Ibotyporanga Mello-Leitão, 1944

Huber, Bernhard A., Meng, Guanliang, Král, Jiří, Ávila Herrera, Ivalú M. & Carvalho, Leonardo S., 2024, Diamonds in the rough: Ibotyporanga (Araneae, Pholcidae) spiders in semi-arid Neotropical environments, European Journal of Taxonomy 963, pp. 1-169 : 11-29

Genus Ibotyporanga Mello-Leitão, 1944

Ibotyporanga Mello-Leitão, 1944: 6 . Type species: I. naideae Mello-Leitão, 1944 .

Ibotyporanga – Huber 2000: 94 . — Huber & Brescovit 2003: 17. — Huber & Villarreal 2020: 62 .

Diagnosis

Small short-legged pholcids with eight eyes and globular abdomen ( Figs 25 , 55 , 73 ), similar in size and body shape to other New World Ninetinae in the genera Pholcophora , Papiamenta Huber, 2000 , and Gertschiola Brignoli, 1981 . Males are easily distinguished from these and from all other pholcids by single median process on chelicerae (e.g., Figs 29 , 34 , 40 ). Corresponding pockets on female epigyna (e.g., Figs 30 , 35 , 41 ) are diagnostic for females, but less unique among Pholcidae (similar median pockets occur in other subfamilies, e.g., in Mesabolivar ). Males further distinguished by unique light prolateral band of unknown function on procursus (e.g., Figs 28 , 33 , 37 ).

Description

Male

MEASUREMENTS. Total body length 1.5–2.7; carapace width 0.6–1.0. PME diameter 50–100 µm; AME diameter 30–80 µm. Leg 1 length 2.8–7.4; tibia 1 length 0.7–2.0; tibia 4 in most species longer than tibia 1, only in the relatively long-legged I. xique sp. nov. and I. xakriaba sp. nov., as well as in I. itatim sp. nov., shorter (tibia 4 / tibia 1 length: 0.94-0.98); leg femora diameters 0.13–0.22; leg tibiae diameters 0.09–0.12. Tibia 1 L/d in most species 8–15, only in the long-legged I. xique and I. xakriaba higher: 20 and 18, respectively.

COLOUR. In ethanol mostly ochre-yellow to light brown, carapace medially and ocular area often darker, legs usually with darker rings on femora (subdistally) and tibiae (proximally and subdistally); abdomen pale gray, usually with darker internal marks dorsally and laterally; ventrally with light ochre to brown plates in front of gonopore and in front of spinnerets. Live specimens often with slightly reddish or orange prosoma and legs.

BODY. Habitus as in Figs 25 , 55 , and 73. Ocular area slightly raised. Carapace with distinct but shallow thoracic groove ( Figs 2–3 ). Clypeus with sclerotized rim with median notch ( Fig. 107A ). Sternum slightly wider than long, sometimes with pair of low and indistinct anterior processes near coxae 1 ( Fig. 3G ); processes without pores. Abdomen globular. Numerous rimmed pores ( Fig. 18 ) at regular intervals on many body parts, especially on legs, but also on chelicerae and abdomen; outer diameter 2.2–4.2 µm; opening diameter: 0.35–0.45 µm. Gonopore with four epiandrous spigots in variable pattern (evenly spaced or in two pairs; Figs 4–5 ). Spinnerets general arrangement as usual in Pholcidae ( Fig. 6 ); ALS with only two spigots ( Figs 7–8 ): one strongly widened spigot and one long and slender spigot; PMS with two short conical spigots ( Figs 6–8 ); PLS without spigots; AMS area with one or two median hairs and slightly modified sculptured area ( Fig. 7A–B ).

CHELICERAE. With distinctive median frontal apophysis, usually in proximal position (e.g., Figs 34 , 40 ), only in I. ziruma sp. nov. in distal position ( Fig. 29 ); chelicerae width 0.25–0.40. Stridulatory files ( Figs 9–10 ) very fine and poorly visible in dissecting microscope, with ~45–65 ridges, distances between ridges (in eight species studied with SEM) 1.7–3.1 µm, distances proximally in some species smaller than distally (e.g., I. guanambi sp. nov., I. sertao sp. nov.), in other species proximally larger than distally (e.g., I. naideae Mello-Leitão ; I. itatim sp. nov.).

PALPS. Coxa unmodified; trochanter with indistinct ventral protrusion; femur variably long, proximally with retrolateral process sometimes directed towards distal, with prolateral stridulatory pick (modified hair; Fig. 12F ), femur distally widened but unmodified; femur-patella joints usually not (or very slightly) shifted toward prolateral side; patella ranging from very short (e.g., Fig. 47 ) to very long (e.g., Fig. 109 ); tibia with two trichobothria; tibia-tarsus joints usually slightly shifted toward retrolateral side; palpal tarsal organ capsulate with small opening ( Fig. 13 ), outer diameter 6–10 µm, opening diameter 1.3– 2.5 µm; procursus shape highly variable, from short and wide (e.g., Figs 28 , 33 , 39 ) to very long and slender (e.g., Figs 62 , 85 , 102 ), always with light prolateral band of unknown function ( Figs 28 , 33 , 37 ); distally often with tiny side branch ( Fig. 12A–C ), tip often semi-transparent ( Figs 57 , 76 , 114 ); in some species with variably long dorsal side branch (e.g., Figs 81 , 89 , 98 ); genital bulb with proximal sclerite connecting to tarsus, strongly attached to tendon of muscle from tibia (sclerite thus usually breaks when detaching bulb from palp; therefore often not drawn); bulbous part of genital bulb in most species with distinct prolateral sclerite (e.g., Figs 28 , 33 , 37 ), in several species with small ventral tubercles (prominent in I. naideae , Fig. 11E–F ); bulbal process with variably complex tip consisting of sclerotized and membranous elements and presumably carrying sperm duct opening.

LEGS. Without spines but with stronger hairs ventrally on all femora ( Fig. 15A–B ; stronger compared to neighboring leg hairs: diameter ~5 µm versus ~3 µm in neighboring hairs, but similar hairs occur on leg coxae, sternum, palps, carapace, and abdomen). Without curved hairs. Tibia 1 or tibia 1+ tibia 2 with sexually dimorphic short vertical hairs ( Fig. 16 ), in several rows or bands, base diameter 4.1–6.3 µm, length 17–34 µm (usually 17–25 µm), diameter proximally 1.6–2.1 µm, at half-length 0.9–1.0 µm. Distal leg segments (tarsi and metatarsi, also tibiae) with putative chemoreceptors ( Fig. 17 ), in light microscope similar to sexually dimorphic short vertical hairs but with distal side branches, with regular oblique furrows, and flattened (i.e., with oval diameter); base diameter ~4.5–7.5 µm, length 20–40 µm, diameter proximally 1.7–2.8 µm, at half-length with oval cross section, ~1.5×2.5 µm. Femora, tibiae, metatarsi, and tarsi with round or oval cuticular plates ( Fig. 18 ) at regular intervals, diameter ~4.5–7.5 µm. Base of trichobothria round but with distal bulge and usually with pair of variably distinct lateral humps ( Fig. 15 ); outer diameter of base: 11–18 µm; proximal diameter of seta: 1.3–2.1 µm; retrolateral trichobothrium on tibia 1 at 51–66% of tibia length; prolateral trichobothrium absent on tibia 1. Without slender metatarsal hairs (as described in Huber et al. 2023c, 2024c). Tarsi with 3–4 distinct pseudosegments distally (cf. Fig. 21C ), proximally not pseudosegmented. Leg tarsal organs capsulate with small opening ( Fig. 19 ), outer diameter 3.9–7.4 µm, opening diameter 1.0–1.5 µm. Tarsi 4 with two comb hairs distally on prolateral side ( Fig. 20D–F ). Tarsal claws as usual for family ( Figs 20–21 ); main claws with 7–10 teeth; teeth on tarsus 4 slightly different (shorter and directed more towards distal).

Female

In general, similar to males; often larger; coloration often slightly darker; legs in some species longer than in males (especially in short-legged species; Fig. S4 ), in other species shorter (especially in long-legged species); without sexually dimorphic short vertical hairs on tibiae. Sternum and clypeus unmodified. Chelicerae without stridulatory files ( Fig. 9 ). Palpal tarsal organ as in males ( Fig. 14 ), usually oval rather than round, opening diameter 1.4–1.9 µm. Other sensory organs and comb hairs on tarsi 4 as in males. Spinnerets as in males ( Figs 6–8 ). Tip of palp with simple pointed process and subterminal invagination ( Fig. 14 ). Epigynum anterior plate trapezoidal to semicircular, with variably distinct pocket or ridge usually in anterior position (e.g., Figs 35 , 41 ), only in I. diroa in central position ( Fig. 72 ) and in I. ziruma sp. nov. in posterior position ( Fig. 30 ). Posterior epigynal plate relatively large but simple, unmodified. Internal genitalia with distinct pore plates on posterior, weakly sclerotized arc (e.g., Figs 29 , 36 , 41 ), sometimes with distinct median sclerite (e.g., Figs 100 , 104 , 108 ), sometimes with membranous lateral pouches or tubes (e.g., Figs 46 , 63 , 77 , 86 ), and sometimes with large and complex expandable median sac (e.g., Figs 100 , 108 , 112 , 124 , 128 ; cf. Appendix 3, character 24).

Relationships

The latest molecular phylogeny of Pholcidae ( Eberle et al. 2018; Huber et al. 2018) resolved Ibotyporanga as sister to a North and Central American clade including the genera Pholcophora and Papiamenta . This sister group relationship received low support, but it is strongly supported by unpublished UCE data including more representatives of the mentioned genera, and many more representatives of other Ninetinae genera (G. Meng, L. Podsiadlowski, B.A. Huber, unpubl. data).

The monophyly of Ibotyporanga is well supported by several morphological synapomorphies ( Fig. 1 ): median notch on male clypeus (char. 1); reduction of ALS spigots to only two (char. 2); single median apophysis on male chelicerae (char. 3); distinct light band on procursus (char. 10); and median pocket on female epigynal plate (char. 18). In addition, the prolateral sclerite on the bulbous part of the genital bulb (char. 13) might be a further synapomorphy (see Discussion).

Internal relationships appear reasonably well resolved in the morphological cladogram ( Fig. 1 ) but ten of the 15 nodes in this tree are supported by a single character each, and very few groups receive additional support from CO1 sequences (cf. Fig. S2 ). The cladistic analysis suggests that species with a ‘regular’ procursus (i.e., not slender and elongated; char. 6) are plesiomorphic, while those with a slender elongated procursus are derived and monophyletic. Within this clade, species with a ‘split’ procursus, i.e., with a distinct dorsal branch in addition to the main branch (char. 8), are again resolved as a monophylum.

Natural history

Ibotyporanga spiders are typically found in dry habitats, dominated by low and thorny vegetation ( Figs 22–23 ). Some species were found on bare rock outcrops, under rocks lying in the plain sun, while others were found in low and dry forests. They seem to avoid more humid habitats even when available nearby (see I. kanoe sp. nov.). Several species also occupy highly degraded and artificial habitats, such as roadside rocks, piles of construction materials, and pastures; two species are known to enter buildings, and of these, I. naideae has probably extended its original distribution by human transport.

Within these habitats, any sheltered space at ground level seems to be suitable for Ibotyporanga , but the most common microhabitat was under stones. The spiders were also found under and in other objects lying on the ground such as dead branches and tree trunks, cacti, bromeliads, and bark, and rarely even in the leaf litter or ( I. naideae only) in the understory vegetation. Different species do not seem to be specialized to specific microhabitats: several species were found in a variety of different microhabitats.

In some cases, the microhabitat was shared with another species of Pholcidae , usually another representative of Ninetinae . In Brazil, this was usually a representative of Kambiwa ; in northern South America, it was Galapa Huber, 2000 . Highly degraded and artificial habitats were often shared with synanthropic species such as Modisimus culicinus (Simon, 1893) or Physocyclus globosus (Taczanowski, 1874) . In a few cases, we found two species of Ibotyporanga to share a habitat or even a microhabitat. For example, Ibotyporanga emekori was found together with different congeners ( I. diroa ; I. itajubaquara sp. nov.; I. tuxa sp. nov.), apparently in identical microhabitats; I. naideae was found to share localities with I. guanambi sp. nov., I. imale sp. nov., and I. sertao sp. nov.; I. sertao sp. nov. was also found together with I. xakriaba sp. nov.

Upon disturbance, the spiders usually started to run rapidly until finding a new hiding place, often just a small depression in the rock. In some species, however, a sexual dimorphism in this behavior was observed, with males running but females remaining sitting on the rock or feigning death when beaten out of a branch or cactus.

Egg sacs were usually round but slightly flattened ( Figs 25 , 55 ), with a diameter of ~1.5–2.5. They were covered by a very sparse and poorly visible layer of silk. They contained ~10– 40 eggs (in most cases 15–30). Egg diameters ranged from 0.46 to 0.62, with a positive correlation between egg diameter and body size (carapace width) ( Fig. S5 ).

For further details on individual species, see species descriptions below.

Distribution

Ibotyporanga is apparently restricted to South America ( Fig. 24 ). Here, it is largely limited to two areas dominated by dry and semi-arid environments: the northern South American coast, and the Brazilian Cerrado and Caatinga biomes. The genus seems to be absent from the Andes; the highest known record is at 1160 m a.s.l. Ibotyporanga is largely absent from the Amazon biome. The finding of I. kanoe sp. nov. in Rondônia suggests that within the Amazon biome, Ibotyporanga may be limited to small pockets of relatively dry environments (cf. Fig. 22B ). In northern South America, Ibotyporanga is remarkably absent from the Leeward Antilles ( Huber et al. 2024a).

On lost types

It seems that the type specimens (and most non-type specimens) of all four previously described Brazilian Ibotyporanga species are lost. The type series of I. naideae Mello-Leitão, 1944 was apparently destroyed in the fire that burned a large part of the collection of the Museu Nacional do Rio de Janeiro on 2 Sep. 2018 (A. Kury, personal communication, 22 Nov. 2023). All specimens of the three species described in Huber & Brescovit (2003) ( I. diroa , I. emekori , and I. ramosae ) were apparently destroyed in the fire that burned a large part of the scorpion and spider collections at Instituto Butantan (São Paulo) on 15 May 2010 (A.D. Brescovit, personal communication, 1 Sep. 2023). We do not designate neotypes for these species as we see no “exceptional need” (ICZN Art. 75) for doing so.

Composition and species limits

The genus now includes 24 nominal species. Of these, a few show geographical morphological variation that may represent species limits, i.e., they need to be restudied with larger samples and more data (e.g., molecular). In some species, we found unusually high genetic distances among putative conspecifics ( Figs S5–6 , Table S1 ). Particularly problematic in this sense are the type species ( I. naideae ) and I. imale sp. nov.; but also I. walekeru sp. nov., I. guanambi sp. nov., I. emekori , I. kiriri sp. nov., and I. canudos sp. nov.; see individual species descriptions below for evidence suggesting that each of these species may eventually need to be split into two or more species.

Three further probable species are available in collections but not formally described, either because no males are available, or because the material is in very poor condition: (1) I. “Ven18-182” from Venezuela (Falcón, Peninsula de Paraguaná, near Cueva del Guano, 11.9026° N, 69.9456° W; ZFMK Ar 21861; Fig. 26 ) is included in the trees in Figs S2 , S 3 , and S6. The epigynum and internal genitalia of the only available adult specimen were illustrated in Huber & Villarreal (2020: figs 197–199). (2) I. “Br22- 182” from Brazil (Bahia, E of São Félix do Coribe, top of hill, 13.4040° S, 44.1100° W; CHNUFPI 9066 [deposited in ZFMK Br22-182]; Fig. 74 ) could not be sequenced (barcoding failed). The epigynum and internal genitalia of the only available adult specimen are similar to those of I. imale sp. nov. (which was found at the base of the same hill), but the legs are much longer (tibia 1: 1.60; in 54 females of I. imale : 0.95–1.33). (3) I. “Geneve453” from Colombia (Magdalena, Tayrona National Park, Bahia de Gairaca, 11.317° N, 74.108° W; MHNG; Fig. 26 ). The available specimens (3 ♂♂, 3 ♀♀, 1 juv.) are strongly bleached and difficult to study. The shape of the procursus (S-shaped) strongly suggests that it represents an additional species.

Figures

Fig. 25. Ibotyporanga spp., live specimens, part one (cf. Figs 55, 73). A. I. ziruma Huber sp. nov., female with egg sac from Colombia, Magdalena, Santa Marta. B–C. I. walekeru Huber sp. nov., male from Colombia, La Guajira, S of Riohacha, and female with egg sac from Colombia, Cesar, ESE of Pueblo Bello. D–E. I. piojo Huber sp. nov., male and female with egg sac from Colombia, Atlántico, near Piojo. F–G. I. itatim Huber sp. nov., male and female with egg sac from Brazil, Bahia, W of Itatim. H–I. I. xique Huber sp. nov., male and female with egg sac from Brazil, Bahia, S of Xique-Xique.

Fig. 55. Ibotyporanga spp., live specimens, part two (cf. Figs 25, 73), all from Brazil. A–B. I. camarai Huber sp. nov., male and female from Pernambuco, NE of Petrolina. C–D. I. naideae Mello-Leitão, 1944, male and female with egg sac from Bahia, SE of Jacobina. E–F. I. diroa Huber & Brescovit, 2003, male and female with egg sac from Bahia, near Queimada Nova. G–H. I. imale Huber sp. nov., male from Bahia, E of São Félix do Coribe, and female with egg sac from Minas Gerais, NW of Itacarambi.

Fig. 73. Ibotyporanga spp., live specimens, part three (cf. Figs 25, 55), all from Brazil. A. I. sertao Huber sp. nov., male from Pernambuco, W of Orocó. B–C. I. guanambi Huber sp. nov., male and female with egg sac from Bahia, N of Guanambi. D. I. emekori Huber & Brescovit, 2003, male from Bahia, near Toca do Índio.E. I. tuxa Huber sp. nov., male from Bahia, W of Barra do Mendes. F. I. kiriri Huber sp. nov., male from Bahia, SE of Paramirim. G. I. ouro Huber sp. nov., male from Bahia, E of Gentio do Ouro. H. I. canudos Huber sp. nov., male from Bahia, SW of Morro do Chapéu.

Fig. 29. Ibotyporanga ziruma Huber sp. nov., male and female from Colombia, Magdalena, Santa Marta, ZFMK Ar 24349. A–B. Male chelicerae, frontal and lateral views. C. Cleared female genitalia, dorsal view. Abbreviation: pp= pore plate. Scale lines: 0.2 mm.

Fig. 34. Ibotyporanga walekeru Huber sp. nov. A–B. Male chelicerae, frontal and lateral views, from Colombia, Cesar, ESE of Pueblo Bello, ZFMK Ar 24350. C. Cleared female genitalia, dorsal view, female from Colombia, La Guajira, S of Riohacha, ZFMK Ar 24351. Scale lines: 0.2 mm.

Fig. 40. Ibotyporanga piojo Huber sp. nov., male and female from Colombia, Atlántico, near Piojo, ZFMK Ar 24353. A–B. Male chelicerae, frontal and lateral views. C. Cleared female genitalia, dorsal view (bold arrows: internal tubes). Abbreviations: ep=epigynal pocket; pp = pore plate. Scale lines: 0.2 mm.

Fig. 30. Ibotyporanga ziruma Huber sp. nov., female from Colombia, Magdalena, Santa Marta, ZFMK Ar 24349. A. Abdomen, ventral view. B–C. Cleared female genitalia, ventral and dorsal views (asterisk: transversal membranous structure). D. Detail of cleared female genitalia, dorsal view. Abbreviations: ep= epigynal pocket; pp =pore plate. Scale lines: A=0.3 mm; B–D =0.1 mm.

Fig. 35. Ibotyporanga walekeru Huber sp. nov. A–C. Abdomen, ventral view, and cleared female genitalia, ventral and dorsal views, female from Colombia, Cesar, ESE of Pueblo Bello, ZFMK Ar 24350. D–F. Same as for preceding, female from Colombia, La Guajira, S of Riohacha, ZFMK Ar 24351. G–I. Same as for preceding, female from Colombia, La Guajira, near Papayal, ZFMK Ar 24352. Abbreviation: ep =epigynal pocket. Scale lines: 0.2 mm.

Fig. 41. Ibotyporanga piojo Huber sp. nov., female from Colombia, Atlántico, near Piojo, ZFMK Ar 24353. A. Abdomen, ventral view. B–C. Cleared female genitalia, ventral and dorsal views. Abbreviations: ep=epigynal pocket; pp =pore plate. Scale lines: 0.2 mm.

Fig. 28. Ibotyporanga ziruma Huber sp. nov., male from Colombia, Magdalena, Santa Marta, ZFMK Ar 24349. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views (bold arrow: distinctive apophysis on genital bulb). Abbreviations: lb=light band; ps =prolateral sclerite. Scale lines: 0.2 mm.

Fig. 33. Ibotyporanga walekeru Huber sp. nov., male from Colombia, Cesar, ESE of Pueblo Bello, ZFMK Ar 24350. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Abbreviations: lb=light band; ps =prolateral sclerite. Scale lines: 0.2 mm.

Fig. 37. Ibotyporanga bariro Huber, 2020, male holotype from Venezuela, Falcón, SE of Bariro, ZFMK Ar 21862. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Abbreviations: lb= light band; ps =prolateral sclerite. Scale lines: 0.2 mm.

Fig. 2. Ibotyporanga spp., prosoma. A–B. I. ziruma Huber sp. nov., male and female prosomata, frontal views. C. I. itatim Huber sp. nov., male prosoma, oblique frontal view. D. I. walekeru Huber sp. nov., male, frontal view. E–F. I. guanambi Huber sp. nov., female, frontal view and male, frontal-dorsal view. G. I. payaya Huber sp. nov., female, dorsal view. Scale lines: 100 µm.

Fig. 3. Ibotyporanga spp., prosoma and ocular area. A–B. I. payaya Huber sp. nov., female, frontal view and male ocular area, frontal-dorsal view. C–D. I. canudos Huber sp. nov., male and female, frontal views. E–F. I. naideae Mello-Leitão, 1944, female, frontal view and male ocular area, frontal view. G. I. ziruma Huber sp. nov., hump on male sternum. Scale lines: A, C–F =100 µm; B =20 µm; G =10 µm.

Fig. 107. Ibotyporanga tuxa Huber sp. nov., male and female from Brazil, Bahia, W of Barra do Mendes, ZFMK Ar 24378. A. Male ocular area, clypeus, and chelicerae, oblique frontal view (arrow: clypeus notch).B–C. Male chelicerae, frontal and lateral views. D. Cleared female genitalia, dorsal view (asterisk: expandable membranous sac not drawn, cf. Fig. 108E). Scale lines: 0.3 mm.

Fig. 18. Ibotyporanga spp., rimmed pores and cuticular plates. A. I. ziruma Huber sp. nov., left male tibia 1, prolateral view. B. I. itatim Huber sp. nov., left female tibia 1, dorsal view. C. I. itatim, right male chelicera. D. I. guanambiHuber sp. nov., right male tibia 3, retrolateral-ventral view.E. I. guanambi, male abdomen, above spinnerets. F. I. naideae Mello-Leitão, 1944, right female metatarsus 1. G. I. payaya Huber sp. nov., right female tarsus 4, retrolateral view. H. I. canudos Huber sp. nov., right female femur 4, retrolateral view. Abbreviations: pl = cuticular plate; rp = rimmed pore. Scale lines: A–G =10 µm; H =2 µm.

Fig. 4. Ibotyporanga spp., male gonopore and epiandrous spigots. A. I. ziruma Huber sp. nov. B. I. walekeru Huber sp. nov. C. I. itatim Huber sp. nov.D. I. naideaeMello-Leitão, 1944. E. I. guanambi Huber sp. nov. F. I. payaya Huber sp. nov. Scale lines: 10 µm.

Fig. 5. Ibotyporanga spp., male gonopore and epiandrous spigots, and female epigynum. A. I. sertao Huber sp. nov., male gonopore. B. I. canudos Huber sp. nov., male gonopore.C. I. ziruma Huber sp. nov., epigynum, ventral view. D. I. payaya Huber sp. nov., epigynum, ventral view. E. I. sertao Huber sp. nov., epigynal pocket. F. I. canudos Huber sp. nov., epigynal pocket. Abbreviation: ep =epigynal pocket. Scale lines: A–B =10 µm; C–D= 100 µm; E–F =20 µm.

Fig. 6. Ibotyporanga spp., spinnerets. A. I. ziruma Huber sp. nov., female.B. I. guanambi Huber sp. nov., female. C. I. naideae Mello-Leitão, 1944, male. D. I. payaya Huber sp. nov., male. E–F. I. sertao Huber sp. nov., male. Abbreviations: ALS =anterior lateral spinneret; PLS= posterior lateral spinneret; PMS= posterior median spinneret. Scale lines: A, C–E =20 µm; B=100 µm; F =10 µm.

Fig. 7. Ibotyporanga spp., spinnerets. A. I. walekeru Huber sp. nov., female AMS area. B. I. naideae Mello-Leitão, 1944, female AMS area. C–D. I. ziruma Huber sp. nov., male and female ALS and PMS. E. I. walekeru Huber sp. nov., male ALS. F. I. itatim Huber sp. nov., male ALS. Scale lines: 10 µm.

Fig. 8. Ibotyporanga spp., spinnerets. A–B. I. naideae Mello-Leitão, 1944, male ALS and female ALS and PMS. C–D. I. guanambi Huber sp. nov., male ALS and female ALS and PMS. E–F. I. payaya Huber sp. nov., male and female ALS. Scale lines: 10 µm.

Fig. 9. Ibotyporanga spp., chelicerae, frontal-lateral views. A–B. I. ziruma Huber sp. nov., male and female.C–D. I. canudosHuber sp. nov., male and female.E. I. sertao Huber sp. nov., male.F. I. guanambi Huber sp. nov., female. Scale lines: A–B, D =20 µm; C, E–F =100 µm.

Fig. 10. Ibotyporanga spp., stridulatory files on male chelicerae, lateral views (A, B, E are mirror images of left chelicerae). A. I. ziruma Huber sp. nov. B. I. walekeru Huber sp. nov. C. I. itatim Huber sp. nov. D. I. naideae Mello-Leitão, 1944. E. I. guanambi Huber sp. nov. F. I. sertao Huber sp. nov. Scale lines: A–D, F =10 µm; E= 20 µm.

Fig. 12. Ibotyporanga spp., male palp. A. I. payaya Huber sp. nov., right procursus tip, distal view; arrow points at tiny side branch. B. I. payaya, tiny side branch (arrow) of left procursus, prolateraldorsal view. C. I. canudos Huber sp. nov., tiny side branch (arrow) on main branch of right procursus. D. I. canudos, tip of dorsal branch of right procursus. E. I. sertao Huber sp. nov., right procursus, dorsal view; arrow points at longitudinal slit. F. I. guanambi Huber sp. nov., stridulatory pick on left femur. Abbreviation: p=procursus. Scale lines: A=20 µm; B–E =10 µm; F =2 µm.

Fig. 47. Ibotyporanga xakriaba Huber sp. nov., male from Brazil, Piauí, Parque Nacional da Serra das Confusões, CHNUFPI 1134. Left palp, prolateral, dorsal, and retrolateral views (arrow: distinctive dorsal process on femur). Scale line: 0.3 mm.

Fig. 109. Ibotyporanga atikum Huber sp. nov., male from Brazil, Bahia, SE of Bom Jesus da Lapa, ‘site 2’, ZFMK Ar 24379. Left palp, prolateral, dorsal, and retrolateral views. Scale line: 0.3 mm.

Fig. 13. Ibotyporanga spp., male palpal tarsal organs. A. I. ziruma Huber sp. nov. B. I. itatim Huber sp. nov. C. I. naideae Mello-Leitão, 1944. D. I. guanambi Huber sp. nov. E. I. payaya Huber sp. nov. F. I. sertao Huber sp. nov. Abbreviation: to =tarsal organ. Scale lines: 10 µm.

Fig. 39. Ibotyporanga piojo Huber sp. nov., male from Colombia, Atlántico, near Piojo, ZFMK Ar 24353. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views (arrow: distinctive apophysis on embolus). Scale lines: 0.2 mm.

Fig. 62. Ibotyporanga naideae Mello-Leitão, 1944, male from Brazil, Bahia, SE of Jacobina, ZFMK Ar 24357. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views (arrow: distinctive prolateral process proximally on procursus). D–F. Left genital bulb, prolateral, dorsal, and retrolateral views (bold arrow: distinctive apophysis on embolus). Abbreviation: ps= prolateral sclerite. Scale lines: 0.2 mm.

Fig. 85. Ibotyporanga sertao Huber sp. nov., male from Brazil, Pernambuco, NE of Lagoa Grande, ZFMK Ar 24367. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Scale lines: 0.2 mm.

Fig. 102. Ibotyporanga payaya Huber sp. nov., male from Brazil, Bahia, SE of Bom Jesus da Lapa, ‘site 1’, ZFMK Ar 24377. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Abbreviation: ps =prolateral sclerite. Scale lines: 0.2 mm.

Fig. 57. Ibotyporanga camarai Huber sp. nov., male from Brazil, Pernambuco, NE of Petrolina, ZFMK Ar 24356. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views (arrows: distinctive apophysis on embolus). Scale lines: 0.2 mm.

Fig. 76. Ibotyporanga imale Huber sp. nov. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views (arrow: distinctive bend of dorsal branch of procursus), male from type locality, Brazil, Bahia, E of São Félix do Coribe, ZFMK Ar 24361. D–F. Left tarsi and procursi, retrolateral views (arrows: distinctive bend of dorsal branch of procursus), males from Brazil, Minas Gerais, SE of Monte Rei (D), CHNUFPI 5910; from Minas Gerais, Parque Nacional Cavernas do Peruaçu, near visitor center (E), ZFMK Ar 24364; and from Minas Gerais, Parque Nacional Cavernas do Peruaçu, near guest house (F), ZFMK Ar 24366. Scale line: 0.3 mm (all figures at same scale).

Fig. 114. Ibotyporanga kiriri Huber sp. nov., male from Brazil, Bahia, SE of Paramirim, ZFMK Ar 24380. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views (arrow: dorsal hump on tarsus). D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Scale lines: 0.2 mm.

Fig. 81. Ibotyporanga ramosae Huber & Brescovit, 2003, male from Brazil, Bahia, São Desiderio, near Gruta da Passagem, CHNUFPI 3730. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Abbreviation: db =dorsal branch of procursus. Scale lines: 0.2 mm.

Fig. 89. Ibotyporanga guanambi Huber sp. nov. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views (bold arrow: tiny side branch on main branch of procursus), male from Brazil, Bahia, N of Guanambi, ZFMK Ar 24370. D. Left tarsus and procursus, retrolateral view (bold arrow: tiny side branch on main branch of procursus), male from Bahia, NE of Marcolino Moura; ZFMK Ar 24371; at same scale as A–C. E–G. Left genital bulb, prolateral, dorsal, and retrolateral views; same specimen as in A–C. Abbreviation: db =dorsal branch of procursus. Scale lines: 0.3 mm.

Fig. 98. Ibotyporanga emekori Huber & Brescovit, 2003, male from Brazil, Bahia, W of Queimada Nova, ZFMK Ar24374. A–C. Left tarsus and procursus, prolateral, dorsal, and retrolateral views. D–F. Left genital bulb, prolateral, dorsal, and retrolateral views. Abbreviations: db=dorsal branch of procursus; ps =prolateral sclerite. Scale lines: 0.3 mm.

Fig. 11. Ibotyporanga spp., male palp. A. I. ziruma Huber sp. nov., tip of left procursus (and mechanoreceptive hair), distal view. B. I. itatim Huber sp. nov., tips of right procursus and embolus, dorsal view. C. I. guanambi Huber sp. nov., left embolus, dorsal-distal view. D. I. guanambi, right procursus and embolus, dorsal view; bold arrow points at longitudinal slit. E. I. naideae Mello-Leitão, 1944, left bulb and embolus, retrolateral view.F. Detail from preceding figure.Abbreviations: db =dorsal branch of procursus; e =embolus; p =procursus. Scale lines: A–C, F =10 µm; D–E=100 µm.

Fig. 15. Ibotyporanga spp., leg femora and trichobothria. A. I. ziruma Huber sp. nov., right male femur 1, retrolateral view; arrows point at slightly stronger ventral hairs. B. I. guanambi Huber sp. nov., right male femur 3, retrolateral view; arrow points at slightly stronger ventral hair. C. I. ziruma, retrolateral trichobothrium on right male tibia 3. D. I. walekeru Huber sp. nov., retrolateral trichobothrium on right female tibia 2. E. I. itatim Huber sp. nov., retrolateral trichobothrium on right female tibia 2. F. I. naideae Mello-Leitão, 1944, trichobothrium on right female metatarsus 1. Scale lines: 10 µm.

Fig. 16. Ibotyporanga spp., sexually dimorphic short vertical hairs on male tibiae. A. I. ziruma Huber sp. nov., right tibia 1, retrolateral view. B–C. I. itatim Huber sp. nov., right tibia 1, retrolateral views. D. I. guanambi Huber sp. nov., right tibia 1, retrolateral view. E. I. payaya Huber sp. nov., left tibia 1, prolateral view. F. I. canudos Huber sp. nov., left tibia 1, prolateral view. Scale lines: A, C–F =10 µm; B=2 µm.

Fig. 17. Ibotyporanga spp., putative chemoreceptors on male and female legs. A–B. I. ziruma Huber sp. nov., right female tarsus 1 and right male metatarsus 1. C. I. walekeru Huber sp. nov., left female metatarsus 1, near tip, prolateral view. D–E. I. itatim Huber sp. nov., right female metatarsus 2 and right female metatarsus 3. F. I. canudos Huber sp. nov., left female tarsus 4. Scale lines: A, E–F =2 µm; B–D =10 µm.

Fig. 21. Ibotyporanga spp., right leg tarsi, retrolateral views. A. I. walekeru Huber sp. nov., female tarsus 1. B. I. itatim Huber sp. nov., female tarsus 2. C. I. naideae Mello-Leitão, 1944, female tarsus 2 (arrows: membranous areas between pseudosegments). D. I. itatim, female tarsus 3. E. I. itatim, female tarsus 4. F. I. naideae, male tarsus 4. Scale lines: 10 µm.

Fig. 19. Ibotyporanga spp., leg tarsal organs. A. I. guanambi Huber sp. nov., male tarsus 1. B. I. sertao Huber sp. nov., female tarsus 1. C. I. ziruma Huber sp. nov., male tarsus 2. D. I. naideae Mello-Leitão, 1944, male tarsus 2. E. I. sertao, female tarsus 2. F. I. itatim Huber sp. nov., female tarsus 3. G. I. itatim, male tarsus 4. H. I. naideae, male tarsus 4. Scale lines: A, E–G =10 µm; B–D, H =2 µm.

Fig. 20. Ibotyporanga spp., left leg tarsi, prolateral views. A. I. sertao Huber sp. nov., male tarsus 1. B. I. canudosHuber sp. nov., female tarsus 2. C. I. naideaeMello-Leitão, 1944, male tarsus 3. D. I. ziruma Huber sp. nov., female tarsus 4 (arrows: comb-hairs). E. I. walekeru Huber sp. nov., female tarsus 4 (arrows: comb-hairs). F. I. naideae, male tarsus 4 (arrows: comb-hairs). Scale lines: 10 µm.

Fig. 14. Ibotyporanga spp., female palp tip. A. I. ziruma Huber sp. nov. B. I. itatim Huber sp. nov. C. I. naideae Mello-Leitão, 1944. D. I. guanambi Huber sp. nov. E. I. payaya Huber sp. nov. F. I. sertao Huber sp. nov. Abbreviation: to=tarsal organ. Scale lines: 10 µm.

Fig. 72. Ibotyporanga diroa Huber & Brescovit, 2003. A–B. Abdomens, ventral views, females from Brazil, Bahia, near Toca do Índio (A), ZFMK Ar 24360, and from Bahia, W of Queimada Nova (B), ZFMK Ar 24359. C–D. Cleared female genitalia, ventral and dorsal views; same specimen as in A (bold arrows: putative lateral sacs or pockets). Abbreviations: ep= epigynal pocket; es =expandable membranous sac; pp =pore plate. Scale lines: 0.3 mm.

Fig. 36. Ibotyporanga walekeru Huber sp. nov. (A–C) and I. bariro Huber, 2020 (D), details of cleared female genitalia, dorsal views (arrows: internal tubes). A. Colombia, Cesar, ESE of Pueblo Bello, ZFMK Ar 24350. B. Colombia, La Guajira, S of Riohacha, ZFMK Ar 24351. C. Colombia, La Guajira, near Papayal, ZFMK Ar 24352. D. Venezuela, Falcón, SE of Bariro, ZFMK Ar 21863. Abbreviation: pp =pore plate. Scale lines: 0.1 mm.

Fig. 100. Ibotyporanga emekori Huber & Brescovit, 2003.A–C. Abdomens, ventral views, females from Brazil, Bahia, near Toca do Índio (A), ZFMK Ar 24373, from Bahia, W of Queimada Nova (B), ZFMK Ar 24374, and from Bahia, Fazenda Pedra Branca (C), ZFMK Ar 24376. D–E. Cleared female genitalia, ventral and dorsal views; same specimen as in A. Abbreviations: ep =epigynal pocket; es =expandable membranous sac; sc =internal sclerite. Scale lines: 0.3 mm.

Fig. 104. Ibotyporanga payaya Huber sp. nov., female from Brazil, Bahia, SE of Bom Jesus da Lapa, ‘site 1’, ZFMK Ar 24377. A. Abdomen, ventral view. B–C. Cleared female genitalia, ventral and dorsal views. D. Detail of cleared female genitalia, dorsal view. Abbreviations: es = expandable membranous sac; sc =internal sclerite. Scale lines: A–C =0.3 mm; D =0.1 mm.

Fig. 108. Ibotyporanga tuxa Huber sp. nov. A–C. Female abdomens, ventral views, females from Brazil, Bahia, W of Barra do Mendes (A–B), ZFMK Ar 24378, and from Bahia, São Desiderio (C), CHNUFPI 3659. D–E. Cleared female genitalia, ventral and dorsal views, same specimen as in B. F–G. Cleared female genitalia, ventral and dorsal views, same specimen as in C. Abbreviations: es=expandable membranous sac; sc =internal sclerite. Scale lines: 0.3 mm.

Fig. 46. Ibotyporanga itatim Huber sp. nov., females from Brazil, Bahia, W of Itatim, ZFMK Ar 24354. A–B. Abdomens, ventral views. C–D. Left putative receptacles and pore plates in female internal genitalia, dorsal views. E–H. Cleared female genitalia, ventral (E, G) and dorsal (F, H) views. Abbreviation: pp=pore plate. Scale lines: A–B, E–H=0.2 mm; C–D =0.1 mm.

Fig. 63. Ibotyporanga naideae Mello-Leitão, 1944. A–B. Male chelicerae, frontal and lateral views, male from Brazil, Bahia, SE of Jacobina, ZFMK Ar 24357. C–D. Male chelicerae, lateral views, males from Mato Grosso do Sul, Terra Indígena Kadiwéu (C), CHNUFPI 5071, and from Maranhão, Reserva Mansinha (D), CHNUFPI 4205; all chelicerae at same scale. E. Cleared female genitalia, dorsal view, female from Maranhão, Reserva Mansinha, CHNUFPI 4206. F. Left tube in female internal genitalia, female from Maranhão, Reserva Mansinha, CHNUFPI 4205. G–H. Tubes in female internal genitalia, same specimen as in E. Scale lines: A–E=0.3 mm; F–H=0.1 mm.

Fig. 77. Ibotyporanga imale Huber sp. nov., male and female from Brazil, Bahia, E of São Félix do Coribe, ZFMK Ar 24361. A–C. Left genital bulb, prolateral, dorsal, and retrolateral views. D–E. Male chelicerae, frontal and lateral views. F. Cleared female genitalia, dorsal view (asterisk: expandable membranous sac not drawn, cf. Fig. 78D). Scale lines: 0.2 mm.

Fig. 86. Ibotyporanga sertao Huber sp. nov. A–B. Male chelicerae, frontal and lateral views, from Brazil, Pernambuco, NE of Lagoa Grande, ZFMK Ar 24367. C. Cleared female genitalia, dorsal view, female from Pernambuco, NE of Cabrobó, ZFMK Ar 24369. D–E. Schematic drawings of left side and right side tubes, dorsal views, showing that the convolutions are apparently not regular and symmetric; same specimen as in C. Scale lines: A–C=0.2 mm; D–E =0.1 mm.

Fig. 112. Ibotyporanga atikum Huber sp. nov., female from Brazil, Bahia, SE of Bom Jesus da Lapa, ‘site 2’, ZFMK Ar 24379. A. Abdomen, ventral view. B–C. Cleared female genitalia, ventral and dorsal views. D. Detail of cleared female genitalia, dorsal view. Abbreviations: es = expandable membranous sac; pp =pore plate. Scale lines: A–C= 0.3 mm; D =0.1 mm.

Fig. 124. Ibotyporanga itajubaquara Huber sp. nov. A–C. Abdomens, ventral views, females from Brazil, Bahia, N of Itajubaquara (A–B), CHNUFPI 5973 and ZFMK Ar 24383, and from Bahia, NW of Gameleira do Assuruá (C), ZFMK Ar 24384. D–E. Cleared female genitalia, dorsal view, and detail of same figure, same specimen as in B. F–G. Cleared female genitalia, dorsal view, and detail of same figure, same specimen as in C. Abbreviations: es=expandable membranous sac; pp =pore plate. Scale lines: A–D, F =0.3 mm; E, G =0.1 mm.

Fig. 128. Ibotyporanga canudos Huber sp. nov. A–C. Female abdomens, ventral views, females from Brazil, Bahia, SE of Lagoa do Boi (A), ZFMK Ar 24387, Bahia, 25 km WNW of Morro do Chapéu (B), ZFMK Ar 24385, and from Bahia, 4 km SW of Andorinha (C), ZFMK Ar 24388. D–E. Cleared female genitalia, ventral and dorsal views, same specimen as in A. F. Cleared female genitalia, ventral view, same specimen as in B. G. Cleared female genitalia, ventral view, same specimen as in C. Abbreviation: es =expandable membranous sac. Scale lines: 0.3 mm.

Fig. 1. Single most parsimonious cladogram of Ibotyporanga Mello-Leitão, 1944, resulting from analyses of the matrix in Appendix 1. Only unambiguous character changes are shown; black hash marks indicate unique character changes, white hash marks indicate homoplasious changes or reversals; numbers above hash marks indicate character numbers and below character states. Male palpal procursi on the right show one particular result of this tree: it suggests that strongly elongated procursi are derived. Top row: I. piojo Huber sp. nov., I. itatim Huber sp. nov., I. xique Huber sp. nov.; center: I. diroa Huber & Brescovit, 2003; bottom: I. canudos Huber sp. nov.; live spider: I. imale Huber sp. nov., male from Brazil, E of São Félix do Coribe.

Fig. 22. Ibotyporanga spp., typical habitats, part one (cf. Fig. 23): rock outcrops; all localities are in Brazil. A. Bahia, S of Xique-Xique (type locality of I. xique Huber sp. nov.). B. Rondônia, near Jamari (type locality of I. kanoe Huber sp. nov.). C. Bahia, W of Itatim (type locality of I. itatim Huber sp. nov.). D. Bahia, Guanambi (type locality of I. guanambi Huber sp. nov.). E. Bahia, NE of Morro do Chapéu (I. canudos Huber sp. nov.). F. Pernambuco, NE of Cabrobó (I. sertao Huber sp. nov.). Photos BAH.

Fig. 23. Ibotyporanga spp., typical habitats, part two (cf. Fig. 22); all localities except for A are in Brazil. A. Colombia, Santa Marta (type locality of I. ziruma Huber sp. nov.). B. Bahia, E of São Félix do Coribe (type locality of I. imale Huber sp. nov.). C. Bahia, SE of Bom Jesus da Lapa (type locality of I. atikum Huber sp. nov.). D. Bahia, W of Bom Jesus da Lapa (I. emekori Huber & Brescovit, 2003). E. Bahia, Paramirim (type locality ofI. kiriri Huber sp. nov.). F. Pernambuco, NE of Lagoa Grande (type locality of I. sertao Huber sp. nov.). G. Bahia, near Toca do Índio (I. emekori Huber & Brescovit, 2003 and I. diroa Huber & Brescovit, 2003). H. Bahia, SW of Morro do Chapéu (I. canudos Huber sp. nov.). Photos BAH.

Fig. 24. Known geographic distribution of Ibotyporanga Mello-Leitão, 1944. Green: taxa with short plesiomorphic procursus; blue: taxa with long, derived procursus (cf. Fig. 1). For detailed distribution maps of individual species groups, see Figs 26, 42, 60, 74, and 96.

Fig. 26. Known geographic distribution of Northern SouthAmerican species.Abbreviation:A=Atlántico.

Fig. 74. Known geographic distribution of Brazilian species with long procursus and with dorsal branch on procursus, but without median sclerite in female internal genitalia. Abbreviations: BA=Bahia; CE =Ceará; GO =Goiás; MG = Minas Gerais; PE =Pernambuco; PI =Piauí.