Guaranita Huber, 2000

Huber, Bernhard A., Meng, Guanliang, Král, Jiří, Ávila Herrera, Ivalú M. & Izquierdo, Matías A., 2023, Revision of the South American Ninetinae genus Guaranita (Araneae, Pholcidae), European Journal of Taxonomy 900 (1), pp. 32-80 : 37-41

Genus Guaranita Huber, 2000

Guaranita Huber, 2000: 96 .

Type species

Guaranita goloboffi Huber, 2000 .

Diagnosis

Small (body length ~ 1mm) short-legged pholcids with globular abdomen ( Fig. 2 ), distinguished from most other genera of Ninetinae by dorsal flap on procursus (e.g., Figs 4F , 9F , 12D ); from Galapa , which shares a dorsal process on the procursus (cf. Huber 2000: figs 383, 387), by pair of prominent apophyses on male chelicerae (e.g., Figs 4A – C , 11A – B ; absent in Galapa ) and by unmodified fangs of male chelicerae (with processes in Galapa ).

Description

Male

MEASUREMENTS. Total body length 0.9–1.1, carapace width 0.4–0.5. Legs relatively short, tibia 1 0.5–0.6; tibia 1 L/d 7 – 9; leg formula 4-1-2-3; metatarsus 1 shorter than tibia 1 or same length (metatarsus 1/ tibia 1: 0.95 – 1.00); tibia 2 much shorter than tibia 4 (tibia 2 /tibia 4: 0.65–0.75).

COLOUR. Live specimens reddish brown ( Fig. 2 ); abdomen without or with very indistinct marks; legs without dark or light bands. Color in ethanol similar but paler, ochre-yellow.

BODY. Ocular area barely raised, eight eyes ( Figs 6A , 11A – D , 17A , 27A ),AME relatively large (diameter: 25–30 µm, i.e., 50–60% of PME diameter). Carapace without thoracic groove ( Figs 6A , 11A – D , 17A , 27A ). Clypeus usually unmodified, only in G. dobby with distinct median process ( Torres et al. 2016: fig. 9). Sternum slightly wider than long, with pair of rounded anterior processes near leg coxae 1, processes apparently without pores. Abdomen globular; four (rarely five) epiandrous spigots arranged in two pairs ( Figs 11E – F , 17F , 27F ); ALS with seven spigots each (as in female, cf. Figs 6C , 11H , 17B – D , 27C – E ): one strongly widened spigot, one long pointed spigot, and five cylindrical spigots (one of which is unusually large); PMS with two short, pointed spigots (as in female, cf. Figs 6D , 27D ); PLS without spigots ( Figs 17B , 27C – D ).

CHELICERAE. With pair of long frontal apophyses (e.g., Figs 4A – C , 11A – B ); with stridulatory files on relatively small lateral patches ( Figs 12A , 18A – B , 27G ), with ~15 – 25 stridulatory ridges each.

PALPS. Coxa unmodified; trochanter without or with very indistinct ventral projection; femur cylindrical, slightly widened distally, proximally without or with very low retrolateral hump, with prolateral stridulatory pick (modified hair; Fig. 27H ); patella short; tibia oval to globular, with two trichobothria; palpal tarsal organ raised, capsulate ( Figs 13A , 28A – B ), with small opening (diameter of opening ~1.1– 1.5 µm); procursus with distinctive dorsal flap, large semi-transparent ventral membrane, and complex tip bent towards dorsal (e.g., Figs 4D – F , 12C – F ); genital bulb with simple proximal sclerite, distinct distal (main) sclerite, and variably complex ‘embolar division’ consisting of membranous and sclerotized elements ( Figs 4G – I , 29 ).

LEGS. Without spines and curved hairs; with ‘short vertical hairs’ in 1 – 2 rows on tibia 1 ( Figs 13D , 19A – C , 30A – B ; length of hairs ~10 – 15 µm). Trichobothria in usual arrangement: three on each tibia (except tibia 1: prolateral trichobothrium absent), one on each metatarsus, slightly feathered (as in female, cf. Fig. 28E – F ); length of trichobothria ~60 µm; retrolateral trichobothrium of tibia 1 in very distal position (at ~55–65% of tibia length). Tibiae and metatarsi with tiny pores with cuticular rim (diameter of opening ~0.6 µm; as in female, cf. Figs 6E – F , 19F , 31A ). Metatarsi 3 and 4 with ~1 – 5 slender hairs ventrally ( Figs 13E , 19G – H , 31C ), with bases as in regular hairs but shafts reminding of trichobothria (i.e., feathered and small proximal diameter: ~2 µm; regular leg hair proximal diameter: 3 – 4 µm). Tarsus 1 with 5–6 pseudosegments, poorly visible in dissecting microscope; tarsus 4 distally with one comb-hair on prolateral side (as in female, cf. Fig. 31D ); leg tarsal organs very small, not raised, capsulate ( Fig. 13C ), with small opening (diameter of opening ~0.8–1.0 µm); three claws, superior claws with 8 – 11 tines (as in female, cf. Figs 7F , 13F – G , 20D – H , 31D – F ).

Female

In general, similar to male but chelicerae without stridulatory files ( Figs 12B , 18C ), sternum without pair of anterior humps, palpal tarsal organ only weakly raised ( Figs 19E , 28C – D ), and tibia 1 with usual low number of short vertical hairs; legs either slightly shorter than in males or of same length [only G. goloboffi Huber, 2000 , G. munda ( Gertsch, 1982) , and G. yaculica Huber, 2000 with reasonable sample sizes: male/female tibia 1 length: 1.00 – 1.08]. Spinnerets, leg pores, leg tarsal organs, and combhairs as in male. Main (anterior) epigynal plate usually trapezoidal, only in G. dobby Torres et al., 2016 rather triangular, weakly protruding (e.g., Figs 5A , 10A , 16A ); posterior plate simple, short but wide. Internal genitalia very simple, usually with distinct median structure (poorly developed in G. dobby ), sometimes with membranous median sac (receptacle?) (e.g., Figs 5C – D , 10C – D , 32 ); apparently with very small pore plates (arrows in Fig. 32 ). The “pair of receptacles” mentioned and illustrated in Torres et al. (2016: 10, fig. 14) is a misinterpretation either of the book lungs or of a pair of silk glands.

Relationships

The molecular analysis of Eberle et al. (2018) included only a single species of Guaranita ( G. yaculica ), which was placed (with moderate support) as sister to the South American Ninetinae genera Pemona and Kambiwa . Preliminary analyses of molecular (UCE) data (G. Meng, B.A. Huber, L. Podsiadlowski, unpubl. data) support the close relationship among these three genera and add Galapa to this clade, a genus not included in Eberle et al. (2018). Our new SEM data confirm the position of Guaranita among Ninetinae (in particular the small opening of the tarsal organs; cf. character 57 in Huber 2000). Within Guaranita , our CO1 data suggest that the morphologically distinct G. dobby is sister to the other species, a topology that is also supported by preliminary analyses of UCE data.

Natural history

While Guaranita auadae Huber sp. nov., G. dobby , and G. goloboffi were found in relatively arid environments with cacti and low bushes ( Fig. 34A, D – F ), G. munda and G. yaculica were collected in dry to humid forests ( Fig. 33B – C ). In arid environments, the specimens were collected by turning stones and rocks; in more humid environments by shaking dead bromeliads lying on the ground and by sifting leaf litter. Guaranita munda was collected by turning stones of a loosely built wall situated in a low forest ( Fig. 33B ). When disturbed by turning a rock, the spiders ran rapidly a few centimeters over the rock surface but seemed reluctant to drop to the ground. Webs were not seen in the field but the spiders quickly built flimsy webs in small glass vials. We never found more than one species of Guaranita at one locality. Other pholcid spiders sharing the microhabitats of Guaranita were Gertschiola macrostyla (Mello-Leit„o, 1941) and Nerudia spp. (Ninetinae) , and several small undescribed representatives of Modisiminae Simon, 1893. Eggs sacs were carried under the prosoma and contained 5 – 8 eggs arranged in a single layer ( Fig. 2A – B, D, H ); they are thus among the smallest egg-sacs known in pholcids ( Huber & Eberle 2021).

Distribution

Guaranita is widespread in northern Argentina and reaches into Paraguay and southern Brazil (and probably southern Bolivia and Uruguay) ( Fig. 33 ). It does not seem to cross the Andes into Chile.

Composition

The genus now includes five described species, all of which are treated below. Our limited genetic data support the species limits ( Table 2 ): intraspecific K2P distances (N = 4) range from 0.2–3.0% (mean 0.9%); interspecific distances within Guaranita (N = 24) range from 13.6–21.7% (mean 17.1%).

Figures

Fig. 2. Guaranita Huber, 2000, live specimens. A–B. G. dobby Torres et al., 2016; females with egg-sacs from NW of Campo Quijano. C–D. G. munda (Gertsch, 1982); male and female with eggsac from E of Nono. E–F. G. yaculica Huber, 2000; male and female from Calilegua National Park. G–H. G. auadae Huber sp. nov.; male and female with egg-sac from between San Salvador and Purmamarca. I–J. G. goloboffi Huber, 2000; male and female from NW of Chumbicha.

Fig. 4. Guaranita dobby Torres et al., 2016; male from NW of Campo Quijano (ZFMK Ar 24121). A–C. Chelicerae, frontal, lateral, and ventral views. D–F. Left procursus, prolateral, dorsal, and retrolateral views. G–I. Left genital bulb, prolateral, dorsal, and retrolateral views. Abbreviations: df = dorsal flap; ds = distal bulbal sclerite; ed = embolar division; ps = proximal bulbal sclerite; vm = ventral membrane. Scale bars = 0.1 mm.

Fig. 9. Guaranita munda (Gertsch, 1982); male from E of Nono (ZFMK Ar 24122). A–C. Chelicerae, frontal, lateral, and ventral views. D–F. Left procursus, prolateral, dorsal, and retrolateral views. G–I. Left genital bulb, prolateral, dorsal, and retrolateral views. Scale bars = 0.1 mm.

Fig. 12. Guaranita munda (Gertsch, 1982); from E of Nono (A–B; ZFMK Ar 24122) and Parque Nacional El Palmar (C–F; MACN Ar 32745). A. Stridulatory file on male chelicera. B. Right female palp and chelicera (note absence of stridulatory file). C–F. Right male pedipalp in retrolateral, retrolateral-dorsal, and dorsal views. Abbreviations: b = genital bulb; p = procursus. Scale bars: A = 10 µm; B–F = 20 µm.

Fig. 11. Guaranita munda (Gertsch, 1982); from E of Nono (A–E, G; ZFMK Ar 24122) and Parque Nacional El Palmar (F; MACN Ar 32745). A–B. Male prosoma, oblique frontal and dorsal views. C–D. Female prosomata, oblique frontal and lateral views. E–F. Male gonopores and epiandrous spigots. G. Female abdomen, ventral view. H. Female ALS and PMS. Scale bars: A–D, G = 100 µm; E–F, H =10 µm.

Fig. 6. Guaranita dobby Torres et al., 2016; female from NW of Campo Quijano (ZFMK Arg187). A. Prosoma, oblique frontal view. B. Abdomen, ventral view. C. ALS spigots. D. PMS spigots. E. Pore (arrow) and cuticular plate (asterisk) among regular hairs on right metatarsus 3. F. Pore on left tibia 4. Abbreviations: aep = anterior (main) epigynal plate; pep = posterior epigynal plate. Scale bars: A–B = 100 µm; C, E = 10 µm; D, F = 2 µm.

Fig. 17. Guaranita yaculica Huber, 2000; from Calilegua National Park (ZFMK Ar 24124, Arg175). A. Female prosoma, frontal view. B. Female spinnerets and anal cone. C. Female ALS. D. Male ALS. E. Female abdomen, ventral view. F. Male gonopore and epiandrous spigots. Scale bars: A, E = 100 µm; B–C, F = 10 µm; D = 2 µm.

Fig. 27. Guaranita goloboffi Huber, 2000; from NW of Chumbicha (ZFMK Ar 24130, Arg220). A. Female prosoma, frontal view. B. Epigynum, ventral view. C. Female spinnerets and anal cone. D. Female ALS and PMS. E. Female ALS. F. Male gonopore with epiandrous spigots. G. Stridulatory file on left male chelicera. H. Stridulatory pick of male palpal femur. Scale bars: A–B = 100 µm; C = 20 µm; D–G = 10 µm; H = 2 µm.

Fig. 18. Guaranita yaculica Huber, 2000; from Calilegua National Park (ZFMK Ar 24124). A. Left male chelicera. B. Stridulatory file on male chelicera. C. Left female chelicera, showing absence of stridulatory file. D–F. Right male palp, retrolateral, retrolateral-dorsal, and dorsal views; arrow: tarsal organ. Abbreviations: b, genital bulb; p, procursus. Scale bars: A, D–F = 20 µm; B–C = 10 µm.

Fig. 13. Guaranita munda (Gertsch, 1982); from Parque Nacional El Palmar (A, C, D; ZFMK Ar 24122) and E Nono (B, E–G; MACN Ar 32745). A. Male palpal tarsal organ. B. Tarsal organ on female right tarsus 2. C. Tarsal organ on male right tarsus 1. D. Short vertical hairs (and regular hairs) on male tibia 1. E. Slender hairs (arrows) on male metatarsus 4. F. Tip of female right tarsus 1. G. Tip of male left tarsus 3. Scale bars: A–B = 2 µm; C = 1 µm; D–G = 10 µm.

Fig. 28. Guaranita goloboffi Huber, 2000; from NW of Chumbicha (ZFMK Ar 24130, Arg220). A. Male left palpal tarsus, showing position of tarsal organ (arrow). B. Male palpal tarsal organ (detail of previous figure). C. Tip of left female palp, dorsal view. D. Female palpal tarsal organ (and short vertical hair). E. Female left palpal tibia, showing two trichobothria. F. Prolateral trichobothrium (and dorsal trichobothrium in the back) on female left tibia 2. Scale bars: A, C, E–F = 10 µm; B, D = 2 µm.

Fig. 29. Guaranita goloboffi Huber, 2000; male from NW of Chumbicha (ZFMK Ar 24130, Arg220). A–C. Right tarsus, procursus, and bulb, in retrolateral-dorsal, retrolateral, and retrolateral-ventral views. D–F. Right genital bulb (and neighboring elements of male palp), in prolateral, prolateral-dorsal, and dorsal views. Abbreviations: b = genital bulb; p = procursus. Scale bars = 20 µm.

Fig. 19. Guaranita yaculica Huber, 2000; from Calilegua National Park (ZFMK Ar 24124, Arg175). A. Male right tibia 1, retrolateral view. B. Detail of previous figure, showing short vertical hairs (among regular hairs). C. Short vertical hairs on male left tibia 1, prolateral-ventral view. D. Chemoreceptive hairs (arrows) on female left tarsus 3. E. Sensory organs on left female palpal tarsus (arrow: short vertical hair). F. Pore with cuticular rim (arrow) on right female metatarsus 1. G. Slender hairs (arrows) on female metatarsus 4. H. Slender hair (arrow) on male left metatarsus 4. Abbreviations: sl = slit sense organ; to = tarsal organ. Scale bars: A = 100 µm; B–E, G–H = 10 µm; F = 2 µm.

Fig. 30. Guaranita goloboffi Huber, 2000; from NW of Chumbicha (ZFMK Ar 24130, Arg220). A. Male right tibia 1, retrolateral view, showing two rows of short vertical hairs. B. Two short vertical hairs (and basis of regular hair) on male right tibia 1. C. Female metatarsus 4, showing slender hair (arrow) among regular hairs. D. Dorsal lyriform organ distally on female metatarsus 1. E. Chemoreceptive hairs distally on female left metatarsus 2. F. Female tarsus 4; note tarsal organ (arrow) and chemoreceptive hair (upper right). Scale bars = 10 µm.

Fig. 31. Guaranita goloboffi Huber, 2000; from NW of Chumbicha (ZFMK Ar 24130, Arg220).A. Pore and cuticular plate on female right tibia 2. B. Tarsal organ on female right tarsus 4. C. Male right metatarsus 4, showing two slender hairs (arrows) and regular hairs. D. Tip of female tarsus 4, prolateral view, showing claws and comb-hair (arrow). E. Claws on female right tarsus 1, retrolateral view. F. Claws of female left tarsus 3, prolateral view. Scale bars: A = 2 µm; B = 1 µm; C–F = 10 µm.

Fig. 7. Guaranita dobby Torres et al., 2016; female from NW of Campo Quijano (ZFMK Arg187). A. Trichobothrium on left metatarsus 1.B. Bases of regular hair and of trichobothrium on left metatarsus 1. C. Slender hair (arrow) among regular hairs on right metatarsus 4. D. Tarsal organ on right tarsus 3. E. Tarsal organ on right tarsus 2. F. Tip of right tarsus 1. Scale bars: A, C, F = 10 µm; B, D–E = 2 µm.

Fig. 20. Guaranita yaculica Huber, 2000; from Calilegua National Park (ZFMK Ar 24124, Arg175). A. Tarsal organ on female tarsus 2. B. Tarsal organ on female tarsus 3. C. Tarsal organ on female tarsus 4. D. Tip of female left tarsus 2, prolateral view. E. Tip of female left tarsus 1, prolateral view. F. Tip of male right tarsus 2, retrolateral view. G. Tip of female left tarsus 3, prolateral view. H. Tip of female right tarsus 4, retrolateral view. Scale bars: A–B = 2 µm; C = 1 µm; D–H = 10 µm.

Fig. 5. Guaranita dobby Torres et al., 2016; females from NW of Campo Quijano (ZFMK Ar 24121). A. Abdomen, ventral view. B. Cleared epigynum, ventral view. C–D. Cleared epigyna of two specimens, dorsal views. Scale bars = 0.1 mm (B–D at same scale).

Fig. 10. Guaranita munda (Gertsch, 1982); females from E of Nono (ZFMK Ar 24122). A. Abdomen, ventral view. B. Cleared epigynum, ventral view. C–D. Cleared epigyna of two specimens, dorsal views. Scale bars: 0.1 mm (B–D at same scale).

Fig. 16. Guaranita yaculica Huber, 2000; females from Calilegua National Park (ZFMK Ar 24124). A. Abdomen, ventral view. B. Cleared epigynum, ventral view. C–D. Cleared epigyna of two specimens, dorsal views. Scale bars = 0.1 mm (B–D at same scale).

Fig. 32. Guaranita Huber, 2000, female internal genitalia; arrows point at possible pore plates. A. G. dobby Torres et al., 2016; from NW of Campo Quijano. B. G. munda (Gertsch, 1982); from E of Nono. C. G. yaculica Huber, 2000; from Calilegua National Park. D. G. auadae Huber sp. nov., from between San Salvador and Purmamarca. E. G. goloboffi Huber, 2000; from NW of Chumbicha. Scale bar = 0.1 mm (all at same scale).

Fig. 34. Typical habitats of species of Guaranita Huber, 2000 in Argentina. A. NW of Campo Quijano (Salta), G. dobby Torres et al., 2016. B. E of Nono (Córdoba), G. munda (Gertsch, 1982). C. Calilegua National Park (Jujuy), G. yaculica Huber, 2000. D. Between San Salvador and Purmamarca (Jujuy), type locality of G. auadae Huber sp. nov. E. SW of Alemanía (Salta), G. goloboffi Huber, 2000. F. W of Cafayate (Salta), G. goloboffi.

Fig. 33. Known distribution of Guaranita Huber, 2000. Inset: map of South America, showing all known records. A–B. Known distributions of individual species. Question marks denote dubious outliers (Torres et al. 2015, 2016; not restudied) that need confirmation.