Urupelma, Kaderka, Lüddecke, Rezac, Rezacova and Hüsser, 2023

Radan Kaderka, Tim Lüddecke, Milan Řezáč, Veronika Řezáčová & Martin Hüsser, 2023, Revision of the Peruvian tarantula Homoeomma peruvianum (Chamberlin, 1916): description of a new genus with eleven new species and insights to the evolution of montane tarantulas (Araneae: Theraphosidae: Theraphosinae), Journal of Natural History 57 (41 - 44), pp. 1710-1824 : 1725-1728

Genus Urupelma gen. n.

Type species

Urupelma peruvianum ( Chamberlin, 1916) , comb. n.

Etymology

Urupelma (feminine) is a composition of two words: ̍Urubamba̾ referring to Urubamba River in Peru, and the Greek word ̍pelma̾, which means ̍sole of the foot̾. In the Quechua language, Urubamba (or Urupampa) also means ̍a plateau of spiders̾.

Diagnosis

Urupelma gen. n. differs from all other Theraphosinae genera in the presence of short and stout embolus of the bulb carrying more than two prolateral keels, PI keel well developed, A keel weakly developed, and R keel developed, in the combination with the following characters: abdominal urticating setae of type III or III+IV located in one central patch, two unequal subapical apophyses are present on male tibia I, metatarsus I when flexed touches retrolateral side of retrolateral tibial apophysis, except for U. veronicae sp. n. whose metatarsus I touches the apex of retrolateral branch. Spermathecae with two oval seminal receptacles with sub-basal constriction, not heavily sclerotised, separate (in U. sanctitheresae sp. n., U. sanctimariae sp. n., U. awanqay sp. n., U. machiguenga sp. n., U. veronicae sp. n.) or basally fused (in U. ashaninka sp. n. and U. johannae sp. n.). PS keel, if present, is located mainly on tegulum of male palpal bulb, sometime reaching basal half of embolus. SA keel, which can be considered an extension of A keel, is present in some species. PA absent.

Distribution

Peru ( Departments of Cusco, Apurímac, Junín and Pasco), at altitudes between 387 and 4450 m a .s .l. ( Figures 76 , 77 ).

Affinities

Urupelma gen. n. shares with Agnostopelma Pérez-Miles and Weinmann, 2010 , Aguapanela Perafán and Cifuentes, 2015 , Anqasha Sherwood and Gabriel, 2022 , Bumba Pérez-Miles et al., 2014 , Euathlus Ausserer, 1875 , Grammostola Simon, 1892 , Homoeomma Ausserer, 1871 , Kochiana Fukushima, Nagahama and Bertani, 2008 , Melloleitaoina Gerschman de Pikelin and Schiapelli, 1960 , Munduruku Miglio et al., 2013 , Thrixopelma Schmidt, 1994 and Tmesiphantes Simon, 1892 the presence of urticating setae of type III or III+IV (sensu Kaderka et al., 2019) located in one central patch and bipartite spermathecae in females consisting of two separate or basally fused oval seminal receptacles without spiral necks or spherical nodules or ventral loops.

Urupelma gen. n. differs from Agnostopelma in the presence of scopula on tarsi IV and short and stout embolus of male palpal bulb, carrying more than two prolateral keels (only PS and PI in Agnostopelma ). Females differ in spermathecae with flat and weakly sclerotised oval seminal receptacles, which are subparallel with sub-basal constriction (strongly sclerotised and divergent in Agnostopelma ).

Urupelma gen. n. differs from Aguapanela and Bumba in the presence of short and stout embolus of male palpal bulb, carrying more than two prolateral keels (only PS and PI in Aguapanela and Bumba ). Females of Urupelma gen. n. differ from Aguapanela and Bumba in the presence of slightly rounded spermathecal lobes.

Urupelma gen. n. differs from Anqasha in the presence of sigmoidly curved PI keel in male palpal bulb, and in the morphology of spermathecae (flat, oval receptacles in Urupelma gen. n. and tubular receptacles in Anqasha ).

Urupelma gen. n. differs from Euathlus in the presence of short and stout embolus of male palpal bulb, carrying more than two prolateral keels (only PS and PI in Euathlus ), and in subparallel seminal receptacles (strongly divergent in Euathlus ).

Urupelma gen. n. differs from Grammostola in the absence of stridulatory organ between coxae of palps and legs I, and in the presence of short and stout embolus of male palpal bulb, carrying more than two prolateral keels (only PS and PI in Grammostola ).

Urupelma gen. n. differs from Homoeomma and Tmesiphantes (including Melloleitaoina as a junior synonym of Tmesiphantes Fabiano-da-Silva et al., 2019) in the presence of short and stout embolus of male palpal bulb, carrying more than two prolateral keels (only PS and PI in Homoeomma and Tmesiphantes ). Females of Urupelma gen. n. differ from Tmesiphantes by having spermathecae with oval seminal receptacles with sub-basal constriction, separate or basally fused, and a non-incrassate femur, whereas Tmesiphantes displays spermathecae with long, slender receptacles with subapical constriction, and an incrassate femur in leg III (not all females, but all males).

Urupelma gen. n. differs from Kochiana in the presence of short and stout embolus of male palpal bulb, carrying more than two prolateral keels (only PS and PI complemented by prolateral accessory keel in Kochiana ), and in subparallel seminal receptacles (strongly divergent in Kochiana ).

Urupelma gen. n. differs from Munduruku in the presence of well-developed PI keel and R keel in male palpal bulb morphology. Females of Urupelma gen. n. differ from Munduruku by featuring an unpatterned abdomen and spermathecae with oval seminal receptacles with sub-basal constriction. In contrast, Munduruku exhibits a patterned abdomen and spermathecae with spheroid distal receptacles and a straight funnelshaped neck bearing a sclerotised area.

Urupelma gen. n. differs from Thrixopelma in spermathecae with flat and weakly sclerotised seminal receptacles with sub-basal constriction (strongly sclerotised and hypertrophied in Thrixopelma ). Male palpal bulb of Urupelma gen. n. differ in PI keel more developed on embolus and absent or weakly developed on tegulum (weakly developed only in U. awanqay sp. n.). In Thrixopelma the PI keel is less developed on embolus but extends to tegulum.

General description

The genus Urupelma gen. n. comprises small to medium-sized spiders, with total length 14.3 to 38.7 mm, exluding chelicerae and spinnerets. Carapace oval, uniformly coloured. Caput moderately domed. Ocular tubercle oval, moderately elevated, distinctly wider than long, with eight eyes, anterior eye row procurved, posterior row recurved in dorsal view. Clypeus indistinct to narrow. Fovea transverse, slightly procurved to procurved, deep. Chelicerae without rastellum and stridulatory bristles, with teeth on promargin (10–17), first basal teeth are complemented with granulation. Labium domed, wider than long, with 13–91 cuspules in anterior third to half, maxillae with 57–277 cuspules in basal half on ventral side. Ventral maxillae without short spiniform setae. Maxillary lobe pronounced into conical process. Labiosternal groove distinct, shallow and flat, with two joined elongated sigilla. Sternum oval, with three pairs of small, oval sigilla located near coxae III, coxae II and coxae I, posteriorly separated from the margin approximately by their own 0.7–2.0 diameters. Legs uniformly hirsute. Leg formula (from longest to shortest): IV> I> II> III. Leg segments: generally uniform, incrassate femora III in males.

Dense scopulae on ventral side of all tarsi, metatarsi I 25–100%, II 30–70% scopulate, posterior metatarsi partly scopulate (0–40%), scopulae more extended on anterior than posterior legs. Tarsal scopulae I usually undivided or divided by a longitudinal line or band of setae, on tarsi II undivided or divided by a longitudinal line or band of setae, on tarsi III, IV usually divided by longitudinal band or wide band of setae. Retrolateral side of femur IV and prolateral side of femur I without pad of plumose setae. Maxillary and trochanteral stridulatory setae or bristles absent. Spination as in species descriptions. Dorsal face of all tarsi and cymbium with two irregular longitudinal rows of very short claviform trichobothria. Paired tarsal claws with teeth (up to seven), third claw absent in all tarsi. Claw tufts dense, bilobate, present on all tarsi.

Abdomen uniformly coloured, with one dorsal patch of urticating setae. Urticating setae of type III, along with the setae of intermediate morphology between III and IV, or with types III+IV ( U. ashaninka sp. n., U. megantonianum sp. n., U. atarraz sp. n.), or type IV only (in the female of U. johannae sp. n., the type III setae or setae of intermediate morphology are supposed to be present in males). Four spinnerets present. PLS composed of three digitiform segments. PMS digitiform, mono-segmented.

Male palpal organ with a short and stout embolus, projecting slightly retrolateraly ( Figures 4 , 13 , 19 , 20 , 28 , 35 , 41 , 44 , 48 , 57 , 65 , 74 ; Table 24 ), with well-developed PI, weakly developed A and developed to well-developed R keel. If present, PS keels is located mainly on tegulum ( U. peruvianum comb. n., U. sanctitheresae sp. n., U. pampas sp. n., U. veronicae sp. n., U. dianae sp. n.) or may be extended to the embolar part, in which, however, the PS can be weakly developed ( U. peruvianum comb. n., U. veronicae sp. n., U. dianae sp. n.) or distinctly separated from its tegular part ( U. peruvianum comb. n., U. sanctitheresae sp. n.). The SA is present in the palpal bulb morphology of U. sanctimariae sp. n., U. pampas sp. n., U. awanqay sp. n., U. megantonianum sp. n. Sperm pore is located between PI and A keel as in the majority of theraphosines. Tegulum with short basal conical protuberance, more or less developed. Cymbium retrolateraly or dorsally with short spiniform setae, cymbial lobes of unequal size. Palpal tibia with distinct ( U. peruvianum comb. n., U. veronicae sp. n., U. dianae sp. n.) or indistinct retrolateral process ( U. awanqay sp. n., U. megantonianum sp. n., U. atarraz sp. n., U. ashaninka sp. n.) or without such process ( U. sanctitheresae sp. n., U. sanctimariae sp. n., U. pampas sp. n., U. machiguenga sp. n.), the retrolateral subapical area of short spiniform setae may be present. Two unequal tibial apophyses are present on tibia I: a longer ventral tibial apophysis, with short or long apical spine, and a shorter prolateral tibial apophysis with single, retrolateral spine at base, of similar length to prolateral apophysis or shorter. Both branches may be separated or basally fused. Metatarsus I not sigmoidly curved, without basal or median protuberance on retrolateral face. When flexed, it contacts the retrolateral side of retrolateral tibial apophysis, except for U. veronicae sp. n. whose metatarsus I touches the apex of retrolateral branch when flexed.

Females with spermathecae composed of two oval seminal receptacles with sub-basal constriction, which may be separated ( Figures 24 , 32 , 52 , 68 , 75 ) or fused (only in U. ashaninka sp. n. and U. johannae sp. n.) ( Figures 38 , 61 ).

Remarks

In congeners, the seminal receptacles are separated, with only one exception which is one ontogenetic stage with fused receptacles found in the female paratype of U. sanctimariae sp. n. ( MUSM-ENT 0513025 ) ( Figure 24E ). Until the females of U. megantonianum sp. n. and U. atarraz sp. n. with urticating setae types III+IV are collected, examined and sequenced, both species will remain in the proposed new genus.

Figures

Figure 4. A–D. Urupelma peruvianum (Chamberlin, 1916), right palpal bulb of male holotype (MCZ 15413) from Peru, Huadquina. A. Prolateral view. B. Prolaterally dorsal view. C. Dorsal view. D. Retrolateral view. E–H. Urupelma peruvianum (Chamberlin, 1916), left palpal bulb of male (MCZ 15415) from Peru, Tincochaca. E. Prolateral view. F. Prolaterally dorsal view. G. Dorsal view. H. Retrolateral view, arrow shows subapical tooth on R keel. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 13. A–D. Urupelma peruvianum (Chamberlin, 1916), right palpal bulb of male (MUSM-ENT 0510457) from Peru, Cusco, Santa Teresa. A. Prolateral view. B. Retrolateral view. C. Ventral view. D. Prolaterally posterior view, arrows show subapical tooth on R keel. E–H. Urupelma sanctitheresae sp. n., right palpal bulb of male holotype (MUSM-ENT 0513021) from Peru, Cusco, Santa Teresa. E. Prolateral view. F. Retrolateral view. G. Ventral view. H. Prolaterally posterior view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 19. Urupelma sanctimariae sp. n., male holotype (MUSM-ENT 0513026) from Peru, Cusco,Santa María, morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. C. Ventral view. D. Dorsal view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 20. Urupelma sanctimariae sp. n., male paratype (MUSM-ENT 0513024) from Peru, Cusco,Santa María, morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. C. Ventral view. D. Dorsal view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 24. A–D. Urupelma sanctitheresae sp. n., spermathecae with two separated seminal receptacles. A. Female (MUSM-ENT 0513020) from Peru, Cusco, Convención, Santa Teresa. Spermathecae dissected from dried female, ventral view. B. Immature female paratype (MUSM-ENT 0513022) from Peru, Cusco, Convención, Santa Teresa. Spermathecae dissected from exuvia, ventral view. C. Immature female paratype (MUSM-ENT 0513022) from Peru, Cusco, Convención, Santa Teresa. Spermathecae dissected from exuvia, ventral view. D. Female paratype (MUSM-ENT 0513022) from Peru, Cusco,Convención, Santa Teresa.Spermathecae dissected from specimen in alcohol, dorsal view. E–]. Urupelma sanctimariae sp. n., spermathecae with two separated or basally joined seminal receptacles. E. Female paratype (MUSM-ENT 0513025) from Peru, Cusco, Convención, Santa María. Spermathecae with basally joined seminal receptacles, dissected from exuvia, ventral view. F. Female paratype (MUSM-ENT 0513025) from Peru, Cusco, Convención, Santa María. Spermathecae dissected

Figure 28. Urupelma awanqay sp. n., male holotype (MUSM-ENT 0511135) from Peru, Apurímac, Abancay, 2818 m a.s.l., morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. C. Ventral view. D. Ventrally prolateral view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 32. Urupelma awanqay sp. n., spermathecae with two separated seminal receptacles. A. Immature female (MUSM-ENT 514037) from Peru, Apurímac, Abancay, ventral view. B. Mature female paratype (MUSM-ENT 0511169) from Peru, Apurímac, Abancay, ventral view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 35. A–B. Urupelma ashaninka sp. n., male holotype (MUSM-ENT 0511137) from Peru, Junín, Puerto Ocopa, 654 m a.s.l., morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. C– D. Urupelma ashaninka sp. n., male paratype (MUSM-ENT 0511176) from Peru, Junín, Puerto Ocopa, 654 m a.s.l., morphology of right palpal bulb. C. Prolateral view. D. Retrolateral view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 38. Urupelma ashaninka sp. n., female spermathecae with fused seminal receptacles,terminating with two oval lobes, dorsally slightly concave. A–B. Female paratype (MUSM-ENT 0511139) from Peru, Junín, Puerto Ocopa. Spermathecae dissected from specimen in alcohol. A. Dorsal view. B. Ventral view. C–D. Female paratype (MUSM-ENT 0511140) from Peru, Junín, Puerto Ocopa. Spermathecae dissected from specimen in alcohol. C. Dorsal view. D. Ventral view. E. Female (MUSM-ENT 0513573) from Peru, Junín, Puerto Ocopa. Spermathecae dissected from exuvia. F. Free. G–H. Female (MUSM-ENT 0513573) from Peru, Junín, Puerto Ocopa. Spermathecae dissected from exuvia. G. Dorsal view. H. Ventral view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 41. Urupelma atarraz sp. n., male holotype (MUSM-ENT 0508428) from Peru, Pasco, Villa Rica, Quebrada Atarraz, 1598 m a.s.l., morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 44. Urupelma megantonianum sp. n., male holotype (MUSM-ENT 0510523) from Peru, Cusco, Echarate, 410 m a.s.l., morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 48. Urupelma machiguenga sp. n., male holotype (MUSM-ENT 0513029) from Peru, Cusco, La Convención, Echarati, 1075 m a.s.l., morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. C. Ventral view. D. Dorsal view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 52. Urupelma machiguenga sp. n., female spermathecae with two seminal receptacles, terminating with rounded lobes. A. Female paratype (MUSM-ENT 0513030) from Peru, Cusco, Echarati. Spermathecae dissected from exuvia (car. 9.15). B. Female paratype (MUSM-ENT 0513030) from Peru, Cusco, Echarati. Spermathecae dissected from exuvia (car. 9.79). C. Female paratype (MUSM-ENT 0513027) from Peru, Cusco, Echarati. Spermathecae dissected from exuvia (car. 7.86). D. Female paratype (MUSM-ENT 0513027) from Peru, Cusco, Echarati. Spermathecae dissected from exuvia (car. 9.00). E–F. Female paratype (MUSM-ENT 0513027) from Peru, Cusco, Echarati. Spermathecae dissected from exuvia (car. 12.06). E. Dorsal view. F. Ventral view. G. Female paratype (MUSM-ENT 0513028) from Peru, Cusco, Echarati. Spermathecae dissected from exuvia (car. 8.59). H. Female paratype (MUSM-ENT 0513027) from Peru, Cusco, Echarati. Spermathecae dissected from specimen in alcohol, dorsal view. Scale bar = 1 mm. Photo by R. Kaderka.

Figure 57. Urupelma pampas sp. n., male holotype (MUSM-ENT 0514012) from Peru, Apurimac province, between Ocros and Chincheros, morphology of left palpal bulb. A. Retrolateral view. B. Prolateral view. C. Almost prolateral view (bulb slightly tilted dorsally and posteriorly). Scale bar = 1 mm. Photo by R. Kaderka.

Figure 61. Urupelma johannae sp. n., female holotype (MUSM-ENT 0514015) from Peru, Cusco, Quincemil. Spermathecae with fused seminal receptacles, terminating with two oval lobes, dorsally concave, ventral view. A. Dissected from exuvia. B. Dissected from specimen in alcohol. Scale bar = 1 mm. Photo: R. Kaderka.

Figure 65. Urupelma veronicae sp. n., male holotype (MUSM-ENT 0513019) from Peru, Cusco, Abra Málaga near Nevado Veronica, 4428 m a.s.l., morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. C. Dorsal view. Large arrow shows sperm pore, short arrow shows small oval lobe at the distal end of PI keel. D. Ventral view. Scale bar = 1 mm. Photo: R. Kaderka.

Figure 68. A. Urupelma veronicae sp. n., female paratype (MUSM-ENT 0511156) from Peru, Cusco, Nevado Veronica. Spermathecae with separate seminal receptacles, terminating with rounded lobes, dissected from exuvia. Scale bar = 1 mm. Photo: R. Kaderka.

Figure 74. Urupelma dianae sp. n., male holotype (MUSM-ENT 0513017) from Peru, Cusco, between Phiry near Ollantaytambo and Abra Málaga., 3868 m a.s.l., morphology of right palpal bulb. A. Prolateral view. B. Retrolateral view. C. Dorsal view. Large arrow shows sperm pore, short arrow shows small oval lobe at the distal end of PI keel. D. Ventral view. Scale bar = 1 mm. Photo: R. Kaderka.

Figure 75. Urupelma dianae sp. n. from Peru, Cusco, between Phiry near Ollantaytambo and Abra Málaga., 3868 m a.s.l. A–F. Morphology of female spermathecae dissected from exuviae collected in situ. Scale bar = 1 mm. Photo: R. Kaderka.

Figure 76. Distribution map of Urupelma gen. n. from Peru. Prepared by M. Hüsser.

Figure 77. Distribution map of Urupelma spp. from Urubamba River Valley, Cusco, Peru, showing highly diversified populations and so-called ‘high mountain insularisation̍ caused by geographical barriers and restricted gene flow. Red square = Urupelma peruvianum (Chamberlin, 1916) and Urupelma sanctitheresae sp. n. from Santa Teresa, Cusco; white square = Urupelma peruvianum (Chamberlin, 1916) and Urupelma sanctimariae sp. n. from Santa María, Cusco; yellow square = Urupelma machiguenga sp. n. from Echarati, Cusco; green square = Urupelma veronicae sp. n. from Abra Málaga near Nevado Veronica, Cusco; pink square = Urupelma dianae sp. n. from between Phiry near Ollantaytambo and Abra Málaga, Cusco. Scale bar: 10 km. Prepared by R. Kaderka.