Tolteca Huber, 2000

Huber, Bernhard A., Meng, Guanliang, Valdez-Mondragón, Alejandro, Král, Jiří, Ávila Herrera, Ivalú M. & Carvalho, Leonardo S., 2023, Short-legged daddy-long-leg spiders in North America: the genera Pholcophora and Tolteca (Araneae, Pholcidae), European Journal of Taxonomy 880 (1), pp. 1-89 : 46-49

Genus Tolteca Huber, 2000

Tolteca Huber, 2000: 117 .

Type species: Pholcophora hesperia Gertsch, 1982 .

Diagnosis

Easily distinguished from only other North American Ninetinae genus Pholcophora by small male cheliceral apophyses originating distally ( Figs 38A–B , 44 ; in Pholcophora large and originating proximally); also by absence of stridulatory ridges on male chelicerae ( Figs 41A , 46F ); most species (except T. sinnombre sp. nov.) also by knob-shaped structure between epigynum and pedicel ( Figs 43A , 45A , 49A ); from most species (except P. tehuacan sp. nov.) also by smaller size (body length ~1.1–1.4; in Pholcophora ~1.7–3.1) and shorter legs (tibia 1 <0.7, in Pholcophora >1.0). From other geographically close genera ( Papiamenta , Galapa ) also by simple rod-shaped procursus ( Figs 38C , 51C ; much shorter in Papiamenta ; with dorsal process in Galapa ), by presence of humps on male sternum (absent in Papiamenta ), and by unmodified male cheliceral fangs (with processes in Galapa ).

Description

Male

MEASUREMENTS. Total body length 1.1–1.4, carapace width 0.45–0.55. Legs very short, tibia 1 ~0.45– 0.60; tibia 1 ~1.0–1.1 x carapace width; tibia 1 L/d 7–9; tibia 2 much shorter than tibia 4 (tibia 2 / tibia 4: 0.6–0.7).

COLOUR. Live specimens reddish brown ( Fig. 36 ); carapace usually monochromous, sometimes with very indistinct darker median line widening anteriorly behind ocular area; abdomen colour slightly variable, usually monochromous, sometimes with indistinct dorsal marks; legs without dark or light bands. Colour in ethanol similar but paler, rather yellowish.

BODY. Ocular area barely raised, eight eyes, AME relatively large, diameter ~25–30 µm, ~60–80% of PME diameter; Carapace without thoracic groove or with very indistinct low median indentation (visible in frontal view only). Clypeus unmodified or with rim slightly more sclerotized than in female. Sternum wider than long, with pair of distinct anterior processes near leg coxae 1. Abdomen globular; presence of epiandrous spigots unclear (reported as present in Huber 2000: fig. 126; not seen in two newly examined males of T. hesperia and T. manzanillo Huber sp. nov.; Fig. 46G ); ALS with seven spigots each ( Fig. 54C ): 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. 54C ); PLS without spigots.

CHELICERAE. With one pair of simple frontal apophyses ( Figs 38A–B , 41A–B ); without stridulatory files ( Figs 41A , 46F ).

PALPS. Coxa unmodified; trochanter barely modified (indistinct ventral projection); femur cylindrical, slightly widened distally, proximally without retrolateral hump; patella short; tibia globular, with two trichobothria; palpal tarsal organ raised, capsulate with small opening ( Figs 41E–F , 46C–D ; diameter of opening ~1.1–1.3 µm); procursus simple and straight ( Figs 38C , 46A–B ), without dorsal flap, not strongly elongated; genital bulb large (compared to palp size), with complex distal system of sclerites and folds, partly only visible in SEM ( Figs 41C–D , 46A–B ).

LEGS. Without spines and curved hairs; with very short vertical hairs in higher density on tibia 1 ( Fig. 42A–B ; 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. 54D ); length of dorsal trichobothrium on tibia 1: ~100 µm; retrolateral trichobothrium of tibia 1 in very distal position (at 59–65%). Tarsus 1 with 4–6 distinct pseudosegments; tarsus 4 distally with one comb-hair on prolateral side (cf. Fig. 54H ); leg tarsal organs very small, capsulate with small opening ( Fig. 46E ; diameter of opening ~1.0–1.4 µm); three claws ( Fig. 42H ).

Female

In general (size, colour) similar to male ( Fig. 36 ) but sternum without pair of anterior humps, palpal tarsal organ less strongly raised ( Figs 41G , 54E ), and leg tibia 1 with usual low number of short vertical hairs; legs either slightly shorter than in males or of same length (only T. oaxaca sp. nov. with reasonable sample size: male/female tibia 1 length: 1.06). Spinnerets and comb-hairs as in male. Epigynum main (anterior) plate transversal band-shaped to crescent-shaped, weakly protruding in lateral view; posterior plate often indistinct, short but wide, usually with median anterior projection. Usually with distinct knob-shaped structure between epigynum and pedicel ( Figs 43A , 45A , 49A , 54A–B ; absent in T. sinnombre sp. nov.). Internal genitalia very simple, with pair of distinct transversal sclerites and pair of membranous sacs originating medially ( Fig. 55 ), sacs very short in T. oaxaca (9–13 µm) and in T. jalisco ( Gertsch, 1982) (18 µm), very long in T. sinnombre (85 µm); in other species ~42–48 µm; apparently without pore plates (possibly with very indistinct tiny groups of pores near median line).

Relationships

The genus Tolteca was not included in the molecular analysis of Eberle et al. (2018). Our new molecular data mostly suggest that Tolteca is sister to a clade consisting of true Pholcophora and a Caribbean clade ( Papiamenta Huber, 2000 + Caribbean ‘ Pholcophora ’). This supports a monophyletic North American-Caribbean clade of Ninetinae ( Fig. 1 ; see also general results of molecular analyses above). The latter clade is also strongly supported in our preliminary analyses of UCE data (G. Meng, L. Podsiadlowski, B.A. Huber, unpubl. data), but in that case with Papiamenta as sister to Tolteca + true Pholcophora (no Caribbean ‘ Pholcophora ’ is included in the UCE dataset).

Within Tolteca , our unpartitioned analysis suggests that the most southern species ( T. oaxaca sp. nov.) is sister to all other species; among those, T. hesperia is sister to T. jalisco + ( T. manzanillo sp. nov. + T. huahua Huber sp. nov.). Our unpublished UCE dataset does not include T. huahua but otherwise it supports the same intrageneric relationships.

Distribution

The genus appears restricted to the Pacific Lowlands and Baja Californian biogeographic provinces of Mexico, as delimited in Morrone et al. (2017) ( Fig. 35 ). The female specimens originating from San Luis Potosí and Puebla provinces tentatively identified as Tolteca and briefly mentioned in Huber (2000: 120) are possibly not Tolteca .

Natural history

Most newly collected specimens were found in low, relatively dry forests ( Fig. 56 ). Here they occupied the thin layers of leaf litter and sometimes the spaces under small stones and pebbles. They often shared the microhabitat with other species of Pholcidae , but Tolteca appeared largely restricted to the dryer areas while other genera (mostly Modisimus ) seemed to prefer slightly more humid leaf litter. No webs were observed in the field, but the spiders built tiny silk mats in the glass vials. When disturbed, they ran rapidly and barely slowed down for several minutes. Females carried their disc-shaped egg-sacs under the prosoma ( Fig. 36 ); egg-sacs usually contained 5 or 6 eggs, each with a diameter of ~ 0.35–0.45 mm ( Huber & Eberle 2021). Some females had a genital plug (cf. Fig. 49A–B ).

Composition

The genus now includes six nominal species, all of which are treated below.

Figures

Fig. 38. Tolteca hesperia (Gertsch, 1982), male from Mexico, Sinaloa, S of Rosario (ZFMK Ar 23953). A–B. Chelicerae, frontal and lateral views. C. Left palpal tarsus and procursus, retrolateral view. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Scale bars = 0.1 mm.

Fig. 44. Tolteca spp., male chelicerae, frontal and lateral views, and left male palpal tarsi and procursi, retrolateral views. A–C. T. manzanillo Huber sp. nov., paratype from Mexico, Colima, E of Manzanillo (ZFMK Ar 23958).D–F. T. sinnombre Huber sp. nov., holotype from Mexico, Colima, S of Coquimatlán (LATLAX). G–I. T. huahua Huber sp. nov., paratype from Mexico, Michoacán, W of Huahua (ZFMK Ar 23957). Scale bars = 0.1 mm.

Fig. 41. Tolteca hesperia (Gertsch, 1982), male and female from Mexico, Sinaloa, 3 km S of Rosario (ZFMK Ar23953 and Mex253). A. Male chelicerae, lateral view. B. Male right chelicera, frontal view. C. Left male palp, distal view. D. Right genital bulb (and procursus), dorso-distal view. E. Detail of male palpal tarsus showing position of tarsal organ (arrow). F. Male palpal tarsal organ. G. Female palpal tarsal organ. H. Male ALS and PMS. Abbreviations: b = genital bulb; fe = femur; p = procursus. Scale bars: A, C–D = 20 µm; B, E–F = 10 µm; G = 1 µm; H = 2 µm.

Fig. 46. Tolteca manzanillo Huber sp. nov.; male from Mexico, Colima, 17 km E of Manzanillo (ZFMK Ar 23958). A. Right palp, retrolateral-distal view. B. Left palp, retrolateral-dorsal view. C–D. Palpal tarsal organ.E. Tarsal organ on tarsus 2.F. Lateral face of right chelicera, showing absence of stridulatory file. G. Gonopore. H. ALS. Abbreviations: b = genital bulb; p = procursus. Scale bars: A–B, F = 20 µm; C–D = 2 µm; E = 1 µm; G–H = 10 µm.

Fig. 43. Tolteca jalisco (Gertsch, 1982), females from Mexico, Jalisco, N of La Quemada (ZFMK Ar 23956). A–D. Epigyna, ventral (A, C) and lateral (B, D) views; arrows point at anterior knob-shaped structure. E–G. Cleared female genitalia, ventral (E) and dorsal (F) views, and detail of median internal structures (G); arrows point at membranous sacs (cf. Fig. 55C). Abbreviations: ep = epigynum (main epigynal plate); pep = posterior epigynal plate. Scale bars: A–D = 0.2 mm; E–F = 0.1 mm; G = 0.05 mm.

Fig. 45. Tolteca manzanillo Huber sp. nov., females from Mexico, Colima, E of Manzanillo (ZFMK Ar 23958). A–D. Epigyna, ventral (A, C) and lateral (B, D) views. E–G. Cleared female genitalia, ventral (E) and dorsal (F) views, and detail of median internal structures (G). Arrows point at anterior epigynal knob. Asterisk: membranous sac (cf. Fig. 55E). Scale bars: A–D = 0.2 mm; E, F = 0.1 mm; G = 0.05 mm.

Fig. 49. Tolteca huahua Huber sp. nov., females from Mexico, Michoacán, W of Huahua (ZFMK Ar 23957). A–D. Epigyna, ventral (A, C) and lateral (B, D) views. E–G. Cleared female genitalia, ventral (E) and dorsal (F) views, and detail of median internal structures (G). Arrows point at knobshaped structure. Asterisk: membranous sac (cf. Fig. 55D). Abbreviation: gp = genital plug. Scale bars: A–D = 0.2 mm; E–F = 0.1 mm; G = 0.05 mm.

Fig. 51. Tolteca oaxaca Huber sp. nov., male paratype from Mexico, Oaxaca, N of San Pedro Totolapa (ZFMK Ar 23961). A–B. Chelicerae, frontal and lateral views. C. Left palpal tarsus and procursus, retrolateral view. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Scale bars = 0.1 mm.

Fig. 36. Tolteca Huber, 2000 live specimens. A–B. T. hesperia (Gertsch, 1982), male and female with egg-sac from Mexico, Sinaloa, S of Rosario. C–D. T. jalisco (Gertsch, 1982), females with egg-sacs from Mexico, Jalisco, N of La Quemada. E–F. T. manzanillo Huber sp. nov., male and female with egg-sac from Mexico, Colima, E of Manzanillo. G–H. T. sinnombre Huber sp. nov., male and female from Mexico, Colima, S of Coquimatlán. I–J. T. huahua Huber sp. nov., male and female from Mexico, Michoacán, W of Huahua. K–L. T. oaxaca Huber sp. nov., male and female with egg-sac from Mexico, Oaxaca, NW of Tehuantepec.

Fig. 54. Tolteca oaxaca Huber sp. nov., female from Mexico, Oaxaca, 3 km N of San Pedro Totolapa (ZFMK Mex362).A. Epigynum, ventral view (arrow: knob-shaped structure). B. Knob-shaped structure between epigynum and pedicel. C. ALS and PMS. D. Prolateral trichobothrium on tibia 3. E. Palpal tarsal organ. F. Tarsal organ on tarsus 2. G. Tip of left tarsus 2, retrolateral view. H. Tip of right tarsus 4, prolateral view, showing comb hair (arrow). Abbreviations: ep = epigynum (main epigynal plate); pep = posterior epigynal plate. Scale bars: A = 100 µm; B–D, G–H = 10 µm; E–F = 2 µm.

Fig. 42. Tolteca hesperia (Gertsch, 1982), male and female from Mexico, Sinaloa, 3 km S of Rosario (ZFMK Ar23953 and Mex253). A. Male tibia 1, prolateral view. B. Male tibia 1, retrolateral view. C. Sexually dimorphic short vertical hair on male tibia 1. D. ‘Regular’ short vertical hair on male metatarsus 1. E. Tip of ‘regular’ short vertical hair, detail of preceding image. F. Male metatarsustarsus 1 joint, retrolateral-dorsal view. G. Tarsal organ on female tarsus 1. H. Tip of left male tarsus 2, retrolateral view. Scale bars: A–B, F, H = 10 µm; C–D, G = 2 µm; E = 1 µm.

Fig. 55. Tolteca Huber, 2000 internal female genitalia, median section of main transversal internal sclerite. A–B. T. hesperia (Gertsch, 1982); females from Mexico, Sinaloa, S of Rosario (ZFMK Ar 23953). C. T. jalisco (Gertsch, 1982), female from Mexico, Jalisco, N of La Quemada (ZFMK Ar 23956). D. T. huahua Huber sp. nov., female from Mexico, Michoacán, W of Huahua (ZFMK Ar 23957). E. T. manzanillo Huber sp. nov., female from Mexico, Colima, E of Manzanillo (ZFMK Ar 23958). F. T. sinnombre Huber sp. nov., female from Mexico, Colima, S of Coquimatlán (ZFMK Ar 23960). G–H. T. oaxaca Huber sp. nov., females from Mexico, Oaxaca, N of San Pedro Totolapa (ZFMK Ar 23961). Scale bar = 0.1 mm.

Fig. 1. Relationships of Pholcophora Banks, 1896 and Tolteca Huber, 2000 derived from analysis of molecular data using IQ-Tree and the ClipKIT gappy trimming strategy. Newly generated CO1 sequences (codes starting with N) were combined with data from Eberle et al. (2018) (all other codes). Numbers on the branches are SH-aLRT supports (%). The tree shows only the ingroups and the Ninetinae outgroups. Clade colours are as in Figs 2 and 35. For the complete tree, and for clade support using different alignment strategies, see Supp. file 1: Fig. S76. Photos on the right from top to bottom: Tolteca manzanillo Huber sp. nov.; Papiamenta savonet Huber, 2000; Pholcophora papanoa Huber sp. nov.

Fig. 35. Known distribution of Tolteca Huber, 2000.

Fig. 56. Representative sample of habitats of Tolteca Huber, 2000 in Mexico. A. Sinaloa, 3 km S of Rosario (T. hesperia (Gertsch, 1982)). B. Colima, 17 km E of Manzanillo (type locality of T. manzanillo Huber sp. nov.; showing collection method). C–D. Oaxaca, 3 km N of San Pedro Totolapa (type locality of P. oaxaca Huber sp. nov.; overview and spot with high abundance of specimens).