Araneae: Corinnidae) Endemic to the Brazilian Atlantic Rainforest 127-159 74 (2): 127 – 159 10.10.2016

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Zeitschrift/Journal: Arthropod Systematics and Phylogeny

Jahr/Year: 2016

Band/Volume: 74

Autor(en)/Author(s): Magalhaes Ivan L. F., Fernandes Luciu R., Ramirez E., Ramirez Martin J., Bonaldo Alexandre Bragio

Artikel/Article: Phylogenetic position and taxonomic review of the Ianduba spiders (Araneae: Corinnidae) endemic to the Brazilian Atlantic rainforest 127-159 74 (2): 127 – 159 10.10.2016

Phylogenetic position and taxonomic review of the Ianduba spiders (Araneae: Corinnidae) endemic to the Brazilian Atlantic rainforest

Ivan L.F. Magalhaes *, 1, 2, Lúciu R. Fernandes 3, Martín J. Ramírez 1 & Alexandre B. Bonaldo 3

1 División Aracnología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” – CONICET, Av. Ángel Gallardo 470, C1405DJR. Buenos Aires, Argentina; Ivan L.F. Magalhaes * [[email protected]] — 2 Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, Av. Antônio Carlos 6627, 31270-901. Belo Horizonte, Minas Gerais, Brazil. — 3 Museu Paraense Emílio Goeldi, Coordenação de Zoologia, Laboratório de Aracnologia, Av. Perimetral 1901, 66077-830. Belém, Pará, Brazil — * Corresponding author

Accepted 04.v.2016. Published online at www.senckenberg.de/arthropod-systematics on 21.ix.2016.

Editor in charge: Lorenzo Prendini

Abstract The spider genus Ianduba is known from seven species, all restricted to the Brazilian Atlantic rainforest, a biodiversity hotspot. The genitalic morphology of these spiders is rather peculiar and they have been considered incertae sedis in Corinnidae. We present novel morphological data for the genus, including scanning electronic microscopy images for several somatic and genitalic features, and test their phylogenetic position by including one Ianduba species in a large morphological matrix of dionychan spiders. Our results suggest that Ianduba is a true corinnid, possibly belonging in a clade sister to Corinninae. Furthermore, we here describe eight new species of the genus: I. acaraje sp.n. (Bahia), I. apururuca sp.n. (Minas Gerais to Espírito Santo), I. capixaba sp.n. (Espírito Santo), I. dabadu sp.n. (Espírito Santo), I. beaga sp.n. (Minas Gerais), I. benjori sp.n. (Rio de Janeiro), I. liberta sp.n. (Minas Gerais) and I. angeloi sp.n. (Minas Gerais to São Paulo). Six of the new species seem to be closely related to I. varia, previously considered an aberrant species. Thus, we divide the genus into two morphological groups. All species from the varia group (except for I. varia, which is synanthropic in southeastern Brazil) appear to be restricted, or more common, at altitudes of at least 800 m above sea level. We argue that unsampled montane rainforest areas from southeastern Brazil are likely to yield new records or even undescribed species of Ianduba, and that montane species are likely to be under threat of extinction. New records for previously known species are provided, the female of I. caxixe Bonaldo, 1997 is described and illustrated for the first time, and distribution maps and an identification key for the fifteen known species are provided.

Key words Dionycha, Espinhaço range, conservation hotspot, new species, Quadrilátero Ferrífero, Serra da Mantiqueira, Serra do Mar, taxonomy.

1. Introduction

The Brazilian Atlantic rainforest is considered one of the nomic revisions of Atlantic rainforest-associated groups world’s biodiversity hotspots, as it is highly species-rich continue to yield descriptions of several new taxa (e.g. and threatened (Myers et al. 2000; Ribeiro et al. 2009). Rheims 2010; Brescovit et al. 2012; Pinto-da-Rocha et Also, it is the best-known Brazilian region regarding its al. 2014; Rodrigues & Bonaldo 2014; Huber 2015), in- spider fauna, being the region which has been sampled dicating that a picture of the arachnid fauna of this region and studied the most, and also the one from which more is still far from complete. species have been recorded (Oliveira 2011). Yet, taxo-

ISSN 1863-7221 (print) | eISSN 1864-8312 (online) 127 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 1. Living female of Ianduba varia (Keyserling, 1891) from São Paulo, Brazil. Note white-colored distal end of tibia I. Photo by A. Anker.

Although it has a continuous distribution, the Atlan- in typical corinnids, such as the presence of a male pal- tic rainforest is not homogeneous, and three main cent- pal median apophysis and courses of the sperm duct that ers of endemism are recognized: the Pernambuco center, do not match the patterns found in either Corinninae or the Bahia center and the South/Serra do Mar center (see Castianeirinae. Thus, they were considered as Corinni- Carnaval & Moritz 2008; Silva et al. 2012; Carnaval dae incertae sedis by Bonaldo (1997, 2000). Indeed, a et al. 2014; Oliveira et al. 2015). In particular, the Serra recent analysis has found no evidence for a monophyletic do Mar center is located in a region where several moun- Corinnidae including these genera possessing a median tain ranges can be found. Associated with these are sev- apophysis, which were joined under the informal name eral endemic birds (Vasconcelos 2008), plants (Safford “Pronophaea group” (Ramírez 2014). However, Ian- 2007), spiders (Polotow & Brescovit 2006) and harvest- duba was not among the terminals analyzed, and thus men (Pinto-da-Rocha & Silva 2005), amongst others. its phylogenetic position remains untested – although The genus Ianduba Bonaldo was described in 1997 to Bonaldo (2000) suggested it might be the sister group include Castianeira varia Keyserling, 1891 and four new to the Corinninae. We here describe the fine morphology species. After that, two more species were described by of I. varia and I. caxixe in detail and include the genus Bonaldo et al. (2007). Except for Ianduba varia (Fig. 1), in Ramírez’s (2014) matrix to test its phylogenetic posi- which is distributed from southeastern Brazil to Argen- tion. A better understanding of which groups are closely tina and is known to be synanthropic at least in southeast- related to Ianduba would be essential for polarizing char- ern Brazil, the other six species are restricted to the Bahia acters for inferring the internal phylogeny of the genus endemism center of the Atlantic rainforest, occurring at and interpreting the rather bizarre genitalic morphology relatively northern latitudes. In addition to that, I. varia of its representatives. has until now been considered a species with a deviant In this paper, we describe eight new species of Iandu- genitalic morphology, bearing little resemblance to its ba from the Brazilian Atlantic rainforest. Seven of these congeners (Bonaldo 1997). new species occur south of Bahia state, and six of these The phylogenetic position of Ianduba within Corinni- seem to be closely related to Ianduba varia. All species dae has been a matter of doubt since the original descrip- herein described are narrowly distributed, and six of tion of the genus. Bonaldo (1997) argued that this genus them are recorded from montane regions in the Atlantic and the African Mandaneta Strand, Procopius Thorell rainforest (at least 800 m above sea level). Additionally, and Pseudocorinna Simon share a distinct trichoboth- we test the phylogenetic position of the genus, presenting rial morphology with Corinnidae. However, Ianduba evidence that it is a true corinnid, possibly allied to the and these African genera possess characters not found Corinninae.

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2. Material and methods Morphological structures. A – epigynal apodemes; Ac – acini- form gland spigot; ALE – anterior lateral eyes; ALS – anterior lateral spinnerets; aMA – apical sector of the median apophysis; AME – anterior median eyes; AP – apical process of the lobes of Specimens were examined completely immersed in 75% the RTA; AS – apical spur of the RTA; AT – anal tubercle; AtP – ethanol in an Olympus SZ40 stereoscopic microscope. atrial pockets of the epigynum; BP – basal process of the lobes of The internal structure of female genitalia was examined the RTA; C – conductor; CB – conductor base; CD – copulatory after digestion in a pancreatin solution prepared accord- ducts; Co – colulus; CO – copulatory openings; Cy – cylindrical gland spigot; DL – dorsal lobe of the RTA; DP – dorsal piece of ing to Álvarez-Padilla & Hormiga (2008). Drawings the vulva; DPP – dorsal piece process; E – embolus; EB – embolic were made on a Leica M205C stereoscopic microscope base; EP – embolic process; FD – fertilization ducts; FDM – ferti- with a coupled camera lucida. The description format fol- lization duct membrane; hpMA – ‘hematodocha’ of the pMA; IS – lows Bonaldo et al. (2007). Leg spines are counted in the inframammilary sclerite; MA – median apophysis; mAP – minor dorsal (d), prolateral (p), ventral (v) and retrolateral (r) ampullate gland spigot; MAP – major ampullate gland spigot; MP – median plate of the epigynum; MPr – median process of the surfaces, generally on the proximal, median, and distal lobes of the RTA; N – nubbin; P – petiole; Pi – piriform gland spig- thirds of each surface (Platnick & Shadab 1975), but the ot; PLE – posterior lateral eyes; PLS – posterior lateral spinnerets; number of longitudinal sectors counted in each surface pMA – prolateral sector of the median apophysis; PME – posterior may vary according to spine distribution. All measure- median eyes; PMS – posterior median spinnerets; rMA – retro- ments are in millimeters and were preferentially taken on lateral sector of the median apophysis; RTA – retrolateral tibial apophysis; S – primary spermathecae; Sp – spinnerets; SS – sec- the left side of specimens. Specimens without coordinates ondary spermathecae; T – tartipore; TM – tegular modified area; were georeferenced post-hoc using the GeoLoc tool in the TP – tegular projection; Tr – posterior tracheae; TrS – posterior SpeciesLink website (http://splink.cria.org.br/geoloc; last tracheal spiracle; VL – ventral lobe of the RTA. accessed May 4th, 2014). Original label information on coordinates is referred to between parentheses; post-hoc coordinates are referred to between square brackets. Alti- tude information for all specimens was extracted from an 4. Results and discussion elevation raster using ArcMap10 software (ESRI). We prepared specimens for scanning electron microscopy as described by Ramírez (2014), except that Phylogenetic position and genital morphology. Ian- specimens were dehydrated using hexamethyldisilazane duba has been recovered as sister to Allomedmassa mae instead of critical-point dried. Parts were mounted on Dankittipakul & Singtripop (labeled as “cf. Medmassa individual stubs, sputter-coated in gold-palladium and THA” in Ramírez 2014), and this group is part of the examined using a FEI XL 30 TMP scanning electron mi- “true” Corinnidae under all weighting schemes (Fig. 2). croscope. However, its position within Corinnidae varies (sister to, For the phylogenetic analysis, we follow the same or part of, Castianeirinae with k values lower or equal procedures for analyzing the data as explained in Ramírez to 7; sister to Corinninae with k values higher than 8). (2014), using the same automated scripts mentioned We follow Ramírez (2014), presenting the results of the there. As discussed in that work, results are sensitive to analysis with k = 9 (length = 3015, fit = 176.04, CI = analysis parameters, so we explored the phylogenetic po- 0.153, RI = 0.513), and an alternative resolution under sition of Ianduba under equal and implied weights (test- equal weights (Fig. 2). The supports are generally low for ing k concavity constants from 1 to 9). All analyses were most of the groupings, and of all the synapomorphies of run using TNT 1.1 (Goloboff et al. 2008). Corinnidae listed in Table 1, Ianduba only presents a scu- The distribution of the varia group was modelled us- tum on the male abdomen (although characters 100, 182 ing records of all species in this group, except the widely and 351 have ambiguous scorings for Ianduba). Bonaldo distributed I. varia. The model was generated in MaxEnt (2000) comments on some similarities between Ianduba 3.3.1 (Phillips et al. 2006) using the 19 climatic variables and Corinninae, namely a T-shaped tegulum, a sperm from the WorldClim database (Hijmans et al. 2005), an duct restricted to the tegulum, and the shape of the ATR. aridity index (Zomer et al. 2008) and altitudinal data. These characters have not been included in the dataset we analyzed, but they would support the results we found under higher k values, where Ianduba + Allomedmassa Dankittipakul & Singtripop are a clade sister to Corin- 3. Abbreviations ninae. At any rate, the pivotal position of Ianduba near the base of the subfamilies Corinninae and Castianeirinae Collections. Examined specimens are deposited in the following makes the genus an important representative in future collections (curators in brackets): CHNUFPI – Coleção de História studies of corinnid phylogeny. Natural da Universidade Federal do Piauí, Floriano, Piauí, Bra- We here present novel morphological data for Iandu- zil (L.S. Carvalho); IBSP – Instituto Butantan, São Paulo, Brazil ba, including SEM images of genitalic (Figs. 5 – 8) and so- (A.D. Brescovit); MNRJ – Museu Nacional, Rio de Janeiro, Bra- matic (Figs. 9, 10) structures. The structure of the median zil (A.B. Kury); MPEG – Museu Paraense Emílio Goeldi, Belém, Pará, Brazil (A.B. Bonaldo); UFMG – Coleções Taxonômicas da apophysis in Ianduba is quite unique. It is known that Universidade Federal de Minas Gerais, Belo Horizonte, Minas Ge- the median apophysis has a highly homoplasious evo rais, Brazil (A.J. Santos). lution (consistency index is only 0.03 in Ramírez 2014,

129 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 2. Least-rejected cladogram obtained under implied weights (concavity constant k = 9). Values above branches are clade numbers as referred in Table 1, below branches are Bremer support (values multiplied by 100) and jackknifing frequencies (only values higher than 50 are shown). Inset shows an alternative resolution under equal weights, where part of the Pronophaea group appears as sister to classical corinnids. Photo of a Ianduba varia male by J.P. Burini.

see his character 356). Bonaldo & Brescovit (2005) dis- of the conductor, since in some species this sclerite is covered that the Attacobiini tegular process (as defined fused with the median apophysis at its base (see Figs. 5C, by Bonaldo & Brescovit 1998) is actually loosely at- 6B). Even among species of Ianduba, the homology of tached to the tegulum and thus could be considered ho- genitalic structures is frequently unclear. Throughout this mologous to the median apophysis of other dionychan paper, we have applied the same names to structures that spiders. These authors stated that other tegular sclerites seem to be homologous across species, such as the tegu- in Corinninae (as those found in Xeropigo O. Pickard- lar projections, the lobes of the RTA and the sectors of Cambridge and some Corinna C.L. Koch) also could be the median apophysis. However, their homology should considered as such. Likewise, Ramírez (2014) states that be reassessed when a greater diversity of corinnids and the Corinna tegular process (PTC) in Bonaldo (2000: potential relatives are known. figs. 80 – 92) might be homologous to the median apophysis in other Dionycha. The structure in Ianduba could Ianduba species groups. Bonaldo (1997) noted that I. va be homologous to the PTC in Corinna, to the PTX in ria has a palpal morphology differing in many aspects Xeropigo and, especially, to the median apophysis in from that of the type species of the genus, I. vatapa Bo Attacobius Mello-Leitão, which is divided in sectors as naldo, 1997. We here describe the first species ofIanduba in most Ianduba. This view might be supported by the that seem to be closely related to I. varia on the basis close relationship between Ianduba and the Corinninae. of genitalic morphology, particularly the retrolaterally Another possibility is that the ‘median apophysis’ in Ian- curved embolus and the conductor positioned ventrally duba has been misinterpreted and that it is an elaboration to the median apophysis. We erect two morphological

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Table 1. List of synapomorphies for Corinnidae and internal groups, and the Pronophaea group. Clade numbers refer to Fig. 2, character numbers are the same as in Ramírez (2014).

Pronophaea group, clade 219 tarsal cuticle texture (100): fingerprint→ smooth trochanter distal ventral margin notch (105): deep at least legs I-II → shallow or absent scales (= feathery scales and all bent setae) (157): present → absent Corinnidae, clade 223 tarsal cuticle texture (100): fingerprint→ smooth trichobothria distal plate transversal ridge (182): distal plate embedded below transversal ridge → distal ridge continuous in a closed alveolus dorsal scutum on male abdomen (205): absent → present embolus attachment (351): flexibly attached→ fixed MA (356): present → absent epigynum lobes (365): LL+MF delimited by furrows or sutures → undivided plate suture not visible Castianeirinae, clade 228 cymbial tip ventral groove (323): absent → present cymbium dorsal chemosensory patch (324): present → absent conductor (359): present → absent SP2 lumen (372): SP2 blind sac with defined lumen → SP2 a pore field without its own lumen Clade 222 male epigastric sclerite (207): absent → present Ianduba + Allomedmassa, clade 226 dorsal scutum on female abdomen (201): absent → present female inframammillary sclerite (210): absent → present male inframammillary sclerite (211): absent → present cymbial tip apical thick setae (327): absent → present Corinninae, clade 221 MaAm number in male (258): one plus nubbin (generally with Ta visible) → two (generally with Ta visible) male PMS MiAm number (274): one plus nubbin (generally with Tp visible) → two (generally with Tp visible) sperm duct distal thickness (346): gradually tapering, or thinned before embolus → thick sclerotized apical bulb sperm duct spiral meander in ventral tegulum (348): absent → present cuticular glands on epigyne (376): absent → present groups to respectively accommodate species related to species are narrow endemics of single mountain ranges I. vatapa and to I. varia. Although both are potentially (Levi 1988; Polotow & Brescovit 2006; Pinto-da- monophyletic, this should be tested in a future phyloge- Rocha et al. 2014). Our results from distribution mod- netic analysis of the genus. Interestingly enough, the two eling suggest that Ianduba of the varia group are asso- groups we propose have distributions that are largely co- ciated with these montane regions (Fig. 4). However, it incident with endemism areas of the Atlantic forest: the also reveals that much of this potentially suitable habitat, vatapa group is associated with the Bahia center, and the as of yet, has no records of Ianduba spiders – mainly the varia group is associated with the South/Serra do Mar Mantiqueira, Espinhaço, Canastra, Brigadeiro, Caparaó center (Fig. 21; see Carnaval & Moritz 2008; Silva et and Serra dos Órgãos ranges (Fig. 4). Most of these have al. 2012; Oliveira et al. 2015). not been adequately sampled for spiders yet. It may be that further collecting in these areas may yield new re- Biogeography. Among Ianduba species, the only one cords of the known species – and considering that most that is widely distributed is I. varia, ranging from Minas Ianduba have narrow ranges, it would not be surpris- Gerais, Brazil to Corrientes, Argentina (Fig. 21). All oth- ing to find new, endemic species from these unexplored er species are very rare, being known from five localities mountaintops. at most, and often from a single one. In addition to that, all species of the varia group have been recorded from Conservation. Huber (2015) described several pholcid montane Atlantic rainforest regions. Although I. varia spiders known from single localities in the Atlantic forest, has been recorded from sea-level altitudes, and I. dabadu and suggested that this area harbours a higher proportion sp.n. has been recorded at 200 m, both can reach alti- of narrowly distributed species than other regions in the tudes up to 1150 m, and the other five species range from Neotropics (e.g. the Amazon). Ianduba apparently has a 800 m to 1200 m above sea level (Fig. 3). As pointed similar pattern, with most species occurring in one or a out above, the montaintops of eastern Brazil are rich in few localities in the Atlantic forest. The fact that Ianduba endemic taxa (Pinto-da-Rocha & Silva 2005; Safford species are so rare and narrowly distributed raises con- 2007; Vasconcelos 2008); in the case of arachnids, some cerns about the conservation status of these spiders. They

131 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 3. Known altitudinal range of Ianduba species (mean, maximum and minimum). Numbers between parentheses indicate the number of unique records for each species.

Fig. 4. Distribution model for Ianduba group varia (except I. varia). Highlighted areas have suitability greater than 0.7. Small dots re present records for I. varia, other symbols indicate records for other species: squares = I. beaga, circle = I. liberta, crosses = I. angeloi, triangle = I. benjori, losangles = I. dabadu, stars = I. capixaba.

are strictly associated with the Atlantic forest, a highly list of endangered species due to habitat loss (Machado deteriorated and fragmented region with only 8 – 14% of et al. 2008; but see Ministério do Meio Ambiente 2014). the original cover still standing (Ribeiro et al. 2009). This In the case of the newly described montane species, there has been noticed before, as four of the seven previously are two additional threats. First, montane taxa are espe- known species had been included in the Brazilian official cially prone to extinction due to global warming (Sorte

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Fig. 5. Scanning electron microscopy, Ianduba angeloi sp.n., male (UFMG). A: Left palpal tibia and bulb, ventral view. B: Same, retrolateral view. C: Apical area of bulb, ventral view. D: Palpal tibia, retrolateral view. E: Same, ventral view. F: Apex of chelicerae, retromarginal view (note absence of cheliceral apophysis). See chapter 3 for abbreviations. Scale bars = 0.2 mm, except C = 0.1 mm.

& Jetz 2010). Second, some of the mountain ranges in of Ianduba highlights the vast amount of undescribed southeastern Brazil are under severe pressure due to ore taxa that are present, even in well-known regions such as exploration, particularly the Quadrilátero Ferrífero range the southeastern Atlantic rainforest. We hope that further (Jacobi et al. 2007) – a region where four species of Ian- studies on this fascinating genus will reveal interesting duba have been recorded, two of which are endemic. evolutionary and biogeographical patterns. This calls for special conservation attention not only for these species, but for southeastern Brazil mountain ranges as a whole. The discovery of eight new species

133 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 6. Scanning electron microscopy, Ianduba caxixe Bonaldo, 1997 (IBSP 63907), male. A: Left bulb, prolateral view. B: Same, apical view. C: Same, retrolateral view. D: Left palpal tibia, subapical view. E: Same, retrolateral view. See chapter 3 for abbreviations. Scale bars = 0.1 mm.

5. Taxonomy which is covered by a membrane that is posteriorly connected to the fertilization ducts (Figs. 8, 13, 14; DP).

5.1. Ianduba Bonaldo, 1997 Description. See Bonaldo (1997).

Ianduba Bonaldo, 1997: 168. Type species by original designation I. vatapa Bonaldo, 1997. 5.2. Identification key

Emended diagnosis. All Ianduba species have the distal 1 Males ...... 2 end of tibia I white to whitish yellow in color, contrasting 1’ Females ...... 16 with the rest of the leg, in both sexes (Fig. 1). In males, the metatarsus might be lightly colored as well. Males Males are further distinguished from other Neotropical corin- 2 Embolus prolaterally inserted (Fig. 6A) nids by the combined presence of an uncoiled tegular ...... 3 (vatapa group) duct and median apophysis in the palp (Figs. 5 – 7, 13, 2’ Embolus retrolaterally inserted (Fig. 7C) 14; MA); the petiole is large and its edge is always visible ...... 10 (varia group) in retrolateral view (Figs. 13, 14; P); the retrolateral tibial 3 Conductor large; tegular projection absent (Fig. apophysis is always bifurcate, and the ventral lobe might 11A) ...... Ianduba acaraje sp.n. or might not have a prolaterally directed apical spur. Fe- 3’ Conductor small; tegular projection present (Fig. males are distinguished by the dorsal piece in the vulva, 13A) ...... 4

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Fig. 7. Scanning electron microscopy, Ianduba varia (Keyserling, 1891) (UFMG 6055), male. A: Left palpal tibia, retrolateral view. B: Same, ventral view. C: Left bulb, ventral view. D: Same, retrolateral view. See chapter 3 for abbreviations. Scale bars = 0.1 mm.

4 Tegular projection short, lamellar and undulated (Bo- 6’ Tegular projection inserted basally (Bonaldo et al. naldo 1997: fig. 15) ...... Ianduba vatapa 2007: fig. 1)...... Ianduba mugunza 4’ Tegular projection long and curved (Fig. 13A) ..... 5 7 Tegular projection tubular (Bonaldo et al. 2007: fig. 5 Tegular projection inserted prolaterally (Bonaldo 26) ...... Ianduba abara 1997: fig. 23)...... 6 7’ Tegular projection concave, with anterior surface 5’ Tegular projection inserted medially or retrolaterally excavated (Fig. 13B; Bonaldo 1997: fig. 26) ...... 8 (Fig. 13A; Bonaldo et al. 2007: fig. 27) ...... 7 8 Tegular projection with an additional, longitudinal 6 Tegular projection inserted apically (Bonaldo 1997: prolateral excavation (Bonaldo 1997: fig. 26) fig. 23) ...... Ianduba caxixe ...... Ianduba paubrasil

135 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 8. Scanning electron microscopy, female genitalia, pancreatin digested. A–C: Ianduba varia (Keyserling, 1891) (UFMG 6055). A: Ventral view. B: Dorsal view. C: Dorsolateral view. D–F: Ianduba caxixe Bonaldo, 1997 (IBSP 63907). D: Ventral view. E: Dorsal view. F: Dorsolateral view. See chapter 3 for abbreviations. Scale bars = 0.1 mm.

8’ Tegular projection without additional excavation 10’ Short prolateral tegular projection absent; ventral lobe ...... 9 of RTA not displaced dorsally (Fig. 19A – B) ...... 14 9 Dorsal lobe of RTA tubular, pMA basal projection 11 Basal process of ventral lobe of the RTA with a blunt short (Fig. 13A) ...... Ianduba apururuca sp.n. apex (Fig. 14A) ...... Ianduba capixaba sp.n. 9’ Dorsal lobe of RTA with a basal constriction; pMA 11’ Basal process of ventral lobe of the RTA hook- basal projection large (Bonaldo 1997: fig. 21) shaped (Fig. 15A) ...... 12 ...... Ianduba patua 12 Dorsal lobe of the RTA with a long median process, 10 Short prolateral tegular projection present; ventral with a small spine at its blunt apex (Fig. 15B) lobe of RTA displaced dorsally (Fig. 15A – C) .... 11 ...... Ianduba dabadu sp.n.

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Fig. 9. Scanning electron microscopy, Ianduba varia (Keyserling, 1891) (UFMG 6055), spinnerets. A–D: Female. A: Spinnerets, ventral. B: Right ALS, ventral. C: Left PMS, ventral. D: Left PLS, ventral. E,F: Male. E: Spinnerets, ventral. F: Right PMS, ventral. See chapter 3 for abbreviations. Scale bars: A,E = 0.1 mm; B, D = 0.05 mm, C, F = 0.01 mm.

12’ Median process of RTA’s dorsal lobe represented by 14 Median apophysis without long hair-like projec- an unsclerotized median area, without spine (Fig. tions; ventral lobe of the RTA hirsute (Fig. 18A) . 17A) ...... 13 ...... Ianduba benjori sp.n. 13 Tegular projection unsclerotized and pointing ven- 14’ Median apophysis with hair-like projections; ventral trally (Fig. 16A) ...... Ianduba liberta sp.n. lobe of the RTA not particularly hisute (Figs. 7C, 13’ Tegular projection sclerotized and pointing to the 19A) ...... 15 embolus (Fig. 17A) ...... Ianduba angeloi sp.n. 15 Dorsal lobe of the RTA small and digitiform (Fig. 19B) ...... Ianduba beaga sp.n.

137 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 10. Scanning electron microscopy, somatic morphology. A–H: Ianduba varia (Keyserling, 1891) (UFMG 6055), female. A: Left tarsus I, apical view. B: Left palp, claw, subapical view. C: Left tarsus I, trichobothrial socket. D: Left leg I, tarsal organ. E: Left tarsus I, claw tuft setae. F: Left metatarsus I, scopula setae. G: Left metatarsus I, retrolateral view. H: Left metatarsus I, metatarsus stopper, dorsal view. I: Ianduba angeloi sp.n. (UFMG), male, eye region, anterior. Scale bars: A, H = 0.05 mm; B–D, E–F = 0.01 mm; G = 0.5 mm; I = 0.1 mm.

15’ Dorsal lobe of the RTA large and club-shaped (Fig. 17 Copulatory ducts extend anteriorly to the spermathe- 7A) ...... Ianduba varia cae; epigynal median plate distinctly raised from epigynal area (Fig. 13C – D) Females (those of I. patua are unknown) ...... 18 (remaining members of vatapa group) 16 Secondary spermathecae (visible ventrally) large, 17’ Copulatory ducts not extending anteriorly to the globose (Fig. 11D) ...... Ianduba acaraje sp.n. spermathecae; epigynal median plate at the same 16’ Secondary spermathecae absent or inconspicuous, plane of epigynal area (Fig. 19C – D) not visible ventrally (Figs. 13D, 19D) ...... 17 ...... 23 (varia group)

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18 Atrial pockets extended medially (Fig. 13C) ...... 19 (Fig. 13; CD). (Potential synapomorphies of the group 18’ Atrial pockets restricted to anterior half of epigynum marked with an asterisk *.) (Fig. 12A) ...... 21 19 Epigynal median plate notably expanded posteriorly Notes. Ianduba acaraje sp.n. has a rather deviant palpal (Bonaldo et al. 2007: fig. 28) ...... Ianduba abara morphology, lacking many of the putative apomorphic 19’ Epigynal median plate not strongly expanded poste- features found in either the vatapa and varia groups. We riorly (Fig. 13C) ...... 20 have decided to allocate it in the vatapa group because it 20 Anterior margins of epigynal median plate strongly shares some morphological characters, notably the fold- incised (Fig. 13C) ...... Ianduba apururuca sp.n. ed RTA with an apical spur in the ventral lobe and large 20’ Anterior margins of epigynal median plate not strong- copulatory ducts that extend anteriorly beyond the sper- ly incised (Bonaldo 1997: fig. 27) mathecae. Also, its area of distribution is closer to that of ...... Ianduba paubrasil members of the vatapa group. However, its taxonomic 21 Epigynal plate wide, almost wider than long (Bo position would be better tested by means of a phyloge- naldo 1997: fig. 17) ...... Ianduba vatapa netic analysis of the genus. 21’ Epigynal plate narrow ...... 22 22 Anterior margins of epigynal plate arched (Bonaldo Distribution. Southern Bahia endemism center of the et al. 2007: fig. 3) ...... Ianduba mugunza Brazilian Atlantic rainforest (Fig. 21). 22’ Anterior margins of epigynal plate nearly straight (Fig. 12A) ...... Ianduba caxixe Included species. I. abara Bonaldo & Brescovit, 2007, 23 Epyginal plate longer than wide; atrial pockets con- I. acaraje sp.n., I. apururuca sp.n., I. caxixe Bonaldo, vergent (Fig. 8A) ...... Ianduba varia 1997, I. mugunza Bonaldo & Brescovit, 1997, I. patua 23’ Epigynal plate and atrial pockets otherwise ...... 24 Bonaldo, 1997, I. paubrasil Bonaldo, 1997, I. vatapa Bo 24 Epigynal plate strongly notched posteriorly; atrial naldo, 1997. pockets wide, posteriorly placed (Fig. 14C) ...... Ianduba capixaba sp.n. 5.3.1. Ianduba acaraje sp.n. 24’ Epigynal plate otherwise; atrial pockets, if wide, anteriorly placed ...... 25 Fig. 11 25 Atrial pockets wide, anteriorly placed (Figs. 18C, 19C) ...... 26 Type material. Male holotype from Serra da Jibóia, Santa Teres- 25’ Atrial pockets inconspicuous, posteriorly placed inha, Bahia, Brazil (–39.50145, –12.82132, 803 m), G.C. Ferreira col., 24 – 27/IV/2009 (IBSP 146285). Paratypes: two females with (Figs. 16C, 17C) ...... 27 the same collecting data (IBSP 146283, IBSP 146287); one male 26 Atrial pockets openings anteriorly directed (Fig. from the same locality, 8/XI/2010, L.S. Carvalho col. (CNHUFPI 19C) ...... Ianduba beaga sp.n. 160). 26’ Atrial pockets openings laterally directed (Fig. 18C) ...... Ianduba benjori sp.n. Derivatio nominis. The Brazilian Portuguese word “aca- 27 Epigynal plate with M-shaped reinforced posterior rajé” refers to a typical dish from the cuisine of State of margin; DPP absent (Fig. 15D) Bahia, consisting of fried fradinho-bean pasta filled with ...... Ianduba dabadu sp.n. dried shrimp, vatapá, caruru and chilli. The name should 27’ Epigynal plate without reinforced posterior margin; be treated as a noun in apposition. DPP present (Figs. 16D, 17D) ...... 28 28 Dorsal plate oval; spermathecae not covered by DP Differential diagnosis. This is the most distinct spe- (Fig. 17D) ...... Ianduba angeloi sp.n. cies of Ianduba. Males have a short, prolaterally in- 28’ Dorsal plate shield-shaped; spermathecae partially serted embolus with an enlarged base, but no processes; covered by DP (Fig. 16D) ...... Ianduba liberta sp.n. a single-pieced median apophysis folded upon itself; a large, hyaline conductor; and no tegular projections (Fig. 11A,B). Females have two round, black atrial pockets at 5.3. The vatapa group the epigynum, and the secondary spermathecae are large and extend anteriorly to the primary spermathecae (Fig. Diagnosis. Males of this group can be recognized by the 11C,D). prolaterally inserted, straight to slightly sinuous embolus; by the large, often sculptured, embolic base* (Figs. Description. Male holotype: Carapace dark brown. Che 6, 13; EB); by the reduced conductor* (large in I. aca- licerae, endites, labium and sternum brown. Leg I dark raje sp.n.); and by the complex and folded RTA, with an brown, except for white tibia and tarsus. Legs II – III yel- apical spur in the ventral lobe that seems to be detached low, except for brown femora and patellae. Leg IV brown, from the rest of the apophysis* (Figs. 6, 13; VL, AS). Fe- except for yellow metatarsus. Abdomen browish gray, males can be recognized by the large and raised median with dark brown scutum. Total length 6.48. Carapace plate of the epigynum (Figs. 8D, 13; MP) and by the long 3.08 long, 2.12 wide, 0.81 high. Clypeus 0.31 high. Ante- copulatory ducts extending anteriorly to the spermathe- rior eye row 0.69 wide, posterior eye row 0.81 wide. Eye cae*; these are folded* in all species but I. acaraje sp.n. diameters and interdistances: AME 0.13, ALE 0.13, PME

139 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 11. Ianduba acaraje sp.n. A,B: Male holotype (IBSP 146285). A: Palp, ventral view. B: Palp, retrolateral view. C,D: Female paratype (IBSP 146283). C: Epigynum, ventral view. D: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Scale bars = 0.1 mm.

0.13, PLE 0.10, AME – AME 0.13, AME – ALE 0.10, 1.02, 2.36, 2.75, 1.32. Leg spination: I– tibia v2-2-2, me PME – PME 0.15, PME – PLE 0.18, ALE – PLE 0.05. Me- tatarsus v2-2. II– tibia v2-2-2, metatarsus v2-2. III– tibia dian ocular quadrangle length 0.36, anterior width 0.33, d2-2, v1-2-2, metatarsus d2-2-2, v2-2-3. IV(right leg)– posterior width 0.36. Chelicerae long, with three promar- tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-2. Palp (Fig. ginal teeth, two retromarginal teeth and setal tufts on pro- 11A,B). Retrolateral tibial apophysis with two-branched, margins; no ventral tooth-like apophysis. Sternum 1.58 folded ventral lobe, with apical spur with rounded apex; long, 1.14 wide. Abdomen 2.36 long, 1.42 wide. Leg dorsal lobe spatulate. Tegulum without any apophyses, measurements: I– femur 2.31, patella 0.99, tibia 2.17, with incision near insertion of embolus and small, whit- metatarsus 1.96, tarsus 1.30. II– 2.01, 0.84, 1.78, 1.68, ish bump on retrolateral edge. Median apophysis made of 1.17. III– 1.81, 0.84, 1.50, 1.83, 1.04. IV(right leg)– 2.49, single piece with rounded, blunt margin that folds onto

140 ARTHROPOD SYSTEMATICS & PHYLOGENY — 74 (2) 2016

itself. Conductor large, rounded, hyaline, arising dorsally 5.3.2. Ianduba caxixe Bonaldo, 1997 to median apophysis and positioned in apposition to em- Figs. 6, 8D – F, 12 bolus. Embolus short, prolaterally inserted, with enlarged base and acute apex, without any processes. Ianduba caxixe Bonaldo, 1997: 175, figs. 12, 23, 24. Female paratype (IBSP 146283): Carapace and chelicerae dark brown. Endites, labium and sternum brown. Type material. Male holotype from Matiapã Farm, Camacan, Ba- Leg I dark brown, except for whitish yellow distal end hia, Brazil, deposited in Museu de Ciências Naturais da Fundação Zoobotânica in Porto Alegre, Brazil (MCN 27666), examined. of tibia and tarsus. Legs II – III light brown, except for dark brown femora and patellae. Leg IV light brown, Additional material examined. BRAZIL: Bahia: Ilhéus, CE- PLAC [–39.0333, –14.8167, 8 m], 7/IV/1998, A.D. Brescovit col., except for dark brown femur, patella and tibia. Abdomen 1♂ (IBSP 19364); Una, Estação Ecológica de Una (–39.07444, browish gray. Total length 8.60. Carapace 3.92 long, –15.29667, 31 m), 2003, M.F. Dias col., 1♂ (IBSP 62562), 1♂ 2.58 wide, 1.19 high. Clypeus 0.38 high. Anterior eye 1♀ (IBSP 62563), 1♂ (IBSP 62564), 1♂ (IBSP 62565), 1♂ (IBSP row 0.81 wide, posterior eye row 0.99 wide. Eye dia 62566); same locality and collector, X/1999–IX/2000, 1♂ (IBSP meters and interdistances: AME 0.15, ALE 0.15, PME 63901), 1♂ (IBSP 63904), 2♀ (IBSP 63906), 2♂ 1♀ (IBSP 63907), 1♂ (IBSP 63915), 1♀ (IBSP 64254), 1♀ (IBSP 64285), 1♀ (IBSP 0.13, PLE 0.13, AME – AME 0.08, AME – ALE 0.1, 64373), 1♀ (IBSP 64409), 1♂ (IBSP 64413), 1♀ (IBSP 64430), PME – PME 0.15, PME – PLE 0.23, ALE – PLE 0.08. 1♀ (IBSP 64432), 1♂ (IBSP 64461), 1♂ (IBSP 64462), 1♂ (IBSP Median ocular quadrangle length 0.46, anterior width 64470), 2♂ (IBSP 64479), 1♂ (IBSP 64518), 1♀ (IBSP 64563), 1♀ 0.41, posterior width 0.41. Chelicerae 1.53 long, with (IBSP 65092), 1♀ (IBSP 65146), 1♀ (IBSP 65154), 1♂ 1♀ (IBSP three promarginal teeth, two retromarginal teeth and se- 65180), 1♂ (IBSP 65191), 1♀ (IBSP 65195), 1♂ (IBSP 65204), 1♂ (IBSP 65243), 1♂ (IBSP 65253), 1♂ (IBSP 65332), 1♀ (IBSP tal tufts on promargins; no ventral tooth-like apophysis. 65336), 1♀ (IBSP 65349), 1♀ (IBSP 65355), 1♂ (IBSP 65358), Sternum 1.96 long, 1.47 wide. Abdomen 4.50 long, 2.35 1♂ (IBSP 65368), 1♂ (IBSP 65371), 1♀ (IBSP 65403), 1♂ (IBSP wide. Leg measurements: I– femur 2.64, patella 1.37, 65406), 1♂ 1♀ (IBSP 65455), 1♂ (IBSP 65461), 1♂ (IBSP 65470). tibia 2.54, metatarsus 2.19, tarsus 1.45. II– 2.21, 1.04, 2.14, 2.08, 1.37. III– 1.83, 1.12, 1.83, 2.19, 1.17. IV– Differential diagnosis. Male diagnosed by Bonaldo 3.02, 1.27, 2.87, 3.39, 1.5. Leg spination: I– tibia v2-2-2, (1997). Females are most similar to those of I. mugunza metatarsus v2-2. II– tibia v2-2-2, metatarsus v2-2. III– in having a posteriorly widened median plate of the epi tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-3. IV– tibia gynum. They can be distinguished by the more angulose d2-2, v2-2-2, metatarsus d2-2-2, v2-2-3. Genitalia (Fig. atrial pockets of the median plate, by the less convoluted 11C,D). Epigynum with wide, subquadrate median plate copulatory ducts, and by the much narrower dorsal piece between two black, rounded excavated areas that con- of the vulva (Fig. 12A,B). tinue to two posterior incisions. Vulva with short dorsal piece which is widest anteriorly and notched anteriorly; Description. Male described by Bonaldo (1997). secondary spermathecae huge, extending anteriorly and Female from Una, Bahia (IBSP 63906): Carapace and covering primary spermathecae dorsally; primary sper- chelicerae brown. Endites, labium and sternum light mathecae small, oval, with fertilization ducts directed brown. Leg I light brown, except for whitish yellow dis- anteriorly. tal end of tibia and yellow tarsus. Legs II – IV light brown, except for yellow metatarsi and tarsi. Abdomen browish Variation. Two males: total length 6.48 – 7.15; carapace gray, with tiny brown dorsal scutum on anterior end. Total length 3.08 – 3.47; femur I 2.31 – 2.47. Two females: to- length 8.51. Carapace 3.86 long, 2.67 wide, 1.37 high. tal length 8.6 – 9.65; carapace length 3.92 – 3.99; femur I Clypeus 0.37 high. Anterior eye row 0.83 wide, poste- 2.64 – 2.87. rior eye row 0.95 wide. Eye diameters and interdistances: AME 0.17, ALE 0.15, PME 0.13, PLE 0.13, AME – AME Natural history. This species has been manually col- 0.08, AME – ALE 0.10, PME – PME 0.17, PME – PLE lected from a secondary montane Atlantic rainforest frag- 0.17, ALE – PLE 0.07. Median ocular quadrangle length ment in Bahia, at 800 m above sea level. The lowlands 0.38, anterior width 0.42, posterior width 0.42. Chelicer- surrounding the Serra da Jibóia range, where the species ae 1.53 long, with three promarginal teeth, four retromar- was found, are dominated by Caatinga (a strongly xero- ginal teeth and setal tufts on promargins; no ventral tooth- morphic dry forest) and are heavily degraded (L.S. Car- like apophysis. Sternum 1.83 long, 1.40 wide. Abdomen valho, pers. comm.). 4.26 long, 2.10 wide. Leg measurements: I– femur 2.73, patella 1.40, tibia 2.77, metatarsus 2.30, tarsus 1.53. II– Note. The species is sympatric with I. patua, which is 2.50, 1.33, 2.40, 2.17, 1.50. III– 2.33, 1.00, 2.07, 2.27, known only from the male. Although there is a possibility 1.20. IV– 3.07, 1.40, 2.90, 3.59, 1.63. Leg spination: I– that the female here described might belong to I. patua, tibia v2-2-2, metatarsus v2-2. II– tibia v2-2-2, metatarsus we believe that our matching is more likely since both v2-2. III– tibia d2-2, v1-2-2, metatarsus d2-2-2, v2-2-3. sexes have a genital morphology quite distinct from other IV– tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-3. Pos- species of the vatapa group. terior tracheae as in Fig. 12C,D. Genitalia (Fig. 12A,B). Epigynum with posteriorly widened median plate flanked Distribution. Known from a single locality in eastern by two narrow atrial pockets. Vulva with trapezoidal Bahia, Brazil (Fig. 23). dorsal piece much longer than wide; sclerotized copula-

141 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 12. Ianduba caxixe Bonaldo, 1997. A,B: Female from Una, Bahia (IBSP 65455). A: Epigynum, ventral view. B: Epigynum, dorsal view, cleared. C,D: Posterior tracheae, dorsal view. See chapter 3 for abbreviations. Scale bars: A, B = 0.1 mm; C = 0.5 mm; D = 0.25 mm. tory ducts extend anteriorly; primary spermathecae small, 5.3.3. Ianduba apururuca sp.n. oval, deeply attached to surrounding copulatory ducts, Fig. 13 with fertilization ducts directed mesally. Type material. Male holotype from Fazenda Montes Claros, Es- Variation. Three females: total length 8.11 – 8.51; cara- tação Biológica de Caratinga, Caratinga, Minas Gerais, Brazil pace length 3.30 – 3.86; femur I 2.23 – 2.73. [–41.79222, –19.72417, 457 m], L.G. Dias col., 25/II/2000 (UFMG 1172). Paratype: one male from Estação de Preservação e Desen- Natural history. Specimens were collected in pitfall volvimento Ambiental de Peti, Santa Bárbara, Minas Gerais, Brazil (–43.36667, –19.88333, 768 m), G.H.F. Azevedo et al. col., 8 – 9/ traps in coastal Atlantic forest. XII/2012 (UFMG 13116). Additional material examined. BRAZIL: Espírito Santo: Santa Note. Several males and females were collected in the Teresa, Reserva Biológica Augusto Ruschi [–40.6003, –19.9356, same locality in Una, Bahia throughout the years of 756 m], T. Souza, T. Bernabé & E. Soveiro col., IV – VIII/2006, 1♀ 1999 – 2000. 1 juvenile (IBSP 121252).

Distribution. Known from southern Bahia, Brazil (Fig. Note. Males and females have not been collected in the 23). same localities. They have been matched because they

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Fig. 13. Ianduba apururuca sp.n. A–C: Male holotype (UFMG 1172). A: Palp, ventral view. B: Palp, retrolateral view. C: Palpal tibia, ventral view. D,E: Female (IBSP 121252). D: Epigynum, ventral view. E: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Scale bars = 0.1 mm. are the only records of the vatapa group occurring south with denticles (Fig. 13A,B). Females differ by the medi- of the Doce river. The association is uncertain. ally widened median plate of the epigynum, by the short dorsal piece of the vulva, and by the anteriorly elongated Derivatio nominis. The Brazilian Portuguese expression copulatory ducts (Fig. 13D,E). “à pururuca” is used to refer to a piglet that is whole- roasted and then bathed in hot oil, so that its skin be- Description. Male holotype: Carapace, chelicerae, en- comes crispy while the interior is soft and tender. This dites, labium and sternum dark brown. Leg I dark brown is a typical dish from the cuisine of Minas Gerais, the except for yellow tibia and tarsus. Legs II and III light type locality. The name should be treated as a noun in brown except for dark brown coxae, trochanters and fem- apposition. ora. Leg IV with dark brown coxa, trochanter and femur, light brown patella, tibia and metatarsus, and yellow tar- Differential diagnosis. Males differ from other Ian- sus. Abdomen dark brown. Total length 7.83. Carapace duba species with a large, curved tegular projection by 3.84 long, 2.68 wide, 1.42 high. Clypeus 0.38 high. Ante- the large and conical dorsal lobe of the RTA, and by the rior eye row 0.81 wide, posterior eye row 0.94 wide. Eye elongated MA with a short, trapezoidal prolateral sector diameters and interdistances: AME 0.18, ALE 0.15, PME

143 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

0.10, PLE 0.13, AME – AME 0.10, AME – ALE 0.10, Santa Bárbara has transverse posterior gray markings on PME – PME 0.18, PME – PLE 0.20, ALE – PLE 0.08. the posterior dorsum of the abdomen. Median ocular quadrangle length 0.46, anterior width 0.41, posterior width 0.43. Chelicerae 1.22 long, with Natural history. Specimens have been collected in areas two promarginal and two retromarginal teeth. Ventral of semi-deciduous Atlantic forest. Male specimens have retrolateral surface with short distal tooth-like apophysis. been collected in the rainy season (between December Sternum 1.88 long, 1.40 wide. Abdomen 4.26 long, 1.98 and February). wide. Leg measurements: I– femur 2.81, patella 1.25, tibia 2.54, metatarsus 2.26, tarsus 1.63. II– 2.42, 1.17, Distribution. Eastern Minas Gerais and western Espírito 2.08, 1.96, 1.40. III– 2.16, 0.99, 1.83, 2.06, 1.14. IV– Santo states, Brazil (Fig. 22). 2.95, 1.19, 2.75, 3.29, 1.58. Leg spination. I– tibia v2-2- 2, metatarsus v2-2, II– tibia v2-2-2, metatarsus v2-2, III– tibia v2, metatarsus v2-2, d2-2-2, IV– tibia v1-2-2, d2-2, 5.4. The varia group metatarsus v2-2, d2-2-2. Palp (Fig. 13A,B). Retrolateral tibial apophysis with short, single-branched ventral lobe Diagnosis. Males of this group can be recognized by the with apical spur, a large, conical, single-branched dorsal curved embolus, which starts going retrolaterally and lobe, and folded area prolaterad to dorsal lobe. Tegulum then has a gentle curve to meet the conductor ventrally with deep incision occupied by membranous area and (Fig. 14; E)*; by the lack of a sculptured embolic base large, curved tegular projection. Median apophysis elon- (except in I. varia and I. beaga sp.n., visible in the ex- gated and roughly triangular, with digitiform aMA and panded palp); and by the large, membranous conductor rMA and short, trapezoidal pMA with denticles. Conduc- (also present in I. acaraje sp.n.); the conductor arises tor hidden by MA in ventral view, with basal prolateral ventrally to the median apophysis (Figs. 5, 7, 14; C)*. sclerotized spur. Embolus long and slightly sinuous, pro- Females can be recognized by the epigynum with very laterally inserted, with enlarged base, ventral digitiform inconspicuous openings, with an unraised median plate process and prolateral, folded process. at the same height as the lateral plates, and by the normal Female from Santa Tereza (IBSP 121252): Carapace copulatory ducts, not extending anteriorly to the sper- and chelicerae brown. Endites, labium and sternum light mathecae (Figs. 8, 14; CO, CD). (Potential synapomor- brown. Leg I light brown, except for whitish yellow phies of the group marked with an asterisk *.) distal end of tibia and yellow tarsus. Legs II – III light brown. Leg IV light brown, except for yellow metatar- Distribution. Serra do Mar and Southern Brazil ende sus and tarsus. Abdomen brownish gray, with light gray mism centers of the Brazilian Atlantic rainforest (Fig. venter and dark transverse dorsal markings. Total length 21). 8.50. Carapace 4.16 long, 2.87 wide, 1.47 high. Clypeus 0.36 high. Anterior eye row 0.92 wide, posterior eye Included species. I. angeloi sp.n., I. beaga sp.n., I. ben- row 1.04 wide. Eye diameters and interdistances: AME jori sp.n., I. capixaba sp.n., I. dabadu sp.n., I. liberta 0.18, ALE 0.18, PME 0.13, PLE 0.13, AME – AME 0.13, sp.n., I. varia (Keyserling, 1891). AME – ALE 0.08, PME – PME 0.20, PME – PLE 0.25, ALE – PLE 0.08. Median ocular quadrangle length 0.46, 5.4.1. Ianduba capixaba sp.n. anterior width 0.48, posterior width 0.46. Chelicerae 1.70 long, with three promarginal teeth, three retromarginal Fig. 14 teeth and setal tufts on promargins; no ventral tooth-like apophysis. Sternum 2.11 long, 1.65 wide. Abdomen 2.16 Type material. Male holotype from near Reserva Biológica Au- long, 2.34 wide. Leg measurements: I– femur 2.98, patel- gusto Ruschi, Santa Teresa, Espírito Santo, Brazil (–40.57667, –19.86583, 896 m), 5 – 6/VII/2013, P.H. Martins & M.T.T. Santos la 1.45, tibia 2.81, metatarsus 2.21, tarsus 1.58. II– 2.70, col. (UFMG 13977). Paratypes: one female from Santa Teresa, Es- 1.37, 2.44, 2.11, 1.50. III– 2.47, 1.25, 2.06, 2.31, 1.30. tação Biológica de Santa Lúcia (–40.53222, –19.96194, 935 m), 7/ IV– 3.25, 1.45, 3.15, 3.71, 1.53. Leg spination: I– tibia VII/2013, P.H. Martins & M.T.T. Santos col. (UFMG 14029); two v2-2-2, metatarsus v2-2. II– tibia v2-2-2, metatarsus v2- males, one male and one female from Santa Teresa, Reserva Biológi- ca Augusto Ruschi (–40.6003, –19.9356, 756 m), IV – VIII/2006, T. 2. III– tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-3. IV– Souza, T. Bernabé & E. Soveiro col. (IBSP 121336, IBSP 121651, tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-3. Genitalia IBSP 121582); two males from Santa Teresa (–40.51667, –19.95, (Fig. 13D,E). Epigynum with medially widened median 729 m), IV – IX/2005, T. Souza col. (IBSP 132693). plate flanked by two atrial pockets.Vulva with short trapezoidal dorsal piece divided longitudinally; long, folded Note. Males and females have been matched because copulatory ducts extend anteriorly, with tiny secondary both have been collected in two independent expeditions spermathecae directed posteriorly; primary spermathe- to Santa Teresa, Espírito Santo. cae small, oval, with fertilization ducts directed laterally, inserted in internal surface of spermathecae. Derivatio nominis. The Brazilian Portuguese word “ca pixaba” is used to refer to people born in the State of Es- Variation. Two males: total length 7.50 – 7.83; carapace pírito Santo, where the type locality is placed. The name length 3.32 – 3.84; femur I 2.58 – 2.81. The male from should be treated as a noun in apposition.

144 ARTHROPOD SYSTEMATICS & PHYLOGENY — 74 (2) 2016

Fig. 14. Ianduba capixaba sp.n. A,B: Male holotype (UFMG 13977). A: Palp, ventral view. B: Palp, retrolateral view. C,D: Female paratype (IBSP 121582). C: Epigynum, ventral view. D: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Dot line indicating DL. Scale bars = 0.1 mm.

Differential diagnosis. Males share a simple median Description. Male holotype: Carapace, chelicerae and apophysis and a short prolateral tegular projection with labium dark brown. Endites and sternum light brown. I. liberta sp.n. and I. angeloi sp.n.; they can be distin- Leg I yellow, except for dark brown femur and white tib- guished by the enlarged and sclerotized rMA, which is ia and tarsus. Legs II – III yellow. Leg IV yellow, except fused to the base of the conductor, and by having a broad, for dark brown and white markings on tibia and metatar- striate and transparent plate-shaped median process in sus. Abdomen brown, with light gray venter and trans- the dorsal lobe of the RTA (Fig. 14A,B). Females can be verse dorsal markings. Total length 6.00. Carapace 2.98 distinguished by the strong and sinuous copulatory ducts, long, 2.14 wide, 1.07 high. Clypeus 0.31 high. Anterior almost as wide as the primary spermathecae, visible in eye row 0.71 wide, posterior eye row 0.86 wide. Eye di- ventral and dorsal views; by the short median piece, and ameters and interdistances: AME 0.13, ALE 0.13, PME by the wide posterior piece of the vulva (Fig. 14C,D). 0.13, PLE 0.13, AME – AME 0.10, AME – ALE 0.08,

145 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

PME – PME 0.15, PME – PLE 0.20, ALE – PLE 0.08. Variation. Three males: total length 6.00 – 6.61; cara- Median ocular quadrangle length 0.41, anterior width pace length 2.98 – 3.25; femur I 2.49 – 2.67. Two females: 0.36, posterior width 0.38. Chelicerae 1.25 long, with total length 7.17 – 7.25; carapace length 2.98 – 3.05; fe- three promarginal teeth, two retromarginal teeth and se- mur I 2.29 – 2.36. The shape of the dorsal piece of the tal tufts on promargins; no ventral tooth-like apophysis. vulva is variable (some specimens have the sides slightly Sternum 1.63 long, 1.32 wide. Abdomen 2.92 long, 1.40 indented). wide. Leg measurements: I– femur 2.49, patella 1.07, tibia 2.59, metatarsus 2.29, tarsus 1.58. II(right leg)– 2.21, Natural history. This species has been manually collect- 1.02, 2.01, 1.88, 1.30. III– 1.91, 0.92, 1.65, 1.93, 1.04. ed in montane Atlantic rainforest, at altitudes between IV– 2.67, 0.93, 2.70, 3.19, 1.37. Leg spination: I– tibia 750 and 930 m above sea level. All adult specimens have v2-2-2, metatarsus v2-2. II– tibia v2-2-2, metatarsus v2- been collected in the dry season (April to September). 2. III(right leg)– tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-2. IV– tibia d2-2, v1-2-2, metatarsus d2-2-2, v2-2- Distribution. Known from three localities in Santa Te- 2. Palp (Fig. 14A,B). Retrolateral tibial apophysis with resa, Espírito Santo, Brazil (Fig. 22). ventral lobe displaced dorsally, dorsal lobe trifid, delimited and slightly detached from cuticle of tibia by membra- 5.4.2. Ianduba dabadu sp.n. nous strip; with simple basal process with truncate apex, and simple apical process with acuminate apex; these Figs. 15, 22 processes are separated by a transparent, striate and wide median process. Tegulum with small prolateral apophysis Type material. Male holotype from Reserva Estadual de Duas Bocas, Cariacica, Espírito Santo, Brazil (–40.47667, –20.27294, and projected, whitish bump on its retrolateral edge. Me- 237 m), 13 – 14/X/2005, A.Giupponi col. (IBSP 133310). dian apophysis two-pieced, with two sectors: pMA with Additional material examined. BRAZIL: Espírito Santo: Do- truncate apex, and sculptured rMA fused to base of con- mingos Martins, São Paulo do Aracê, private área near Parque ductor. Conductor large, rounded, membranous, arising Estadual de Pedra Azul (–41.02, –20.4325, 1148 m), 12/VII/2013, ventrally to median apophysis, positioned in apposition P.H. Martins & M.T.T.Santos col., 2♀ (UFMG 14211). to embolus. Embolus long, filiform, curved to meet conductor retrolaterally, without any processes. Note. Males and females have not been collected in the Female paratype (IBSP 121582): Carapace and cheli- same locality; they have been matched because both oc- cerae brown, endites and labium dark brown. Sternum cur in Espírito Santo, and both resemble Ianduba capixa- light brown. Leg I light brown, except for whitish yel- ba sp.n. in genitalic morphology. The association is un- low distal end of tibia and dark brown femur. Legs II – certain: the male has been collected at 230 m above sea III light brown, except for dark brown femora. Leg IV level, while the females were collected at 1150 m. light brown, except for dark brown femur and tibia; tibia with light brown markings. Abdomen brownish gray. Derivatio nominis. The specific name is a combination Total length 7.25. Carapace 2.98 long, 2.24 wide, 1.14 of letters, which, in conjunction with the generic name, high. Clypeus 0.23 high. Anterior eye row 0.71 wide, are intended to sound like the famous yell of Fred Flint- posterior eye row 0.81 wide. Eye diameters and inter- stone, a fictional Hanna-Barbera’s character. distances: AME 0.13, ALE 0.13, PME 0.13, PLE 0.13, AME – AME 0.10, AME – ALE 0.10, PME – PME 0.15, Differential diagnosis. Males share a short prolateral PME – PLE 0.18, ALE – PLE 0.08. Median ocular quad- tegular projection with I. capixaba sp.n., I. liberta sp.n. rangle length 0.41, anterior width 0.36, posterior width and I. angeloi sp.n.; they can be distinguished by the 0.38. Chelicerae 1.22 long, with three promarginal teeth, two-pieced median apophysis; by the enlarged base of two retromarginal teeth and setal tufts on promargins; the embolus; and by the distinctive RTA: the dorsal lobe no ventral tooth-like apophysis. Sternum 1.65 long, 1.27 is trifid, having a simple basal process, a very large medi- wide. Abdomen 4.21 long, 2.54 wide. Leg measure- an process with a spine in the apex, and a slender, curved, ments: I– femur 2.29, patella 1.19, tibia 2.36, metatarsus digitiform apical process embracing the base of the me- 1.96, tarsus 1.35. II– 2.16, 1.12, 2.03, 1.81, 1.25. III– dian process (Fig. 15A,B). Females are distinguished 1.93, 0.94, 1.60, 1.88, 1.02. IV– 2.61, 1.14, 2.54, 2.98, by the very wide copulatory ducts (wider than the sper- 1.27. Leg spination: I– tibia v2-2-2, metatarsus v2-2. mathecae); by the rectangular median piece, which is II– tibia v2-2-2, metatarsus v2-2. III– tibia d2-2, v1-2-2, longer than wide; and by the M-shaped posterior piece of metatarsus d2-2-2, v2-2-2. IV– tibia d2-2, v1-2-2, meta- the vulva (Fig. 15C,D). tarsus d2-2-2-2, v2-2-2. Genitalia (Fig. 14C,D). Epigy- num lightly sclerotized, with two narrow, median copu- Description. Male holotype: Carapace and chelicerae latory openings, posterior piece wide. Vulva with short brown, endites dark brown. Labium and sternum light dorsal piece, which is widest posteriorly and anteriorly brown. Leg I light brown, except for whitish yellow tibia connected to spermathecae; copulatory ducts almost as and dark brown femur. Legs II – III light brown. Leg IV wide as primary spermathecae, sinuous and posteriorly light brown, except for dark brown markings on femur, directed; primary spermathecae small, oval, with fertili- patella and tibia. Abdomen brown with light gray venter zation ducts directed mesally. and transverse dorsal markings. Total length 6.14. Cara-

146 ARTHROPOD SYSTEMATICS & PHYLOGENY — 74 (2) 2016

Fig. 15. Ianduba dabadu sp.n. A–C: Male holotype (IBSP 133310). A: Palp, ventral view. B: Palp, retrolateral view. C: Palpal tibia, dorsal view. D,E: Female (UFMG 14211). D: Epigynum, ventral view. E: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Dot line indicating DL. Scale bars = 0.1 mm. pace 3.08 long, 2.14 wide, 0.94 high. Clypeus 0.31 high. with three promarginal teeth, two retromarginal teeth and Anterior eye row 0.71 wide, posterior eye row 0.89 wide. setal tufts on promargins; no ventral tooth-like apophy- Eye diameters and interdistances: AME 0.15, ALE 0.13, sis. Sternum 1.58 long, 1.30 wide. Abdomen 3.05 long, PME 0.13, PLE 0.13, AME – AME 0.08, AME – ALE 1.46 wide. Leg measurements: I– femur 2.49, patella 0.05, PME – PME 0.18, PME – PLE 0.20, ALE – PLE 1.07, tibia 2.54, metatarsus 2.21, tarsus 1.50. II– 2.21, 0.05. Median ocular quadrangle length 0.38, anterior 1.04, 2.03, 1.91, 1.30. III– 1.93, 0.89, 1.68, 1.96, 1.04. width 0.36, posterior width 0.41. Chelicerae 1.32 long, IV– 2.67, 1.12, 2.70, 3.22, 1.37. Leg spination: I– tibia

147 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

v2-2-2, metatarsus v2-2. II– tibia v2-2-2, metatarsus v2- 5.4.3. Ianduba liberta sp.n. 2. III– tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-3. IV– Fig. 16 tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-3. Palp (Fig. 15A,B). Retrolateral tibial apophysis with bifid ventral Type material. Male holotype from Parque Estadual do Itacolomi, lobe displaced dorsally, and trifid dorsal lobe delimited Ouro Preto, Minas Gerais, Brazil (–43.50906, –20.43481, 1343 m), by membranous strip, composed of digitiform basal pro- K.P. Santos et al. col., 13/IV/2008 (UFMG 2371). One male paratype, same collecting data (MPEG 24752); one female and two cess, slender finger-like apical process, which embraces male paratypes, same locality and collector, 2 – 4/XI/2007 (UFMG base of median process, which is large, digitiform and 2068, UFMG 2069, UFMG 2070). bears spine near its apex. Tegulum with small prolateral apophysis and projected, whitish bump at retrolateral Derivatio nominis. The name is a Latin adjective mean- edge. Median apophysis made of two pieces: sharp, tri- ing “freed” or “set free”, and is an allusion to the fact that angular retrolateral sector and digitiform prolateral sec- the Inconfidência Mineira, one of the separatist move- tor. Conductor large, rounded, membranous, arising ven- ments that eventually led to Brazilian independency from trally to median apophysis, positioned in apposition to Portugal in 1822, had its main headquarters in Ouro Pre- embolus. Embolus long, ribbon-shaped, curved to meet to, the type locality of this species. conductor retrolaterally, with enlarged base but without any processes. Differential diagnosis. Males share a short prolateral Female from Domingos Martins (UFMG 14211): tegular projection with I. capixaba sp.n., I. dabadu sp.n. Carapace and chelicerae dark brown. Endites, labium and I. angeloi sp.n.; they are most similar to I. angeloi and sternum light brown. Leg I light brown, except for sp.n., as both possess a membranous and short median whitish yellow tibia and dark brown femur. Legs II – III process of the DL of the RTA. I. liberta sp.n. can be dis- light brown, except for dark brown femora. Leg IV light tinguished by the longer, unsclerotized tegular projection brown, except for dark brown markings on femur, tibia pointing away from the embolus, by the broad median and metatarsus. Abdomen brown with light gray trans- apophysis with two pointed edges (one ventral and one verse dorsal markings. Total length 8.25. Carapace 3.08 retrolateral), and by the slender and shorter apical pro- long, 2.24 wide, 1.22 high. Clypeus 0.25 high. Anterior cess of the dorsal lobe of the RTA (Fig. 16A,B). Females eye row 0.81 wide, posterior eye row 0.99 wide. Eyes are most similar to I. angeloi sp.n. in having antero-lat- diameters and interdistances: AME 0.15, ALE 0.15, eral projections in the dorsal piece of the vulva; they can PME 0.15, PLE 0.13, AME – AME 0.10, AME – ALE be distinguished by the less sclerotized epigynum, by the 0.10, PME – PME 0.20, PME – PLE 0.18, ALE – PLE spermathecae being partially covered by the dorsal piece, 0.08. Median ocular quadrangle length 0.43, anterior and by the pointed shape of the dorsal piece projections width 0.41, posterior width 0.46. Chelicerae 1.32 long, (Fig. 16C,D). with three promarginal teeth, two retromarginal teeth and setal tufts on promargins; no ventral tooth-like apophy- Description. Male holotype: Carapace, chelicerae dark sis. Sternum 1.60 long, 1.40 wide. Abdomen 4.77 long, brown. Endites, labium dark brown, with light brown 2.75 wide. Leg measurements: I– femur 2.36, patella anterior borders. Sternum light brown. Leg I with dark 1.19, tibia 2.31, metatarsus 1.91, tarsus 1.25. II– 2.19, brown coxa, trochanter and femur, light brown patella 1.09, 1.98, 1.68, 1.12. III– 1.93, 0.94, 1.70, 1.81, 0.92. and metatarsus, whitish yellow tibia and tarsus. Legs II IV– 2.58, 1.14, 2.61, 2.98, 1.19. Leg spination: I– tibia and III with dark brown femora, other articles yellow. v2-2-2, metatarsus v2-2. II– tibia v2-2-2, metatarsus v2- Leg IV with yellow coxa, trochanter and tarsus, other 2. III– tibia d2-2, v1-2-2, metatarsus d2-2-2, v2-2-3. IV– articles dark brown with proximal and distal yellow bor- tibia d2-2, v1-2-2, metatarsus d2-2-2, v2-2-3. Genitalia ders. Abdomen dark brown. Total length 7.50. Carapace (Fig. 15C,D). Epigynum lightly sclerotized, with two 3.51 long, 2.42 wide, 1.07 high. Clypeus 0.36 high. An- narrow, median copulatory openings. Vulva with rectan- terior eye row 0.76 long, posterior eye row 0.97 long. gular dorsal piece, which is wider than long; copulatory Eye diameters and interdistances: AME 0.15, ALE 0.13, ducts wider than spermathecae, straight and posteriorly PME 0.15, PLE 0.15, AME – AME 0.10, AME – ALE directed; primary spermathecae small, oval, with fertili- 0.08, PME – PME 0.20, PME – PLE 0.23, ALE – PLE zation ducts directed mesally; M-shaped posterior piece 0.05. Median ocular quadrangle length 0.43, anterior of epigynum present. width 0.38, posterior width 0.43. Chelicerae 1.60 long, with three promarginal teeth, two retromarginal teeth and Variation. Two females: total length 7.83 – 8.25; cara- setal tufts on promargins. Sternum 1.73 long, 1.45 wide. pace length 3.08 – 3.32; femur I 2.36 – 2.55. Abdomen 3.76 long, 1.88 wide. Leg measurements: I– femur 2.81, patella 1.17, tibia 2.70, metatarsus 2.39, tar- Natural history. Specimens have been collected in low- sus 1.60. II– 2.34, 1.02, 2.06, 1.96, 1.35. III– 2.01, 0.94, land and montane Atlantic rainforest, at altitudes between 1.73, 1.88, 1.02. IV– 2.78, 1.09, 2.67, 3.05, 1.32. Leg 200 and 1150 m above sea level. spination: I– tibia v2-2-2, metatarsus v2-2, II– tibia v2- 2-2, metatarsus v2-2, III– tibia v2-2-2, metatarsus v2-2, Distribution. Southern Espírito Santo state, Brazil d2-2-2, IV– tibia v1-2-2, metatarsus v2-2. Palp (Fig. (Fig. 22). 16A,B). Retrolateral tibial apophysis with ventral lobe

148 ARTHROPOD SYSTEMATICS & PHYLOGENY — 74 (2) 2016

Fig. 16. Ianduba liberta sp.n. A–C: Male holotype (UFMG 2371). A: Palp, ventral view. B: Palp, retrolateral view. C: Palpal tibia, ventral view. D,E: Female paratype (UFMG 2069). D: Epigynum, ventral view. E: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Dot line indicating DL. Scale bars = 0.1 mm. displaced dorsally; dorsal lobe delimited by membranous Female paratype (UFMG 2069): Carapace, chelicerae strip, with simple, hook-shaped basal process, short and dark brown. Endites, labium dark brown, with light brown membranous median process, and apical process slender anterior borders. Sternum dark brown. Leg I dark brown, and slightly bent near apex. Tegulum with prominent pro- except for light brown metatarsus and tarsus, whitish yel- lateral apophysis pointing away from embolus and small low tibia. Legs II and III with dark brown femora, other clear bump in its retrolateral edge. Median apophysis articles yellow. Leg IV light brown, with yellow bands made of single concave piece with two pointy edges, one on proximal ends of articles. Abdomen dark brown. Total ventral and one retrolateral. Conductor large, rounded, length 7.25. Carapace 2.95 long, 2.10 wide, 0.98 high. membranous, arising ventrally to median apophysis and Clypeus 0.23 high. Anterior eye row 0.66 wide, poste- positioned in apposition to embolus. Embolus long, ret- rior eye row 0.81 wide. Eye diameters and interdistances: rolaterally inserted, without any processes. AME 0.13, ALE 0.10, PME 0.13, PLE 0.13, AME – AME

149 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

0.10, AME – ALE 0.08, PME – PME 0.15, PME – PLE Derivatio nominis. The species name is a patronym in 0.13, ALE – PLE 0.08. Median ocular quadrangle length honor of Dr. Angelo Barbosa Monteiro Machado, a bril- 0.36, anterior width 0.33, posterior width 0.36. Chelicer- liant doctor, professor, neuroanatomist, Odonata taxono- ae 1.35 long, with three promarginal teeth, two retromar- mist, conservationist and writer, in recognition for his ginal teeth and setal tufts on promargins. Sternum 1.60 decades of efforts on researching, teaching generations long, 1.37 wide. Abdomen 4.16 long, 2.38 wide. Leg of students, and helping to conserve Brazilian biodiver- measurements: I– femur 2.36, patella 1.07, tibia 2.16, sity. metatarsus 1.78, tarsus 1.17. II– 2.08, 0.99, 1.93, 1.65, 1.07. III– 1.86, 0.86, 1.45, 1.63, 0.94. IV– 2.52, 1.07, Differential diagnosis. Males share a short prolateral 2.42, 2.75, 1.30. Leg spination: I– tibia v2-2-2, meta- tegular projection with I. capixaba sp.n., I. dabadu sp.n. tarsus v2-2. II– tibia v2-2-2, metatarsus v2-2. III– tibia and I. liberta sp.n.; they are most similar to I. liberta d1-2, v2-2, metatarsus d2-2-2, v2-2-2. IV– tibia d2-2, v2- sp.n., as both possess a membranous and short median 2-2, metatarsus d2-2-2, v1-2-2. Genitalia (Fig. 16C,D). process of the DL of the RTA. I. angeloi sp.n. can be Epigynum unsclerotized, with wide median plate; curved distinguished by the short, curved and sclerotized tegular copulatory ducts seen by transparency, delimiting an in- projection pointing towards the embolus, by the slender verted heart-shaped area; copulatory openings narrow, median apophysis, and by the blade-like apical process slit-shaped, posterior. Vulva with large, as wide as long of the dorsal lobe of the RTA (Figs. 17A,B). Females are dorsal piece, with two pointed projections directed ante- most similar to I. liberta sp.n. in having antero-lateral ro-laterally, dorsal piece covering spermathecae almost projections in the dorsal piece of the vulva; they can be completely; copulatory ducts S-shaped; spermathecae distinguished by the strongly sclerotized, oval epigynal small, oval, with fertilization ducts directed medially. median plate, by the spermathecae not being covered by the dorsal piece, and by the membranous, wrinkled ap- Variation. Four males: total length 6.92 – 7.92; carapace pearance of the dorsal piece projections (Figs. 17C,D). length 3.25 – 3.96; femur I 2.81 – 3.08. Some specimens have five short, transverse light patches in the posterior Description. Male holotype: Carapace, chelicerae, en- portion of the abdominal dorsum. dites, labium dark brown. Sternum light brown. Leg I with light brown coxae, whitish yellow tibiae and tarsi, Natural history. Specimens were collected in pitfall other articles dark brown. Legs II – III with dark brown traps in a semi-deciduous Atlantic forest and a secondary femora, other articles brownish yellow. Leg IV with semi-deciduous Atlantic forest area that was occupied by browish yellow coxae, whitish yellow tarsus, other arti- Eucalyptus plantations around 50 years ago. Adult speci- cles dark brown with yellow markings. Abdomen brown- mens were obtained in both rainy and dry seasons. ish gray, with white transverse markings near posterior end of dorsum. Total length 6.82. Carapace 3.18 long, Distribution. Known only from Ouro Preto in central 2.29 wide, 1.12 high. Clypeus 0.36 high. Anterior eye Minas Gerais state, Brazil (Fig. 22). row 0.74 wide, posterior eye row 0.86 wide. Eye dia meters and interdistances: AME 0.15, ALE 0.13, PME 5.4.4. Ianduba angeloi sp.n. 0.13, PLE 0.18, AME – AME 0.08, AME – ALE 0.05, PME – PME 0.15, PME – PLE 0.18, ALE – PLE 0.05. Figs. 5, 10I, 17 Median ocular quadrangle length 0.38, anterior width 0.36, posterior width 0.38. Chelicerae 1.35 long, with Type material. Male holotype from Serra do Ouro Branco, Ouro three promarginal teeth, two retromarginal teeth and se- Branco, Minas Gerais, Brazil (–43.60139, –20.50056, 1095 m), Y. Antonini col., 23/III/2007 (UFMG 12683). Seven male and tal tufts on promargins. Sternum 1.63 long, 1.35 wide. three female paratypes from the same locality, several collecting Abdomen 3.29 long, 1.58 wide. Leg measurements: I– dates (UFMG 12680, UFMG 12681, UFMG 12682, UFMG 12684, femur 2.55, patella 1.12, tibia 2.61, metatarsus 2.31, tar- UFMG 12685, UFMG 12686, UFMG 12687, MPEG 24568, MPEG 24569, MPEG 24570). sus 1.50. II– 2.26, 0.97, 2.03, 1.86, 1.27. III– 1.93, 0.89, Additional material examined. BRAZIL: Minas Gerais: Ouro 1.63, 1.91, 0.99. IV– 2.70, 1.07, 2.34, 3.08, 1.37. Leg Preto, Parque Estadual do Itacolomi (–43.50906, –20.43481, spination: I– tibia v2-2-2, metatarsus v2-2. II– tibia v2- 1343 m), 02 – 04/XI/2007, K.P. Santos et al. col., 1♀ (UFMG 2-2, metatarsus v2-2. III– tibia d2-2, v2-2-2, metatarsus 2071); Santa Bárbara, RPPN Santuário do Caraça, Pico do Sol (–43.50548, –20.05885, 1127 m), 16/I/2010, L.N. Perillo col., 1♂ d2-2-2, v2-2-2. IV– tibia d2-2, v1-2-2, metatarsus d2-2, (UFMG 6558). São Paulo: Jundiaí, Parque Estadual da Serra do v2-2-2. Palp (Fig. 17A,B). Retrolateral tibial apophysis Japi (–46.98333, –23.28333, 1133 m), 21/IV/2008, J. Sobjack col., with ventral lobe displaced dorsally; dorsal lobe delim- 1♀ (UFMG 6449); São Luís do Paraitinga, Núcleo Santa Virgínia, Parque Estadual da Serra do Mar (–45.14583, –23.33583, 945 m), ited by membranous strip with simple, hook-shaped ba- 21/XI/2004, M. Uehara-Prado col., 1♂ (IBSP 143300), same local- sal process and large, blade-like apical process; median ity and collector, 18/II/2005 1♀ (IBSP 143316). process of dorsal lobe short, membranous. Tegulum with sclerotized, curved prolateral apophysis pointing towards Note. Males and females have been matched because embolus and small clear bump in its retrolateral edge. they have been collected repeatedly at the type locality. Median apophysis comprising single slender piece with Both sexes have also been recorded in São Luís do Pirait- blunt apex. Conductor large, rounded, membranous, aris- inga, São Paulo. ing prolaterally to median apophysis, positioned in appo-

150 ARTHROPOD SYSTEMATICS & PHYLOGENY — 74 (2) 2016

Fig. 17. Ianduba angeloi sp.n. A,B: Male holotype (UFMG 12683). A: Palp, ventral view. B: Palp, retrolateral view. C,D: Female (UFMG 2071). C: Epigynum, ventral view. D: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Dot line indicating DL. Scale bars = 0.1 mm. sition to embolus. Embolus long, retrolaterally inserted, light brown patch. Abdomen brownish gray. Total length without any processes. 7.42. Carapace 3.25 long, 2.31 wide, 1.30 high. Clypeus Female (UFMG 2071): Carapace, chelicerae dark brown. 0.31 high. Anterior eye row 0.71 wide, posterior eye Endites, labium, sternum light brown. Leg I light brown, row 0.92 wide. Eye diameters and interdistances: AME except for whitish yellow distal end of tibia and dark 0.13, ALE 0.13, PME 0.13, PLE 0.10, AME – AME 0.10, brown femur. Legs II – III light brown, except for dark AME – ALE 0.08, PME – PME 0.15, PME – PLE 0.18, brown femora. Leg IV light brown, except for dark ALE – PLE 0.08. Median ocular quadrangle length 0.41, brown femur and tibia; femur with retrolateral, median anterior width 0.36, posterior width 0.41. Chelicerae 1.27

151 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

long, with three promarginal teeth, two retromarginal Description. Male holotype: Carapace, chelicerae dark teeth and setal tufts on promargins. Sternum 1.65 long, brown. Endites, labium, sternum light brown. Leg I 1.45 wide. Abdomen 4.21 long, 2.65 wide. Leg meas- brownish yellow, except for whitish yellow distal end urements: I– femur 2.36, patella 1.14, tibia 2.39, meta- of tibia and dark brown femur, patella and proximal end tarsus 1.96, tarsus 1.27. II– 2.29, 1.09, 2.14, 1.73, 1.19. of tibia. Legs II – III brownish yellow, except for dark III– 1.96, 0.89, 1.70, 1.81, 1.02. IV– 2.64, 1.09, 2.61, brown femora. Leg IV light brown, except for dark brown 2.87, 1.32. Leg spination: I– tibia v2-2-2, metatarsus tibia with yellow proximal and distal extremities. Abdo- v2-2. II– tibia v2-2-2, metatarsus v2-2. III– tibia d2-2, men purplish gray, with dark brown abdominal scutum v2-2-2, metatarsus d2-2-2, v2-2-2. IV– tibia d2-2, v1-2- and gray, transverse markings on dorsum. Total length 2, metatarsus d2-2-2, v2-2-2. Genitalia (Figs. 17C,D). 6.34. Carapace 3.07 long, 2.21 wide, 0.97 high. Clypeus Epigynum heavily sclerotized, with wide median plate; 0.31 high. Anterior eye row 0.71 wide, posterior eye copulatory openings narrow, slit-shaped and posterior. row 0.86 wide. Eye diameters and interdistances: AME Vulva with large dorsal piece, wider than long, not cover- 0.15, ALE 0.15, PME 0.13, PLE 0.13, AME – AME 0.13, ing spermathecae, with two blunt projections directed an- AME – ALE 0.05, PME – PME 0.15, PME – PLE 0.20, teriorly; copulatory ducts S-shaped; spermathecae small, ALE – PLE 0.05. Median ocular quadrangle length 0.41, oval, with fertilization ducts directed medially. anterior width 0.38, posterior width 0.41. Chelicerae 1.19 long, with three promarginal teeth, two retromarginal Variation. Six males: total length 6.08 – 7.33; carapace teeth and setal tufts on promargins. Sternum 1.58 long, length 3.12 – 3.59; femur I 2.50 – 3.02. Three females: to- 1.30 wide. Abdomen 3.15 long, 1.65 wide. Leg meas- tal length 7.30 – 7.60, carapace length 3.12 – 3.25, femur urements: I– femur 2.47, patella 1.14, tibia 2.39, meta- I 2.36 – 2.47. The shape and size of the tegular projection tarsus 2.08, tarsus 1.32. II– 2.19, 1.09, 2.03, 1.83, 1.22. on the male palp is slightly variable, as is that of the dor- III– 2.01, 0.99, 1.70, 1.88, 0.92. IV– 2.72, 1.07, 2.55, sal piece projections on the female genitalia. 2.58, 1.25. Leg spination: I– tibia v2-2-2, metatarsus v2- 2. II– tibia v2-2-2, metatarsus v2-2. III– tibia d2-2, v2- Natural history. Specimens were collected in pitfall 2-2, metatarsus d2-2-2, v2-2-2. IV– tibia d2-2, v1-2-2, traps in semi-deciduous Atlantic forest in Ouro Branco metatarsus d2-2, v2-2-2. Palp (Fig. 18A,B). Retrolateral and Ouro Preto. All adult specimens were collected in the tibial apophysis with simple, hirsute, blunt ventral lobe, rainy season (from November to April). All records come and simple, blade-like dorsal lobe. Tegulum without pro- from areas with altitudes above 800 m above sea level. lateral apophysis, with modified, blade-like retrolateral area. Median apophysis two-pieced, with large prolateral Distribution. Central Minas Gerais to Jundiaí, São Pau- sector which folds itself around small, pointed retrolat- lo, Brazil (Fig. 22). eral sector. Conductor large, fan-shaped, membranous, arising prolaterally to median apophysis, positioned in 5.4.5. Ianduba benjori sp.n. apposition to embolus. Embolus short, retrolaterally inserted. Fig. 18 Female paratype (MNRJ 13691): Carapace, chelicerae dark brown. Endites, labium, sternum light brown. Leg I Type material. Male holotype from Morumbeca, Parque Es- light brown, except for whitish yellow distal end of tibia tadual do Desengano, Santa Maria Madalena, Rio de Janeiro, and dark brown femur, patella and proximal end of tibia. Brazil [–41.91861, –21.87722, 1243 m], A. Chagas et al. col., 13 – 17/V/2008 (MNRJ 13956). Two female paratypes, same col- Legs II – III light brown, except for dark brown femora. lecting data (MNRJ 13950, MNRJ 13961). Leg IV light brown, except for dark brown femur and tibia; femur with retrolateral, distal light brown patch. Derivatio nominis. The name is a patronym in honour Abdomen brownish gray. Total length 8.33. Carapace of Jorge Duílio Lima Meneses, better known as Jorge 3.51 long, 2.55 wide, 0.94 high. Clypeus 0.28 high. An- Ben Jor, a Brazilian samba singer born in Rio de Janeiro, terior eye row 0.79 wide, posterior eye row 1.04 wide. Brazil. Eye diameters and interdistances: AME 0.15, ALE 0.15, PME 0.13, PLE 0.15, AME – AME 0.10, AME – ALE Differential diagnosis. Males of I. benjori sp.n. share a 0.05, PME – PME 0.18, PME – PLE 0.15, ALE – PLE globose tegulum, without prolateral tegular projection, 0.03. Median ocular quadrangle length 0.46, anterior with I. varia and I. beaga sp.n. It can be diagnosed by the width 0.41, posterior width 0.46. Chelicerae 1.50 long, fan-shaped conductor, which is prolaterally inserted, by with three promarginal teeth, two retromarginal teeth and the prolateral sector of the median apophysis embracing setal tufts on promargins. Sternum 1.83 long, 1.47 wide. its retrolateral sector, by the retrolateral, apical blade-like Abdomen 4.82 long, 3.22 wide. Leg measurements: I– modification in the tegulum, by the hirsute ventral lobe of femur 2.61, patella 1.29, tibia 2.47, metatarsus 2.08, tar- the RTA, and by the simple, blade-like dorsal lobe of the sus 1.27. II– 2.42, 1.27, 2.21, 1.91, 1.25. III– 2.19, 1.12, RTA (Fig. 18A,B). Females are recognized by the wider 1.73, 2.03, 1.02. IV– 2.92, 1.30, 2.16, 3.29, 1.32. Leg than long epigynum with a narrow median plate and by spination: I– tibia v2-2-2, metatarsus v2-2. II– tibia v2- the rectangular, much wider than long dorsal piece of the 2-2, metatarsus v2-2. III– tibia d2-2, v2-2-2, metatarsus vulva (Fig. 18C,D). d2-2-2, v2-2-2. IV– tibia d2-2, v1-2-2, metatarsus d2-2-

152 ARTHROPOD SYSTEMATICS & PHYLOGENY — 74 (2) 2016

Fig. 18. Ianduba benjori sp.n. A,B: Male holotype (MNRJ 13956). A: Palp, ventral view. B: Palp, retrolateral view. C,D: Female paratype (MNRJ 13961). C: Epigynum, ventral view. D: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Scale bars = 0.1 mm.

2, v2-2-2. Genitalia (Fig. 18C,D). Epigynum with narrow Variation. Two females: total length 8.33 – 8.60, cara- bottle-shaped median plate between lip-shaped margins pace length 3.30 – 3.51, femur I 2.58 – 2.61. of atrial pockets. Vulva with short dorsal plate, which is much wider than long; copulatory ducts straight, parallel Natural history. Specimens were collected in montane to body axis; secondary spermathecae short, digitiform, Atlantic forest at altitudes of approximately 1100 m laterally directed, visible by transparency through dorsal above sea level. piece; primary spermathecae small, oval, with fertilization ducts directed mesally; two small apodemes located Distribution. Known from a single locality in northern in cuticle near spermathecae. Rio de Janeiro, Brazil (Fig. 22).

153 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

5.4.6. Ianduba beaga sp.n. v2-2-2. Palp (Figs. 19A,B, 20A). Retrolateral tibial apophysis with two lobes: two-branched ventral lobe, with Figs. 19, 20 blunt basal process and acuminate apical process; and Type material. Male holotype from Pico do Sol, Reserva Particu- narrow, digitiform dorsal lobe. Tegulum globose, without lar do Patrimônio Natural Serra do Caraça, Santa Bárbara, Minas any apophyses, with shallow incision near insertion of Gerais, Brazil (-43.50547, -20.05883, 1000 m) L.N. Perillo col., embolus. Median apophysis made of three pieces: short, 7/X/2010 (UFMG 6655). Paratypes: one male, same collecting pointed retrolateral sector; globular, unsclerotized apical data (MPEG 24571); one female, same locality and collector, 15/ IX/2010 (UFMG 6654); one female, Serra do Baú, Santa Bárbara, sector bearing spine-like projections, and large, laminar Minas Gerais, Brazil (-43.42887, -19.96764, 767 m), B.V.S. Pi- prolateral sector bearing long hair-like projections (short- menta & M.W.F. Faria col., 25/X – 4/XI/2008 (UFMG 8421). er near apex), with hematodocha-like membranous area Additional material examined. BRAZIL: Minas Gerais: Belo at base. Conductor large, rounded, membranous, arising Horizonte, Estação Ecológica da UFMG (-43.96667, -19.86667, 817 m), XI/2012, A.A.S. Monteiro & R.C. Digue-Loja col., 3♂ prolaterally to median apophysis, positioned in apposi- (UFMG 13237), same locality and collector, XI – XII/2012, 3♂ tion to embolus. Embolus short and retrolaterally insert- (UFMG 13238). ed, with process visible in expanded palp. Female paratype (UFMG 8421): Carapace, chelicerae Derivatio nominis. “Beagá” is a commonly used nick- dark brown. Endites, labium, sternum light brown. Leg I name for Belo Horizonte, one of the localities where the light brown, except for whitish yellow distal end of tibia species is known to occur and also the homeland of the and dark brown femur, patella and proximal end of tibia. first author. The name should be treated as a noun in ap- Legs II – III light brown, except for dark brown femora. position. Leg IV light brown, except for dark brown femur and tibia; femur with a retrolateral, median light brown patch. Differential diagnosis. Males are similar to those of Abdomen brownish gray, with gray transverse markings I. varia and I. benjori sp.n. by lacking a prolateral tegu- on dorsum. Total length 7.25. Carapace 3.50 long, 2.54 lar projection; they are easily distinguished by the lami- wide, 1.07 high. Clypeus 0.31 high. Anterior eye row 0.84 nar prolateral sector of the median apophysis (pMA), wide, posterior eye row 1.04 wide. Eye diameters and in- which bears many long hair-like projections, and by the terdistances: AME 0.18, ALE 0.18, PME 0.15, PLE 0.18, short, digitiform dorsal lobe of the RTA (Figs. 19A,B, AME – AME 0.13, AME – ALE 0.08, PME – PME 0.18, 20). Females are distinguished by the trapezoidal dorsal PME – PLE 0.23, ALE – PLE 0.08. Median ocular quad- piece of the epigynum, which is widest posteriorly and rangle length 0.43, anterior width 0.41, posterior width covers the spermathecae partially; by the laterally pro- 0.46. Chelicerae 1.35 long, with three promarginal teeth, jecting secondary spermathecae, visible laterally to the two retromarginal teeth and setal tufts on promargins. dorsal piece; and by the large and strongly sclerotized Sternum 1.70 long, 1.50 wide. Abdomen 3.86 long, 2.34 apodemes just anterior to the vulva (Fig. 19C,D). wide. Leg measurements: I– femur 2.70, patella 1.17, tibia 2.61, metatarsus 2.11, tarsus 1.47. II– 2.47, 1.12, Description. Male holotype: Carapace, chelicerae, en- 2.29, 1.98, 1.35. III– 2.16, 1.07, 1.83, 2.03, 1.12. IV– dites, labium dark brown. Sternum light brown. Leg I 2.98, 1.25, 2.98, 3.35, 1.47. Leg spination: I– tibia v2-2- dark brown, except for light brown metatarsus and yel- 2, metatarsus v2-2. II– tibia v2-2-2, metatarsus v2-2. III– lowish white tarsus and distal half of tibia. Legs II – III tibia d2-2, v2-2-2, metatarsus d2-2-2, v2-2-2. IV– tibia light brown, except for dark brown femora. Leg IV light d2-2, v1-2-2, metatarsus d2-2-2, v2-2-2. Genitalia (Fig. brown, except for dark brown femur, tibia and metatar- 19C,D). Epigynum very lightly sclerotized, with wide- sus; femur with retrolateral, median light brown patch. ly separated copulatory openings positioned medially. Abdomen dark brown, with gray, transverse markings Vulva with short, trapezoidal dorsal piece, which is wid- on posterior end of dorsum. Total length 6.61. Carapace est posteriorly, with an anterior notch; copulatory ducts 3.36 long, 2.52 wide, 1.35 high. Clypeus 0.36 high. An- thin; secondary spermathecae partially visible laterally to terior eye row 0.81 wide, posterior eye row 1.04 wide. dorsal piece in dorsal view; primary spermathecae small, Eye diameters and interdistances: AME 0.18, ALE 0.18, oval, partially covered by dorsal piece; fertilization ducts PME 0.15, PLE 0.15, AME – AME 0.13, AME – ALE directed medially, with apex bended anteriorly. With two 0.08, PME – PME 0.18, PME – PLE 0.20, ALE – PLE strong, sclerotized apodemes near apex of dorsal piece. 0.10. Median ocular quadrangle length 0.43, anterior width 0.41, posterior width 0.46. Chelicerae 1.68 long, Variation. Five males: total length 6.61 – 7.10; carapace with three promarginal teeth, two retromarginal teeth and length 3.36 – 3.72; femur I 2.75 – 2.99. Two females: to- setal tufts on promargins. Sternum 1.73 long, 1.53 wide. tal length 7.25 – 7.40, carapace length 3.50 – 3.47, femur Abdomen 3.12 long, 1.86 wide. Leg measurements: I– I 2.70 – 3.87. The palpal morphology is distinct between femur 2.75, patella 1.22, tibia 2.31, metatarsus 2.42, tar- males from Santa Bárbara and Belo Horizonte: the latter sus 1.68. II– 2.42, 1.12, 2.24, 2.08, 0.89. III– 2.19, 1.04, have a less slender palpus, and the shape of the retro- 1.73, 2.06, 1.17. IV– 2.95, 1.22, 2.42, 3.39, 1.53. Leg lateral tibial apophysis is slightly different (Fig. 20B,C). spination: I– tibia v2-2-2, metatarsus v2-2. II– tibia v2-2- We have not considered this rather small variation as 2, metatarsus v2-2. III– tibia d2-2, v2-2-2, metatarsus d2- sufficient to recognize these populations as distinct spe- 2-2, v2-2-2. IV– tibia d2-2, v1-2-2, metatarsus d2-2-2, cies, as closely related species of Ianduba have strikingly

154 ARTHROPOD SYSTEMATICS & PHYLOGENY — 74 (2) 2016

Fig. 19. Ianduba beaga sp.n. A,B: Male holotype (UFMG 6655). A: Palp, ventral view. B: Palp, retrolateral view. C,D: Female paratype (UFMG 8421). C: Epigynum, ventral view. D: Epigynum, dorsal view, cleared. See chapter 3 for abbreviations. Scale bars = 0.1 mm. different palpal morphologies. As more populations are Distribution. Known from three nearby localities in cen- discovered, it will become clearer whether this is a clinal tral Minas Gerais, Brazil (Fig. 22). variation or these populations are independent evolutionary lineages.

Natural history. Label data indicate that specimens were 6. New records collected in Möricke traps at altitudes of approximately 1000 m, and in pitfall traps at 767 – 834 m above sea level. All records are located in areas of secondary semi-de- During the course of this work, several new records of cidual Atlantic forest. Adults were collected in the rainy previously described species were found. They are listed season (October to November). below.

155 Magalhaes et al.: Taxonomy of Atlantic rainforest spiders

Fig. 20. Variation in palpal tibia morphology in Ianduba beaga sp.n. A: Male holotype (UFMG 6655), ventral view. B,C: Male from Belo Horizonte (UFMG 13237). B: Ventral view. C: Lateral view. See chapter 3 for abbreviations. Scale bars = 0.1 mm.

Ianduba abara Bonaldo & Brescovit, 2007 (Fig. 23). BRAZIL: 06476); Nova Iguaçu, Reserva Biológica do Tinguá (–43.4511, Bahia: Porto Seguro, Estação Vera Cruz Celulose [–39.17194, –22.7592, 69 m), II/2002, E.F.Ramos col., 1♀ (IBSP 118091), –16.39194, 63 m], 2006, J.P. Souza-Alves col., 1♂ (IBSP 70175), III/2002, E.F. Ramos et al. col., 1♀ (IBSP 119267); Barrelão 1♂ (IBSP 70177), 1♂ (IBSP 70179). [–43.4511, –22.7592, 69 m], 2002, E. Folly col., 1♂ (IBSP 125353), 1♂ (IBSP 125354); Rio de Janeiro, Parque Nacional da Tijuca Ianduba mugunza Bonaldo & Brescovit, 2007 (Fig. 23). BRA- [–43.1667, –22.9, 7 m], 2♀ 6 juveniles (MNRJ 6669), 1♂ 2♀ ZIL: Bahia: Ilhéus, CEPLAC [–39.0333, –14.8167, 8 m], 7/IV/ (MNRJ 6670), 4♂ (MNRJ 6671), 2♀ (MNRJ 6672), 2♂ 4 juveniles 1998, A.D. Brescovit col., 1♀ (IBSP 19429). (MNRJ 6673), Pedra Bonita, 1♂ 2 juveniles (MNRJ 6674), Ianduba patua Bonaldo, 1997 (Fig. 23). BRAZIL: Bahia: Santa Cochrano, 1♂ (MNRJ 6675); Santa Maria Madalena, Parque Es- Teresinha, Serra da Jibóia (–39.48111, –12.0375, 253 m), 24 – 27/ tadual do Desengano, Morumbéca [–41.91861, –21.87722, 1243 IV/2009, G.C. Ferreira col., 1♂ (IBSP 146296), 08/XI/2010, L.S. m], 1♀ (MNRJ 6771); Teresópolis, Parque Nacional da Serra dos Carvalho col., 1♂ (CHNUFPI 120). Órgãos [–42.9833, –22.4333, 1001 m], 1♀ (MNRJ 6677); Volta Ianduba paubrasil Bonaldo, 1997 (Fig. 22). BRAZIL: Bahia: Redonda, Floresta Cicuta, Trilha da Pedra do Sino [–44.1167, Prado, Cumuruxatiba (–39.18186, –17.10073), 1♂ (UFMG 17859). –22.5333, 399 m], 11/III/2002, E. Folly col., 1♂ (IBSP 39602). Espírito Santo: Conceição da Barra, Floresta Nacional do Rio Santa Catarina: Urussanga, Rio Molha, Gruta da Santa [–49.3167, Preto [–39.8493, –18.37159, 47 m], 19/X/2005, T. Souza et al. col., –28.5167, 90 m], 19 – 26/VII/2007, R.A. Teixeira col., 1♀ (IBSP 1♀ (IBSP 135186). 133875), 23/XI/2007, 1♂ (IBSP 132860). São Paulo: Amparo, Fa- zenda São Bento da Lapa [–46.7644, –22.7011, 745 m], 09/ Ianduba varia (Keyserling, 1891) (Figs. 1, 7, 8A – C, 9, 10A – H, VIII/2004, L.S. Uchoa Jr. col., 1♀ (IBSP 50876); Bertioga, Trilha 21). BRAZIL: Minas Gerais: Belo Horizonte, Estação Ecológica do rio Itatinga [–46.1386, –23.8544, 17 m], VII/2000, M.S.C. da UFMG (–43.96667, –19.86667, 817 m), 29/III/2000, E.S.S. Morini col. col., 1♀ (IBSP 62356); Biritiba Mirim, Barragem do Álvares & C.S.Azevedo col., 1♂ (IBSP 27211), 1♂ 1 juvenile Rio Biritiba [–45.8667, –23.6333, 905 m], V/2003, Equipe IBSP (IBSP 27213, IX/1999 – II/2001, 1♂ (IBSP 32260), VII/1999 – col., 1♂ 1♀ (IBSP 122418); Campos do Jordão, Gruta dos Crioulos II/2001, 1♂ 3♀ (IBSP 32332), 27/III/2000, 1♂ 1♀ (UFMG 3193), [–45.5833, –22.7333, 1656 m], X/2001, P. Gnaspini col., 1♂ (IBSP III/2001, E.S.S. Álvares col., 8♂ 7♀ (UFMG 3199), I – III/2001, 31893); Catanduva, Usina São Domingos [–48.9728, –21.1378, 8♂ 8♀ (UFMG 6055); Belo Horizonte, Parque Municipal das Man- 526 m], 03/X/2001, C. Rheims col., 1♂ (IBSP 31897); Cotia, Cau- gabeiras (–43.90532, –19.95413, 1035 m), 20/I/2005, H.H. Santos caia do Alto, Laboratório de Artrópodes [–46.9192, –23.6039, 792 col., 1♂ (UFMG 8495); Brumadinho, Condomínio da Aldeia do m], 02/V/2004, F.S. Cunha col., 1♀ (IBSP 43560), 25/II/1998, M. Rio das Pedras [–44.1997, –20.1433, 777 m], 01/I/1998, A.J. San- Magro col., 1♀ (IBSP 16268), IV/1973, V. Haddad col., 1♂ 1♀ tos col., 1♂ (IBSP 36231); Ouro Branco, Serra do Ouro Branco (IBSP 2630); Cubatão, Mata de Encosta da Copebrás (–46.39778, (–43.60139, –20.50056, 1095 m), 25/IV/2007, Y. Antonini col., 1♀ –23.835, 38 m), VI, IX, XII/2008, A.A. Nogueira col., 1♀ (IBSP (UFMG 12672), 26/III/2007, 1♂ (UFMG 12673), 9/V/2007, 1♂ 141872), 1♂ (IBSP 141873), 1♂ (IBSP 141874), 1♂ (IBSP (UFMG 12674), 22/X/2007, 1♂ (UFMG 12675), 21/X/2007, 1♀ 141875); Guarujá, Ilha da Moela [–46.2667, –24,6 m], 17 – 19/ (UFMG 12676), 25/III/2007, 1♂ (UFMG 12677), 24/X/2007, 1♀ VIII/2009, R.P. Indicatti & G.P. Perroni col., 1♀ (IBSP 145308), (UFMG 12678), 20/VII/2007, 1♀ (UFMG 12679); Rio Acima, 29 – 31/III/2009, R.P. Indicatti & F.U. Yamamoto col., 1♂ 1♀ Condomínio Cachoeiras do Tangará [–43.79806, –20.09639, 871 (IBSP 134479); Itapevi, Condomínio Trensurb, Praia do Tombo m], I/2005, T.S. Moreira col., 1♀ (UFMG 1073); Rio Preto [–46.9342, –23.5489, 785 m], 29/IX/2000, V. Onofrio & D.M. Bat- [–43.8275, –22.0864, 521 m], 1♀ 2 juveniles (MNRJ 06676). Par- testi col., 1♂ (IBSP 123526); Mogi das Cruzes, Parque das Nebli- aná: Fenix, Parque Estadual de Vila Rica do Espírito Santo [–51.95, nas (–46.16222, –23.74778, 777 m), 20/III/2006, M. Uehara-Prado –23.9, 328 m], 2001 – 2003, Equipe Embrapa col., 1♀ (IBSP col., 1♂ (IBSP 142159), 1♂ (IBSP 142161), 01/XII/2005, 1♀ 124262), 24 – 29/IV/2003, 2♂ (IBSP 124265); Londrina, Parque (IBSP 142162), 03/I/2006, 1♀ (IBSP 142163); Núcleo Santa Estadual Mata dos Godoys [–51.15, –23.3, 552 m], 13/IV/1999, J. Vírginia, Parque Estadual Serra do Mar [–45.1253, –23.3444, 863 Lopes col., 2♂ (IBSP 38241). Rio de Janeiro: Ilha Marambaia m], 13/IV/2005, M.V. Prado col., 2♂ (IBSP 58538), 14/III/2005, [–41.7833, –22.3833, 8 m], 27/IV/2001, E.F. Ramos col., 1♀ (IBSP 1♂ (IBSP 58541), 17/II/2005, 1♀ (IBSP 58549), 14/III/2005, 1♂ 28229), 28/IV/2001, 1♀ (IBSP 28400); Macaé, Parque Nacional (IBSP 58552); Peruíbe, Estação Ecológica Juréia-Itatins Restinga de Jurubatiba, Fazenda São Lázaros [–41.78291, (–47.01758, –24.38711, 146 m), 21 – 26/IV/2012, G.H.F. Azevedo –22.38257, 8 m], 27/III/2010, R.L.C. Baptista col., 1♂ 1♀ (MNRJ & J.P.P. Pena-Barbosa col., 1♀ (UFMG 13011); Salesópolis, Es-

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Fig. 21. Left: distribution of Ianduba species groups: varia (gray circles) and vatapa (black squares). Arrow indicates the mouth of the Doce River. Right: distribution of Ianduba varia (Keyserling, 1891).

Fig. 22. Left: distribution of Ianduba beaga sp.n. (crosses), I. liberta sp.n. (circle), I. benjori sp.n. (square), I. dabadu sp.n. (triangles) and I. capixaba sp.n. (stars). Right: distribution of I. paubrasil Bonaldo, 1997 (triangles), I. apururuca sp.n. (circles) and I. angeloi sp.n. (crosses).

Fig. 23. Left: distribution of Ianduba patua Bonaldo, 1997 (circles) and I. mugunza Bonaldo & Brescovit, 2007 (crosses). Right: distribution of I. acaraje sp.n. (square), I. vatapa Bonaldo, 1997 (cross), I. caxixe Bonaldo, 1997 (circles) and I. abara Bonaldo & Brescovit, 2007 (triangle).

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tação Biológica de Boracéia (–45.88889, –23.65278, 883 m), 27/ an anonymous reviewer for a thorough revision of the manuscript. III/2006, M. Uehara-Prado col., 1♀ (IBSP 142145), 1♂ (IBSP We thank the Willi Hennig Society for sponsoring TNT. ILFM has 142146), 12/I/2006, 1♀ (IBSP 142147), 27/III/2006, 1♂ (IBSP been supported by Conselho Nacional de Desenvolvimento Cientí- 142148), 08/XII/2005, 1♀ (IBSP 142150), 09/XII/2005, 1♀ (IBSP fico e Tecnológico (CNPq; 132086/2011-5 and 372279/2013-9) 142152), 20/II/2006, 1♀ (IBSP 142153), 28/IV/2006, 1♂ 1♀ (IBSP and Consejo Nacional de Investigaciones Científicas y Técnicas 142154), 25/V/2006, 1♂ (IBSP 142155); Santo André, Reserva Bi- (CONICET). ABB is supported by CNPq (PQ grant 304965/2012- ológica do Alto da Serra de Paranapiacaba (–46.30556, –23.76667, 0), and ILFM and MJR by CONICET (PIP 2012-00943) and FON- 806 m), 20/III/2007, 1♀ (IBSP 142628), 17/XI/2006, 1♀ (IBSP CyT (PICT 2011-1007). 142630), 18/XII/2006, 1♂ (IBSP 142636); São Bernardo do Cam- po, Parque Estoril [–46.55, –23.7, 771 m], 12 – 19/III/2008, C.V. Janini et al. col., 1♂ (IBSP 119064), 05 – 10/IV/2006, B.Távora col. col., 1♂ (IBSP 72250); São Luiz do Paraitinga, Parque Estadual da 7. References Serra do Mar (–45.14583, –23.33583, 945 m), 20/XI/2004, M. Uehara-Prado col., 1♀ (IBSP 143315); São Paulo, Campus da USP [–46.71667, –23.55, 745 m], 28/V/2007, E.O. Machado col., 1♂ Álvarez-Padilla F., Hormiga G. 2008. A protocol for digesting in- (IBSP 116743) 2000, R.Willemart col., 1♀ (IBSP 27270), V/2001, ternal soft tissues and mounting spiders for scanning electron Alunos USP col., 1♂ (IBSP 41740), Mata do Cuaso, 1♂ 1♀ (IBSP microscopy. – Journal of Arachnology 35: 538 – 542. 41742), 2♂ (IBSP 41746), 1♀ (IBSP 41796), 16 – 23/IV/1999, D.F. Bonaldo A.B. 1997. On the new Neotropical spider genus Ianduba Candiani col., 1♀ (IBSP 42217), 16 – 23/XI/1999, 1♀ (IBSP (Araneae, Corinnidae). – Iheringia, Série Zoologia 83: 165 – 108. 42226), 1♀ (IBSP 42227), V/2000 – II/2001, 9♂ 14♀ (IBSP Bonaldo A.B. 2000. Taxonomia da subfamília Corinninae (Arane- 76254); São Paulo, Campus IBSP, Mata da Cidade Universitária ae, Corinnidae) nas regiões neotropical e neártica. – Iheringia, Armando de Salles Oliveira [–46.71667, –23.55, 745 m], 22/ Série Zoologia 89: 3 – 148. II/1999, A.D. Brescovit col., 1♂ (IBSP 21448), 06/XI/2000, R. Bonaldo A.B., Brescovit A.D. 1998. On Ecitocobius, a new genus Martins col., 1♂ (IBSP 27351); São Paulo, Horto Florestal Oswal- from central Amazonia with comments on the tribe Attacobiini do Cruz, [–46.6333, –23.6, 781 m], 16 – 23/II/2000, D.F. Candiani (Arachnida, Araneae, Corinnidae, Corinninae). – Spixiana 21: col., 1♂ (IBSP 42210); São Paulo, Ilha Parque dos Eucaliptos, 165 – 172. 06 – 12/XI/1999, R.P. Indicatti col., 1♀ (IBSP 131676), 09 – 15/ Bonaldo A.B., Brescovit A.D. 2005. On new species of the IX/1999, 1♂ (IBSP 131677); São Paulo, Jardim Angela, 1♀ (IBSP Neotropical genus Attacobius Mello-Leitão, 1923 (Araneae, 131678), 1♂ (IBSP 131679), 06 – 12/XI/1999, 1♀ (IBSP 131680); Corinnidae, Corinninae), with a cladistic analysis of the tribe São Paulo, Parque Alfrodo Volpi, 19 – 23/III/2005, A. Bagio col., Attacobiini. – Insect Systematics & Evolution 36: 35 – 56. 1♀ (IBSP 59303); São Paulo, Parque Burle Marx, 18 – 25/VII/2005, Bonaldo A.B., Brescovit A.D., Rocha P.L.B., Costa M.G.C. 1♀ (IBSP 59307), 1♀ (IBSP 59310); São Paulo, Parque do Estado, 2007. Two new species of ant-mimicking spiders of the genus 28/II – 06/III/2002, J.R. Valvassori col., 1♀ (IBSP 59202); São Ianduba Bonaldo (Araneae, Corinnidae) from southern Bahia, Paulo, Parque do Ibirapuera, 06/IX/2007, R. Pando col., 1♀ (IBSP Brazil. – Revista Brasileira de Zoologia 24: 817 – 824. 123974); São Paulo, Parque Estadual da Cantareira, Núcleo Águas Brescovit A.D., Bonaldo A.B., Santos A.J., Ott R., Rheims C.A. Claras, 14 – 19/XI/2004, F.U. Yamamoto col., 1♀ (IBSP 66363), 2012. The Brazilian goblin spiders of the new genus Preda- 02 – 07/II/2005, 1♂ (IBSP 66364), 14 – 19/XI/2004, 1♂ (IBSP toroonops (Araneae: Oonopidae). – Bulletin of the American 66366); São Paulo, Reservatório Guarapiranga, 06 – 12/XI/1999, Museum of Natural History 370: 1 – 68. R.P. Indicatti et al. col., 1♂ (IBSP 132304), 07 – 13/IV/2005, I. Carnaval A.C., Moritz C. 2008. Historical climate modeling pre- Cizauskas & C.R.M Garcia col., 1♀ (IBSP 61558), 1♂ (IBSP dicts patterns of current biodiversity in the Brazilian Atlantic 61561), 07 – 13/X/2004, 1♀ (IBSP 61566); São Paulo, 07/VI/1999, forest. – Journal of Biogeography 35: 1182 – 1201. P.E.S. Cantagalo col., 1♂ (IBSP 23830); 25/VII/2006, P.C.E. Berti Carnaval A.C., Waltari E., Rodrigues M.T., Rosaeur D., Van- col., 1♂ (IBSP 75823); São Paulo, Bairro Butantã, VI/2001, H.F. DerWal J., Damasceno R., Prates I., Strangas M., Spanos Japyassú et al. col., 1♂ (IBSP 28919), Campus da USP, Jardim Z., Rivera D., Pie M.R., Firkowski C.R., Bornschein M.R., Rizzo, 23/II/2000, F.S. Cunha col., 1♂ (IBSP 32930); São Roque, Ribeiro L.F., Moritz C. 2014. Prediction of phylogeographic Reserva Florestal Cuaso [–47.7272, –23.135, 544 m], 29/IV/1996, endemism in an environmentally complex biome. – Proceed- J.C.C. Korte col., 1♂ (IBSP 10989); São Sebastião, Ilha de Alc- ings of the Royal Society of London B 281: 1 – 8. atrazes [–45.6942, –24.10179, 1 m], XII/1996, R. Bertani col., 1♀ Goloboff P.A., Farris J.S., Nixon K.C. 2008. TNT, a free program (IBSP 7893); Sorocaba, Mata de Eucalipto da Mata da UNIP for phylogenetic analysis. – Cladistics 24: 774 – 786. [–47.4581, –23.5017, 598 m], 14/VI/2008, I. Cizauskas et al. col., Hijmans R.J., Cameron S.E., Parra J.L., Jones P.G., Jarvis A. 1♂ (IBSP 99042), 1♀ (IBSP 99044); Ubatuba, Parque Estadual da 2005. Very high resolution interpolated climate surfaces for Ilha Anchieta (–45.05, –23.53333, 51 m), 23 – 30/VII/2001, Equipe global land areas. – International Journal of Climatology 25: Biota col., 1♀ (IBSP 56425), 1♀ (IBSP 56428), 1♀ (IBSP 56429), 1965 – 1978. 1♂ (IBSP 56430), 1♂ (IBSP 56433), 1♀ (IBSP 56434). Huber B.A. 2015. Small scale endemism in Brazil’s Atlantic For- est: 14 new species of Mesabolivar (Araneae, Pholcidae), each known from a single locality. – Zootaxa 3942: 1 – 60. Jacobi C.M., Carmo F.F., Vincent R.C., Stehmann J.R. 2007. Plant 7. Acknowledgments communities in ironstone outcrops: a diverse and endangered Brazilian ecosystem. – Biodiversity and Conservation 16: 2185 – 2200. We are grateful to all curators and institutions which have made Levi H.W. 1988. The Neotropical orb-weaving spiders of the genus material available for this study. B.T. Faleiro and G.H.F. Azeve- Alpaida (Araneae: Araneidae). – Bulletin of the Museum of do sorted specimens from the MNRJ collection, and J.L. Chavari Comparative Zoology 151: 365 – 487. helped with locality data from IBSP specimens. We thank L.N. Machado A.B.M., Drummond G.M., Paglia A.P. 2008. Livro Ver- Perillo, M.T.T. Santos, P.H. Martins and L.S. Carvalho for collect- melho da Fauna Brasileira Ameaçada de Extinção. – Ministério ing some of the specimens studied. Fabián Tricárico kindly helped do Meio Ambiente, Brasília, 1420 pp. acquiring images at the MACN SEM laboratory. A. Anker and J.P. Ministério do Meio Ambiente 2014. Listas das Espécies da Fauna Burini authorized publication of their excellent pictures. We are in- Brasileira Ameaçadas de Extinção. – Available from: http:// debted to A.J. Santos and U. Oliveira for comments on earlier ver- www.icmbio.gov.br/portal/biodiversidade/fauna-brasileira/ sions of this text, and to K.-D. Klass, L. Prendini, J. Bosselaers and lista-de-especies.html. Last accessed January 16, 2016.

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Myers N., Mittermeier R.A., Mittermeier C.G., Fonseca G.A., Rheims C.A. 2010. Caayaguara, a new genus of huntsman spiders Kent J. 2000. Biodiversity hotspots for conservation priori- from the Brazilian Atlantic Forest (Araneae: Sparassidae). – ties. – Nature 403: 853 – 858. Zootaxa 2630: 1 – 29. Oliveira U. 2011. Diversidade e biogeografia de aranhas do Brasil: Ribeiro M.C., Metzger J.P., Martensen A.C., Ponzoni F.J., Hirota esforço amostral, riqueza potencial e áreas de endemismo. – M.M. 2009. The Brazilian Atlantic Forest: how much is left, Unpublished Master’s Thesis, Universidade Federal de Minas and how is the remaining forest distributed? Implications for Gerais, Belo Horizonte, 103 pp. conservation. – Biological Conservation 142: 1141 – 1153. Oliveira U., Brescovit A.D., Santos A.J. 2015. Delimiting areas Rodrigues B.V.B., Bonaldo A.B. 2014. Taxonomic revision of of endemism through Kernel interpolation. – PLoS ONE 10: the species group rubripes of Corinna Koch, 1842 (Araneae; e0116673. Corinnidae). – Zootaxa 3815: 451 – 493. Phillips S.J., Anderson R.P., Schapire R.E. 2006. Maximum entro- Safford H.D. 2007. Brazilian Páramos IV. Phytogeography of the py modeling of species geographic distributions. – Ecological campos de altitude. – Journal of Biogeography 34: 1701 – 1722. Modelling 190: 231 – 259. Silva S.M., Moraes-Barros N., Ribas C.C., Ferrand N., Mor- Pinto-da-Rocha R., Silva M.B. 2005. Faunistic similarity and his- gante J.S. 2012. Divide to conquer: a complex pattern of bio- toric biogeography of the harvestmen of southern and south- diversity depicted by vertebrate components in the Brazilian eastern Atlantic rain forest of Brazil. – Journal of Arachnology Atlantic Forest. – Biological Journal of the Linnean Society 33: 290 – 299. 107: 39 – 55. Pinto-da-Rocha R., Bragagnolo C., Tourinho A.L. 2014. Three Sorte F.A.L., Jetz W. 2010. Projected range contractions of new species of Thrasychiroides Soares & Soares, 1947 from montane biodiversity under global warming. – Proceedings Brazilian mountains (Opiliones, Eupnoi, Neopilionidae). – of the Royal Society of London B, Biological Sciences 277: Zootaxa 3869: 469 – 482. 3401 – 3410. Platnick N.I., Shadab M.U. 1975. A revision of the spider genus Vasconcelos M.F. 2008. Mountaintop endemism in eastern Brazil: Gnaphosa (Araneae, Gnaphosidae) in America. – Bulletin of why some bird species from campos rupestres of the Espinhaço the American Museum of Natural History 155: 1 – 66. Range are not endemic to the Cerrado region? – Revista Bra- Polotow D., Brescovit A.D. 2006. Revision of the Neotropical sileira de Ornitologia 16: 348 – 362. spider genus Itatiaya Mello-Leitão (Araneae: Ctenidae) with Zomer R.J., Trabucco A., Bossio D.A., van Straaten O., Verchot considerations on biogeographic distribution of species. – Re- L.V. 2008. Climate change mitigation: a spatial analysis of vista Brasileira de Zoologia 23: 429 – 442. global land suitability for clean development mechanism af- Ramírez M.J. 2014. The morphology and phylogeny of dionychan forestation and reforestation. – Agriculture, Ecosystems and spiders (Araneae: Araneomorphae). – Bulletin of the American Environment 126: 67 – 80. Museum of Natural History 390: 1 – 374.

Electronic Supplement File at http://www.senckenberg.de/arthropod-systematics (“Contents”)

File 1: magalhaes&al-iandubaspiders-asp2016-electronicsupple ment.rar. – Zipped file containing S1A) character matrix (.tnt format), S1B) tree files obtained under different weighting strengths (.tre format), S1C) list of synapomorphies for all clades and accom- panying graphic key (.txt and .wmf formats), and S1D) graphs with Bremer supports and jackknifing frequencies for clades obtained with concavity constant = 9 (.wmf format).

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