Emblyna Chamberlin, 1948

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) : -

Genus Emblyna Chamberlin, 1948 s.s.

Type species: Dictyna completa Chamberlin and Gertsch, 1929 , currently Emblyna completa (Chamberlin and Gertsch, 1929) .

Type material: USA, Utah, Moab , 15.iv.1928, Coll. A.M. Woodbury (Holotype, 1 female, AMNH) .

Material examined: Emblyna altamira : USA, Massachusetts, Duxbury , 4.vi.21, Coll. E.B. Bryant (1male, MCZ-IZ20606 ) . Emblyna andesiana : Ecuador, Piñán , 1.i–32. xii.1903, Coll. P. Rivet (Lectotype, 1 male, MNHN-AR-AR5292) . Ecuador, Tungurahua Prov., Ambato , 9–14.vi.1943, Coll. H. Exline-Frizzell., D.L. Frizzell (3 males, 19 females, CASENT9118818 ) . Emblyna angulata : USA, Massachusetts, Hyde Park (no verbatim date data), probably 1900–15, Coll. James H. Emerton (Syntype, 1 male, MCZ-IZ20279 ) . Emblyna annulipes : USA, Washington, Olympia (no verbatim date data) (1 male, MCZ-IZ29974 ) . Emblyna borealis cavernosa : USA, Washington, Spokane , vi.45, Coll. E.D. Parmer (Holotype, 1 male, MCZ-IZ20743 ) . Emblyna completa : USA, California, Plumas Co., Sierra Valley Marsh, Elev. 4900 ft., pitfall traps, 12.viii.1976, Coll. K. Richardson (1 male, CASENT9118615 ) . USA, Nevada, Reno , vi.1940, Coll. R. V. Chamberlin (1 male, AMNH) . USA, Nevada, Reno , vi.1940, Coll. R. V. Chamberlin = (1 female, AMNH) . Emblyna cruciata : USA, Connecticut, New Haven , pre-1888 publication date, Coll. James H. Emerton (Syntype, 1 male, MCZ-IZ151460 ) . Emblyna hentzi : USA, Michigan, St. Clair , 3.viii.36, Coll. M.H. Hatch (Exline Coll.) (1 male, MCZ-IZ20983 ) . USA, Michigan, St. Clair , 3.viii.36, Coll. M.H. Hatch (Exline Coll.) (1 female, MCZ-IZ24761 ) . Emblyna maxima : USA, New York, Ithaca , 1888–90 (from publication), Coll. Nathan Banks (Holotype, 1 female, MCZ-IZ22012 ) . Emblyna olympiana : USA, Washington, Edmunds Lake , 2.vi.35, Coll. M.H. Hatch (Exline Coll.) (1 female, MCZ-IZ24762 ) . USA, Washington, Edmunds Lake , 2.vi.35, Coll. M.H. Hatch (Exline Coll.) (1 male, MCZ-IZ21254 ) . USA, Washington, Olympia , [no verbatim date data] (Holotype, 1 male, MCZ-IZ29975 ) . Emblyna roscida : USA, Connecticut, New Haven. 15.v.pre-1888 publication date, Coll. James H. Emerton (1 male, MCZ-IZ22935 ) .

Diagnosis: Males of Emblyna s.s. ( Figs 58D, 59A–F) can be distinguished from other Dictynidae s.s. by having a thick palpal embolus that is distally modified ( Figs 58D, 59E, F; Chamberlin and Gertsch 1958: pl. 22, figs 2, 3, 8, 12) (embolus thin and unmodified in Dictyna s.s.); the dictynid process conspicuous ( Fig. 59E, F; Chamberlin and Gertsch 1958: pl. 42, fig. 4) (dictynid process absent in Tivyna and Mallos ); conductor lower arm with a paraconductor process in addition to the conductor scaly tip, coiled no more than 360° ( Figs 58D, 59E). Females can be distinguished by the wide copulatory duct, usually up to four times longer than the primary spermatheca, connected to an enlarged membranous sac; the primary spermathecae are reduced, as wide as or slightly wider than the widest section of the copulatory duct.

List of included species: 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. peragrata (Bishop and Ruderman, 1946) , E. pinalia ( Chamberlin and Gertsch, 1958) , E. piratica (Ivie, 1947) , 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) .

Genus Eriena Cala-Riquelme, Crews, Esposito gen. n.

u:lsid:zoobank.org:act:A178E265-F734-4 CA 1-rnA42E- 1A3215EA6CE6.

Type species: Dictyna minuta Emerton, 1888 .

Type material: USA, Rhode Island, Providence , 21.viii.1871, Coll. James H. Emerton (Holotype, 1 male, MCZ-IZ22095 ) .

Etymology: The generic epithet refers to the Indigenous Erie people, also known as the Eriechronon, Yenresh, Erielhonan, Eriez, Nation du Chat, and Riquéronon, historically living on the south shore of Lake Erie in the Northeastern USA; gender feminine.

Diagnosis: The males of Eriena gen. n. can be distinguished from other Dictynidae s.s. by having a short, unmodified embolus that originates prolateral distal on the bulb ( Chamberlin and Gertsch 1958: pl. 19, fig.7) (embolus thick and distally modified in Emblyna s.s.); the dictynid process with tubercle inconspicuous or absent ( Chamberlin and Gertsch 1958: pl. 19, figs 5, 6) (dictynid process absent in Tivyna , Mallos , Ajmonia , Dictynomorpha , Nigma , and Califorenigma gen. n.); conductor lower arm three or more times longer than the conductor upper arm, coiled no more than 180°, and with a long and thin conductor scaly tip, ( Chamberlin and Gertsch 1958: pl. 19, fig. 4). Females can be distinguished by having the primary spermathecae slightly wider to two times wider than the copulatory duct length, connected to a membranous sac two times as wide as the primary spermathecae.

List of included species: Eriena minuta (Emerton, 1888) comb. n. and E. mora ( Chamberlin and Gertsch, 1958) comb. n.

Genus Khalotyna Cala-Riquelme, Alequín, Esposito gen. n.

urn:lsid:zoobank.org:act:43122FE3-C797-48C6-8291-DC4C1EDB6CD6 .

Type species: Dictyna calcarata Banks, 1904 .

Type material: USA, California, Los Angeles Co., San Pedro, pre- 1904 publication date, Coll. T . D.A. Cockerell, Nathan Banks (Holotype, 1 male, MCZ-IZ20638 ) .

Material examined: Dictyna calcarata : USA, Texas, Brewster Co., Big Bend, Boquillas Canyon , 9.vii.2021, Coll. M. Pleisher (6 females, CASENT9087232 ) . USA, Arizona, Pinal Co., Gila R. Management Area ‘ The Shores’ , 2 km N Winkelman, 33°10.26′N 110°44.32′W, Elev. 600 m, dry outwash, malaise, 17.iv.2015, Coll. M.E. Irwin (1 male, CASENT9087249 ) . USA, California, Los Angeles Co., 1–31.x.1965, Coll. F. Delsue (1 male, CASENT9118632 ) . USA, California, Los Angeles Co., Long Beach , 25.vii.1963, Coll. W.D. Stockton (1 female, CASENT9118640 ) . Mexico, Baja California, Cañon de Guadalupe, 32°09.28′N 115°47.42′W, Elev. 370 m, upper canyon near pool with reeds, palms, 23–31.iii.2014, Coll. M.E. Irwin, M.J. Sharkey (1 male, 4 females, 1 imm., CASENT9118644 ) . Same as previous, Elev. 380 m, malaise, upper canyon, damp sandy wash (3 males, 3 females, 5 imm., CASENT9118654 ) . USA, California, Kern Co., Delano , 35°46'12.0"N 119°12'36.0"W, 18.iv.2010, Coll. D. P. Carroll (1 male, CASENT9056276 ) . USA, Arizona, Cochise Co., 1429 Franklin Street , 31°24′23′′N 109°55′57′′S, Elev. 1585 m, malaise, 7–18.v.2014, Coll. A.S. Menke (1 male, CASENT9058902 ) . USA, Arizona, Pima Co., Vail, Mountain Creek Ranch , 32°04.99′N 110°39.56′W, Elev. 1100 m, malaise, small dry wash, 18–30.iv.2014, Coll. M.E. Irwin (1 male, CASENT9118569 ) . Mexico, Yucatan, Izamal , 1–31.vii.1981, Coll. C. Gold (1 female, CASENT9118814 ) .

Etymology: The genus epithet is named in honour of Magdalena Carmen Frida Khalo y Calderon (Frida Khalo), a Mexican painter known for employing a naïve folk art style to explore questions of identity in Mexican society. It is combined with – yna, a rhyming morph of the last two syllables of Dictyna where the type species was originally placed; gender feminine.

Diagnosis: Males of Khalotyna gen. n. ( Fig. 64A–F) can be distinguished from other Dictynidae s.s. by having the conductor upper arm three or more times longer than the retrolateral arm, following the path of the tegulum prolaterally, extending well beyond it apically ( Fig. 64D, F; Chamberlin and Gertsch 1958: pl. 17, fig. 1) (present in Purplecorna gen. n.; absent in Dictyna s.s.; Emblyna s.s.); embolus thin, originating from the proximal to slightly prolateral part of the bulb, coiled 360° ( Fig. 64D; Chamberlin and Gertsch 1958: pl. 17, fig. 1, 3); conductor scaly tip directed retrolaterally ( Fig. 64D; Chamberlin and Gertsch 1958: pl. 17, fig. 1); the dictynid process with tubercle two or more times longer than the tibia ( Fig. 64D, E; Chamberlin and Gertsch 1958: pl. 17, figs 1, 4). Females can be distinguished from other congeners by having a looped copulatory duct that is ten or more times longer than the primary spermathecae width; SS/AG conspicuous.

List of included species: Khalotyna calcarata (Banks, 1904) comb. n.

Genus Nopalityna Cala-Riquelme and Esposito gen. n.

urn:lsid:zoobank.org:act:69F50B66-7676-403A-B08D-DED9A01D1869 .

Type species: Theridion sublatum Hentz, 1850, currently Emblyna sublata (Hentz, 1850) .

Type material: USA, Virginia, Falls Church (no verbatim date data), Coll. Nathan Banks (Neotype, 1 male, 1 female, MCZ-IZ23186 ) .

Material examined: Dictyna francisca : USA, California, Siskiyou Co., Juanita Lake Campground, 23.55 km SW Dorris, 41°49.077'0"N 122°07.440'0"W, Elev. 1500 m, dry second growth coniferous forest, 9.viii.2008, Coll. F. Álvarez Padilla, A. Carmichael, D. Dimitrov., C. Griswold, G. Hormiga and A. Saucedo (1 female, CASENT9031870 ). Emblyna sublata : USA, New York, Ithaca, 1888–1890, April and in summer (from publication), Coll. Nathan Banks (1 male, 3 females, MCZ-IZ21022 ) . USA, New York, Ithaca , 1888–90 (from publication), Coll. Nathan Banks (1 female, MCZ-IZ21132 ). Emblyna uintana : USA, Utah, Uintah Mountains, Chalk Creek , pre-1919 publication date, Coll. R. V . Chamberlin (1 male, 1 female, MCZ-IZ25183 ) . USA, Utah, Uintah Mountains, Chalk Creek , pre-1919 publication date, Coll. Ralph V . Chamberlin (Holotype, 1 female, MCZ-IZ15154 ) .

Etymology: The generic name refers to nopalitos, diced prickly pear cactus, a gastronomic treasure of Mexico, with the addition of the – yna suffix, a rhyming morph of the last two syllables of Emblyna where the type species was originally placed; gender feminine.

Diagnosis: Males of Nopalityna gen. n. ( Figs 29A–C, 58F) can be distinguished from other Dictynidae s.s. by the embolus with one or two long branches, usually flattened, that converge inside the conductor, unmodified at the tip ( Figs 29A–C, 58F; Chamberlin and Gertsch 1958: pl. 39, figs 3, 4, 7, 8, 11, 12) (embolus, thick and distally modified in Emblyna s.s.); a wide conductor, usually inflated at the base ( Figs 29A–C, 58F; Chamberlin and Gertsch 1958: pl. 39, figs 1, 5, 6, 9, 10, 13); and the dictynid process with a short to inconspicuous tubercle ( Chamberlin and Gertsch 1958: pl.39, figs 3, 4, 7, 8, 11, 12) (dictynid process absent in Tivyna , Mallos , Ajmonia , and Shango ). Females can be distinguished by having a developed, globose, membranous extension of the proximal copulatory duct ( Figs 16D–F; Chamberlin and Gertsch 1958: pl. 38, figs 7, 8); the copulatory duct uncoiled; and the primary spermathecae elongated.

List of included species: Nopalityna 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., N. uintana (Chamberlin, 1919) comb. n.

Genus Pangunus Cala-Riquelme gen. n.

urn:lsid:zoobank.org:act:7E62AF72-3860-48F9-8D50-BB9A06D04A55 .

Type species: Emblyna kaszabi Marusik and Koponen, 1998 .

Material examined: Emblyna kaszabi : Mongolia, Uldzit Somon , 23.viii.1975, Coll. J. Halgos (Holotype, 1 male, SMF332 ; Paratype, 3 female, SNM) .

Etymology: The generic epithet refers to Pangu (Pan-Koo), a Chinese mythology and Taoism figure; gender masculine.

Diagnosis: Males of Pangunus gen. n. can be distinguished from other Dictynidae s.s. by the embolus coiled approximately 720°, originating proximal to proximal-prolateral from the bulb ( Marusik and Koponen 1998: 80, figs 10, 11, Marusik et al. 2006: 355, fig. 6).

List of included species: Pangunus kaszabi ( Marusik and Koponen 1998) comb. n., P. umai (Tikader, 1966) comb. n., P. xizangensis (Hu and Li, 1987) comb. n.

Genus Purplecorna Cala-Riquelme and Esposito gen. n.

urn:lsid:zoobank.org:act:6B4199FC-50F8-44B5-875F-

484F841FF54A.

Type species: Dictyna incredula Gertsch and Davis, 1937 .

Material examined: Dictyna incredula : Peru, Paita Prov., Amotape, Chira R. Valley , 21.x.1938, Coll. D.L. Frizzell, H. Exline-Frizzell (2 males, 7 females, 5 imm., CASENT9118816 ) . Peru, Piura Region, Sullana, N Mallares, Chira R, 4.i.1942, Coll. D. Frizzell, H. Frizzell (10 males, 9 females, 8 imm., CASENT9118828 ) . Ecuador, from bananas, 4.i.1942, Coll. unknown (1 male, 1 female, CASENT9118830 ) . Peru, Piura Region, Sullana, Mallares , 31.xi.1941, Coll. D.L. Frizzell., H. Exline-Frizzell (2 males, 6 females, CASENT9118832 ) . Ecuador, Guayas Prov., Guayaquil , 15.iii.1943, Coll. Carvallio (from Mello-Leitao) (1 male, CASENT9118851 ) . Dictyna lecta : Panama, El Volcan , viii.1950, Coll. Arthur M. Chickering (Holotype, 1 male, MCZ-IZ21815 ) . Dictyna meditata : Cuba, La Habana Province, Havana, 1880-publication data, Coll. Baker, Nathan Banks (1 male, 1 female, MCZ-IZ21321 ) . Dictyna miniata : Mexico, Baja California Sur, Gulf of California , Monserrate Island, 25.v.1921, Coll.Joseph C. Chamberlin, California Academy of Science Expedition (1 male, MCZ-IZ15596 ) . Dictyna terrestris : USA, New Hampshire, Lake Winnipesaukee, Three Mile Island , 1.vi.1909, Coll. James H. Emerton (Syntype, 1 male, MCZ-IZ23261 ) .

Etymology: The generic name refers to purple corn (in Spanish: maiz colorado or maiz morado), a variety originating in South America; gender feminine.

Diagnosis: Males of Purplecorna gen. n. ( Figs 54A–C; 55A, B) can be distinguished from other Dictynidae s.s. by the cymbium narrow at the base, long and thin, sometimes strongly curved ventrally, and narrowed apically to a slender finger-like structure ( Figs 54D–F, 55C, D; Chamberlin and Gertsch 1958: pl. 18, figs 6, 7); the conductor upper arm longer than the conductor lower arm, following the cymbium prolaterally, extending well beyond it apically ( Figs 54D–F, 55C, D; Chamberlin and Gertsch 1958: pl. 18, figs 6, 7); embolus thin with the tip unmodified, coiled 360°, originating proximally to slightly prolaterally from the bulb ( Figs 54D–F, 55C, D). The female can be distinguished from other genera by the copulatory duct usually with many loops or coils, thin, but gradually widening near the primary spermathecae, more than 10 times longer than the primary spermathecae width; with the membranous sac as long as or slightly longer than the complete copulatory duct.

List of included species: Purplecorna gloria (Chamberlin and Ivie, 1944) comb. n., P. guerrerensis (Gertsch and Davis, 1937) comb. n., P. incredula (Gertsch and Davis, 1937) comb. n., P. lecta (Chickering, 1952) comb. n., P. meditata (Gertsch, 1936) comb. n., P. miniata ( Banks, 1898) comb. n., P. terrestris (Emerton, 1911) comb. n.

Genus Shikibutyna Cala-Riquelme, Gorneau, Esposito gen. n. urn:lsid:zoobank.org:act:5D966098-FBF3-4A39-ACF2-2E806B896EE1 .

Type species: Dictyna felis Bösenberg and Strand, 1906 .

Material examined: Dictyna felis : Japan, Yokohama , 15.x.1945, Coll. T . Aarons (1 female, CASENT9118672 ) .

Etymology: The generic epithet is in honour of Murasaki Shikibu, a female Japanese writer and poet known worldwide for being the author of Genji Monogatari , the first novel in Japanese history and the first modern novel in the world, written in the early 11th century. The name includes the suffix – yna referring to the last two syllables of Dictyna , the genus in which the type species was originally placed; gender feminine.

Diagnosis: The male of Shikibutyna gen. n. ( Figs 56A–F, 57A) resembles Purplecorna gen. n. by having a narrow, long, and thin cymbium ( Figs 56D–F, 57A); the embolus tip is unmodified, coiled 360°, originating proximally to slightly prolaterally on the bulb ( Figs 56D–F, 57A); however, it can be distinguished from Purplecorna gen. n. and other congeners by having a dictynid process ( Fig. 56D–F) (dictynid process absent in Tivyna , Mallos , Ajmonia , and Shango ); the embolus thick, following a curved path without loops ( Figs 56D–F, 57A) (embolus thin and forming a loop close to the embolus base in Purplecorna gen. n., embolus thin in Dictyna s.s.); the conductor upper arm two or three times longer than the conductor lower arm ( Figs 56D, E, 57A). The female can be distinguished from other congeners by having the copulatory duct between the copulatory opening and SS/AG thin and uncoiled; and the copulatory duct between the SS/AG and primary spermathecae wider than the primary spermathecae.

List of included species: Shikibutyna felis (Bösenberg and Strand, 1906) comb. n., S. foliicola (Bösenberg and Strand, 1906) comb. n., S. guanchae (Schmidt, 1968) comb. n., S. mongolica ( Marusik and Koponen, 1998) comb. n., S. procerula (Bösenberg and Strand, 1906) comb. n., S. schmidti (Kulczyński, 1926) comb. n., S. szaboi (Chyzer, 1891) comb. n., S. wangi (Song and Zhou, 1986) comb. n., S. zherikhini (Marusik, 1988) comb. n.

Genus Simziella Cala-Riquelme and Alequín , gen. n.

urn:lsid:zoobank.org:act:8F3C1BE1-313C-4830-BFEC-

47 DBBB 87B2F2.

Type species: Dictyna major Menge, 1869 .

Material examined: Dictyna major : USA, Washington, Olympia (no verbatim date data), Coll. Nathan Banks (1 male, MCZ-IZ15970 ) . USA, California, Claremont , ~1920, Coll. Unknown (2 males, 3 females, CASENT9129252 ) . USA, Idaho, NE Fruitland, 30.vi.1943, Coll. Wilton Ivie (12 males, 24 females, CASENT9129369 ) . Dictyna tridentata : USA, Colorado, Long’s Peak Inn, Canadian Zone , vii.1919, Coll. Theodore D.A. Cockerell, Nathan Banks (1 male, MCZ-IZ20427 ) .

Etymology: The generic epithet refers to the Romanian fairy tale Ileana Simziana known in English as The Princess Who Would Be a Prince (1872–86) by Petre Ispirescu; gender feminine.

Diagnosis: Males of Simziella gen. n. ( Figs 29D–F, 67A) resemble Arethyna gen. n. and Kharitonovia by the embolus coiled 275°, originating proximo prolateral from the bulb ( Figs 29E, 68D, F); however, it can be distinguished by the conductor upper arm close to the tegulum ( Figs 29D, E; 68D); the conductor lower arm with the tip directed retrolateral or slightly retrolateral-proximal [e.g. S. sancta (Gertsch, 1946) comb. n.], without a paraconductor, and by the conductor locking mechanism shorter than conductor tip ( Figs 29F, 68D, E). Females ( Figs 15A–F, 67B–F, 68A–C) resemble Arethyna gen. n., Emblyna s.s., and Nopalityna gen. n. by having the membranous sac as long as/or longer than the copulatory duct; however, it can be distinguished from these and other congeners by the C-shape of the copulatory duct close to the copulatory opening, and the digitiform SS/AG.

List of included species: Simziella 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.

Genus Spagnius Cala-Riquelme and Crews , gen. n.

urn:lsid:zoobank.org:act:1 EACFB 82-2BAD-4185-9871-

39C23B05FEF7.

Type species: Theridion foliaceum Hentz, 1850, currently Dictyna foliacea (Hentz, 1850) .

Material examined: Dictyna foliacea : USA, New York, Tompkins Co., Mundy Wildflower Garden, Cornell University , 42°27'00.5"N 76°28'08.1"W, 23.vi.2021, Coll. J. Gorneau (1 female, CASENT9103498 ) . Russia, Primorskiy Krai, Ussuriysk Dist, Gornotayozhnoye , 43°39'36.0"N 132°15'00.0"E, pitfall traps, yellow pan trap, 11–12.vi.2000, Coll. M. V . Michailovskaya (1 male, CASENT9118689 ) . USA, Alabama, pre-1947 publication date, Coll. Banks, wild caught (Neotype, 1 male, 1 female, MCZ-IZ21350 ) . USA, Connecticut, New Haven , 1881, Coll. James H. Emerton (1 male, MCZ-IZ21401 ) .

Etymology: The generic epithet is in honour of our late friend and colleague Dr. Joseph Spagna, without whom this project would not have been possible; gender masculine.

Diagnosis: Males of Spagnius gen. n. resemble Eriena gen. n. by having a short, unmodified embolus that originates prolateral distal on the bulb ( Chamberlin and Gertsch 1958: pl. 19, fig. 10); and the dictynid process with the tubercle inconspicuous ( Chamberlin and Gertsch 1958: pl. 19, fig. 13); however, it can be distinguished from Eriena gen. n. and other Dictynidae s.s. by the tibia three times longer than patella length ( Chamberlin and Gertsch 1958: pl. 19, fig. 13); conductor lower arm as long as the conductor upper arm, coiled no more than 180°, and with a conductor scaly tip, long and thin with a paraconductor process at the base; dictynid process with the ctenidia variable in size, the largest hook-shaped ( Chamberlin and Gertsch 1958: pl. 19, fig. 13). Females can be distinguished by having the copulatory duct approximately as wide as the primary spermathecae, folded once, close to the copulatory opening, and three to four times longer than the primary spermatheca width (shorter in Eriena gen. n.).

List of included species: Spagnius albopilosa (Franganillo, 1936) comb. n., Sp. foliacea (Hentz, 1850) comb. n., Sp. jacalana (Gertsch and Davis, 1937) comb. n., Sp. nebraska (Gertsch, 1946) comb. n.

Genus Tolkienus Cala-Riquelme, Crews, Esposito gen. n. urn:lsid:zoobank.org:act:5C0D5A64-21F0-4C93-862F-B04400A9F196 .

Type species: Dictyna longispina Emerton, 1888 .

Type material: USA, Connecticut, Meriden, pre-1888 publ. date, Coll. James H. Emerton (Syntype, 1 male, 1 female, MCZ-IZ151459 ) .

Material examined: Dictyna bellans : USA, Alabama, Auburn , pre-1947 publication date, Coll. Baker, Nathan Banks (1 female, MCZ-IZ24751 ) . USA, Alabama, Auburn , pre- 1947 publication date, Coll. Baker, Nathan Banks (1 male, MCZ-IZ21258 ) . USA, Alabama, Auburn , pre-1947 publication date, Coll. Baker, Nathan Banks (2 males, 2 females, MCZ-IZ24752 ) . USA, Mississippi, Canton , pre-1919 publication date, Coll. R. V. Chamberlin (Holotype, 1 male, MCZ-IZ15152 ) . Dictyna bellans hatchi : USA, Oregon, Roseburg , 14.vi.1938, Coll. M.H. Hatch (Exline Coll.) (Syntype, 1 male, MCZ-IZ24764 ) . USA, Oregon, Roseburg , vii.1939, Coll. M.H. Hatch (Exline Coll.) (Syntype, 1 female, MCZ-IZ21526 ) . Dictyna longispina : USA, Arkansas, Washington Co., University farm, low vegetation, open, 11.vii.1955, Coll. W. Peck (1 male, 2 females, 1 imm., CASENT9118614 ) . USA, Arkansas, Washington Co., University farm, low vegetation open field, 12–13.vi.1965, Coll. W. Peck (1 female, CASENT9118618 ) .

Etymology: The generic epithet is in honour of John Ronald Reuel Tolkien, an English writer known worldwide for The Hobbit (1937) and The Lord of the Rings (1954–55); gender masculine.

Diagnosis: Males of Tolkienus gen. n. ( Figs 57C, 60A–F, 62A–D; Marusik and Koponen 2017: fig. 1C, D) resemble Khalotyna gen. n. and Phantyna by having the dictynid process with a tubercle as long as or longer than the tibia length ( Figs 57C, 60D, 62A– D; Chamberlin and Gertsch 1958: pl. 16, figs 1, 3, Marusik and Koponen 2017: fig. 2B, C); however, it can be distinguished from other Dictynidae s.s. by the thin embolus originating distally to slightly prolaterally on the bulb, coiled no more than 180° ( Fig. 57C; 62A; Chamberlin and Gertsch 1958: pl. 16, fig. 1); conductor lower arm longer than upper arm, coiled twice, with a conspicuous conductor paraterminal process in addition to the conductor scaly tip ( Figs 57C, 60B). The females ( Fig. 61A–F; Marusik and Koponen 2017: fig. 1A, B) can be distinguished by the AG/SS wider than the copulatory duct, copulatory duct folded once close to the copulatory opening ( Fig.61E, F; Marusik and Koponen 2017: fig. 1F, G), up to three times longer than the primary spermathecae width; and the membranous sac as long as or slightly longer than the complete copulatory duct.

List of included species: Tolkienus armatus (Thorell, 1875) comb. n., T. bellans (Chamberlin, 1919) comb. n., T. bellans hatchi (Jones, 1948) comb. n., T. estoc sp. n., T. longispinus (Emerton, 1888) comb. n., T.ottoi ( Marusik and Koponen 2017) comb. n.

Tolkienus estoc Cala-Riquelme and Al-Jamal sp. n.

urn:lsid:zoobank.org:act:ECA1AA36-C947-426E-BA32-

90269208EFE2.

Type material: Male holotype ( CASENT9118701 ) from Equatorial Guinea, Bioko Prov., Moka Wildlife Sanctuary, 3°21′46′′N 8°39′52′′E, Elev. 1400 m, 2–10.x.1998, Coll. D.K. Dabney, D. Ubick, beating/sweeping foliage; 4 males and 1 female paratypes ( CASENT9118701 ), same data as holotype . Male paratype ( CASENT9118720 ), same data as holotype .

Material examined: Bioko prov. , Moka Wildlife Sanctuary , 3°22'0"N 8°39'57′"E, 1500 m, beating/sweeping foliage, 6–10.x.1998, Coll. D.K. Dabney, D. Ubick (3 males, 1 female, CASENT9118702 ). Bioko prov. , Moka Wildlife Sanctuary , 3°22'0"N 8°39'57"E, 1500 m, beating/sweeping foliage, 6–10.x.1998, Coll. D.K. Dabney, D. Ubick (3 males, 1 female, CASENT9118702 ). Bioko prov., Moka Wildlife Sanctuary, Elev. 1403–1419 m, 26.x–1.xi.2007, Coll. R . H. Pine (1 male, FMNH-INS0000085077 ) .

Etymology: The species’ epithet is a noun in apposition referring to an estoc sword (tuck in English), a French variation of the longsword used from the 14th to the 17th centuries.

Diagnosis: Males of Tolkienus estoc sp. n. ( Figs 57C, 60A–F, 62A–D) can be distinguished from other Tolkienus gen. n. by having the conductor tip directed proximally and as long as the paraconductor process ( Figs 57C, 62B). Females ( Fig. 61A–F) can be distinguished from other congeners by having the membranous sac as wide as the primary spermathecae, and the SS/ AG in an ectal-medial position ( Fig. 61F).

Description: Male ( CASENT 9118701, Equatorial Guinea): Carapace ( Fig. 60A–D) gallstone yellow, anteriorly directed white setae around eyes, darker branching patterns on lateral surfaces of prosoma. Chelicerae, labium, endites, and sternum honey yellow. Legs pale honey yellow. Abdomen ( Fig. 60A) dorsum broccoli brown, with snow white guanine crystals in fused medial chevron pattern, and a large, square patch of similar crystals at posterior end, venter pale broccoli brown. Sternum slender and squared-off anteriorly, slightly recurved. Total length 2.25. Carapace length 0.88, width 0.80, height 0.50. Clypeus height 0.3. Eye diameters and interdistances: AME 0.03, PME 0.04, ALE 0.05, PLE 0.03, AME–AME 0.08, PME– PME 0.11. Sternum length 0.53, width 0.45. Palp: femur 0.25, tibia 0.14 (dictynid process 0.38). Leg I: femur 0.95, patella 0.25, tibia 0.80, metatarsus 0.59, tarsus 0.40. II: 0.90, 0.25, 0.75, 0.59, 0.40. III: 0.64, 0.25, 0.50, 0.50, 0.31. IV: 0.76, 0.25, 0.56, 0.54, 0.26. Abdomen: length 1.33, width 0.68. Male palp ( Figs 57C, 62A–D) with femur straight, shorter than patella + tibia length; patella two times shorter than tibia; tibia shorter than dictynid process, with a small proximal retrolateral dorsal process in addition to the RTA; conductor lower arm two times longer than conductor upper arm length, coiled 360°, with a paraconductor process in addition to the conductor scaly tip; embolus, connected prolateral distally to the tegulum, and coiled no more than 180°.

Female ( CASENT 9118701, Equatorial Guinea): Carapace ( Fig. 61A–D) pale honey yellow, anteriorly directed white setae around eyes. Chelicerae, labium, endites and sternum pale honey yellow. Legs cream-yellow. Abdomen ( Fig. 61B–D) dorsum cream-yellow but slightly darker dorsolaterally, with snow white guanine crystals distributed broadly and visible laterally, though obscured in dorsal view due to condition of specimen, venter cream-yellow. Sternum squared-off anteriorly, slightly procurved. Total length 1.95. Carapace length 0.78, width 0.58, height 0.33. Clypeus height 0.19. Eye diameters and interdistances: AME 0.04, PME 0.03, ALE 0.05, PLE 0.03, AME–AME 0.02, PME–PME 0.05. Sternum length 0.45, width 0.38. Palp: femur length 0.21, tibia length 0.10. Leg I: femur 0.64, patella 0.21, tibia 0.45, metatarsus 0.39, tarsus 0.31. II: 0.61, 0.25, 0.38, 0.35, 0.28. III: 0.49, 0.21, 0.30, 0.33, 0.26. IV: 0.58, 0.21, 0.40, 0.43, 0.26. Abdomen: length 1.28, width 0.93. Cribellum length 0.2. Epigyne ( Fig. 61E, F): copulatory opening ectal, separated by six times the diameter of SS/AG, anterior to the primary spermathecae; membranous sac eight or nine times longer and three times wider than spermathecae diameter; copulatory duct uncoiled, with copulatory duct receptacle as wide as primary spermathecae, and connected ectal proximal with the primary spermathecae; SS/AG reduced but conspicuous, as long as copulatory duct receptacle; fertilization duct located mesal proximally.