Novalena Chamberlin & Ivie 1942

Maya-Morales, Julieta & Jiménez, María Luisa, 2017, Revision of the funnel-web spider genus Novalena (Araneae: Agelenidae), Zootaxa 4262 (1), pp. 1-88 : 5-7

Genus Novalena Chamberlin & Ivie 1942

Type species. Novalena intermedia (Chamberlin & Gertsch 1930) by original designation ( Chamberlin & Ivie 1942).

Composition. Fifty-three species: N. annamae (Gertsch & Davis 1940) , N. approximata (Gertsch & Ivie 1936) , N. attenuata (F.O. Pickard-Cambridge 1902) , N. bipartita (Kraus 1955) , N. bipunctata Roth 1967 stat. rev., N. calavera Chamberlin & Ivie 1942 , N. costata (F.O. Pickard-Cambridge 1902) , N. intermedia (Chamberlin & Gertsch 1930) , N. laticava (Kraus 1955) , N. lutzi (Gertsch 1933) , N. orizaba (Banks 1898) , N. shlomitae (García- Villafuerte 2009), N. simplex (F.O. Pickard-Cambridge 1902) , N. ajusco sp. nov., N. alamo sp. nov., N. alvarezi sp. nov., N. atzimbo sp. nov., N. bosencheve sp. nov., N. chamberlini sp. nov., N. cieneguilla sp. nov., N. cintalapa sp. nov., N. clara sp. nov., N. comaltepec sp. nov., N. creel sp. nov., N. dentata sp. nov., N. divisadero sp. nov., N. durango sp. nov., N. franckei sp. nov., N. garnica sp. nov., N. gibarrai sp. nov., N. irazu sp. nov., N. iviei sp. nov., N. ixtlan sp. nov., N. jiquilpan sp. nov., N. leonensis sp. nov., N. mexiquensis sp. nov., N. oaxaca sp. nov., N.

paricutin sp. nov., N. perote sp. nov., N. plata sp. nov., N. poncei sp. nov., N. popoca sp. nov., N. prieta sp. nov., N. puebla sp. nov., N. punta sp. nov., N. rothi sp. nov., N. saltoensis sp. nov., N. sinaloa sp. nov., N. tacana sp. nov., N. triunfo sp. nov., N. valdezi sp. nov., N. victoria sp. nov., and N. volcanes sp. nov.

Diagnosis. The genus is diagnosed by the following characteristics in combination: the male pedipalp with a short and simple curved embolus ( Figs. 3 b, 13a, 18a, 32a, 43d); conductor with two projections ( Figs. 3 c, 10a, 14c, 27a, 41f); tegular median process present ( Figs. 10 a, 13a, 26a, 34d, 38a); RTA with distal and dorsal projections, ventral projection usually present ( Figs. 3 d, 11b, 16m, 30e, 33e) (absent in N. shlomitae , N. simplex , N. chamberlini , N. cintalapa , N. gibarrai , N. oaxaca , and N. tacana , Figs. 15 b, 18b, 32b); atrium posteriorly located on the epigynal plate, wider than long ( Figs. 5 a, 13c, 19c, 24d, 40a), and usually lacking a division (if divided [ N. calavera and N. lutzi ] then copulatory openings visible in posterior view, Figs. 7 e, 12e); atrial spurs usually present ( Figs. 5 d, 15c, 24a, 33a, 42c) (absent in N. calavera , N. lutzi , and N. garnica , Figs. 7 c, 12c, 26c); short and strongly sclerotized copulatory ducts, straight or curved ( Figs. 5 b, 14d, 20b, 36a, 38d), connected ectally to longer than wide primary spermathecae ( Figs. 5 e, 10b, 15d, 25d, 31d); secondary spermathecae blind-ended receptacles with primary pores and each connected to the union of copulatory duct and primary spermathecae by short diverticulum ( Figs. 10 f, 14f, 30g, 36b, 44e); and short fertilization ducts ( Figs. 2 d, 13d, 22d, 31d, 39d).

Novalena differs from Eratigena Bolzern et al. 2013 and Tegenaria Latreille 1804 in having strongly procurved eye rows in frontal view; from Agelenopsis Giebel 1869 , Barronopsis Chamberlin & Ivie 1941 , Melpomene , and Tortolena Chamberlin & Ivie 1941 in lacking both a strongly modified embolus and projecting tegular lateral process in males (= tegular apophysis, Stocks 2009: fig. 39), and lacking long, membranous and plicate copulatory ducts in females (as in Stocks 2009: figs. 52, 61); from Calilena Chamberlin & Ivie 1941 in lacking a membranous fulcrum (as in Chamberlin & Ivie 1941: figs. 68, 70, 74) (only present in N. annamae ) and an epigynal scape (as in Chamberlin & Ivie 1941: figs. 48–60); from Hololena (Chamberlin & Gertsch 1929) in lacking a fulcrum supporting the embolus (as in Chamberlin & Ivie 1942: figs. 6, 11, 12) and an anteriorly located epigynal atrium (as in Chamberlin & Ivie 1942: fig. 10); from Rothilena Maya-Morales & Jiménez 2013 in lacking three projections on conductor (as in Maya-Morales & Jiménez 2013: fig. 18) and spherical primary spermathecae (as in Maya-Morales & Jiménez 2013: fig. 29); from Rualena in lacking a membranous fulcrum (as in Maya- Morales & Jiménez 2016: fig. 43) and an atrium as long as wide (as in Maya-Morales & Jiménez 2016: fig. 30); and from Hoffmannilena in lacking a basal projection on RTA (as in Fig. 45 b, c) and a strongly sclerotized epigynal plate (as in Maya-Morales & Jiménez 2016: fig. 116).

Description. Medium-sized spiders, 4–14 mm total length. Eight eyes in two rows strongly procurved in frontal view ( Fig. 8 b). Carapace with two symmetrical longitudinal dark bands intensified by feathery scales and a black band around the border of thoracic region ( Fig. 1 c, i, r). Chelicerae with three promarginal teeth and two to four retromarginal teeth ( Fig. 8 c). Sternum longer than wide ( Fig. 1 f). Feathery scales present on carapace, opisthosoma, pedipalps, and legs ( Fig. 8 a, b). Opisthosoma oval with dorsal foliate pattern and/or posterior chevrons ( Fig. 1 b, k, s). Colulus divided with few hairs on each side ( Fig. 9 c). PLS longest with distal segment 0.5– 1.25 times basal ( Figs. 1 f, 9a). Legs formula usually 4123. Legs usually annulated ( Fig. 1 b–d, k). Patella I and II with two dorsal spines and one prolateral spine, patella III and VI with two dorsal spines, one prolateral spine, and one retrolateral spine. Leg tarsi with five to eight trichobothria. Capsulate tarsal organ distad of trichobothrial row ( Fig. 8 f). Pedipalp femur with two to four dorsal spines. Female pedipalp tibia usually with one + two prolateral spines. Male pedipalp tibia with two to four spines and with prolateroventral protuberance and retrolateroventral groove ( Fig. 10 e). RTA of male pedipalp with distal and dorsal projections; ventral projection usually present ( Fig. 21 d). Embolus a short and simple curve ( Fig. 14 a). Conductor with two short projections ( Fig. 3 c). Median apophysis spoon-shaped ( Fig. 10 a) (only reduced in N. alamo , Fig. 16 d). Tegular median process present ( Fig. 21 b). Fulcrum absent (only present in N. annamae , Fig. 2 a). Female epigynal plate wider than long ( Fig. 25 c). Atrium on posterior part of plate, usually wider than long ( Fig. 30 f). Atrial spurs usually present ( Fig. 5 a) (absent in N. calavera , N. lutzi , and N. garnica ). Internal genitalia with copulatory ducts strongly sclerotized, straight ( Fig. 23 a) or curved ( Fig. 14 d). Spermathecae bipartite ( Fig. 36 ); primary spermathecae longer than wide ( Fig. 32 d), secondary spermathecae blind-ended receptacles with primary pores ( Bennett 1992) and each connected to the union of copulatory duct and primary spermathecae by short diverticulum ( Fig. 14 b). Fertilization ducts short, originating from the posterior part of spermathecae ( Fig. 10 d).

Habitat. In North America, Novalena spiders are usually found in webs under loose bark of standing trees and stumps in dense coniferous forest, and occasionally in webs on the ground in shaded areas ( Roth & Brame 1972).

In tropical montane cloud forests of southeastern Mexico, they are found in webs within epiphytes and on understory vegetation and the ground.

Distribution. Novalena is widely distributed in the western Nearctic and northern Neotropics from southwestern Canada to Costa Rica and Trinidad and Tobago (Maps 1–8).

Figures

FIGURE 1. Novalena spp., habitus: a, e, N. approximata (Gertsch & Ivie 1936), female; b, f, N. alvarezi sp. nov., female; c, g, N. chamberlini sp. nov., male; d, h, N. clara sp. nov., female; i, m, N. dentata sp. nov., male; j, n, N. irazu sp. nov., female; k, o, N. iviei sp. nov., male; l, p, N. leonensis sp. nov., female; q, u, N. mexiquensis sp. nov., male; r, v, N. poncei sp. nov., female; s, w, N. puebla sp. nov., female; t, x, N. rothi sp. nov., female. a – d, i – l, q – t, dorsal view; e – h, m – p, v – x, ventral view. Scale bars: a, d, j, q, s, t = 4 mm; b, i, l, r = 3 mm; c, k = 2 mm.

FIGURE 2. Novalena annamae (Gertsch & Davis 1940): a, male pedipalp, ventral view; b, same, retrolateral view; c, epigynum, ventral view; d, internal genitalia, dorsal view; e, epigynum, posterior view. Abbreviations: A, atrium; C, conductor; CD, copulatory ducts; CO, copulatory openings; E, embolus; F, fulcrum; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process. Scale bars: a = 0.5 mm; e = 0.25 mm.

FIGURE 3. Novalena approximata (Gertsch & Ivie 1936), male pedipalp (SEM): a, ventral view; b, prolateral view; c, d, retrolateral view. Abbreviations: Cd, dorsal projection of conductor; Cv, ventral projection of conductor; E, embolus; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; St, subtegulum; T, tegulum; TMP, tegular median process.

FIGURE 5. Novalena spp.: a – c, Novalena attenuata (F. O. Pickard-Cambridge 1902); d – f, Novalena bipunctata Roth 1967 stat. rev. a, d, epigynum, ventral view; b, e, internal genitalia, dorsal view; c, f, epigynum, posterior view. Abbreviations: A, atrium; CD, copulatory ducts; CO, copulatory openings; FD, fertilization ducts; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae. Dashed line indicates distance between anterior and posterior margins. Scale bars: 0.5 mm.

FIGURE 7. Novalena spp. genitalia: a – e, Novalena calavera Chamberlin & Ivie 1942; f – h, Novalena costata (F. O. Pickard- Cambridge 1902). a, male pedipalp, ventral view; b, same, retrolateral view; c, f, epigynum, ventral view; d, g, internal genitalia, dorsal view; e, h, epigynum, posterior view. Abbreviations: A, atrium; C, conductor; CD, copulatory ducts; CO, copulatory openings; E, embolus; FD, fertilization ducts; H, hoods; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process. Scale bars: a = 1 mm; e, h = 0.5 mm.

FIGURE 8. Novalena intermedia (Chamberlin & Gertsch 1930), female (SEM): a, carapace; b, eyes; c, cheliceral fang; d, serrula; e, tarsus II; f, tarsal organ. a, d, f, dorsal view; b, frontal view; c, apical view; e, lateral view.

FIGURE 9. Novalena intermedia (Chamberlin & Gertsch 1930), female (SEM): a, spinnerets; b, anal tubercle; c, colulus and tracheal spiracle; d, ALS, details. Abbreviations: Cl, colulus; ALS, anterior lateral spinnerets; AT, anal tubercle; MAS, major ampullate glans spigots; PGS, piriform gland spigots; PLS, posterior lateral spinnerets; PMS, posterior median spinnerets; TH, thick hairs; TS, tracheal spiracle.

FIGURE 10. Novalena intermedia (Chamberlin & Gertsch 1930) (SEM): a, c, e, male pedipalp; b, d, f, internal female genitalia. a, c, ventral view; b, d, dorsal view; e, prolateral view, arrow to prolateral protuberance; f, anterior view. Abbreviations: CD, copulatory ducts; Cd, dorsal projection of conductor; Cv, ventral projection of conductor; E, embolus; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process.

FIGURE 14. Novalena simplex (F. O. Pickard-Cambridge 1902) (SEM): a, c, e, male pedipalp; b, d, f, internal female genitalia. a, ventral view; b, dorsal view; c, e, retrolateral view; d, f, anterior view. Abbreviations: CD, copulatory ducts; Cd, dorsal projection of conductor; Cv, ventral projection of conductor; E, embolus; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process.

FIGURE 15. Novalena simplex (F. O. Pickard-Cambridge 1902): a, male pedipalp, ventral view; b, same, retrolateral view; c, epigynum, ventral view; d, internal genitalia, dorsal view; e, epigynum, posterior view. Abbreviations: A, atrium; C, conductor; CD, copulatory ducts; E, embolus; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process. Scale bars: a = 0.5 mm; e = 0.25 mm.

FIGURE 16. Novalena spp. genitalia: a – c, Novalena ajusco sp. nov.; d, e, Novalena alamo sp. nov.; f – h, Novalena alvarezi sp. nov.; i – k, Novalena atzimbo sp. nov.; l, m, Novalena bosencheve sp. nov. a, f, i, epigynum, ventral view; b, g, j, internal genitalia, dorsal view; c, h, k, epigynum, posterior view; d, l, male pedipalp, ventral view; e, m, same retrolateral view. Abbreviations: A, atrium; C, conductor; CD, copulatory ducts; CO, copulatory openings; E, embolus; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process. Dashed line indicates distance between anterior and posterior margins. Scale bars: c, h, k = 0.25 mm; d, l = 0.5 mm.

FIGURE 21. Novalena dentata sp. nov., male pedipalp (SEM): a, c, ventral view; b, prolateral view; d, retrolateral view. Abbreviations: C, conductor; E, embolus; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; T, tegulum; TMP, tegular median process.

FIGURE 23. Novalena divisadero sp. nov. (SEM), internal female genitalia, dorsal view. Abbreviations: CD, copulatory ducts; FD, fertilization ducts; S 1, primary spermathecae; S 2, secondary spermathecae.

FIGURE 25. Novalena franckei sp. nov.: a, male pedipalp, ventral view; b, same, retrolateral view; c, epigynum, ventral view; d, internal genitalia, dorsal view; e, epigynum, posterior view. Abbreviations: A, atrium; C, conductor; CD, copulatory ducts; E, embolus; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process. Scale bars: a = 0.5 mm; e = 0.25 mm.

FIGURE 30. Novalena spp. genitalia: a – c, Novalena jiquilpan sp. nov.; d – h, Novalena leonensis sp. nov. a, f, epigynum, ventral view; b, g, internal genitalia, dorsal view; c, h, epigynum, posterior view; d, male pedipalp, ventral view; e, same, retrolateral view. Abbreviations: A, atrium; C, conductor; CD, copulatory ducts; CO, copulatory openings; E, embolus; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; RTAv, ventral projection of RTA; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process. Dashed line indicates distance between anterior and posterior margins. Scale bars: c, h = 0.25 mm; d = 0.5 mm.

FIGURE 32. Novalena oaxaca sp. nov.: a, male pedipalp, ventral view; b, same, retrolateral view; c, epigynum, ventral view; d, internal genitalia, dorsal view; e, epigynum, posterior view. Abbreviations: A, atrium; C, conductor; CD, copulatory ducts; E, embolus; FD, fertilization ducts; MA, median apophysis; RTAd, distal projection of RTA; RTAo, dorsal projection of RTA; S, spurs; S 1, primary spermathecae; S 2, secondary spermathecae; T, tegulum; TMP, tegular median process. Scale bars: a = 0.5 mm; e = 0.25 mm.

FIGURE 36. Novalena puebla sp. nov. (SEM), internal female genitalia: a, ectal view; b, dorsal view. Abbreviations: CD, copulatory ducts; S 1, primary spermathecae; S 2, secondary spermathecae.

FIGURE 45. Hoffmannilena nova (O. Pickard-Cambridge 1896) comb. nov., male pedipalp. a, prolateral view; b, ventral view; c, retrolateral view. Abbreviations: C, conductor; E, embolus; MA, median apophysis; RTAb, basal projection of RTA; RTAd, distal projection of RTA; T, tegulum; TMP, tegular median process. Scale bar: 0.25 mm.