. oasa (Ivie, 1947) , E. palomara Chamberlin, 1948 , E. pinalia ( Chamberlin and Gertsch, 1958) , E. piratica (Ivie, 1947) , E. peragrata (Bishop and Ruderman, 1946) , E. reticulata (Gertsch and Ivie, 1936) , E. roscida (Hentz, 1850) , E. saylori (Chamberlin and Ivie, 1941) , E. scotta Chamberlin, 1948 , E. seminola ( Chamberlin and Gertsch, 1958) , E. shasta ( Chamberlin and Gertsch, 1958) , E. shoshonea ( Chamberlin and Gertsch, 1958) , E. stulta (Gertsch and Mulaik, 1936) , E. sublatoides (Ivie and Barrows, 1935) , E. suwanea (Gertsch, 1946) , and E. zaba (Barrows and Ivie, 1942) ], Eriena gen. n. [ Er. minuta (Emerton, 1888) comb. n., Er. mora ( Chamberlin and Gertsch, 1958) comb. n.], Helenactyna Benoit, 1977 , Khalotyna gen. n. [ K. calcarata (Banks, 1904) comb. n.], Kharitonovia Esyunin , et al., 2017, Mallos O. Pickard-Cambridge, 1902 , Marilynia Lehtinen, 1967 , Mashimo Lehtinen, 1967 , Mexitlia Lehtinen, 1967 , Myanmardictyna Wunderlich, 2017 , Nigma Lehtinen, 1967 , Nopalityna gen. n. [ N. francisca (Bishop and Ruderman, 1946) comb. n., N. jonesae (Roewer, 1955) comb. n., N. orbiculata (Jones, 1947) comb. n., N. sublata (Hentz, 1850) comb. n., N. suprenans ( Chamberlin and Ivie, 1935) comb. n., and N. uintana (Chamberlin, 1919) comb. n.], Paradictyna Forster, 1970 , Pangunus gen. n. [ Pangunus kaszabi ( Marusik and Koponen, 1998) comb. n., Pangunus umai (Tikader, 1966) comb. n., and Pangunus xizangensis (Hu and Li, 1987) comb. n.], Penangodyna Wunderlich, 1995 , Phantyna Chamberlin, 1948 [ Ph. agressa (Ivie, 1947) comb. n., Ph. bicornis (Emerton, 1915) , Ph. estebanensis (Simon, 1906) , Ph. formidolosa (Gertsch and Ivie, 1936) comb. n., Ph. mandibularis (Taczanowski, 1874) , Ph. meridensis (Caporiacco, 1955) , Ph. micro (Chamberlin and Ivie, 1944) , Ph. mulegensis (Chamberlin, 1924) , Ph. pixi ( Chamberlin and Gertsch, 1958) , Ph. provida (Gertsch and Mulaik, 1936) , Ph. remota (Banks, 1924) , Ph. rita (Gertsch, 1946) , Ph. segregata (Gertsch and Mulaik, 1936) , Ph. terranea (Ivie, 1947) , Ph. varyna miranda ( Chamberlin and Gertsch, 1958) , and Ph. varyna ( Chamberlin and Gertsch, 1958) ], Purplecorna gen. n. [ Pu. gloria (Chamberlin and Ivie, 1944) comb. n., Pu. guerrerensis (Gertsch and Davis, 1937) comb. n., Pu. incredula (Gertsch and Davis, 1937) comb. n., Pu. lecta (Chickering, 1952) comb. n., Pu. meditata (Gertsch, 1936) comb. n., Pu. miniata ( Banks, 1898) comb. n., and Pu. terrestris (Emerton, 1911) comb. n.], Rhion O. Pickard-Cambridge, 1871 , Shango Lehtinen, 1967 , Shikibutyna gen. n. [ Sh. felis (Bösenberg and Strand, 1906) comb. n., Sh. foliicola (Bösenberg and Strand, 1906) comb. n., Sh. guanchae (Schmidt, 1968) comb. n., Sh. mongolica ( Marusik and Koponen, 1998) , comb. n., Sh. procerula (Bösenberg and Strand, 1906) comb. n., Sh. schmidti (Kulczyński, 1926) comb. n., Sh. szaboi (Chyzer, 1891) comb. n., Sh. wangi (Song and Zhou, 1986) comb. n., and Sh. zherikhini (Marusik 1988) comb. n.], Simziella gen. n. [ Si. annexa (Gertsch and Mulaik, 1936) comb. n., Si. canadas (Wunderlich, 2022) comb. n., Si. cebolla (Ivie, 1947) comb. n., Si. dunini (Danilov, 2000) comb. n., Si. major (Menge, 1869) comb. n., Si. palmgreni ( Marusik and Fritzén, 2011) comb. n., Si. paramajor (Danilov, 2000) comb. n., Si. sancta (Gertsch, 1946) comb. n., Si. sotnik (Danilov, 1994) comb. n., Si. sylvania (Chamberlin and Ivie, 1944) comb. n., Si. tridentata (Bishop and Ruderman, 1946) comb. n., Si. tucsona (Chamberlin, 1948) comb. n., Si. tyshchenkoi (Marusik, 1988) comb. n., Si. tyshchenkoi wrangeliana (Marusik, 1988) comb. n., and Si. teideensis (Wunderlich, 1992) comb. n.], Spagnius gen. n. [ Sp. albopilosa (Franganillo, 1936) comb. n., Sp. foliaceus (Hentz, 1850) comb. n., Sp. jacalana (Gertsch and Davis, 1937) comb. n., and Sp. nebraska (Gertsch, 1946) comb. n.], Sudesna Lehtinen, 1967 , Tahuantina Lehtinen, 1967 , Thallumetus Simon, 1893 , Tivyna Chamberlin, 1948 [ Ti. moaba (Ivie, 1947) , Ti. pallida (Keyserling, 1887) , Ti. petrunkevitchi (Gertsch and Mulaik, 1940) , Ti. sonora (Gertsch and Davis, 1942) comb. n., and Ti. spatula (Gertsch and Davis, 1937) ], Tolkienus gen. n. [ To. armatus (Thorell, 1875) comb. n., To. bellans (Chamberlin, 1919) comb. n., To. bellans hatchi (Jones, 1948) comb. n., To. estoc sp. n., To. longispina (Emerton, 1888) comb. n., and To. ottoi ( Marusik and Koponen, 2017) comb. n.], and Viridictyna Forster, 1970 .

Genus Arethyna Cala-Riquelme gen. n. urn:lsid:zoobank.org:act:28788E69-D7C0-4F87-8527-0129BC37E9DA .

Type species: Dictyna volucripes Keyserling, 1881 .

Material examined: Dictyna coloradensis : USA, Colorado, El Paso Co., Colorado Springs, pre-1919 publication date, Coll: R. V . Chamberlin (Holotype, 1male, MCZ-IZ15151 ) . USA, Colorado, Electra Lake , 1.vii.1919, Coll. F.E. Lutz (1 female, AMNH). Dictyna secuta : Mexico, Baja California, Gulf of California, San Estaban Island, 19.iv.1921, Coll. Joseph C. Chamberlin, California Academy of Sciences Expedition (Holotype, MCZ-IZ15594 ). Dictyna volucripes : USA, Massachusetts, [no specific locality data], pre-1881 publication date, [(type status needs research), 1 male, MCZ-IZ23523 ] . USA, Massachusetts (no specific locality data), pre-1881 publication date, [(type status needs research), 1 female, MCZ-IZ43955 ] .

Etymology: The generic epithet is derived from ‘Areth-’, in honour of Magdalena Aretha Louise Franklin (Aretha Franklin), an American singer, songwriter and pianist with over 75 million records sold worldwide and 112 singles on the US Billboard charts, and -yna, a rhyming morph of the last two syllables of Dictyna , another genus in the family; gender feminine.

Diagnosis: Males of Arethyna gen. n. ( Figs 65 A-F 66A-C) resemble Dictyna s.s. by the thin embolus without distal modifications that converge inside the conductor ( Figs 65D–F, 66D–E; Chamberlin and Gertsch 1958: pl. 26, figs 7, 10), and Simziella gen. n. by the embolus coiled 275°, originating proximo-prolateral from the bulb ( Fig. 65D–F, 66D–E; Chamberlin and Gertsch 1958: pl. 26, figs 1, 5, 8, 10, 12); however, it can be distinguished from Dictyna s.s., Simziella gen. n., and other Dictynidae s.s. by the conductor upper arm longer than the conductor lower arm, thin, coiled no more than 90°, and separated from the tegulum by its width ( Fig. 65D–F; Chamberlin and Gertsch 1958: pl. 26, figs 1, 5, 8, 10, 12); the conductor lower arm coiled no more than 45°, with the tip directed retrolateral-proximal, the paraconductor process conspicuous ( Fig. 66F), and with the conductor locking mechanism as long as conductor tip; dictynid process conspicuous, shorter than the tibia length ( Fig. 65D–F; Chamberlin and Gertsch 1958: pl. 26, figs 3, 6, 9, 13). Females can be distinguished by having a conspicuous membranous sac; the copulatory duct forming loops close to the copulatory opening, and the SS/AG.

List of included species: Arethyna coloradensis (Chamberlin, 1919) comb. n., Ar. idahoana (Chamberlin and Ivie, 1933) comb. n., Ar. osceola ( Chamberlin and Gertsch, 1958) comb. n., Ar. peon ( Chamberlin and Gertsch, 1958) comb. n., Ar. personata (Gertsch and Mulaik, 1936) comb. n., Ar. saltona ( Chamberlin and Gertsch, 1958) comb. n., Ar. secuta (Chamberlin, 1924) comb. n., Ar. sierra (Chamberlin, 1948) comb. n., Ar. ubsunurica ( Marusik and Koponen, 1998) comb. n., Ar. volucripes (Keyserling, 1881) comb. n., and Ar. volucripes volucripoides (Ivie, 1947) comb. n.

Genus Califorenigma Cala-Riquelme, Gorneau, Esposito gen. n.

urn:lsid:zoobank.org:act:88E0C01A-AE03-4F70-A1E3-5896FAE6A299 .

Type species: Heterodictyna linsdalei Chamberlin and Gertsch, 1958 , currently Nigma linsdalei ( Chamberlin and Gertsch, 1958) .

Type material: Nigma linsdalei : USA, California, Monterey Co., Hastings Natural History Reserve , 16.vii.1940, Coll. J.M. Linsdale (Holotype, 1 male, AMNH) . USA, California, Monterey Co., Hastings Natural History Reserve , 16.vii.1940, Coll. J.M. Linsdale (Allotype, 1 female, AMNH) .

Material examined: USA, California, Monterey Co., Hastings Natural History Reserve , 36°22'N 121°33'W, date unknown, Coll.J.M.Linsdale (1male, 1 female, 1imm., CASENT9122646 ) . USA, California, Alameda Co., Berkeley, Lawrence Radiation Laboratory , 4.v.1964, Coll. P. R. Craig (1 male, CASENT9122758 ) . USA, California, Alameda Co., Berkeley, Lawrence Radiation Laboratory , 3.vi.1964, Coll. P. R. Craig (1 male, CASENT9122590 ) . USA, California, Santa Barbara Co., Channel Islands Research Station , 14.vi.1964, Coll. D. Rentz, D. Weissman (1 male, CASENT9122674 ) . USA, California, Santa Barbara Co., Channel Islands Research Station , 21.v.1964, Coll. D. Rentz, D. Weissman (1 male, CASENT9122730 ) . USA, California, San Mateo Co., Edgewood Park (NE), 26.iv.1992, Coll. D. Ubick, mixed woodland, shaking oak foliage (1 male, CASENT9122730 ) . USA, California, San Francisco Co., San Francisco, Golden Gate Park 37°46'14"N 122°27'47"W, 19.v.2023. Coll. A.M. Al-Jamal, J.A. Gorneau, K. O. Montana (1 male, CASENT9112276 ) .

Etymology: The genus epithet is a combination of California, where the genus is endemic, and ‘enigma’, a play on words referencing the enigmatic features of this species as well as Nigma , the name of the genus in which the type species was formerly placed; gender feminine.

Diagnosis: The male of Califorenigma gen. n. ( Figs 27A–F, 48A–D, 49A–D, 50A–F, 51A, B) resembles Nigma by lacking the dictynid process ( Figs 28F, 51A, B); however, it can be distinguished from Nigma and other Dictynidae s.s. by having a spermaphor loop ( Chamberlin and Gertsch 1958: pl. 10, fig. 12) (spermaphor loop absent in Ajmonia and Nigma ); end of the conductor coiled in the same plane, perpendicular to the longitudinal axis of the bulb, with the conductor scaly tip three times longer than wide ( Figs 28A–D, 50E; Chamberlin and Gertsch 1958: pl.10, fig. 12); cymbium with a dorso-posterior extension, conformed by the cavity of a dorsal extension of the cymbium ( Figs 27E, 49B–D, 51A, B; Chamberlin and Gertsch 1958: pl. 10, fig. 9) (present in Dictynomorpha , see: Esyunin et al. 2017); patella convex with a conspicuous, long lobe on the retrolateral side ( Figs 27E, 49A, C, D, 50E, 51B; Chamberlin and Gertsch 1958: pl. 10, figs 9, 12, Griswold et al. 2005: fig. 176A). Females can be distinguished by having a slightly elevated area on the epigyne with a pair of round, very shallow and indistinct atria separated by a broad septum ( Chamberlin and Gertsch 1958: pl. 10, fig. 11, Esyunin et al. 2017).

Remarks: The genus Nigma is a substitute name for Heterodictyna Dahl, 1924 . Esyunin et al. (2017), based on Wunderlich (2011), argued that N. conducens (O. Pickard-Cambridge, 1876) , N. laeta (Spassky, 1952) , N.longipes (Berland, 1914) , and N. linsdalei have been incorrectly placed within the genus Nigma . We found sufficient evidence to conclude that N. linsdalei is a new genus here proposed by us as Califorenigma gen. n. Heterodictyna linsdalei was initially considered Ajmonia or Dictynomorpha by Lehtinen (1967: 210). Some morphological traits of the male palp of C. linsdalei ( Chamberlin and Gertsch, 1958) comb. n. are similar to those of Ajmonia capucina (Schenkel, 1936) , A. lehtineni Marusik and Koponen, 1998 , and A. psittacea (Schenkel, 1936) ; however, these can be differentiated by the shape and configuration of the patellar process (see: Chamberlin and Gertsch 1958: 48, figs 9, 12, Griswold et al. 2005: fig. 176 A, Esyunin et al. 2017), the modification of the palpal tibial apophysis ( Esyunin et al. 2017), and the presence of a developed tegular spermaphor loop. Brignoli (1983: 515) did not accept Lehtinen’s proposal and transferred the species to Nigma . Although the WSC (2024) formally accepts N. linsdalei as a valid name, Esyunin et al. (2017) indicated certain morphological characteristics that differentiate the species from other congeners of Nigma s.s.. As mentioned in the etymology section, this monotypic genus is endemic to California, generally found within 75 km of the coast. The majority of endemic arachnids in California are either Opiliones Sundevall, 1833 (especially the Phalangodidae Simon, 1879 ) or spiders in Mygalomorphae Pocock, 1892 ( Harrison 2013). This work adds Califorenigma to the endemic Araneomorphae genera in California, which also includes the endemic genera Lutica Marx, 1891 ( Zodariidae Thorell, 1881 ) and Titiotus Simon, 1897 ( Zoropsidae Bertkau, 1882 ) ( Harrison 2013).

List of included species: Califorenigma linsdalei ( Chamberlin and Gertsch, 1958) comb. n.