Nerudia ola, Huber & Meng & Král & Ávila Herrera & Izquierdo & Carvalho, 2023

Huber, Bernhard A., Meng, Guanliang, Král, Jiří, Ávila Herrera, Ivalú M., Izquierdo, Matias A. & Carvalho, Leonardo S., 2023, High and dry: integrative taxonomy of the Andean spider genus Nerudia (Araneae: Pholcidae), Zoological Journal of the Linnean Society 198, pp. 534-591 : 566-570

NERUDIA OLA HUBER SP. NOV.

( FIGS 1G , 27–30 )

Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: BF76CDC2-9B0C-4B69-8DB9-DE6925D99DD4.

Nerudia Mich 20: Eberle et al., 2018 (molecular data: 28S). Huber et al., 2018: fig. 2.

D i a g n o s i s: E a s i l y d i s t i n g u i s h e d f r o m k n o w n congeners by armature of male chelicerae ( Fig. 27G, H ; distinctive pair of long frontal apophyses) and by shape of procursus ( Fig. 27A–C ; with bifid tip consisting of slender dorsal process and wider ventral membranous part); also by bulbal processes ( Fig. 27D–F ; ventral apophysis distally slender, curved towards ventral, same length as embolus) and by epigynum and female internal genitalia ( Figs 27I , 28 ; epigynal plate without posterior indentation; internal genitalia simple, with barely visible transparent ‘receptacle’).

Type material: ARGENTINA – San Juan: • ♂ holotype; San Agustín de Valle Fértil ; 30.6366° S, 67.4863° W; 880 m a.s.l.; under rocks near river; 5 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; LABRE-Ar 591 • 5 ♂♂, 2 ♀♀, paratypes; same data as holotype; ZFMK Ar 23897 .

Other material examined: ARGENTINA – San Juan: • 12 ♂♂, 8 ♀♀, 5 juvs, in pure ethanol (two males and two females used for SEM; three female prosomata used for molecular study); same data as holotype; ZFMK Arg 141 • 9 ♀♀, 3 juvs; same data as holotype; LABRE-Ar 560 • 1 ♂, in pure ethanol; same locality as holotype; 22 Jan. 2012; J. M. A. Navarro leg.; ZFMK Mich 20 • 3 ♂♂, 2 ♀♀; ~ 7.5 km S Astica; 31.0223° S, 67.2976° W; 865 m a.s.l.; under rocks near river; 4 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23898 • 1 ♂, 3 ♀♀, 5 juvs, in pure ethanol; same data as preceding; ZFMK Arg135, 136 • 1 ♂, 1 ♀; same data as preceding; LABRE-Ar 528 • 5 ♂♂, 7 ♀♀, 1 juv., in pure ethanol; same data as preceding; LABRE-Ar 529, 547, 548, 552 • 1 ♂, 1 ♀; Parque Provincial Ischigualasto; 30.1839° S, 67.9026° W; 1425 m a.s.l.; under rocks; 5 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23899 • 2 ♀♀, in pure ethanol; same data as preceding; ZFMK Arg144 • 1 ♀, 2 juvs; same data as preceding; LABRE-Ar 531 • 1 ♀ with eggs, in pure ethanol; same data as preceding; LABRE-Ar 554 • 1 ♂; Parque Provincial Ischigualasto; 30.1821° S, 67.9010° W; 1425 m a.s.l.; in dry riverbed, hand collecting at night; 19 Dec. 2018; M. Izquierdo, F. Bollatti, A. Albin, and C. Mattoni leg.; LABRE-Ar 445 • 1 ♂; same data as preceding; LABRE-Ar 444, prep. code MAI-4709 • 1 ♀; Baldecitos; 30.2232° S, 67.6942° W; 1255 m a.s.l.; stone wall of buildings, hand collecting at night; 21 Dec. 2018; M. Izquierdo, F. Bollatti, A. Albin, and C. Mattoni leg.; LABRE-Ar 448, prep. code MAI-4710 .

Assigned tentatiƲely (see Male Ʋariation beloae): La Rioja: • 1 ♂, 1 ♀; Cuesta de Miranda, ‘site 1’; 29.3511° S, 67.7924° W); 1960 m a.s.l.; under rocks; 8 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23900 • 2 ♀♀, in pure ethanol; same data as preceding; ZFMK Arg 152 • 2 ♂♂, 1 ♀; Cuesta de Miranda, ‘site 2’; 29.3468° S, 67.7205° W; 1660 m a.s.l.; under rocks; 8 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23901 • 2 ♀♀, in pure ethanol; same data as preceding; ZFMK Arg154 • 1 ♂, 2 ♀♀; same data as preceding; LABRE-Ar 533 • 1 ♀, in pure ethanol; same data as preceding; LABRE-Ar 553 • 1 ♀; between Chilecito and Famatina; 29.0027° S, 67.4855° W; 1300 m a.s.l.; 9 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23902 • 1 ♀; same data as preceding; LABRE-Ar 541 • 1 ♀ with eggs, in pure ethanol; same data as preceding; LABRE-Ar 546 • 4 ♂♂, 2 ♀♀; SE Aimogasta, ‘site 2’; 28.9015° S, 66.6538° W; 755 m a.s.l.; under rocks; 10 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23903 • 3 ♀♀, in pure ethanol; same data as preceding; ZFMK Arg 162 • 1 ♂, 1 ♀, 1 juv.; same data as preceding; LABRE-Ar 532 • 1 ♀, in pure ethanol; same data as preceding; LABRE-Ar 567 . Catamarca: • 2 ♂♂, 4 ♀♀; ~ 5 km NW Chumbicha, near Balneario El Caolín , ‘site 2’; 28.8109° S, 66.2500° W; 640 m a.s.l.; steep rock field in forest; 28–29 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23904 • 7 ♀♀, 3 juvs, in pure ethanol (three female prosomata used for molecular study); same data as preceding; ZFMK Arg218 • 2 ♂♂, 6 ♀♀, 8 juvs, in pure ethanol; same data as preceding; LABRE-Ar 530, 544, 561, 563, 566 • 3 ♀♀ with eggs, in pure ethanol; same locality as preceding, ‘site 1’; 28.8152° S, 66.2478° W; 605 m a.s.l.; M. A. Izquierdo and B. A. Huber leg.; LABRE-Ar 556 • 2 ♀♀, 8 juvs, in pure ethanol; same data as preceding; LABRE-Ar564 • 3 ♂♂, 2 ♀♀ (one male used for µ-CT study); ~ 10 km N Belén; 27.5641° S, 67.0058° W; 1380 m a.s.l.; in pile of stones; 25 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23905 • 5 ♀♀, 1 juv., in pure ethanol; same data as preceding; ZFMK Arg213 • 1 ♂, 7 ♀♀, 1 juv., in pure ethanol; same data as preceding; LABRE-Ar 545 • 11 ♂♂, 2 ♀♀ (three males and one female used for µ-CT study; two male abdomens used for karyotype study); near Nacimientos; 27.1559° S, 66.6925° W; 2120 m a.s.l.; under rocks on arid slope; 25 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23906 • 4 ♀♀, 3 juvs, in pure ethanol; same data as preceding; ZFMK Arg 211 .

Assigned tentatiƲely (only females aƲailable): ARGENTINA – La Rioja: • 2 ♀♀, 1 juv.; Cuesta de Miranda ; 29°21’ S, 67°43’ W; 1700 m a.s.l.; Aug. 1994; M. Ramírez leg.; MACN Ar 20055 • 1 ♀, in pure ethanol; SE Aimogasta , ‘site 1’; 28.8069° S, 66.6635° W; 915 m a.s.l.; under rocks; 10 Mar. 2019; B. A. Huber and M.A. Izquierdo leg.; ZFMKArg161 . Catamarca: • 1 ♀; ~ 14 km W Fiambalá ; 27.6590° S, 67.7607° W; 2000 m a.s.l.; under rocks; 26 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23907 • 2 ♀♀; Chumbicha ; 28.87° S, 66.23° W; 400 m a.s.l.; Aug. 1994; M.J. Ramírez leg.; MACN Ar 20012 . San Juan: • 1 ♀; ~ 35 km W Las Flores ; 30.3967° S, 69.5576° W; 2910 m a.s.l.; under rocks; 6 Mar. 2019; B. A. Huber and M. A. Izquierdo leg.; ZFMK Ar 23908 .

Etymology: The species epithet ola (Spanish for ‘wave’) is taken from Pablo Neruda’s poem ‘Poema 12’; noun in apposition.

Description

Male (holotype). Measurements: Total body length 1.55, carapace width 0.66. Distance PME–PME 70 µm; diameter PME 65 µm; distance PME–ALE 30 µm; distance AME–AME 20 µm; diameter AME 40 µm. Leg 1: 5.64 (1.48 + 0.25 + 1.45 + 1.78 + 0.68), tibia 2: 1.18, tibia 3: 1.03, tibia 4: 1.53; tibia 1 L/d: 22.

Colour (in ethanol): Prosoma and legs mostly pale ochre-yellow; carapace with light brown Y-mark behind ocular area; sternum whitish; legs without dark rings; abdomen monochromous pale grey.

Body: Habitus as in Figure 1G . Ocular area barely raised. Carapace with indistinct thoracic groove. Clypeus unmodified (only rim slightly more sclerotized than in female). Sternum wider than long (0.46/0.38), with pair of distinct anterior processes near coxae 1. Abdomen globular.

Chelicerae: As in Figure 27G, H ; with distinctive pair of long frontal apophyses, tips slightly bent downwards, obtuse in frontal view, with some ventral hairs directed downwards ( Fig. 29F ); distance between tips of apophyses: 280 µm; stridulatory files on distinct lateral protrusions ( Fig. 29C ).

Palps: In general, similar to N. colina (cf. Fig. 4 ) but femur slightly slenderer (length/width 2.27) and tibia slightly thicker (length/width 0.92); procursus simple, in lateral view slightly directed towards dorsal, with bifid tip consisting of slender dorsal process and wider ventral membranous process ( Fig. 27A–C ); genital bulb similar to N. colina , with slender ventral apophysis curved towards ventral ( Fig. 27D–F ).

Legs: Without spines and curved hairs; with vertical hairs in higher than usual density on proximal half of tibia 1 only, in two dorsal rows ( Fig. 30A ); retrolateral trichobothrium of tibia 1 at 58%; prolateral trichobothrium absent on tibia 1; tarsus 1 with six to seven pseudosegments, distally distinct.

Variation (male): Tibia 1 in 36 males (including holotype): 1.04–1.60 (mean 1.33). The distance between the tips of the cheliceral apophyses varies considerably (235–340 µm), but also within localities (e.g. Nacimientos: 240–310 µm). In northern specimens (from La Rioja and Catamarca), the cheliceral apophyses are slightly straighter and longer. This difference is minimal and some males appear intermediate. In specimens from Chumbicha , the procursus is slightly shorter. Abdomen variably dark. COI data indicate a deep split between northern and southern specimens, and a phylogenetic analysis of the molecular data (Supporting Information, Fig. S2 ) even questions the monophyly of the northern + southern clade. However , deep splits also occur among southern specimens and among northern specimens ( Fig. 2 ). As a result, a preliminary ASAP analysis (Supporting Information, Figs S3 , S 4 ) favours the existence of up to seven ‘species’ (lowest scores of 2.0 and 3.0). What is here interpreted as one species may thus in fact be several more or less cryptic species .

Female: In general, similar to male but sternum without pair of anterior humps and tibia 1 with usual low number of short vertical hairs. Tibia 1 in 57 females: 0.97–1.55 (mean 1.25). Epigynum ( Figs 28A , 30F ) anterior plate weakly protruding, without posterior indentation; posterior plate large, simple. Internal genitalia ( Figs 27I , 28B–D ) simple, with barely visible transparent receptacle. No difference could be seen between northern and southern specimens.

Distribution: Widely distributed in Argentina, Provinces San Juan, La Rioja and Catamarca ( Fig. 3 ). Note that specimens from La Rioja and Catamarca are assigned tentatively.

Natural history: All specimens were found by turning rocks. The spiders started to run rapidly when disturbed but then stopped suddenly and did barely ever drop from the rock. They were found in a variety of habitats, ranging from a relatively humid block field in a low forest at 640 m a.s.l. near Chumbicha to an exposed arid hill with dry ravines at 2120 m a.s.l. near Nacimientos. At Valle Fértil , the spiders seemed to have a patchy distribution, with up to eight adult specimens on a single large rock in suitable places (with some shade and leaf litter among the rocks). They shared the microhabitat with a superficially similar species of Metagonia Simon, 1893 (M. ‘MACN79’; undescribed; Valle Fértil ; Chumbicha), with Gertschiola macrostyla (Astica; Valle Fértil ; Ischigualasto; SE Aimogasta ‘site 2’), with Chibchea araona Huber, 2000 (?) (Cuesta de Miranda, both sites) and, in at least one case, with another species of Nerudia ( N. poma ; Chumbicha).

Figures

Figure 1. Nerudia, live specimens from Argentina. A, N. colina sp. nov., female from Jujuy, between San Salvador and Purmamarca (type locality). B, N. poma sp. nov., male from Salta, NW Campo Quijano (type locality). C, D, N. centaura sp. nov., male and female from Catamarca, E Paso de San Francisco (type locality). E, N. trigo sp. nov., female from Salta, SW Alemania. F, N. guirnalda sp. nov., male from Catamarca, El Rodeo (type locality). G, N. ola sp. nov., male from San Juan, San Agustin de Valle Fertil (type locality). H, N. nono sp. nov., male from Cordoba, E Nono (type locality).

Figure 2. NJ COI tree of all sequenced Nerudia and Gertschiola specimens. Species groups on the right are working hypotheses. Photo: female of Nerudia colina sp. nov. For complete tree, and for tree resulting from analysis with IQ-TREE, see Supporting Information, Figures S1, S2.

Figure 3. Known distribution of Nerudia in South America (inset) and of individual species. Regions in Chile: AT, Atacama; COQ, Coquimbo. Provinces in Argentina: CA, Catamarca; COR, Córdoba; JU, Jujuy; LR, La Rioja; SA, Salta; SJ, San Juan.

Figure 4. Nerudia colina sp. nov.; paratype male from Argentina, Jujuy, between San Salvador and Purmamarca (ZFMK Ar 23883). Left palp, prolateral (A) and retrolateral (B) views. Abbreviations: b, genital bulb; c, coxa; e, embolus; f, femur; p, procursus; pa, patella; ta, tarsus; ti, tibia; tr, trochanter; va, ventral apophysis. Scale line: 0.3 mm.

Figure 27. Nerudia ola sp. nov.; paratype male and paratype female from Argentina, San Juan, San Agustín de Valle Fértil (ZFMK Ar 23897). A–C, left male palpal tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F, left male genital bulb, prolateral, dorsal, and retrolateral views. G, H, male chelicerae, frontal and lateral views. I, cleared female genitalia, dorsal view. Scale lines: 0.2 mm.

Figure 28. Nerudia ola sp. nov.; paratype female from Argentina, San Juan, San Agustín de Valle Fértil (ZFMK Ar 23897). A, abdomen and epigynum, ventral view. B, median structures in internal genitalia. C, D, cleared genitalia, ventral and dorsal views. Scale lines: 0.2 mm (A, C, D), 0.1 mm (B).

Figure 29. Nerudia ola sp. nov.; male and female from Argentina, San Juan, San Agustín de Valle Fértil (ZFMK Arg141). A, female prosoma, frontal view. B, left procursus tip, prolateral-dorsal view. C, stridulatory file on male chelicera. D, lateral view of female chelicera, showing absence of stridulatory file. E, stridulatory pick (modified hair) on male palpal femur. F, right male cheliceral apophysis, frontal view. G, processes on male sternum. H, male gonopore. Scale lines: 100 µm (A, G), 10 µm (B, D, F, H), 20 µm (C), 2 µm (E).

Figure 30. Nerudia ola sp. nov.; male and female from Argentina, San Juan, San Agustín de Valle Fértil (ZFMK Arg141). A, male right tibia 1, retrolateral view. B, male metatarsus 4 with trichobothrium. C, male palpal tarsal organ. D, tip of right female palp, dorsal view; arrow points at tarsal organ. E, tarsal organ on male tarsus 2. F, epigynum, ventral view. G, male ALS and PMS. H, female ALS. Scale lines: 10 µm (A, B, D, G, H), 2 µm (C, E), 100 µm (F).