Nerudia, HUBER, 2000

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 : 540-543

publication ID 10.1093/zoolinnean/zlac100

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NERUDIA HUBER, 2000 View in CoL View at ENA

Nerudia Huber, 2000: 87 View in CoL . Type species: Nerudia atacama Huber, 2000 View in CoL .

Diagnosis: Small (total body length 1.4–1.9) eighteyed pholcids with relatively long legs (compared to most other Ninetinae ; leg 1 usually> 3 × total body length; only shorter in the new species N. centaura ), with simple procursus (without dorsal flap, not strongly elongated), paired male cheliceral modifications (strong hairs or pair of apophyses), with stridulatory files on male chelicerae, with simple main (anterior) epigynal plate and large posterior epigynal plate, without or with simple (i.e. not tube-like) median receptacle-like structure in female internal genitalia.


Male. Measurements: Total body length 1.4–1.9, carapace width 0.6–0.8. Legs relatively long (compared to other Ninetinae ), tibia 1 usually ~1.1–2.0 (longer in the new species N. rocio , single male: 2.5); tibia 1 L/d 14–31; metatarsus 1 length ~1.0–1.2 × tibia 1 length; tibia 2 always shorter than tibia 4 (tibia 2/tibia 4: 0.8–0.9).

Colour: Live specimens pale ochre-grey to light brown ( Fig. 1 View Figure 1 ); carapace often with darker median mark; abdomen colour variable, often darker than prosoma, only in N. centaura distinctively lighter than prosoma ( Fig. 1C View Figure 1 ), without or with indistinct marks; legs without dark or light bands. Colour in ethanol similar but paler.

Body: Ocular area barely raised, eight eyes, AME relatively large (diameter: 35–50 µm, i.e. 60–85% of PME diameter). Carapace without or with indistinct thoracic groove (cf. Figs 7B View Figure 7 , 11A View Figure 11 , 29A View Figure 29 ). Clypeus usually barely modified, rim slightly more sclerotized than in female, in N. nono also more bulging than in female, in some species with pair of shallow indentations for genital bulbs at rest ( Figs 7A View Figure 7 , 11B View Figure 11 ). Sternum wider than long, usually with pair of variably distinct anterior processes near leg coxae 1, processes apparently without pores ( Fig. 29G View Figure 29 ); sternum processes absent in N. centaura . Abdomen globular; four epiandrous spigots arranged in two pairs ( Figs 11F View Figure 11 , 29H View Figure 29 ); ALS with seven spigots each ( Fig. 30G View Figure 30 ): one strongly widened spigot, one long pointed spigot and five cylindrical spigots (one of which is unusually large); PMS with two short, pointed spigots ( Fig. 30G View Figure 30 ); PLS without spigots (cf. Fig. 12D View Figure 12 ).

Chelicerae: Usually with pair of simple frontal apophyses (e.g. Fig. 5G, H View Figure 5 ; absent in N. trigo ), tip often flat, i.e. wide in frontal view, pointed in lateral view ( Fig. 11C, D View Figure 11 ); often with patches or brushes of stronger hairs; with stridulatory files on variably distinct lateral protrusions ( Figs 11B, C View Figure 11 , 29C View Figure 29 ).

Palps: Coxa unmodified; trochanter with indistinct ventral projection; femur cylindrical, slightly widened distally, proximally without or with low retrolateral hump, with prolateral stridulatory pick (modified hair; Fig. 29E View Figure 29 ); patella short; tibia globular, with two trichobothria; palpal tarsal organ raised, capsulate ( Figs 7E View Figure 7 , 11E View Figure 11 , 30C View Figure 30 ), with small opening (diameter of opening ~1.6–1.9 µm); procursus simple, without dorsal flap, not strongly elongated, straight or bent towards dorsal, tip with complex cuticular microstructure ( Fig. 29B View Figure 29 ); genital bulb with simple ventral apophysis and dorsal embolus (e.g. Fig. 5F View Figure 5 ).

Legs: Without spines and curved hairs; usually with short vertical hairs in higher density on tibia 1 or tibiae 1 and 2 ( Figs 8A, B View Figure 8 , 30A View Figure 30 ; length of hairs ~20 µm). Trichobothria in usual arrangement: three on each tibia (except tibia 1: prolateral trichobothrium absent), one on each metatarsus, slightly feathered ( Fig. 30B View Figure 30 ); length of dorsal trichobothrium on tibia 1: ~80 µm; retrolateral trichobothrium of tibia 1 in distal position (at 58–70% of tibia length). Metatarsi and tarsi with dorsal rows of tiny pores with cuticular rim (diameter of opening ~0.6 µm; cf. Fig. 18C, D View Figure 18 ). Tarsus 1 with six to eight pseudosegments, distally distinct, proximally usually indistinct; tarsus 4 distally with one comb-hair on prolateral side ( Fig. 12F View Figure 12 ); leg tarsal organs small, capsulate ( Fig. 30E View Figure 30 ), with small opening (diameter of opening ~ 1.2–1.6 µm); three claws ( Figs 12E View Figure 12 , 18G View Figure 18 ).

Female: In general (size, colour, body shape), similar to male but chelicerae without stridulatory files ( Figs 7D View Figure 7 , 24F View Figure 24 , 29D View Figure 29 ), sternum without pair of anterior humps, palpal tarsal organ only weakly raised ( Figs 12A, B View Figure 12 , 24B View Figure 24 ), and leg tibiae with usual low number of short vertical hairs; legs either slightly shorter than in males or of same length (only in the new species N. ola with reasonable sample size: male/female tibia 1 length: 1.08). Spinnerets, leg pores, leg tarsal organs and comb-hairs as in male ( Figs 7F View Figure 7 , 8D–F View Figure 8 , 18B, H View Figure 18 , 24H View Figure 24 , 30E, H View Figure 30 ). Main (anterior) epigynal plate rectangular to trapezoidal or semi-circular, weakly protruding, posteriorly straight or with variably distinct median indentation (e.g. Figs 17A View Figure 17 , 32A View Figure 32 ); posterior plate simple, short but wide, usually wider than anterior plate. Internal genitalia simple, sometimes with distinct median receptacle-like structure (e.g. Figs 6B View Figure 6 , 10B View Figure 10 ), apparently without or with indistinct pair of pore plates (arrows in Figs 10B View Figure 10 , 14B View Figure 14 , 23B View Figure 23 ).

Relationships: The molecular analysis of Eberle et al. (2018) included only a single species of Nerudia (‘ N. Mich20 ’ = N. ola ), which was placed (with moderate support) as sister of the geographically close Gertschiola macrostyla (Mello-Leitão, 1941) . Both genera together were placed in a clade with further South American and Old World Ninetinae . The present paper is not primarily focused on phylogeny and our sampling does not include the type species N. atacama , but our COI and karyotype data support the sister-group relationship between Nerudia and Gertschiola (see Fig. 2 View Figure 2 ; Supporting Information, Figs S1 View Figure 1 , S 2 View Figure 2 ; and section on karyology). Preliminary analyses of Ninetinae relationships based on hybrid enrichment data (G. Meng et al., unpublished data) also support this sister-group relationship. Our new SEM data also confirm the position of Nerudia among Ninetinae (in particular the small opening of the tarsal organs).

In addition, the COI data suggest the existence of three species groups within Nerudia that we consider useful as working hypotheses: (1) the ola group, including the new species N. guirnalda , N. nono , N. ola and N. trigo ; (2) the centaura group, including the new species N. centaura , N. rocio and the undescribed N. ‘Arg23a’; and (3) the poma group, including the new species N. hoguera , N. poma and the undescribed N. ‘Arg163’. One new species, N. colina , appears isolated. The three groups appear neither supported nor contradicted by morphology. The type species N. atacama and the new N. flecha cannot be assigned with confidence to any of these groups (no fresh material suitable for sequencing is available of these two species).

Distribution: Nerudia iswidespreadintheArgentinean and Chilean Andes, ranging at least from 24°S to 31°S, and in the east up to the Córdoba mountain ranges in central Argentina ( Fig. 3 View Figure 3 ). See biogeographic analysis for further details and distribution modelling.

Natural history: Most species were found in relatively arid environments ( Fig. 45A–F View Figure 45 ), and some seem to tolerate extremely dry and cold conditions (see Natural history section of N. centaura and biogeographical analysis for further details). In a few cases, Nerudia has been found in relatively humid forested areas, e.g. N. guirnalda at El Rodeo or N. ola and N. poma at Chumbicha (both: Argentina, Catamarca). Most specimens were found by turning stones and rocks. Two species ( N. poma and N. trigo ) were also found on the ceilings of small cave-like shelters composed of large rocks and boulders. Nerudia nono was collected by turning stones of a loosely built wall situated in the sun ( Fig. 45F View Figure 45 ). When disturbed, the spiders either remained more or less motionless or they ran rapidly a few centimetres over the rock surface; they seemed reluctant to drop to the ground. Webs were not seen in the field, but at least N. nono was observed to build flimsy webs in small containers in the laboratory. In most visited localities, only one species of Nerudia was found; in a few cases, two or even three species were found at a single locality, apparently in identical microhabitats. Other pholcid spiders sharing the microhabitats of Nerudia were Gertschiola macrostyla and Guaranita Huber, 2000 spp. (Ninetinae) , Chibchea Huber, 2000 spp. and other small undescribed representatives of Modisiminae, and an undescribed species of Metagonia Simon, 1893 (Pholcinae) . Egg sacs were slightly flattened (but never reduced to a single layer of eggs as in Guaranita ) and consisted of about eight to 18 eggs.

Composition: The genus now includes 11 described species, ten of which are newly described below.













Huber, Bernhard A., Meng, Guanliang, Král, Jiří, Ávila Herrera, Ivalú M., Izquierdo, Matias A. & Carvalho, Leonardo S. 2023


Huber BA 2000: 87
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