Phylogeny of Sicariidae Spiders (Araneae: Haplogynae), with a Monograph on Neotropical Sicarius

Zoological Journal of the Linnean Linnean Society Society,, 2017,2016. 179 With, 767–864. 63 ﬁgures With 63 figures

Phylogeny of Sicariidae spiders (Araneae: Haplogynae), with a monograph on Neotropical Sicarius

IVAN L. F. MAGALHAES1,2*, ANTONIO D. BRESCOVIT3 and ADALBERTO J. SANTOS2

1Division� Aracnolog�ıa, Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Av. Angel� Gallardo 470, C1405DJR, Buenos Aires, Argentina 2Departamento de Zoologia, Instituto de Ciencias^ Biologicas,� Universidade Federal de Minas Gerais, Av. Antonio^ Carlos, 6627, Belo Horizonte, Minas Gerais, CEP 31270-901, Brazil 767 3Laboratorio� Especial de Coleçoes~ Zoologicas,� Instituto Butantan, Av. Vital Brazil, 1500, Sao~ Paulo, SP, CEP 05503-900, Brazil

Received 2 September 2015; revised 29 March 2016; accepted for publication 30 March 2016

Sicariids are an infamous spider family containing two genera: the poorly known Sicarius Walckenaer and the medically important Loxosceles Heineken & Lowe. We present the first broad survey of the morphology of the family from a phylogenetic perspective in order to resolve its relationships. We scored morphological, behavioural and venom feature data for 38 taxa, including New and Old World species of both Sicarius and Loxosceles and three outgroups. Our results point to the monophyly of Sicariidae and its two genera as currently delimited, with the identification of novel synapomorphies for all of them. We present evidence of a group of ‘true’ Loxosceles composed of all members of the genus except those from the spinulosa species group. Sicarius have a very interesting phylogenetic structure, with species from the Americas and Africa forming reciprocally monophyletic groups. Thus, we resurrect Hexophthalma Karsch to accommodate African Sicariinae species. We discuss the evolution of venom proteins, spinning organs and cryptic appearance in Sicariidae. We delimit Sicarius to include only the 20 Neotropical species, and present a taxonomic revision of the genus. We redescribe nine previously known species, elevate one subspecies to species, and describe six new species from Argentina, Bolivia, Brazil and Peru. Sicarius levii sp. nov., from Argentina, is probably the largest known haplogyne spider in both body length and mass. Six specific names are considered junior synonyms of older names. All these species are illustrated and have their distributions mapped. We present an interactive key for identifying Sicarius species.

ADDITIONAL KEYWORDS: crypsis – evolution – morphology – new species – revision – Scytodoidea – Sphingomyelinase D – systematics – taxonomy – venom.

which are moderately diverse (~130 species) and INTRODUCTION more diversiﬁed in Africa and the Americas (World Sicariidae Keyserling stands as one of the most infa- Spider Catalog, 2015). mous spider families, as it contains the medically Sicariids are currently allocated in Scytodoidea, an important genus Loxosceles Heineken & Lowe – the ancient group of true spiders with simple genitalia brown, recluse or violin spiders. The limits of the (Garrison et al., 2016). The monophyly of Sicariidae family have changed since the classical work of has been supported by morphological data (Platnick Simon (1893), and now it is thought to contain only et al., 1991; Ramırez, 2000; Labarque & Ramırez, two genera, the other one being Sicarius Walcke- 2012), although in these analyses only a few exem- naer, known as six-eyed sand spiders. They are hap- plars have been included. The only comprehensive logyne, medium- to large-sized, sedentary spiders, phylogenetic hypothesis for the family is that of Bin- ford et al. (2008), who presented DNA sequence data of a broad sampling of species of both Sicarius and *Corresponding author. E-mail: [email protected] Loxosceles. Their analysis recovered the monophyly

© 20172016 The Linnean Society Society of of London, London, ZoologicalZoological Journal Journal of of the the Linnean Linnean Society Society, 2017,, 2016 179, 767–864 7671 7682 I. L. I. F.L. MAGALHAESF. MAGALHAESET ET AL. AL. SICARIID PHYLOGENY AND REVISION OF SICARIUS 3 of Sicarius and Loxosceles, but showed contentious particles (Duncan, Autumn & Binford, 2007), confer- hexamethyldisilazane (HMDS) or critical-point dried reconnection, retaining up to 1000 trees per replica- support for a monophyletic Sicariidae. As discussed ring a characteristic cryptic appearance to these spi- (fragile parts, such as spinnerets and female genital- tion. Branch supports were estimated by Bremer by them, this result may in part have been caused ders. Additionally, their unique pot-shaped eggcases ia). Dried parts were placed in aluminium stubs (1994) support (using a script that comes with the by the ancient age of the group – Sicariids are are also covered by dirt (Levi & Levi, 1969; Aguilar using adhesive copper tape or carbon stickers, sput- TNT package) and symmetric resampling (Goloboff thought to have started diversifying before the &Mendez,� 1971; Magalhaes~ et al., 2013). Their ter-coated with 10 nm of gold or gold–palladium, and et al., 2003) (estimated based on 1000 pseudorepli- break-up of western Gondwana. Furthermore, Bin- venom, like that of Loxosceles, contains sphin- examined under JEOL JSM-6360L, Quanta 2000 or cates, with a 33% probability of change). Character ford et al. (2008) found evidence that African and gomyelinase D, an enzyme responsible for necrotic Philips FEI XL30 TMP scanning electron micro- optimizations were made in Winclada 1.0.8 (Nixon, Neotropical Sicarius are reciprocally monophyletic. activity. However, the potential activity of the scopes. 1999–2002) and Mesquite 2.75. However, to date no morphological work has been enzyme varies among Sicarius species (Binford et al., Many of the characters used in the analysis have Bayesian inference was carried out in MrBayes 3.2 done to test this hypothesis, or to search for potential 2009; Lopes et al., 2013), and at least some species been taken from the literature (e.g. Platnick et al., (Ronquist et al., 2012) using the Mkv model of evolu- synapomorphies of these clades. This is especially are apparently incapable of causing strong symptoms 1991; Ram�ırez, 2000, 2014; Labarque & Ram�ırez, tion (Lewis, 2001) accounting for rate heterogeneity interesting considering that the first African Sicari- in vertebrates (e.g. Alegre, Meneses & Aguilar, 2012). Morphological scorings were made de novo for among characters (Γ parameter). Two independent inae were described in a separate genus, Hexoph- 1977). They are restricted to xeric habitats, espe- all taxa based on examination of actual specimens, runs of four Markov chains each (one cold and three thalma Karsch, 1879, which was considered a junior cially deserts and tropical dry forests (Magalhaes~ except for L. rufescens (Dufour), scored after the heated) were run for 107 generations, sampling synonym of Sicarius by Simon (1893) and subsequent et al., 2013), and are apparently poor dispersers richly illustrated work of Labarque & Ram�ırez parameters every 103 generations, and discarding authors. Inferring the phylogeny of Sicariidae based (Magalhaes et al., 2014), making them good models (2012); this work was also used to guide the scoring 25% of the initial sampled states as burn-in. Conver- on phenotypic data is desirable for several reasons. for studying the biogeography of these biomes. How- of some cells for the non-sicariid outgroups. Beha- gence of the independent runs and stationarity of First, this would be an independent test of the ever, the lack of taxonomic resolution for the vioural scorings are of our own field observations, the parameters were checked using Tracer 1.6 (Ram- topologies obtained by Binford et al. (2008), and Neotropical species hampers studies about those except when noted. Characters related to venom fea- baut et al., 2014). Branch supports were estimated might improve resolution for some of their clades. interesting aspects of their biology. tures are scored after the findings of Binford et al. as the frequencies of clades in a majority-rule con- Second, morphological phylogenies often reveal Our study has two objectives. We present the first (2009) and Lopes et al. (2013). sensus of the trees sampled in the stationary phase synapomorphies that help in placing taxa for which phylogeny of Sicariidae based on phenotypic data, of the Markov chains. For this analysis, only discrete no molecular data are available, such as fossils; this including morphology, behaviour and venom fea- characters have been used due to limitations of the PHYLOGENETIC ANALYSIS is especially relevant in the context of estimates of tures. This was based on the examination of species Mkv model. divergence times using fossil calibrations. Finally, spanning most of the geographical and morphological Our taxon sampling includes all Neotropical Sicar- phenotypic phylogenies generate a wealth of data on diversity of both Loxosceles and Sicarius, but focus- ius, six African Sicariinae, nine Loxosceles chosen to TAXONOMY morphology as a by-product (see Labarque & ing particularly in the latter genus. The second goal represent the morphological and geographical diver- Ram�ırez, 2012; Ram�ırez, 2014), which might reveal is to present a complete revision of Sicarius from the sity of the genus, and three outgroup taxa in the The format of description follows Magalhaes~ et al. interesting phylogenetic or functional patterns. Neotropical region, redescribing known species, families Drymusidae, Scytodidae and Pholcidae (2013). At least one female per vial was dissected for Due to the medical importance of Loxosceles, this describing new taxa and presenting a survey of the (Appendix 2). We assembled the discrete character examination of genitalia, even when males were pre- genus has been given a good deal of taxonomic atten- morphological diversity of the genus. matrix using Mesquite 2.75 (Maddison & Maddison, sent in the same vial. Males and females were easily tion, with good taxonomic revisions available for both 2013) and exported it as tnt (for parsimony analysis) associated because they are frequently collected New World and African species (e.g. Gertsch, 1967; or nex for MrBayes (for Bayesian inference). Contin- together. Measurements are given in millimetres, Gertsch & Ennik, 1983; Lotz, 2012; see also refer- MATERIAL AND METHODS uous characters were inserted directly into the .tnt and taken preferentially from the left side of speci- ences therein). On the other hand, systematic studies file using a text editor. Intervals for continuous char- mens. Variation of counts/measurements is given as MORPHOLOGY AND CHARACTER SCORINGS on Sicarius have been long neglected. Most Sicarius acters consist of the average value of each measure- minimum–maximum (mean). Drawings were made species have been known only from their original We have examined around 1800 adult specimens of ment for each species two standard errors, as using a Leica M205C stereoscopic microscope (and often incomplete or inadequate) descriptions Sicarius during the course of this work. For separat- suggested by Goloboff, MattoniÆ & Quinteros (2006). equipped with a camera lucida, or based on pho- (e.g. Keyserling, 1880; Mello-Leitao,~ 1936). Until ing the species, the prolateral view of the male palp Parsimony analyses were carried out in TNT 1.1 tographs taken in the same microscope using a very recently, the few restricted revisions and spe- and the ventral view of the female spermathecae pro- (Goloboff, Farris & Nixon, 2008). We used three DFC295 digital camera, and mounted as a single, cies redescriptions available (e.g. Mello-Leitao,~ 1940, vide the most informative characters. Somatic char- approaches: (1) analysis of discrete characters only multi-focal image using Leica Application Suite 3.8. 1941; Gerschman de Pikelin & Schiapelli, 1979) were acters are also of great taxonomic value, and can be under equal weights, (2) analysis of discrete charac- Some female spermathecae were photographed using not based on type-material examination, leading to used to diagnose most groups of closely related spe- ters only under implied weights and (3) analysis of an AxioCam ERc5s digital camera attached to a frequent species misidentifications. Overall, the tax- cies. Exposing the ventral face of the spermathecae discrete and continuous characters under implied Zeiss PrimoStar compound microscope following clar- onomy of Sicarius was chaotic and its species were is somewhat tricky; it is most easily done by digest- weights. We refrained from further analysing a data- ification using clove oil. Geographical coordinates of unrecognizable. This started to change with revisions ing the soft tissues with enzymes, and then cutting set including continuous characters under equal specimens collected by us, or museum specimens of species from Africa (Lotz, 2012) and north-eastern the ventral cuticle of the abdomen, or bending the weights because preliminary runs showed that with coordinates indicated in the original labels, are Brazil (Magalhaes,~ Brescovit & Santos, 2013). How- spermathecae. Generally, we have examined female meristic characters alone (namely the number of given in parentheses. Specimens without coordinates ever, the genus also occurs in arid environments genitalia after digestion using a pancreatin solution spermathecae branches and femoral thorns) heavily were georeferenced using Google Earth and have from Argentina to Central America, from where 14 (see Alvarez-Padilla� & Hormiga, 2008). This was also influenced the analysis, leading to highly deviant their coordinates indicated in square brackets. An species (one of them with two subspecies) have been used for preparing genitalia, and sometimes spin- results. For exploring the sensitivity of the results to interactive key for identifying all Sicarius species named, most of which have never been revised. nerets, for scanning electron microscopy (SEM). the value of k in the implied weight analyses, we was made using the Xper² platform (Ung et al., 2010) Sicarius are unique spiders for several reasons. Parts were dried for SEM using heat from a light used the TNT script by Mirande (2009) to find the and is available as Supporting Information They do not spin webs, but rather stalk their prey bulb (tough parts, such as palps) or by transfer most fit trees under a wide range of k values. Appendix S1. while buried in dry sediment (Reiskind, 1966). Their through a series of solutions of increasing concentra- Searches for optimal trees consisted of 1000 random The examined specimens are deposited in the cuticles have special setae which adhere to soil tion of ethanol, and then either transferred to addition sequences followed by tree bisection and following institutions (curators indicated in

© 2017 The Linnean© 2016 TheSociety Linnean of London, Society Zoological of London, JournalZoological of the JournalLinnean of Society the Linnean, 2017, 179 Society, 767–864, 2016 © 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016 2 I. L. F. MAGALHAES ET AL. SICARIIDSICARIID PHYLOGENY PHYLOGENY AND AND REVISION REVISION OF OFSICARIUSSICARIUS 7693 of Sicarius and Loxosceles, but showed contentious particles (Duncan, Autumn & Binford, 2007), confer- hexamethyldisilazane (HMDS) or critical-point dried reconnection, retaining up to 1000 trees per replica- support for a monophyletic Sicariidae. As discussed ring a characteristic cryptic appearance to these spi- (fragile parts, such as spinnerets and female genital- tion. Branch supports were estimated by Bremer by them, this result may in part have been caused ders. Additionally, their unique pot-shaped eggcases ia). Dried parts were placed in aluminium stubs (1994) support (using a script that comes with the by the ancient age of the group – Sicariids are are also covered by dirt (Levi & Levi, 1969; Aguilar using adhesive copper tape or carbon stickers, sput- TNT package) and symmetric resampling (Goloboff thought to have started diversifying before the &Mendez,� 1971; Magalhaes~ et al., 2013). Their ter-coated with 10 nm of gold or gold–palladium, and et al., 2003) (estimated based on 1000 pseudorepli- break-up of western Gondwana. Furthermore, Bin- venom, like that of Loxosceles, contains sphin- examined under JEOL JSM-6360L, Quanta 2000 or cates, with a 33% probability of change). Character ford et al. (2008) found evidence that African and gomyelinase D, an enzyme responsible for necrotic Philips FEI XL30 TMP scanning electron micro- optimizations were made in Winclada 1.0.8 (Nixon, Neotropical Sicarius are reciprocally monophyletic. activity. However, the potential activity of the scopes. 1999–2002) and Mesquite 2.75. However, to date no morphological work has been enzyme varies among Sicarius species (Binford et al., Many of the characters used in the analysis have Bayesian inference was carried out in MrBayes 3.2 done to test this hypothesis, or to search for potential 2009; Lopes et al., 2013), and at least some species been taken from the literature (e.g. Platnick et al., (Ronquist et al., 2012) using the Mkv model of evolu- synapomorphies of these clades. This is especially are apparently incapable of causing strong symptoms 1991; Ram�ırez, 2000, 2014; Labarque & Ram�ırez, tion (Lewis, 2001) accounting for rate heterogeneity interesting considering that the first African Sicari- in vertebrates (e.g. Alegre, Meneses & Aguilar, 2012). Morphological scorings were made de novo for among characters (Γ parameter). Two independent inae were described in a separate genus, Hexoph- 1977). They are restricted to xeric habitats, espe- all taxa based on examination of actual specimens, runs of four Markov chains each (one cold and three thalma Karsch, 1879, which was considered a junior cially deserts and tropical dry forests (Magalhaes~ except for L. rufescens (Dufour), scored after the heated) were run for 107 generations, sampling synonym of Sicarius by Simon (1893) and subsequent et al., 2013), and are apparently poor dispersers richly illustrated work of Labarque & Ram�ırez parameters every 103 generations, and discarding authors. Inferring the phylogeny of Sicariidae based (Magalhaes et al., 2014), making them good models (2012); this work was also used to guide the scoring 25% of the initial sampled states as burn-in. Conver- on phenotypic data is desirable for several reasons. for studying the biogeography of these biomes. How- of some cells for the non-sicariid outgroups. Beha- gence of the independent runs and stationarity of First, this would be an independent test of the ever, the lack of taxonomic resolution for the vioural scorings are of our own field observations, the parameters were checked using Tracer 1.6 (Ram- topologies obtained by Binford et al. (2008), and Neotropical species hampers studies about those except when noted. Characters related to venom fea- baut et al., 2014). Branch supports were estimated might improve resolution for some of their clades. interesting aspects of their biology. tures are scored after the findings of Binford et al. as the frequencies of clades in a majority-rule con- Second, morphological phylogenies often reveal Our study has two objectives. We present the first (2009) and Lopes et al. (2013). sensus of the trees sampled in the stationary phase synapomorphies that help in placing taxa for which phylogeny of Sicariidae based on phenotypic data, of the Markov chains. For this analysis, only discrete no molecular data are available, such as fossils; this including morphology, behaviour and venom fea- characters have been used due to limitations of the PHYLOGENETIC ANALYSIS is especially relevant in the context of estimates of tures. This was based on the examination of species Mkv model. divergence times using fossil calibrations. Finally, spanning most of the geographical and morphological Our taxon sampling includes all Neotropical Sicar- phenotypic phylogenies generate a wealth of data on diversity of both Loxosceles and Sicarius, but focus- ius, six African Sicariinae, nine Loxosceles chosen to TAXONOMY morphology as a by-product (see Labarque & ing particularly in the latter genus. The second goal represent the morphological and geographical diver- Ram�ırez, 2012; Ram�ırez, 2014), which might reveal is to present a complete revision of Sicarius from the sity of the genus, and three outgroup taxa in the The format of description follows Magalhaes~ et al. interesting phylogenetic or functional patterns. Neotropical region, redescribing known species, families Drymusidae, Scytodidae and Pholcidae (2013). At least one female per vial was dissected for Due to the medical importance of Loxosceles, this describing new taxa and presenting a survey of the (Appendix 2). We assembled the discrete character examination of genitalia, even when males were pre- genus has been given a good deal of taxonomic atten- morphological diversity of the genus. matrix using Mesquite 2.75 (Maddison & Maddison, sent in the same vial. Males and females were easily tion, with good taxonomic revisions available for both 2013) and exported it as tnt (for parsimony analysis) associated because they are frequently collected New World and African species (e.g. Gertsch, 1967; or nex for MrBayes (for Bayesian inference). Contin- together. Measurements are given in millimetres, Gertsch & Ennik, 1983; Lotz, 2012; see also refer- MATERIAL AND METHODS uous characters were inserted directly into the .tnt and taken preferentially from the left side of speci- ences therein). On the other hand, systematic studies file using a text editor. Intervals for continuous char- mens. Variation of counts/measurements is given as MORPHOLOGY AND CHARACTER SCORINGS on Sicarius have been long neglected. Most Sicarius acters consist of the average value of each measure- minimum–maximum (mean). Drawings were made species have been known only from their original We have examined around 1800 adult specimens of ment for each species two standard errors, as using a Leica M205C stereoscopic microscope (and often incomplete or inadequate) descriptions Sicarius during the course of this work. For separat- suggested by Goloboff, MattoniÆ & Quinteros (2006). equipped with a camera lucida, or based on pho- (e.g. Keyserling, 1880; Mello-Leitao,~ 1936). Until ing the species, the prolateral view of the male palp Parsimony analyses were carried out in TNT 1.1 tographs taken in the same microscope using a very recently, the few restricted revisions and spe- and the ventral view of the female spermathecae pro- (Goloboff, Farris & Nixon, 2008). We used three DFC295 digital camera, and mounted as a single, cies redescriptions available (e.g. Mello-Leitao,~ 1940, vide the most informative characters. Somatic char- approaches: (1) analysis of discrete characters only multi-focal image using Leica Application Suite 3.8. 1941; Gerschman de Pikelin & Schiapelli, 1979) were acters are also of great taxonomic value, and can be under equal weights, (2) analysis of discrete charac- Some female spermathecae were photographed using not based on type-material examination, leading to used to diagnose most groups of closely related spe- ters only under implied weights and (3) analysis of an AxioCam ERc5s digital camera attached to a frequent species misidentifications. Overall, the tax- cies. Exposing the ventral face of the spermathecae discrete and continuous characters under implied Zeiss PrimoStar compound microscope following clar- onomy of Sicarius was chaotic and its species were is somewhat tricky; it is most easily done by digest- weights. We refrained from further analysing a data- ification using clove oil. Geographical coordinates of unrecognizable. This started to change with revisions ing the soft tissues with enzymes, and then cutting set including continuous characters under equal specimens collected by us, or museum specimens of species from Africa (Lotz, 2012) and north-eastern the ventral cuticle of the abdomen, or bending the weights because preliminary runs showed that with coordinates indicated in the original labels, are Brazil (Magalhaes,~ Brescovit & Santos, 2013). How- spermathecae. Generally, we have examined female meristic characters alone (namely the number of given in parentheses. Specimens without coordinates ever, the genus also occurs in arid environments genitalia after digestion using a pancreatin solution spermathecae branches and femoral thorns) heavily were georeferenced using Google Earth and have from Argentina to Central America, from where 14 (see Alvarez-Padilla� & Hormiga, 2008). This was also influenced the analysis, leading to highly deviant their coordinates indicated in square brackets. An species (one of them with two subspecies) have been used for preparing genitalia, and sometimes spin- results. For exploring the sensitivity of the results to interactive key for identifying all Sicarius species named, most of which have never been revised. nerets, for scanning electron microscopy (SEM). the value of k in the implied weight analyses, we was made using the Xper² platform (Ung et al., 2010) Sicarius are unique spiders for several reasons. Parts were dried for SEM using heat from a light used the TNT script by Mirande (2009) to find the and is available as Supporting Information They do not spin webs, but rather stalk their prey bulb (tough parts, such as palps) or by transfer most fit trees under a wide range of k values. Appendix S1. while buried in dry sediment (Reiskind, 1966). Their through a series of solutions of increasing concentra- Searches for optimal trees consisted of 1000 random The examined specimens are deposited in the cuticles have special setae which adhere to soil tion of ethanol, and then either transferred to addition sequences followed by tree bisection and following institutions (curators indicated in

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016 ©© 20172016 The The Linnean Linnean Society Society of of London, London, ZoologicalZoological Journal Journal of ofthe the Linnean Linnean Society Society, 2017,, 2016 179, 767–864 7704 I. L. I. F.L. MAGALHAESF. MAGALHAESET ET AL. AL. SICARIID PHYLOGENY AND REVISION OF SICARIUS 5 parentheses): AcAT – Plant Protection Research parsimony analysis found 41 shortest trees Institute, Pretoria, South Africa (A. Dippenaar- [length = 187, consistency index (CI) = 0.583, reten- Schoeman); AMNH – American Museum of Natural tion index (RI) = 0.860, but with a well-resolved History, New York, USA (L. Prendini); CAI – consensus (Fig. 1B). The weighted analyses gave Coleccion� Aracnologica� del Instituto Argentino de identical results either in the absence or in the pres- Investigaciones de las Zonas Aridas,� Mendoza, ence of continuous characters, so we only show the Argentina (S. Lagos); CAS – California Academy of latter results (Fig. 2). Analyses using k values from Sciences, San Francisco, USA (C. Griswold); DNSM – 0.8 to 3.66 gave identical results (three most fit Durban Natural Science Museum, Durban, South trees, length = 283.473, CI = 0.518, RI = 0.815). The Africa (K. Williams); IBSP – Instituto Butantan, Sao~ two higher k values (4.947 and 7.201) differ only in Paulo, Brazil (A.D. Brescovit); DZUB – Universidade the position of Loxosceles of the spadicea group. The de Bras�ılia, Bras�ılia, Brazil (P.S. Motta); KNM – Bayesian inference yielded similar results, although Kwazulu-Natal Museum, Pietermaritzburg, South many of the relationships were unresolved (Fig. 1A). Africa (B. Muller); LEULS – Laboratorio de Ento- Our results consistently recover the monophyly of molog�ıa de la Universidad de La Serena, La Serena, Sicariidae, a group of ‘true’ Loxosceles (all sampled Chile (J. Pizarro-Araya); MACN – Museo Argentino species but L. simillima Lawrence), Sicariinae, de Ciencias Naturales ‘Bernardino Rivadavia’, Bue- Neotropical Sicarius and a group of African Sicarius nos Aires, Argentina (M.J. Ram�ırez & C. Scioscia); including the type species of Hexophthalma, S. hahni MCZ – Museum of Comparative Zoology, Harvard (Karsch) (Figs 1, 2). The only species groups of Lox- University, Cambridge, USA (G. Giribet); MHNS – osceles with more than one terminal included, spadi- Museo de Historia Natural de Santiago, Santiago, cea and rufescens, are recovered in all analyses with Chile (M. Elgueta); MNHN – Museum� National good support. Our results also recover several small d’Histoire Naturelle, Paris, France (C. Rollard); groups of closely related species of Neotropical Sicar- MNRJ – Museu Nacional do Rio de Janeiro, Rio de ius, such as the rugosas group of Nicolet (1849) (see Janeiro, Brasil (A.B. Kury); MUSM – Museo de His- Taxonomy below) or the Sicarius from the Brazilian toria Natural de la Universidad Nacional Mayor de Caatinga (Magalhaes~ et al., 2013). However, relation- San Marcos, Lima, Peru (D. Silva); MZSP – Museu ships among these small species groups remain lar- de Zoologia da Universidade de Sao~ Paulo, Sao~ gely unresolved or unsupported (Fig. 1). The results Paulo, Brazil (R. Pinto da Rocha); NHM – Natural from different methods and weighting schemes are History Museum, London, England (J. Beccaloni); generally similar in topology, disagreeing mainly in NHMB - Naturhistorisches Museum Basel, Basel, unsupported nodes and the shallower relationships Switzerland (A. Hanggi); PAN – Polska Akademia (Figs 1, 2). Nauk, Warsaw, Poland (J. Swedo); RBINS – Royal Figure 1. Phylogenetic relationships among sicariids. A, majority-rule consensus of the trees found through Bayesian Belgian Institute of Natural Sciences, Brussels, Bel- inference under the Mkv + Γ model. Numbers next to nodes are posterior probabilities. The Sicariinae branch was bro- CHARACTER OPTIMIZATIONS gium (L. Baert); TM – Ditsong Museum, Pretoria, ken as it is too long. B, majority-rule consensus of the 41 most parsimonious trees found analysing only the 98 discrete South Africa (A. Ndaba); MUAP – Universidad Sphingomyelinase D has two clades of paralogues, characters under equal weights. Bremer supports are shown above branches, symmetric resampling GC values below Arturo Prat, Iquique, Chile (G. Guzman); UCCC – termed a (ch. 98) and b (ch. 99), which are respec- branches. Nodes with 0 Bremer support are absent from the strict consensus. Scale bar in A indicates the number of Universidad de Concepcion,� Concepcion,� Chile tively recovered as synapormorphies for Sicariidae expected changes per character. (J. Artigas); UFMG – Centro de Coleçoes~ Taxonomi-^ and all Loxosceles but L. simillima. Whether der- cas da Universidade Federal de Minas Gerais, Belo monecrotic activity is present (ch. 100) is ambigu- based on phenotypic (chiefly morphological) data. Sicariidae is the second best supported group of our Horizonte (A.J. Santos); MLP – Facultad de Ciencias ously optimized at the base of the sicariid clade Our results support its monophyly, as well as that of analysis, and rests on eight non-ambiguous, homo- Naturales y Museo, Universidad de La Plata, La (Fig. 3). The diversity of setae and macrosetae in its two subfamilies (Sicariinae and Loxoscelinae), plasy-free synapomorphies. These include three clas- Plata, Argentina (L.A. Pereira); UNSAAC – Museo Sicarius has a mixed signal. Setae with hairlettes for reporting novel synapomorphies for these taxa. We sic or previously known characters: the short and de Historia Natural de la Universidad Nacional de adhering particles are a Sicariinae synapomorphy provide evidence of a strong phylogenetic structure conical setae against the cheliceral promarginal lobe San Antonio Abad del Cusco, Cusco, Peru (R. Orel- (ch. 9), and prolateral black setae on the femora (ch. among Sicarius as presently recognized, with species (ch. 19; Labarque & Ramırez, 2012, fig. 15), the loss lana); USNM – National Museum of Natural 43) are the sole synapomorphy of a large clade from Africa and America forming reciprocally mono- of the inferior tarsal claw (ch. 54; Fig. 4A), and the History, Smithsonian Institution (J. Coddington); within Sicarius (Fig. 3). On the other hand, features phyletic groups, and suggest a subdivision of the large and elongate 3rd abdominal entapophysis (ch. ZMB – Museum fur€ Naturkunde, Berlin, Germany such as multiple rows of macrosetae in the lateral genus. 58, Figs 5F, 6D). We report new synapomorphies, (J. Dunlop). border of the carapace (ch. 10) and white and brown such as the large, detached cheliceral promarginal macrosetae in the femora (ch. 46) seem to have lobe (ch. 17; Labarque & Ramırez, 2012, fig. 15), the MONOPHYLY OF SICARIIDAE evolved multiple times (Fig. 3). behaviour of attaching eggcases to the substrate (ch. RESULTS Our study was not primarily designed to test the 93), and the presence of b clade sphingomyelinase D monophyly of the family, which has been supported in the venom (ch. 99; see Binford et al., 2009). Other PHYLOGENETIC ANALYSES DISCUSSION by previous morphological studies (Platnick et al., characters recovered as synapomorphies, such as the The 38 terminal taxa were scored for 101 phenotypic We here present the first phylogenetic treatment of 1991; Ramırez, 2000; Labarque & Ramırez, 2012), presence of macrosetae on the leg femora (ch. 43) characters (see Appendix 1). The equal weight the medically important spider family Sicariidae and thus we included a limited sample of outgroups. and macrosetae on tibia (ch. 49), are somewhat

Figure 2. Majority-rule consensus of three most-ﬁt trees found analysing 98 discrete and three continuous characters under implied weights using a k value of 3.669 (identical to the majority-rule consensus of trees found using k values of 0.805–7.248). This consensus is identical to the strict consensus, except for relationships within Loxosceles. Values under branches are Bremer supports (top) and GC values of symmetric resampling (bottom). Stars indicate support of branches in different analyses, darker shades indicating higher support. Synapomorphic characters are numbered as in Appendix 1 and are indicated as dots on branches (homoplasy-free ones in black, homoplasious ones in white). Photos, from top to bottom: Loxosceles laeta from Buenos Aires, Argentina (by I.L.F.M.), Hexophthalma sp. from Klapmuts, Wes- tern Cape, South Africa (by G. Van Zyl), Hexophthalma sp. from Namibia (by K. Cramer), Sicarius cariri from Brazil (by P. H. Martins), Sicarius levii holotype from Rıo Negro, Argentina (by I.L.F.M.).

counterintuitive (as macrosetae on femora and tibiae analyses. Also, we consistently recovered a group of are widely present in spiders, and its presence is ‘true’ Loxosceles, including all the species sampled thought to be plesiomorphic) and may be artefacts except for L. simillima. Binford et al. (2008) have due to our limited sample of outgroups. As expected, already presented evidence (based on sequence data) internally Sicariidae is divided into two subfamilies, that Loxosceles of the spinulosa group (including Sicariinae and Loxoscelinae (Fig. 2). L. simillima) are the sister group to other Loxosceles, a result that is supported by our study. In our opin- ion, these ﬁndings might support the allocation of MONOPHYLY AND RELATIONSHIPS AMONG the spinulosa group in a genus of its own. However, LOXOSCELES we are not well acquainted with African Loxosceles, Loxosceles is a large and diverse group, which would and such a taxonomic change must wait for a more certainly deserve a more detailed study. However, densely sampled study focusing on Loxosceles. even our very limited sample of its diversity allows Other points of agreement between our data us to make some considerations about its relation- and those of Binford et al. (2008) are the close ships. This genus is recovered as monophyletic in all relationships between L. amazonica Gertsch and

Figure 3. Optimization of characters related to venom (characters 98, 99 and 100; left) and setae and macrosetae (characters 10, 43 and 46; right). On the right, taxa with their names shaded present soil-adhering setae (character 9, state 1). The coding and optimization of ch. 100 must be taken cautiously, as effects of the venom of most species have never been explored in vivo and our current knowledge of envenomation by sicariids is far from complete.

Figure 4. Scanning electronic microscopy of structures associated to appendages. A, B, Sicarius fumosus (Nicolet, 1849) male, right leg II (IBSP 163716). A, leg tip, retrolateral. B, tibial base, ventral. Note modiﬁed setae (top inset) and chemosensory setae (bottom inset). C, D, Sicarius crustosus (Nicolet, 1849), female, right leg II (MACN 30189). C, femur, dorsal surface. D, tibia, prolateral. Arrows to rows of macrosetae (only ﬁve out of eight are visible). Notice that only the ventralmost bear longer, spiniform macrosetae. CBM, club-shaped macrosetae; M, tibial macrosetae; S, particle-adhering setae. Scale bars, A–B = 0.1 mm; C, 0.2 mm; D, 1 mm.

L. rufescens (Dufour), and among L. laeta (Nicolet), Mello-Leitao,~ and the rufescens group, with L. deserta Gertsch and L. caribbaea Gertsch. On the L. rufescens and L. amazonica) were consistently Figure 5. Scanning electronic microscopy, Sicarius levii sp. nov., male (MACN 30236), spinnerets, setae removed other hand, our results show L. similis Moenkhaus recovered as monophyletic. from the left side. A, spinnerets, ventral view. B, Left anterior lateral spinneret, ventro-posterior view. C, left posterior as sister to L. deserta, an unexpected result in the Unambiguous synapomorphies for Loxosceles (in- median spinneret, ventral view. D, left posterior lateral spinnerets, ventral view. E, epiadrium, ventral view. Inset show light of their distributions and the results of Binford cluding L. simillima) are the slit-like opening of the epiandrium spigots. F, posterior median tracheae, digested, dorsal view. Ac, aciniform gland spigots; ALS, anterior lat- et al. (2008). Unfortunately, our sampling does not major ampullate gland spigot (ch. 69, Figs 7B, 8B, eral spinnerets; MAP, major ampulate gland spigot; Pi, piriform gland spigots; PLS, posterior lateral spinnerets; PMS, allow a test of the monophyly of most of Gertsch’s C), the biﬁd apex of the chelicera lamina (ch. 20, posterior median spinnerets. Scale bars = 0.1 mm. species groups. The only groups represented by two Figs 13C, 14A), the lack of spigots in the posterior species in our sampling (the spadicea group, median spinnerets (ch. 72, Figs 7C, 8D) and the with L. intermedia Mello-Leitao~ and L. hirsuta presence of a setal pit with modiﬁed setae in the

Figure 7. Scanning electron microscopy, Loxosceles hirsuta Mello-Leitao,~ 1931, female (MACN 21483), spinnerets. A, ventral view. B, right anterior lateral spinneret. Inset shows apex of the major ampullate gland spigot. C, posterior median spinnerets. Notice lack of spigots. D, left posterior lateral spinnerets. Ac, aciniform gland spigots; ALS, anterior lat- Figure 6. Scanning electron microscopy, Hexophthalma spatulata (Pocock, 1900) comb. nov., female (TM 13600), eral spinnerets; MAP, major ampulate gland spigot; MS, modiﬁed setae; Pi, piriform gland spigots; PLS, posterior spinnerets. A, ventral view. B, ventral view, close up of region where the posterior median spinnerets should be located. lateral spinnerets; PMS, posterior median spinnerets. Scale bars = A, 0.1 mm; B, 0.02 mm; C–D, 0.05 mm. C, right anterior lateral spinneret, ventral view. D, posterior median tracheae, ventral view. 3rd, posterior median tracheae (= third abdominal entapophysis); Ac, aciniform gland spigots; ALS, anterior lateral spinnerets; MAP, major ampulate gland spigot; Pi, piriform gland spigots; PLS, posterior lateral spinnerets. Scale bars = A, 0.5 mm; B–D, 0.02 mm. the results of Binford et al. (2008), strongly advo- The type species of Sicarius, S. thomisoides Wal- cates for the monophyly of this subfamily. ckenaer, is Neotropical. Currently, Sicarius is con- posterior lateral spinnerets (ch. 73, Figs 7D, 8F). spinnerets covered by black, plumose setae (ch. 64, Both DNA sequences (Binford et al., 2008) and sidered a senior synonym of Hexophthalma The highly branched posterior tracheae are also Figs 9A–C, 10A, 11A, 12C), presence of setae for morphology (this study) support a division of Sicari- (following Simon, 1893), whose type species is exclusive of Loxosceles, but optimization of this char- adhering soil particles (ch. 9, Figs 15, 16), the loss of inae into two clades, one restricted to the Americas S. hahni (Karsch), an African species. Faced with acter is ambiguous because Sicarius lack posterior the serrulae in the chelicera fang and in the endites and one to Africa. Here, the African Sicariinae are the availability of this name and such a phyloge- lateral tracheae. Synapomorphies of ‘true’ Loxosceles (chs 13 and 22, Fig. 14B), the promarginal lobe posi- supported by the loss of the major ampullate gland netic and biogeographical structure, it seems very are the ragged margin of the setae in the setal pit tioned frontally to the apex of the chelicera lamina spigots (ch. 67, Figs 6, 11) and by the loss of the pos- reasonable to resurrect Hexophthalma to accommo- (ch. 74, Fig. 7D) and presence of a clade sphin- (ch. 18, Figs 13, 14B), a membranous sheath at the terior median spinnerets (ch. 70, Figs 6, 11, 12C). date African species. In this way, Sicarius is gomyelinase D in the venom (ch. 98, see Binford apex of the chelicera lamina (ch. 21, Figs 13, 14B, Neotropical Sicarius are supported by the separation restricted to include only Neotropical species et al., 2009). C), a distal crack in the palpal femur (ch. 23, between the lateral eyes (ch. 5, Fig. 12; homopla- (Fig. 25). Furthermore, we found that the African Fig. 43A, B), a proximo-dorsal setae tuft in the sious), the sculptured femoral thorns (ch. 25, clade is subdivided into two lineages. One is com- palpal femur (ch. 26, Fig. 54), loss of the onychium Fig. 18A, C, E, F), a setae tuft anterior to the spin- posed of S. hahni and allied species, and the other MONOPHYLY AND RELATIONSHIPS AMONG SICARIINAE (ch. 51, Fig. 4), straight stalks in the spermathecae nerets in adult females (ch. 61, Fig. 9A), anterior lat- is composed by the aberrant species S. spatulatus The Sicariinae are a very distinct spider group, in (ch. 87, homoplasious), pot-shaped eggcases (ch. 94, eral spinnerets with long setae, covering the Pocock. These two groups are allied through the two our analyses backed by 13 non-ambiguous synapo- Fig. 17), self-burying behaviour (ch. 96) and loss of spinning ﬁeld (ch. 65, Fig. 9A), and loss of the outer synapomorphies listed above, but each individually morphies. These include the anterior lateral the web-building behaviour (ch. 97). This, along with spermathecae (ch. 78, Fig. 19). has several unique characters. The group including

Figure 8. Scanning electron microscopy, Loxosceles spinnerets. A–E, Loxosceles simillima Lawrence, 1927, female (TM Figure 9. Scanning electronic microscopy, Sicarius levii sp. nov., female (MACN 30239), spinnerets, setae removed 10611), ventral view. A, spinnerets. B, left lateral anterior spinneret. C, same, detail of spigot apexes. D, posterior med- from the left side. A, spinnerets, ventral view. B, anterior lateral spinnerets, right side, apex of piriform gland spigots. ian spinnerets. E, right posterior lateral spinneret. F, Loxosceles sp. from South Africa, female (TM 19076) modiﬁed C, anterior lateral spinnerets, close up of black setae. D, close-up of setae from the abdominal ventral setae tuft. E, ante- setae on posterior lateral spinnerets. Ac, aciniform gland spigots; ALS, anterior lateral spinnerets; C, colulus; MAP, rior lateral spinneret, ventro-posterior view. Inset shows detail of putative major ampullate gland spigot. F, posterior major ampullate gland spigot; MS, modiﬁed setae; Pi, piriform gland spigots; PLS, posterior lateral spinnerets; PMS, median spinneret, ventro-posterior view. G, posterior lateral spinneret, ventral view. Ac, aciniform gland spigots; ALS, posterior median spinnerets. Scale bars = A, C, D, 0.1 mm; B, E, 0.05 mm; F, 0.02 mm. anterior lateral spinnerets; BS, black setae; HT, setae tuft; MAP, major ampulate gland spigot; Pi, piriform gland spigots; PLS, posterior lateral spinnerets; PMS, posterior median spinnerets. Scale bars = 0.1 mm, except C (0.01 mm).

Figure 11. Scanning electron microscopy, Hexophthalma sp. 1 (TM 19548), spinnerets. A, ventral view. B, right anterior lateral spinneret, ventral view. C, close-up of region where the posterior median spinnerets should be located, ventral view. Arrows indicate PMS setae. D, right posterior lateral spinnerets. Ac, aciniform gland spigots; ALS, anterior lateral spinnerets; Pi, piriform gland spigots; PLS, posterior lateral spinnerets. Scale bars = A, 0.5 mm; B–D, 0.01 mm. Figure 10. Scanning electron microscopy, Sicarius crustosus (Nicolet, 1849), female (MACN 30189), spinnerets, setae removed from the left side. A, spinnerets, ventral view. B, anterior lateral spinnerets, ventro-posterior view. Inset shows detail of putative major ampullate gland spigot. C, posterior median spinnerets, ventro-posterior view. D, posterior lat- Argentina), to name a few. However, the relation- species, but their sparse sampling and lack of sup- eral spinnerets, ventral view. Ac, aciniform gland spigots; ALS, anterior lateral spinnerets; AT, anal tubercle; MAP, putative major ampullate gland spigot; Pi, piriform gland spigots; PLS, posterior lateral spinnerets; PMS, posterior med- ships among these small clades are less clear. This is port for some relationships (in the latter work) ian spinnerets. Scale bars = 0.1 mm. demonstrated by the large polytomies within Sicar- makes it difﬁcult to compare results with the present ius and by the low support values of many groupings study. We are currently generating sequence data (Figs 1, 2). Despite that, our data indicate that for several species that were not sampled in those S. hahni has a sigmoid embolus (ch. 40, Figs 20A, S. thomisoides and its sister species, S. yurensis stat. works, and expect that this, allied with the pheno- RELATIONSHIPS AMONG NEOTROPICAL SICARIUS C, 21A, C), reduced outer spermathecae (ch. 79 nov., are the sister group to the remaining Sicarius. typic data gathered here, will provide a better pic- Figs 20B, D, 21B, D) and branches of the inner Our results are somewhat disappointing regarding Also, we found evidence of a large clade of species ture of the relationships among Sicarius. spermathecae arising from a single stalk (ch. 81, the internal relationships of Neotropical Sicarius. On with prolateral black setae on the femora (see Fig. 20B, D). Sicarius spatulatus has a long list of the one hand, we recovered several small groups of Fig. 24B–D), including the six species from Brazil, EVOLUTION OF VENOM, SPINNING ORGANS AND autapomorphies, including a broad and blunt embo- obviously related, morphologically similar species, S. peruensis (Keyserling), S. gracilis (Keyserling) CRYPTIC BEHAVIOUR lus (ch. 39, Figs 20E, 22C–E), club-shaped macrose- which also happen to occur in geographical proxim- (both from Peru), S. rugosus (F.O. Pickard-Cam- tae on the leg femora and the abdomen (ch. 47 and ity. This is the case for the six species from Brazil, bridge), S. utriformis (Butler) and the rugosas group Loxosceles have a notorious reputation for the der- 57; see Lotz, 2012) and branched outer spermathe- the rugosas group of Nicolet (S. fumosus, S. crusto- of Nicolet (1849). monecrotic symptoms primarily caused by the action cae (ch. 80, Figs 20F, 23D), making this species an sus and S. lanuginosus, all from Chile), and Binford et al. (2008) and Magalhaes et al. (2014) of a family of proteins homologous with Sphin- interesting lineage for future studies. S. rupestris (Holmberg) + S. mapuche sp. nov. (from presented sequence data on a few Neotropical gomyelinase D (SmaseD), termed SicTox by Binford

Figure 13. Sicariidae chelicerae apex, retromargin. A, Sicarius cariri Magalhaes,~ Brescovit & Santos, 2013 female Figure 12. A, Sicarius thomisoides Walckenaer, 1847, cephalothorax, lateral. Inset showing lateral row of macrosetae. (UFMG 11688) (right chelicerae, mirrored). B, Hexophthalma hahni (Karsch, 1878) comb. rest. female (KNM 24780). B, Sicarius rugosus (F.O. Pickard-Cambridge, 1899), cephalothorax, lateral. Inset showing lateral row of macrosetae. C, C, Loxosceles gaucho Gerstch, 1967 female (IBSP 34933). CL, chelicerae lamina apex; MS, membranous sheath; PL, pro- Hexophthalma spatulata (Pocock, 1900) comb. nov. female spinnerets, ventral view (AcAT 2013/1514). D, Sicarius marginal lobe. Scale bars = 0.1 mm. diadorim Magalhaes,~ Brescovit & Santos, 2013, female third entapophysis, dorsal, digested. ALS, anterior lateral spinnerets; PLS, posterior lateral spinnerets; AT, anal tubercle. Scale bars = A–B, 1 mm; C–D, 0.1 mm.

activity’ as a discrete, absence–presence character is make very little use of silk – we have only observed et al. (2009). These proteins are apparently exclusive appeared at the base of sicariids and was lost in an approach that should be viewed with caution. Neotropical species using it for eggcase construction to Sicariidae among spiders (Binford & Wells, 2003). Neotropical Sicarius (as suggested by the optimiza- Thus, our optimization of this character in Fig. 3 and for building sperm webs. African Sicariinae have Binford et al. (2009) found that the SicTox protein tion by Binford & Wells, 2003), or it appeared con- must be taken conservatively – it is intended to stim- very simplified spinnerets. They lack posterior med- family can be divided into two clades, termed a and vergently in Loxosceles and Hexophthalma gen. ulate future research, and not to be considered as ian spinnerets altogether, and have only one type of b. The a clade is exclusive of ‘true’ Loxosceles, while reval. (Fig. 3). It is noteworthy that the venom of definite evidence that a particular species is or is not spigot in the anterior lateral spinnerets which we the b clade is widespread in sicariids (Fig. 3). Inter- S. spatulatus has never been investigated, and thus harmful to humans. interpret as piriform gland spigots (Figs 6A–C, 11A, estingly, the necrotic activity is present in Loxosceles a description of its composition and effects might The evolution of spinning organs in Sicariidae is B, 12C). On the other hand, Neotropical Sicarius do and African Sicariinae, but not in (most) Neotropical help to clarify this point. Also, the fact that at least very curious. While other scytodoids generally have have posterior median spinnerets with spigots Sicarius, which do express proteins of the SicTox the Neotropical species S. ornatus has SmaseD activ- more plesiomorphic features in their spinnerets (see (Figs 9F, 10C), and distinct, filiform spigots in the family in their venoms (Alegre et al., 1977; New- ity in vitro (Lopes et al., 2013) is yet another Platnick et al., 1991; Labarque & Ram�ırez, 2012), anterior lateral spinnerets that we interpret as lands, 1982; Binford et al., 2009; Zobel-Thropp, Bod- instance of homoplasy (Fig. 3), and thus should be both Loxoscelinae and Sicariinae have modified their major ampullate gland spigots (Figs 9E, 10B). It ner & Binford, 2010). However, Lopes et al. (2013) further investigated. Note, however, that our current spinning apparatus in different directions. Loxosceles would be interesting to investigate what kind of silk reported necrotic activity in the Neotropical S. orna- understanding of the effects of envenomation by have long anterior lateral spinnerets with a unique is spun using these spigots. Interestingly, both Afri- tus Magalhaes,~ Brescovit & Santos. Considering the sicariid bites is still far from complete, especially for morphology of the major ampullate gland spigots, can and Neotropical Sicariinae build modified, sandy relationships among sicariids recovered by Binford Sicariinae spiders. Most species have only been which have a slit-like opening (Figs 7B, 8B, C). This pot-shaped eggcases, despite the fact that only et al. (2008) and our study, an interesting pattern screened using in vitro essays (Binford et al., 2009; allows them to produce ribbon-shaped silk to build Neotropical females have the dense setae tuft anteri- emerges. The presence of the b clade of SmaseD is a Lopes et al., 2013), and the consequences of their their webs (Coddington et al., 2002; Knight & Voll- orly to the spinnerets. These setae are used by the sicariid synapomorphy, but the dermonecrotic activ- bites in humans or other vertebrates are not known. rath, 2002). They also lack spigots in the posterior female as a brush to collect and deposit sand grains ity has an ambiguous optimization. It has either With this in mind, coding ‘SMase-D dermonecrotic median spinnerets. In contrast, Sicariinae seemingly on the eggcase (see Levi & Levi, 1969), and thus we

Figure 14. Scanning electron microscopy, appendages. A, Loxosceles hirsuta Mello-Leitao,~ 1931, female (MACN 21483), right chelicera, posterior view. B, Hexophthalma hahni (Karsch, 1878) comb. rest. female (TM 6511), right chelicera, posterior view. Note absence of fang serrula. C, same, detail of the apex of the cheliceral lamina. D, Hexophthalma spatulata (Pocock, 1900) comb. nov., female (TM 13600), right palp, prolateral. CL, apex of chelicera lamina; F, chelicera fang; FS, fang serrula; FT, femoral thorns; MS, membranous sheath; PL, promarginal lobe of the chelicera. Scale bars: A–C, 0.1 mm; D, 0.5 mm. presume that African Sicarius follow a different rou- paratropidids, and some zodariids and thomisids tine during eggcase construction, which awaits a (Ram�ırez, 2014). In Sicariinae, dry soil particles detailed description. adhere to special setae with fine hairlettes (Duncan One of the most remarkable characteristics of et al., 2007; see Figs 4C, 15, 16; Magalhaes~ et al., Sicariinae is their cryptic appearance and behaviour. 2013, fig. 24). Based on our anedoctal observations, it Many other spiders add materials to their cuticles, seems that some species are able to add thicker layers including homalonychids (Duncan et al., 2007), of dirt to the cuticle, while others have only a fine

Figure 15. Living Sicarius specimens. A, Sicarius thomisoides Walckenaer, 1847, male from Flamenco, Atacama, Chile (IBSP). B, same species, female from Caleta Los Patos, Caldera, Atacama, Chile (IBSP). C, Sicarius crustosus (Nicolet, 1849), male from Freirina, Atacama, Chile (IBSP 165577). D, same species, female from Lorenzo Peralta, Coquimbo, Chile (IBSP 165584). E, Sicarius fumosus (Nicolet, 1849), female from Concon,� Valpara�ıso, Chile (IBSP 163713). F, Sicarius lanuginosus (Nicolet, 1849), male from Punta Choros, Coquimbo, Chile (photo by J. Pizarro-Araya) (LEULS). G, Sicarius rugosus (F.O. Pickard-Cambridge, 1899) from Guanacaste, Costa Rica. Photo by F. Castellucci. H, Sicarius rupestris (Holmberg, 1881), female from Sierra del Tontal, San Juan, Argentina.

Figure 17. Sicarius eggcases. A, Sicarius cariri Magalhaes,~ Brescovit & Santos, 2013 from Campo Formoso, Bahia, Brazil. B, Sicarius fumosus (Nicolet, 1849) from Santiago, Chile. C, Sicarius peruensis (Keyserling, 1880) from Lima, Peru. D, Sicarius thomisoides Walckenaer, 1847 from Los Vilos, Coquimbo, Chile. Note absence of collar. E, Sicarius utriformis (Butler, 1877) from Espanola,~ Galapagos Islands, reproduced from Banks (1902).

layer that barely covers them. The ‘cleaner’ species examination of several specimens from different geo- include S. thomisoides Walckenaer, S. rupestris and graphical regions allowed us to recognize 20 species S. levii sp. nov.; ‘dirtier’ species include S. rugosus, in the New World, of which six are newly described. S. ornatus, S. crustosus and S. spatulatus. We are We redescribe nine poorly known species, elevate not sure of what morphological features allow them to Sicarius terrosus yurensis Strand to the status of add more dirt to their cuticles, but it might be their species and synonymize six species names. The smaller size, higher density of macrosetae and the genus is newly recorded for Bolivia, mainland Ecua- morphology of the hairlettes. Some morphological dor, Nicaragua and Paraguay. For more details, see features, such as the presence of multiple rows of Taxonomy below. macrosetae in the lateral border of the carapace, prolateral black setae on the femora, and white and SPECIES LIMITS IN SICARIUS brown macrosetae on the legs, might help further con- ceal the spiders (see optimization of these characters We have demonstrated before that Sicarius species in Fig. 3). If these characters are indeed correlated are morphologically variable (Magalhaes~ et al., 2013) with a greater capacity of hiding, then highly cryptic and can have strong geographical structure in morphology has evolved independently at least twice: genetic variation (Magalhaes et al., 2014). This is a in Hexophthalma spatulata comb. nov. and in Chilean dangerous combination for a taxonomic study, and species closely related to S. crustosus (Fig. 3B). Curi- suggests that our taxonomic treatment of the group ously, it seems that ‘dirtier’ species tend to remain might be superficial. We cannot help but be motionless when molested, while ‘cleaner’ do not rely reminded of the work of Willis Gertsch on Loxosceles. so much on camouflage and tend to run. It might be After many years working on the group, he provided interesting to further investigate how ‘dirtier’ and a monograph on the North American species ‘cleaner’ species behave when molested, and if there (Gertsch & Ennik, 1983) in which he described doz- Figure 16. Living Sicarius specimens. A, Sicarius levii sp. nov., male paratype from Rıo Negro, Argentina (MACN). are behavioural differences among species with and ens of new species with narrow geographical ranges Photo by D. Proud. B, Sicarius levii sp. nov., female paratype from Rıo Negro, Argentina (MACN). Photo by H. A. without prolateral black setae on the femora. and subtle morphological differences from related Iuri. C, Sicarius peruensis (Keyserling, 1880), male from Cieneguilla, Lima, Peru (IBSP). D, Sicarius saci sp. nov., species, often obscured by great intraspecific varia- male from Morro do Chapeu, Bahia, Brasil (IBSP). Photo by G. F. B. Pereira. E, , tion. However, they are seemingly ‘good’ species, at Sicarius jequitinhonha sp. nov. TAXONOMY female from Espinosa, Minas Gerais, Brasil (IBSP). Photo by P. H. Martins. F, same data, male. G, Hexophthalma cf. least judging by their phylogenetic relationships and hahni comb. rest., male from Blouberg, Limpopo, South Africa. Photo by S. Jacobsz. H, Hexophthalma sp., female from Based on our phylogenetic results, we redelimit genetic distances (Binford et al., 2008). It seems that Klapmuts, Western Cape, South Africa. Photo by G. Van Zyl. Sicarius to include only Neotropical species, and the case with Sicarius might be similar, and that transfer African Sicariinae to Hexophthalma. The the 20 species we recognized are just a coarse picture

Figure 18. Scanning electron microscopy, right male palpi (except H, female), proximal femoral thorn, prolateral view. A, Sicarius thomisoides Walckenaer, 1847 (IBSP 163744). B, Sicarius fumosus (Nicolet, 1849) (IBSP 163716). C, Sicar- ius mapuche sp. nov. (CAI 3092). D, Sicarius peruensis (Keyserling, 1880) (IBSP 162916). E, Sicarius boliviensis sp. nov. (AMNH, from San Antonio de Parapeti, Bolivia). F, Sicarius levii sp. nov. (MACN 30239) (left palp, mirrored). G, Hexophthalma hahni (Karsch, 1878) comb. rest. H, Hexophthalma spatulata (Pocock, 1900) comb. nov. (TM 13600). I, Loxosceles simillima Lawrence, 1927 (left palp, mirrored) (TM 10611). Scale bars = 0.05 mm (except H–I, 0.02). Figure 19. Sicariinae spermathecae, digested, clove oil cleared, ventral views. A, Sicarius thomisoides Walckenaer, 1847 (IBSP 163732). B, Sicarius utriformis (Butler, 1877) (RBINS). C, Hexophthalma hahni (Karsch, 1878) comb. of the true diversity of the genus. Some species are small-scale, regional revisions of those complicated rest. (KNM 24870). D, Hexophthalma spatulata (Pocock, 1900) comb. nov. (AcAT 2013/1514). Inset showing base of notably problematic. Sicarius rupestris, S. thomisoides species. Finally, many of the species are not variable outer spermathecae. IS, inner spermathecae; OS, outer spermathecae; PP, sclerotized pore plate; UE, uterus externus; and S. fumosus have wide geographical ranges and and probably do not represent species complexes, VB, ventral branches. Scale bars = 0.1 mm. are very variable along their entire distributions. such as S. levii sp. nov., S. andinus sp. nov. and Other species have some deviant populations, such S. yurensis stat. nov. Brazil), while the latter occurs in dry forest nuclei in One of the most intriguing issues on the distribu- as S. boliviensis sp. nov. from Cusco or S. gracilis Bolivia and in the Peruvian Andes. These small tion of Sicariinae is their disjunct distribution. from Ecuador and northern Peru. This is stressed by BIOGEOGRAPHY patches of dry forests scattered in inter-Andean val- Regarding their presence in both Africa and the the fact that some populations of two species from leys are probably interesting areas for future sam- Americas, Binford et al. (2008) presented evidence Brazil we had treated before (Magalhaes~ et al., 2013) Sicariinae spiders are largely restricted to dry envi- pling: S. andinus sp. nov. is a micro-endemic species that the subfamily is ancient and was present before are here described as two new species. This does not ronments, namely deserts, xeric scrublands and dry restricted to one of these areas in Huanuco, Peru. the two continents drifted apart. On a finer scale, mean that our taxonomic efforts were unfruitful. forests (Figs 25–29). These environments have a dis- Dry forests in Peru, Colombia, Ecuador and Vene- Sicarius are scattered among different patches of dry First, it seems safe to assume that the 20 species we junct distribution in the Neotropical region (see zuela are probably under-sampled, and collecting environments across the Neotropical region. It would recognize are independent lineages. Second, by rec- Sarkinen€ et al., 2011; Werneck, 2011), and each spe- efforts there might reveal new records and addi- be interesting to investigate if these areas have been ognizing and (re)describing each species, we allow cies is generally restricted to a small region of con- tional, endemic species. In addition to housing those connected in the past, and when, to understand which their identification and provide data on their distri- tiguous dry habitat. Examples of the few species patches of dry forests, the Andes act as an important events led to this disjunct pattern of distribution. The butions. This could form the basis for a finer treat- with disjunct distributions are S. cariri (Fig. 27) and barrier between Argentinean and Chilean species, antiquity of the group (see Binford et al., 2008; ment of each species separately, which would be S. boliviensis sp. nov. (Fig. 26). The former occurs in and the faunas of these two countries do not overlap. Magalhaes et al., 2014) suggests that these events much more feasible than studying the entire genus the Caatinga (a large dry forest in north-eastern Investigating whether there is a correlation between have probably occurred in the Miocene or ealier. Hav- at once from an integrative perspective. Thus, our Brazil) and in small, isolated patches of dry forest the timing of Andean uplift and Sicarius diversifica- ing a dated, complete phylogeny of the group would work can be seen as the starting point to guide within the Cerrado (a savannah formation in central tion might prove an interesting field of research. certainly help to shed light into these questions.

Figure 21. Sicariinae from Africa, genitalia. A, Hexophthalma albospinosa (Purcell, 1908) comb. nov. from Luder-€ itz, Karas, Namibia (KNM 21143), male palp, prolateral. B, same data, female spermathecae, ventral. C, Hexophthalma sp. 2 from Swakopmund, Erongo, Namibia (AcAT 2013/4241), male bulb, prolateral. D, Hexophthalma sp. 2 from 73 km E Walvis Bay, Namibia (AcAT 2013/4239), female spermathecae, ventral. Arrows point to incrassated region of the spermophor. Bd, bulb body; Bs, bulb base; E, embolus; IS, inner spermathecae; OS, outer spermathecae. Scale bars = A, C, Figure 20. Sicariinae from Africa, genitalia. A, Hexophthalma albospinosa (Purcell, 1908) comb. nov. from Luder-€ 1 mm; B, D, 0.2 mm. itz, Karas, Namibia (KNM 21143), male bulb, prolateral. B, Hexophthalma albospinosa (Purcell, 1908) comb. nov. from Luderitz,€ Karas, Namibia (KNM 21143), female spermathecae, ventral. C, Hexophthalma sp. 2 from Swakopmund, Erongo, Namibia (AcAT 2013/4241), male bulb, prolateral. D, Hexophthalma sp. 2 from 73 km E Walvis Bay, Namibia with dorsal macrosetae arranged in a central group (AcAT 2013/4239), female spermathecae, ventral. E, Hexophthalma spatulata (Pocock, 1900) comb. nov. from Mossel SICARIINAE KEYSERLING, 1880 Bay, South Africa (TM 22762), male bulb, prolateral. F, Hexophthalma spatulata (Pocock, 1900) comb. nov. from surrounded by 4–5 pairs of macrosetae lines Port Elizabeth, Eastern Cape, South Africa (AcAT 2013/1514), female spermathecae, ventral. Arrows point to embolus (Figs 22A, 37). Labium longer than wide, tapering keels. Bd, bulb body; Bs, bulb base; E, embolus; IS, inner spermathecae; OS, outer spermathecae. Scale bars = 0.1 mm. Diagnosis: Sicariinae is easily distinguished from distally, partially fused to sternum. Endites with a Loxoscelinae by several characters, of which the white apex, touching in front of labium (Fig. 30D). most readily observable are: presence of special setae Sternum usually cordiform, wider than long and TAXONOMY claws (inferior one absent), both with a single row of to adhere soil particles (Figs 4C, 15, 16); a line of posteriorly truncate, but oval and longer than wide in teeth; laterigrade legs; a large promarginal lobe in several femoral thorns on the palp of males and some Sicarius; with three outer rows of longer SICARIIDAE KEYSERLING, 1880 the chelicerae; and elongated and broad 3rd fused females (Figs 18, 21A, C) (a single thorn in macrosetae near the border, and shorter macrosetae Sicaroidae Keyserling, 1880: 267–268, containing entapophysis which make up the posterior median Loxosceles); and colulus absent (Fig. 12C). in the middle (Fig. 30). Chelicerae fused at the base, Thomisoides Nicolet as the single included trachea. without teeth, with a lamina along the promargin genus. Description: Araneomorphae, ecribellate, haplogyne ending in a pointed, simple apex surrounded by a Included genera: Sicarius Walckenaer, Loxosceles spiders. Six eyes, arranged in three diads, the membranous sheath at the base, without fang serrula Diagnosis: Sicariids can be distinguished from other Heinecken & Lowe and Hexophthalma Karsch gen. anterior median eyes (AME) absent (Fig. 37). (Fig. 14B). Palp without claw, femur with 2–7 large six-eyed haplogyne spiders by having two tarsal reval. Carapace about as wide as long, with one or multiple prolateral thorns and a distal crack in the ventral face rows of macrosetae on lateral borders (Fig. 12), and (Fig. 43A, B). Leg femora covered in the dorsal

Figure 22. Hexophthalma spatulata (Pocock, 1900) comb. nov., male from Mossel Bay, South Africa (TM 22726). A, habitus, dorsal view. B, habitus, ventral view. C, left palp, prolateral view. D, left bulb, prolateral view. E, left bulb, dorsal view. Scale bars = A–B, 2 mm; C, 0.5 mm; D–E, 0.2 mm. surface by macrosetae with a truncate apex, and with with two articles, covered by black setae, apical article a dorsal, longitudinal line without setae (Fig. 24). with numerous piriform gland spigots, and major Tibia with eight rows of pointed macrosetae: two ampullate gland spigots either absent or present dorsal, two prolateral, two retrolateral and two (Figs 6C, 9E, 10B), one or two in number, being long ventral, with large macrosetae intercalated with short and filiform, resembling the piriform gland spigots. macrosetae (Fig. 4D). Tarsus without onychium, with Posterior median spinnerets (PMS) absent, or present two claws (inferior tarsal claw absent), generally with with a few aciniform gland spigots (Figs 6B, 9F, 10C). greatly elongated tips (Fig. 4A). Trichobothria with a Posterior lateral spinnerets (PLS) with only aciniform very narrow socket, a proximal hood and a distal gland spigots (Figs 9A, G, 10A, D, 11D). Palpal ridged or smooth surface (Fig. 31). Tarsal organ bulb simple, pirifom, with three distinguish- Figure 23. Scanning electron microscopy of African Sicariinae genitalia. A, Hexophthalma hahni (Karsch, 1878) capsulate (Labarque & Ramırez, 2012: fig. 10F). able regions: a base, a body and the embolus (Figs 20, comb. rest., female (TM 6511), spermathecae, digested, ventral view. B, same, detail of left spermathecae. Inset shows Abdomen rounded, slightly truncate posteriorly, with 32). Inner spermathecae branched (Fig. 19). Outer a pore. C, same species, male (TM 9904), right palpal bulb, prolateral. D, Hexophthalma spatulata (Pocock, 1900) 3–6 paired transversal rows of macrosetae (Figs 15, spermathecae absent or present (Fig. 19). comb. nov., female (TM 13600), left spermathecae, digested, ventral view. E, same, detail of putative inner spermathe- 16, 22A). Epiandrium composed of a single, dense Spermathecae pores large, with multiple flagella cae. F, same, detail of putative outer spermathecae. Bd, bulb body; Bs, bulb base; E, embolus; IS, inner spermathecae; group of spigots (Fig. 5E). Posterior trachea composed arising from them (Fig. 23B, inset). Subadult females OS, outer spermathecae; UE, uterus externus. Scale bars = A, C, 1 mm; B, D, 0.1 mm; E, 0.05 mm; F, 0.02 mm. of a long and broad 3rd fused entapophysis; lateral with a pre-spermatheca, which is simpler, smaller tracheae absent (Figs 5F, 6D). Colulus absent and without pores (Figs 36H, 39G, 44H, 53E). (Figs 6A, 9A, 10A). Anterior lateral spinnerets (ALS) Carapace, endites, labium, sternum and chelicerae

Figure 25. Known distribution of Sicariinae spiders, including records from Lotz (2012). Left: Hexophthalma gen. reval. (stars, H. spatulata comb. nov., circles, other species). Right: Sicarius. Squares indicate the position of the distribution maps of Sicarius species (see Figs 26, 27). Xeric formations are highlighted in red (deserts and xeric scrublands), yellow (dry forests) and brown (chaco).

the bulb (Fig. 32, Magalhaes~ et al., 2013: fig. 28) (long field, with long and filiform major ampullate gland and with parallel sides in Hexophthalma, Fig. 20A, spigots (one, two or absent) (Figs 9E, 10B). PMS C). Females can be distinguished by the presence of a present, with a few aciniform gland spigots (Figs 9F, dense tuft of setae just anterior to the spinnerets 10C). Inner spermathecae branched, with branches (Fig. 9A, D; Magalhaes~ et al., 2013: fig. 31, ht) (absent inserted throughout the bursa (Figs 19A, B, 33). in Hexophthalma). Both males and females can also Outer spermathecae absent (Figs 19A, B, 33). be distinguished by having posterior median spinnerets (Figs 5A, 9F, 10A, C) (PMS absent in Distribution: Dry regions of South and Central Figure 24. Leg femora. A, Hexophthalma hahni (Karsch, 1878) comb. rest. male, third left leg (KNM 24870), ante- Hexophthalma; Figs 6A, B, 11A, 12C) and by having America. rior view. Inset showing detail of white macrosetae. B, Sicarius fumosus (Nicolet, 1849) female, first right leg (IBSP dense patches of long black setae that partially hide 163741), anterior view (mirrored). C, Sicarius rugosus (F. O. Pickard-Cambridge, 1899) male, second left leg (AMNH), the ALS spinning fields (Fig. 9A, bs; Magalhaes~ et al., THE SICARIUS/THOMISOIDES CONTROVERSY AND anterior view. D, Sicarius utriformis (Butler, 1877) female, second right leg (RBINS), dorso-anterior view (mirrored). E, 2013: fig. 32, bs) (setae much shorter, not hiding the THE NAME OF THE TYPE SPECIES Sicarius yurensis Strand, 1908 stat. nov. female, first left leg (AMNH), dorso-anterior view. White arrows indicate ALS spinning field in Hexophthalma; Fig. 12C). groups of black setae, black arrows indicate white macrosetae. Scale bars = 1 mm. Nicolet (1849) proposed the genus Thomisoides to Description: As for the subfamily, except for the contain seven new Chilean species. While Nicolet’s light brown, reddish brown or dark brown; legs and designated by F.O. Pickard-Cambridge (1899a) following. Cephalic region 0.3–0.5 times as wide as study was still in print, Walckenaer somehow had palpi light brown, yellow, orange, orange brown or (against Simon, 1893, who considered S. terrosus carapace length. Chelicerae lamina with strongly access to the unpublished plates that depicted four of dark brown; abdomen light grey, brownish grey, Nicolet, 1849 to be the type species of Sicarius). sclerotized apex (Fig. 13A). Sternum usually Nicolet’s species. He took advantage of this and, in brownish yellow or light brown. Thomisoides Nicolet, 1849. Type species T. terrosus cordiform, wider than long and posteriorly truncate; his ‘Histoire Naturelle des Insectes: Apteres’� (Walck- Nicolet, 1849, designated by Simon (1893). Synony- oval in S. thomisoides and S. yurensis stat. nov. enaer, 1847), he stated that (1) the four species Included genera: Sicarius Walckenaer and Hexoph- mized by Simon (1893). (Fig. 30). Palp with simple or sculptured femoral depicted in Nicolet’s plate were the same and (2) the thalma Karsch gen. reval. Sicarioides F.O. Pickard-Cambridge, 1899. Type spe- thorns. Leg femora with or without prolateral black name Thomisoides was unsuitable for the genus, as cies by original designation S. rugosus F.O. Pickard- setae, generally with brown macrosetae, rarely with these spiders did not seem to be related to those in Cambridge, 1899. Synonymized by Simon (1903). white and brown macrosetae (Fig. 24). Abdomen the genus Thomisus Walckenaer. Thus, he proposed SICARIUS WALCKENAER, 1847 with a dense tuft of setae anteriorly to the Sicarius as the name of the genus and included a Sicarius Walckenaer, 1847. Type species by mono- Diagnosis: Males can be distinguished from those of spinnerets in adult females (Fig. 9A). ALS covered single species in it, S. thomisoides, considering four typy Sicarius thomisoides Walckenaer, 1847, Hexophthalma gen. reval. by the subconical body of by long, black setae partially covering the spinning of Nicolet’s species as synonyms of it. Paradoxically,

he decided to maintain the same name he rejected levii sp. nov., Sicarius tropicus (Mello-Leitao,~ 1936), for the genus as the specific epithet. As Walckenaer’s Sicarius cariri Magalhaes,~ Brescovit & Santos, 2013, work (1847) was published 2 years before Nicolet’s Sicarius ornatus Magalhaes,~ Brescovit & Santos, (1849), Sicarius has priority over Thomisoides. This 2013, Sicarius diadorim Magalhaes,~ Brescovit & fact did not prevent the use of Thomisoides in subse- Santos, 2013, Sicarius saci sp. nov., Sicarius quent spider species descriptions (Butler, 1877; Key- jequitinhonha sp. nov., Sicarius rugosus (F.O. serling, 1880; Holmberg, 1881; Mello-Leitao,~ 1936), Pickard-Cambridge, 1899) and Sicarius utriformis in some cases with explicit (but unjustified) rejection (Butler, 1877). of the use of Sicarius (Holmberg, 1881; Chamberlin, 1916; Mello-Leitao,~ 1933). This had already been dis- CHILEAN SPECIES, NICOLET’S SEVEN NAMES, AND cussed by Pickard-Cambridge (1899a: 17), who stated THE ABSENCE OF SICARIUS IN VALDIVIA that ‘if we regard Sicarius as the proper generic title [...] we must also regard thomisoides as the proper Chile was arguably the country holding most species specific title [...]’. Thus, he designated in need of revision. Nicolet (1849) described seven S. thomisoides as the type species of Sicarius species from there based on poor diagnostic charac- (against Simon, 1893, who had considered Nicolet’s ters, and Keyserling (1880) added one more species. S. terrosus as the type species), a decision with Despite being one of the most common arthropods in which we agree. However, the arguments of Pickard- the desertic coast of Chile, specific identification was Cambridge (1899a) were largely ignored, and the impossible until now. After our revision, we conclude name S. thomisoides came into disuse until that some species were named several times, and Magalhaes~ et al. (2013) accounted for it. Bonnet reduce the number of species to five (S. thomisoides, (1958) discussed that Sicarius should be regarded as S. fumosus, S. crustosus, S. lanuginosus and the proper generic name, but he argued that the S. yurensis stat. nov., the last being newly recorded name of the species was of lesser importance and thus for the country). he suggested retaining the name terrosus. Following Nicolet described seven species of Thomisoides in him and Simon (1893), catalogues had been listing his seminal work on Chilean spiders (1849) based on S. terrosus as the type of Sicarius (e.g. Roewer, 1942; specimens collected by Claudio Gay. He sorted them Brignoli, 1983; World Spider Catalog, 2015). into two groups, the piriformes (including S. terro- One could argue that S. thomisoides could be con- sus, S. rubripes and S. minoratus) and the rugosas sidered as a nomen oblitum to preserve the stability (including S. fumosus, S. crustosus, S. deformis and of S. terrosus. However, the concept of this species S. lanuginosus). All types that could be found in the has never been clear, and the name S. terrosus has MNHN in Paris (see Ram�ırez, 1989) are immatures, been used to refer to a variety of species – for except for those of S. rubripes and S. nicoleti. Tragi- instance, to S. boliviensis sp. nov. (by Chamberlin, cally, these types (along with that of Thomisoides 1916) and to S. levii sp. nov. (by Mello-Leitao,~ 1941; nicoleti Keyserling) were under study in the Instituto Gerschman de Pikelin & Schiapelli, 1979; Binford Butantan in 2010, when part of its collection was lost et al., 2008, 2009). To our knowledge, the name in a fire, and have not been located after the inci- S. terrosus was very seldom used in the literature to dent. In our view, Nicolet’s piriformes correspond to refer to the correct Chilean species, and thus we see a single species, S. thomisoides. On the other hand, no reason to keep it over the older S. thomisoides. we here recognize three Chilean species which fit in Species included. Sicarius thomisoides Walckenaer, his rugosas group (in which he included four nominal 1847, Sicarius fumosus (Nicolet, 1849), Sicarius species). Basing on Nicolet’s work, it is difficult to crustosus (Nicolet, 1849), Sicarius lanuginosus match his names with the three species we recog- (Nicolet, 1849), Sicarius yurensis Strand, 1908 stat. nize, as the types are immature and the localities nov., Sicarius peruensis (Keyserling, 1880), Sicarius are unspecific (see next paragraph). However, we gracilis (Keyserling, 1880), Sicarius andinus sp. nov., avoided considering his species as doubtful, and pre- Sicarius boliviensis sp. nov., Sicarius rupestris ferred to use somatic morphology to match names to (Holmberg, 1881), Sicarius mapuche sp. nov., Sicarius the species we recognize.

Figure 26. Distribution of Sicarius species. A, Sicarius thomisoides Walckenaer, 1847. B, S. gracilis (Keyserling, 1880) (stars), S. boliviensis sp. nov. (triangles) and S. yurensis Strand, 1908 stat. nov. (circles). C, S. lanuginosus (Nicolet, 1849) (squares), S. fumosus (Nicolet, 1849) (triangles) and S. rupestris (Holmberg, 1881) (circles and stars). Stars show some of the localities where specimens most similar to those from the type locality have been collected. D, S. crustosus (Nicolet, 1849) (triangles), S. levii sp. nov. (circles) and S. mapuche sp. nov. (stars). Xeric formations are highlighted in red (deserts and xeric scrublands), yellow (dry forests) and brown (chaco).

Figure 28. Habitats where Sicarius spiders have been collected. A, xerophilous woodland near Santiago, Chile (S. thomisoides and S. fumosus). B, semi-arid transitional desert in Illapel, Coquimbo, Chile (S. thomisoides). C, coastal desert near Puerto Viejo, Atacama, Chile (S. thomisoides). D, dry forest in Barra Honda National Park, Guanacaste, Costa Rica (S. rugosus), photo by F. Castellucci.

in Valdivia, but nowhere else. Thus, it seems that Thomisoides rubripes Nicolet, 1849: 353, plate 1, fig. Nicolet’s specimens were mislabeled at some point, 10. Female (MNHN 5488) and immature (MNHN and that his localities are unreliable – we cannot say 5491) syntypes from Chile, examined, not found after where they came from, except for the country the Instituto Butantan fire in 2010. SYN. NOV. Figure 27. Distribution of Sicarius species. A, Sicarius saci sp. nov. (white stars), S. jequitinhonha sp. nov. (Chile). Thomisoides minoratus Nicolet, 1849: 354. Two (crosses), S. cariri Magalhaes,~ Brescovit & Santos 2013 (black stars), S. tropicus (Mello-Leitao,~ 1936) (squares), S. orna- immature syntypes from Chile deposited in MNHN tus Magalhaes,~ Brescovit & Santos, 2013 (triangles) and S. diadorim Magalhaes,~ Brescovit & Santos, 2013 (circles). B, 5490, examined, not found after the Instituto Butan- SICARIUS THOMISOIDES WALCKENAER, 1847 S. peruensis (Keyserling, 1880) (circles) and S. andinus sp. nov. (stars). C, S. rugosus (F.O. Pickard-Cambridge, 1899). tan fire in 2010. SYN. NOV. D, S. utriformis (Butler, 1877) (white star indicates type locality). Xeric formations are highlighted in red (deserts and Figures 12A, 15A, B, 17D, 18A, 19A, 26A, 30A, 31B, Thomisoides terrosus Keyserling, 1880: 268, plate 8, xeric scrublands), yellow (dry forests) and brown (chaco). 32A, 33A–C, 34, 35, 36, 37A fig. 148. Sicarius thomisoides Walckenaer, 1847: 379. Thomisoides nicoleti Keyserling, 1880: 270, fig. 149. Description was not based on examination of speci- Male holotype from Chile, deposited in MNHN Nicolet (1849) gave specific localities for only a few found a Sicarius. On the other hand, Sicarius are mens, but rather on the then unpublished illustra- AR5492, examined, not found after the Instituto species (Valdivia for S. terrosus and S. deformis; very common in central and northern Chile – in tions by Nicolet (1849). Butantan fire in 2010. SYN. NOV. Chile for S. rubripes), but labels on the specimens some places, almost every rock has an individual Thomisoides terrosus Nicolet, 1849: 352, plate 1, fig. Sicarius terrosus: Simon, 1893: 271. indicate they all come from Valdivia (see Ram�ırez, lying beneath it. It is known that Gay collected 9. Three immature syntypes from Valdivia, Chile Sicarius nicoleti: Petrunkevitch, 1911: 122. 1989). However, Sicarius seem to be completely throughout most of Chile, including all the places (probably a mislabeling), deposited in MNHN 4159, absent from this region: the closest record of Sicarius where Sicarius are most common (Pizarro, 1944). examined, not found after the Instituto Butantan fire Synonymy: The syntypes of S. terrosus and comes from 400 km to the north, and we and several Having this in mind, to us it seems very unlikely in 2010. SYN. NOV. S. minoratus are immatures, but somatic colleagues collected near Valdivia and have not yet that Gay collected seven different species of Sicarius

Figure 29. Habitats where Sicarius spiders have been collected. A, Canta, Lima, Peru (S. peruensis). B, grassland in Cordoba, Argentina (S. rupestris), photo by G. D. Rubio. C, grassland in Pichi Mahuida, Rıo Negro, Argentina (S. rupestris). D, xeric monte in Paso Cordoba, Rıo Negro, Argentina (S. levii sp. nov.). Figure 30. Male Sicarius, sternum in ventral view. A, S. thomisoides Walckenaer, 1847 from Iquique, Chile (IBSP). B, morphology (namely the shape of the sternum and feature in females to distinguish this species from S. yurensis Strand, 1908 stat. nov. from Yura, Peru (AMNH). C, Sicarius levii sp. nov. from Tucuman, Argentina leg macrosetae) indicates they belong to this species. S. yurensis stat. nov. (AMNH). D, Sicarius crustosus (Nicolet, 1849) from Paposo, Chile (AMNH). Scale bars = 1 mm. The genital morphology of the types of S. nicoleti (Fig. 34C) and S. rubripes (Fig. 36G) falls within the Description: Male (Illapel, Coquimbo, Chile, IBSP Female (Illapel, Coquimbo, Chile, IBSP 165454). Genitalia are variable in both males (Fig. 34) and variation of S. thomisoides (see Variation below). 165453). Coloration: carapace light reddish brown, As in male, except by the following. Total length females (Figs 35, 36). The regions of Coquimbo and The bulb of S. nicoleti’s holotype is expanded from chelicerae, labium and endites reddish brown, 14.85. Carapace length 7.25, anterior width 2.64, Atacama show the highest morphological diversity, the cymbium (Fig. 34C), as can be noted in sternum orange, legs orange, abdomen brownish thoracic region width 7.08. Eye diameters and inter- with males with very long bulbs (Fig. 34F) and Keyserling’s illustration as well (1880, ﬁg. 149). grey. Total length 14.7. Carapace length 7.7, distances: PME 0.18, ALE 0.25, PLE 0.23, PME–PLE aberrant females (Figs 35C, D, 36C, D). Some anterior width 2.52, thoracic region width 7.1. Eye 0.69, ALE–PLE 0.28. Clypeus height 1.12. Sternum females do not have the spermathecae branches Diagnosis: Distinguished from other species, except diameters and interdistances: posterior median eye length 3.63, width 3.18. Femoral thorns: 5. Leg I: bent at the apex. There is a possibility that S. yurensis stat. nov., by the oval, longer-than-wide (PME) 0.2, anterior lateral eye (ALE) 0.25, posterior femur 7.92 long, patella 3.1, tibia 8.3, metatarsus 6, S. thomisoides, as delimited here, represents a sternum (Fig. 30A) (cordiform in other species). It is lateral eye (PLE) 0.23, PME–PLE 0.61, ALE–PLE tarsus 3.3. II: fe 9, ti 8.6. III: fe 8.08, ti 7.5. IV: fe 8, species complex. Its treatment, however, is not easily distinguished from S. yurensis stat. nov. by 0.31. Clypeus height 1.22. Eye tubercles absent. pa 2.75, ti 7.33, mt 5.5, ta 3.08. Abdomen: length trivial, and we prefer to consider it as a single the narrow cephalic region (Fig. 37A). Males are Macrosetae in lateral border of the carapace in a 6.58, width 6.25. Spermathecae branches: 14. Ventral species until further data are available. Subadult further distinguished by the gently curved palpal single row. Femora without prolateral black setae. branches present, one pair, straight (Fig. 35A, B). females have pre-spermathecae with several thin, bulb body, without clear distinction between the bulb Sternum length 3.76, width 3.02, oval. Femoral short, digitiform branches (Fig. 36H). body and the embolus (Fig. 34). Females are further thorns: 5. Leg I: femur 9.6 long, patella 3.7, tibia 10, Variation: Males (N = 5): carapace length 4.72–7.7 distinguished (except from S. yurensis stat. nov.) by metatarsus 8.1, tarsus 4.1. II: fe 11, ti 11.38. III: fe (6.37), femoral thorns 5–6, femur II length 7.42– Natural history: This species occurs in a range of the spermathecae with 8–20 branches usually bent 9.6, ti 9.2. IV: fe 9, pa 2.7, ti 8.7, mt 7.1, ta 3.7. 11.13 (9.92). Females (N = 5): carapace length 5.58– dry habitats in the arid coast of Chile, such as xeric at the apex and without a rounded reservatory Abdomen: length 6.7, width 5.4. Palp as in 8.17 (6.98), femoral thorns 5–6, femur II length woodlands in the Santiago region, semi-arid (Figs 35, 36). We have been unable to ﬁnd a genital Figures 34A and B. 6.75–9.1 (8.2), spermathecae branches 9–19. transition vegetation in the Coquimbo region,

Figure 31. Scanning electron microscopy, metatarsus I, dorsal, distal trichobothria (males, except H, female). A, Sicar- ius fumosus (Nicolet, 1849) (IBSP 163752). B, Sicarius thomisoides Walckenaer, 1847 (IBSP 163744). C, Sicarius peruensis (Keyserling, 1880) (IBSP 162916). D, Sicarius mapuche sp. nov. (CAI 3092). E, Sicarius boliviensis sp. nov. (AMNH, from Marquilla, Bolivia). F, Sicarius levii sp. nov. (AMNH, from Tucuman,� Argentina). G, Sicarius tropicus (Mello-Leitao,~ 1936) (UFMG 11056). H, Hexophthalma spatulata (Pocock, 1900) comb. nov. (AcAT 2013/1514). Scale bars = 0.01 mm, except D (0.05 mm). beaches and the extremely dry deserts in the north Brescovit et al., 12/VII/2012, 1 $ 14 imm. (IBSP of the country. It is usually found beneath boulders, 162896), 1 $ (IBSP 165451); [ 70.31667, 18.48333, À À especially large ones. The eggcases (Fig. 17D) can be 44 m], R. Calderon,� 4/X/1970, 1 # (UCCC); found attached under those boulders. One of those, Camarones, Azapa, Chitita [ 69.66667, 18.8, À À opened in the field, contained 17 eggs; another, that 2681 m], L.E. Pena,~ 10/VI/1986, 1 # 5 $ 42 imm. ecloded in the laboratory, gave rise to 15 juveniles. (AMNH), 1 # 1 imm. (AMNH), 2 # (AMNH); Codpa Dispersing-instar juveniles average 2.2 mg in dry [ 69.73333, 18.81667, 2157 m], M. Codoceor, VII/ À À weight (N = 15). Individuals have been observed in 1955, 1 $ (MHNS); Huara, Caleta Camarones the field feeding on scolopendromorph centipedes, ( 70.2635, 19.19283, 160 m), A.D. Brescovit et al., À À Loxosceles and gnaphosid spiders, solpugids, and on 11/VII/2012, 1 $ 4 imm. (IBSP 162885), 1 $ (IBSP co-especifics. 165476). Region� I de Tarapaca:� Iquique, Huara, Pisagua ( 70.21683, 19.59833), A.D. Brescovit À À Distribution: So far known only from the Chilean dry et al., 11/VII/2012, 5 $ 9 imm. (IBSP 162900), 1 $ coast, from the extreme north of the country to south (IBSP 165490), 1 $ (IBSP 165491); Quebrada de of the Santiago region (Fig. 26A). A record from Tarapaca[� 69.28333, 19.8, 1300 m], L.E. Pena,~ À À Buenos Aires, Argentina (deposited in AMNH), is 25/I/1992, 1 # 3 $ 6 imm. (AMNH); Quebrada certainly a mislabelling. There is no evidence that this Minimi~ ni~ ( 70.006, 19.2035, 798 m), A.D. Figure 32. Scanning electron microscopy, right male palpi, prolateral view. A, Sicarius thomisoides Walckenaer, 1847 À À species occurs in Argentina, and previous records from Brescovit et al., 11/VII/2012, 2 # 1 $ 3 imm. (IBSP (IBSP 163744). B, Sicarius fumosus (Nicolet, 1849) (IBSP 163716) (slightly twisted). C, Sicarius peruensis (Keyserling, this country have been misidentified (see Sicarius 162898); Playa Chanabaya ( 70.13677, 20.88652, 1880) (IBSP 162916). D, Sicarius mapuche sp. nov. (CAI 3092). E, Sicarius boliviensis sp. nov. (AMNH, from San À À levii sp. nov. below). The species probably extends its 73 m), 13/VII/2012, 2 # 1 imm. (IBSP 162893), 1 $ Antonio de Parapeti, Bolivia). F, Sicarius levii sp. nov. (MACN 30239) (left palp, mirrored, slightly twisted). Bd, bulb distribution into southern Peru, but this region is very (IBSP 165446), 1 # (UFMG 15340); Playa Puenta body; Bs, bulb base; E, embolus. Scale bars = 0.1 mm. sparsely sampled and we have not had the chance to Gruesa ( 70.11331, 20.04721, 691 m), 1 $ (IBSP À À examine many Sicarius specimens from there. 162892); Playa Quinteros ( 70.12075, 20.6109, 162880); Punta Patache ( 70.18725, 20.80107, Canchones ( 69.53383, 20.445, 997 m), A.D. À À À À À À 504 m), 2 $ (IBSP 162879); Playa R�ıo Seco 52 m), 1$ 5 imm. (IBSP 162887), 1 # (IBSP 165477), Brescovit et al., 10/VII/2012, 1 $ 15 imm. (IBSP Additional material examined: CHILE. Region� XV ( 70.16417, 21.0023, 514 m), 1 $ 3 imm. (IBSP 1 $ (IBSP 165478); Pica, Altos de Pica [ 68.95, 162899); La Huayca ( 69.54533, 20.43567, 997 m), À À À À À de Arica y Parinacota: Arica, Arica, Playa 162883), 1 # (IBSP 165475); Playa San Marcos 20.36667, 4149 m], M. Ferru, 21/II/2012, 1 $ (IBSP 1 $ 10 imm. (IBSP 162881), 1 $ 10 imm. (IBSP Corazones ( 70.33067, 18.54817, 421 m), A.D. ( 70.12075, 21.10068, 424 m), 1 $ 2 imm. (IBSP 162891);À Pozo Almonte, Centro Experimental 165449); Pampa del Tamarugal, 3 imm. (IBSP À À À À

162910), 1 $ (IBSP 165502); unspecific locality, 1$ 1 Brescovit et al., 16/VII/2012, 1 $ 7 imm. (IBSP imm. (ZMB 7894). Region� II de Antofagasta: 162895), 1 # (IBSP 165483), 1 $ (IBSP 165484), 1 $ Antofagasta, Antofagasta, 57 km N Antofagasta (IBSP 165485), 1 # (IBSP 165486), 1 $ (IBSP [ 70.33333, 23.11667, 435 m], H. Larrain, 5/VI/ 165487), 1 $ (IBSP 165488), 1 # (UFMG 15358), 1 $ À À 1992, 1 # (AMNH); Juan Lopez ( 70.53637, (UFMG 15359); same locality [ 70.4367, 25.26709, À À À 23.50848, 70 m), A.D. Brescovit et al., 14/VII/2012, 252 m], D.A. Masarey, 1910, 1 $ (NHMB 459a); El À 1 imm. (IBSP 162905), 1 $ (IBSP 165494); La Loa, Calama, La Cascada [ 68.91667, 22.45, À À Chimba ( 70.37787, 23.52132, 640 m), 15/VII/ 2321 m], G. Arriagada, 10/III/1984, 1 $ 6 imm. À À 2012, 1 $ 2 imm. (IBSP 162884); La Negra (MHNS 906); X/1982, 1 # 2 imm. (MHNS 596); III/ [ 70.31667, 23.76667, 455 m], H. Larrain, 11/VII/ 1983, 1 $ 3 imm. (MHNS 635); Iquique, Huara, À À 1992, 1 $ 1 imm. (AMNH); Paposo [ 70.45, 25, Quebrada de Tarapaca[� 69.28333, 19.8, 2371 m], À À À À 769 m], L.E. Pena,~ 17–20/IX/1992, 1 $ 2 imm. H. Larrain, 1 # 1 $ 1 imm. (AMNH); Iquique, R�ıo (AMNH); Quebrada La Chimba [ 70.37787, Seco [ 70.11667, 20.25, 432 m], 21/VI/1998, 1 $ À À À 23.52132, 640 m], J.C. Ortiz, 22/VIII/1996, 1 # (AMNH); Tocopilla, Mar�ıa Elena, R�ıo Loa Mouth (UCCC);À Vertedero Municipal ( 70.38565, [ 69.75, 21.43333, 10 m], L.E. Pena,~ 12/II/1992, 3 À À À 23.53375, 126 m), A.D. Brescovit et al., 15/VII/ $ (AMNH); Tocopilla, 20 km N Tocopilla À 2012, 1 # 1 $ 3 imm. (IBSP 162909), 1 # (IBSP [ 70.18333, 22.08333, 263 m], 12/XII/1992, 1 $ 4 165450); Mejillones, Hornitos [ 70.28333, imm.À (AMNH);À Caleta Buena ( 70.2444, 22.28246, À À À 22.91667, 344 m], J.C. Ortiz, 22/VIII/1996, 1 $ 442 m), A.D. Brescovit et al., 14/VII/2012, 4 $ 9 À (UCCC); VIII/1996, 1 $ 1 imm. (UCCC); ( 70.46267, imm. (IBSP 162878), 1 # (IBSP 165467), 1 $ (IBSP À 23.105, 28 m), A.D. Brescovit et al., 14/VII/2012, 1 165468), 1 # (IBSP 165469), 1 # (IBSP 165470), 1 # À $ 14 imm. (IBSP 162894), 1 # (IBSP 165479), 1 $ (IBSP 165471), 1 $ (IBSP 165472), 1 $ (IBSP (IBSP 165480), 1 $ (IBSP 165481), 1 $ (IBSP 165473), 1 # (UFMG 15354), 1 $ (UFMG 15355); 165482), 1 $ (UFMG 15356), 1 $ (UFMG 15367); Caleta Urco Sur ( 70.15312, 21.76992, 285 m), 18/ À À Taltal, 24 km S Paposo [ 70.41667, 25.2, 899 m], VII/2012, 2 imm. (IBSP 162902), 1 # (IBSP 165492); À À N.I. Platnick et al., 11/X/1992, 3 # 2 $ (AMNH), 1 # Playa Consuela ( 70.24829, 22.32645, 524 m), 14/ À À 1 $ (AMNH); same locality [ 70.41667, 25.2, VII/2012, 1 $ 2 imm. (IBSP 162882), 1 $ (IBSP À À 50 m], 3 # 1 $ 1 imm. (AMNH); Caleta Paposo 165474); Playa Punta Arenas ( 70.14537, À ( 70.46602, 24.99828, 45 m), A.D. Brescovit et al., 21.64552), 13/VII/2012, 1 $ 3 imm. (IBSP 162908), À À À 16/VII/2012, 1 $ (IBSP 162904); Cave near road to 1 $ (IBSP 165500); Unspecific province, Camino a Taltal ( 70.44578, 25.34698, 415 m), 17/VII/2012, Paso Jama, 171-178 km, V. Jerez, 11/II/1997, 1 $ À À 1 $ (IBSP 162888); Cifuncho ( 70.64722, 25.65442, (UCCC); Las Vertientes, H. Larrain, 12/VI/1992, 1 $ À À 44 m), 16/VII/2012, 4 imm. (IBSP 162903), 1 $ (AMNH); 1 $ (MHNS). Region� III de Atacama: (IBSP 165493); Paposo [ 70.45, 25, 769 m], J. Chanaral,~ Chanaral,~ Cachinal de la Costa À À Stuardo, 23/VII/1964, 1 # (UCCC); M. Elgueta, 8– [ 70.63333, 26.05, 228 m], J. Stuardo, 13/VII/1964, À À 10/X/1983, 1 # 2 $ (MHNS 767); M. Ferru,� II/2012, 1 # (UCCC); Parque Nacional Pan de Azucar� 1 imm. (IBSP 162907), 1 # (IBSP 165499); Paposo, ( 70.6618, 26.14267, 120 m), A.D. Brescovit et al., À À La Rinconada, G. Arriagada, 25/X/1985, 1 $ 16/VII/2012, 3 imm. (IBSP 162911), 1 $ (IBSP (MHNS); Quebrada del Medano� [ 69.75, 25.28333, 165503), 1 $ (IBSP 165504); same locality À À 2207 m], H. Niemeyer, VII/1985, 1 $ (MHNS 900); ( 70.62823, 26.30238, 242 m), 17/VII/2012, 1 $ 3 À À Reserva Nacional Paposo ( 70.46602, 24.96367, imm. (IBSP 162886), 1 $ (UFMG 15343); Portofino À À 45 m), A.D. Brescovit et al., 16/VII/2012, 1 $ 4 imm. ( 70.70087, 26.52123, 101 m), A.D. Brescovit À À (IBSP 162906), 1 # (IBSP 165495), 1 $ (IBSP et al., 10/XI/2013, 1 # (IBSP 165564), 1 $ (IBSP 165496), 1 $ (IBSP 165497), 1 $ (IBSP 165498), 1 $ 165565), 1 imm. (IBSP 165566); ( 70.63795, À (UFMG 15360); Taltal [ 69.75, 25.28333, 2207 m], 26.35577, 88 m), A.D. Brescovit et al., 17/VII/2012, À À À 1 # (AMNH); ( 70.4367, 25.26709, 252 m), A.D. 8 $ 9 imm. (IBSP 162897), 1 # (IBSP 165489), 1 # À À

Figure 33. Scanning electron microscopy, female genitalia, digested, ventral view. A–C, Sicarius thomisoides Walcke- naer, 1847 (IBSP 163734). A, general view. B, close up, left ventral branch. C, close up, left branches. D–E Sicarius fumosus (Nicolet, 1849) (IBSP 163716). D, general view. E, close up, left branches. F–H, Sicarius peruensis (Keyserling, 1880) (IBSP 162916). F, close up, right ventral branch (note twisted shape). G, general view. H, close up, right pore plate. I, Sicarius mapuche sp. nov. (CAI 3072). J, Sicarius boliviensis sp. nov. (AMNH, from Marquilla, Bolivia). K, Sicarius levii sp. nov. (MACN 30239). Inset showing detail of median branches. L, Sicarius crustosus (Nicolet, 1849) (MACN 30189). PP, pore plate; UE, uterus externus; VB, ventral branch. Scale bars = 0.1 mm (except B, F, H, 0.05 mm).

162910), 1 $ (IBSP 165502); unspecific locality, 1$ 1 Brescovit et al., 16/VII/2012, 1 $ 7 imm. (IBSP imm. (ZMB 7894). Region� II de Antofagasta: 162895), 1 # (IBSP 165483), 1 $ (IBSP 165484), 1 $ Antofagasta, Antofagasta, 57 km N Antofagasta (IBSP 165485), 1 # (IBSP 165486), 1 $ (IBSP [ 70.33333, 23.11667, 435 m], H. Larrain, 5/VI/ 165487), 1 $ (IBSP 165488), 1 # (UFMG 15358), 1 $ À À 1992, 1 # (AMNH); Juan Lopez ( 70.53637, (UFMG 15359); same locality [ 70.4367, 25.26709, À À À 23.50848, 70 m), A.D. Brescovit et al., 14/VII/2012, 252 m], D.A. Masarey, 1910, 1 $ (NHMB 459a); El À 1 imm. (IBSP 162905), 1 $ (IBSP 165494); La Loa, Calama, La Cascada [ 68.91667, 22.45, À À Chimba ( 70.37787, 23.52132, 640 m), 15/VII/ 2321 m], G. Arriagada, 10/III/1984, 1 $ 6 imm. À À 2012, 1 $ 2 imm. (IBSP 162884); La Negra (MHNS 906); X/1982, 1 # 2 imm. (MHNS 596); III/ [ 70.31667, 23.76667, 455 m], H. Larrain, 11/VII/ 1983, 1 $ 3 imm. (MHNS 635); Iquique, Huara, À À 1992, 1 $ 1 imm. (AMNH); Paposo [ 70.45, 25, Quebrada de Tarapaca[� 69.28333, 19.8, 2371 m], À À À À 769 m], L.E. Pena,~ 17–20/IX/1992, 1 $ 2 imm. H. Larrain, 1 # 1 $ 1 imm. (AMNH); Iquique, R�ıo (AMNH); Quebrada La Chimba [ 70.37787, Seco [ 70.11667, 20.25, 432 m], 21/VI/1998, 1 $ À À À 23.52132, 640 m], J.C. Ortiz, 22/VIII/1996, 1 # (AMNH); Tocopilla, Mar�ıa Elena, R�ıo Loa Mouth À(UCCC); Vertedero Municipal ( 70.38565, [ 69.75, 21.43333, 10 m], L.E. Pena,~ 12/II/1992, 3 À À À 23.53375, 126 m), A.D. Brescovit et al., 15/VII/ $ (AMNH); Tocopilla, 20 km N Tocopilla À 2012, 1 # 1 $ 3 imm. (IBSP 162909), 1 # (IBSP [ 70.18333, 22.08333, 263 m], 12/XII/1992, 1 $ 4 165450); Mejillones, Hornitos [ 70.28333, imm.À (AMNH);À Caleta Buena ( 70.2444, 22.28246, À À À 22.91667, 344 m], J.C. Ortiz, 22/VIII/1996, 1 $ 442 m), A.D. Brescovit et al., 14/VII/2012, 4 $ 9 À (UCCC); VIII/1996, 1 $ 1 imm. (UCCC); ( 70.46267, imm. (IBSP 162878), 1 # (IBSP 165467), 1 $ (IBSP À 23.105, 28 m), A.D. Brescovit et al., 14/VII/2012, 1 165468), 1 # (IBSP 165469), 1 # (IBSP 165470), 1 # À $ 14 imm. (IBSP 162894), 1 # (IBSP 165479), 1 $ (IBSP 165471), 1 $ (IBSP 165472), 1 $ (IBSP (IBSP 165480), 1 $ (IBSP 165481), 1 $ (IBSP 165473), 1 # (UFMG 15354), 1 $ (UFMG 15355); 165482), 1 $ (UFMG 15356), 1 $ (UFMG 15367); Caleta Urco Sur ( 70.15312, 21.76992, 285 m), 18/ À À Taltal, 24 km S Paposo [ 70.41667, 25.2, 899 m], VII/2012, 2 imm. (IBSP 162902), 1 # (IBSP 165492); À À N.I. Platnick et al., 11/X/1992, 3 # 2 $ (AMNH), 1 # Playa Consuela ( 70.24829, 22.32645, 524 m), 14/ À À 1 $ (AMNH); same locality [ 70.41667, 25.2, VII/2012, 1 $ 2 imm. (IBSP 162882), 1 $ (IBSP À À 50 m], 3 # 1 $ 1 imm. (AMNH); Caleta Paposo 165474); Playa Punta Arenas ( 70.14537, À ( 70.46602, 24.99828, 45 m), A.D. Brescovit et al., 21.64552), 13/VII/2012, 1 $ 3 imm. (IBSP 162908), À À À 16/VII/2012, 1 $ (IBSP 162904); Cave near road to 1 $ (IBSP 165500); Unspecific province, Camino a Taltal ( 70.44578, 25.34698, 415 m), 17/VII/2012, Paso Jama, 171-178 km, V. Jerez, 11/II/1997, 1 $ À À 1 $ (IBSP 162888); Cifuncho ( 70.64722, 25.65442, (UCCC); Las Vertientes, H. Larrain, 12/VI/1992, 1 $ À À 44 m), 16/VII/2012, 4 imm. (IBSP 162903), 1 $ (AMNH); 1 $ (MHNS). Region� III de Atacama: (IBSP 165493); Paposo [ 70.45, 25, 769 m], J. Chanaral,~ Chanaral,~ Cachinal de la Costa À À Stuardo, 23/VII/1964, 1 # (UCCC); M. Elgueta, 8– [ 70.63333, 26.05, 228 m], J. Stuardo, 13/VII/1964, À À 10/X/1983, 1 # 2 $ (MHNS 767); M. Ferru,� II/2012, 1 # (UCCC); Parque Nacional Pan de Azucar� 1 imm. (IBSP 162907), 1 # (IBSP 165499); Paposo, ( 70.6618, 26.14267, 120 m), A.D. Brescovit et al., À À La Rinconada, G. Arriagada, 25/X/1985, 1 $ 16/VII/2012, 3 imm. (IBSP 162911), 1 $ (IBSP (MHNS); Quebrada del Medano� [ 69.75, 25.28333, 165503), 1 $ (IBSP 165504); same locality À À 2207 m], H. Niemeyer, VII/1985, 1 $ (MHNS 900); ( 70.62823, 26.30238, 242 m), 17/VII/2012, 1 $ 3 À À Reserva Nacional Paposo ( 70.46602, 24.96367, imm. (IBSP 162886), 1 $ (UFMG 15343); Portofino À À 45 m), A.D. Brescovit et al., 16/VII/2012, 1 $ 4 imm. ( 70.70087, 26.52123, 101 m), A.D. Brescovit À À (IBSP 162906), 1 # (IBSP 165495), 1 $ (IBSP et al., 10/XI/2013, 1 # (IBSP 165564), 1 $ (IBSP 165496), 1 $ (IBSP 165497), 1 $ (IBSP 165498), 1 $ 165565), 1 imm. (IBSP 165566); ( 70.63795, À (UFMG 15360); Taltal [ 69.75, 25.28333, 2207 m], 26.35577, 88 m), A.D. Brescovit et al., 17/VII/2012, À À À 1 # (AMNH); ( 70.4367, 25.26709, 252 m), A.D. 8 $ 9 imm. (IBSP 162897), 1 # (IBSP 165489), 1 # À À

Figure 35. Sicarius thomisoides Walckenaer, 1847 from Chile, female genitalia, digested. A, Illapel, Coquimbo (IBSP 165454), dorsal. B, same, ventral. C, Parque Nacional Pan de Azucar,� Atacama (UFMG 15343), dorsal. D, same, ventral. E, Iquique, Tarapaca� (IBSP 165446), dorsal. F, same, ventral. G, Machal�ı, O’Higgins (IBSP 163708), dorsal. H, same, ventral. Abbreviation: VB, ventral branch. Scale bars = 0.1 mm.

(UFMG 15338), 1 $ (UFMG 15339); Diego de (AMNH); Calderilla ( 70.85217, 27.0936, 12 m), À À Almagro, Inca de Oro [ 69.93333, 26.81667, A.D. Brescovit et al., 10/XI/2013, 1 $ (IBSP 165546), À À 1954 m], L.E. Pena,~ 8/X/1980, 3 $ 3 imm. (AMNH); 1 $ (IBSP 165547); Caleta Los Patos ( 70.80013, À Flamenco coastal dunes [ 70.6, 26.61667, 10 m], 26.95912, 11 m), 1 imm. (IBSP 165545), 1 # Figure 34. Sicarius thomisoides Walckenaer, 1847 from Chile, male genitalia in prolateral view. A, Illapel, Coquimbo À À À N.I. Platnick et al., 10/X/1992, 1 (AMNH); 13/X/ (UFMG 15325), 1 (UFMG 15326); E of Puerto (IBSP 165453). B, same, cymbium and bulb. C, holotype of Thomisoides nicoleti Keyserling (bulb expanded and slightly $ $ 1992, 1 $ (AMNH); N of Chanaral~ [ 70.6, Viejo [ 70.78333, 27.35, 152 m], L.E. Pena,~ 9–10/ tilted laterally) (MNHN). D, Chanaral,~ Atacama (UFMG 15338). E, Quebrada Minimi~ ni,~ Tarapaca� (IBSP 162898). F, À À À 26.36667, 204 m], L.E. Pena,~ 10/X/1992, 1 $ X/1980, 1 # 1 imm. (AMNH); Puerto Viejo Caldera, Atacama (AMNH). G, La Laguna, Valpara�ıso (IBSP 163730). H, Iquique, Tarapaca� (UFMG 15340). Scale À (AMNH); Copiapo, Caldera, 25 km SE Caldera ( 70.93197, 27.33708, 63 m), A.D. Brescovit et al., bars = 0.1 mm (except A, 1 mm). À À [ 70.63333, 27.25, 591 m], 27–28/X/1983, 1 # 1 $ 10/XI/2013, 1 # (IBSP 165542), 1 imm. (IBSP À À

Figure 37. Female carapace in dorsal view. A, Sicarius thomisoides Walckenaer, 1847 from Caldera, Atacama, Chile (AMNH). B, Sicarius yurensis Strand, 1908 stat. nov., from Las Yaras, Yura, Peru (AMNH). Scale bar, 1 mm.

27.73333, 700 m], 23–25/X/1983, 2 # 3 imm. 165579), 1 $ (IBSP 165580), 1 $ (IBSP 165581); Los À(AMNH); 56 km N Vallenar [ 70.58333, 28.06667, Pozos ( 71.15447, 28.16075, 198 m), C.J. À À À À 440 m], N.I. Platnick et al., 9/X/1992, 1 $ (AMNH); Grismado et al., 18/VIII/2009, 1 $ 1 imm. (MACN-Ar 60 km S Copiapo[� 70.33333, 27.88333, 1449 m], 32333); Quebrada Talinay [ 71.21667, 28.46667, À À À À L.E. Pena,~ 22/X/1991, 1 $ (AMNH); [ 70.31667, 110 m], L.E. Pena,~ 13/X/1980, 1 $ 2 imm. (AMNH); À 27.36667, 589 m], H. Niemeyer, III/1986, 2 $ 20/III/1972, 1 # (UCCC); Vallenar, 8 km N Vallenar À (MHNS); J. Moreno, 11/X/1982, 1 # 3 $ 4 imm. [ 70.75, 28.5, 568 m], L.E. Pena,~ 6/X/1990, 1 $ À À (AMNH); 20/X/1982, 2 $ 3 imm. (AMNH); Elqui, (AMNH); Domeyko, road to Carrizalillo ( 70.98895, Coquimbo, Tongoy [ 71.26667, 28.4], L.E. Pena,~ 28.97858, 710 m), A.D. Brescovit et al., 11/XI/2013,À À À À 15/X/1990, 1 $ (AMNH); Huasco, Freirina, 3.5 km E 1 # (UFMG 15334), 1 $ (UFMG 15335); S of Carrizalillo ( 71.44536, 29.10379, 66 m), M.J. Vallenar, km 630 [ 70.75, 28.56667, 549 m], L.E. À À À À Ram�ırez et al., 30/X/2011, 1 # 2 $ 6 imm. (MACN- Pena,~ 1/X/1980, 1 $ 5 imm. (AMNH); Unspeciﬁc Ar 32342); 5.5 km N Carrizalillo ( 71.40012, province, 1 km E Domeyko [800 m], N.I. Platnick À 29.05364, 157 m), 1 # (MACN-Ar 32339); Caleta et al., 8/X/1992, 1 # 2 $ 1 imm. (AMNH); Juntas, À Chanaral~ de Aceituno ( 71.48782, 29.0788, 41 m), R�ıo Huasco [1600 m], L.E. Pena,~ 3/X/1980, 1 # À À A.D. Brescovit et al., 11/XI/2013, 1 imm. (IBSP (AMNH); N of Cuesta Pajonales, 8/VIII/1982, 1 $ 4 165551), 1 # (UFMG 15327), 1 $ (UFMG 15328); imm. (AMNH); X/1980, 2 # 1 $ (AMNH); 21–23/X/ same locality ( 71.48759, 29.07202, 41 m), M.J. 1983, 1 # 3 imm. (AMNH). Region� IV de À À Ram�ırez et al., 30/X/2011, 1 # (MACN-Ar 32335); Coquimbo: Choapa, Illapel, Cuesta El Espino Carrizalillo ( 71.40623, 29.10455, 152 m), A.D. ( 71.305, 31.75668, 629 m), A.D. Brescovit & À À À À Figure 36. Sicarius thomisoides Walckenaer, 1847 from Chile, female genitalia, digested. A, Antofagasta, Antofagasta Brescovit et al., 11/XI/2013, 1 # (IBSP 165561), 1 $ I.L.F. Magalhaes,~ 8/IV/2013, 2 $ 7 imm. (IBSP (IBSP 162909), dorsal. B, same, ventral. C, Pan de Azucar,� Coquimbo (IBSP 165589), dorsal. D, same, ventral. E, (IBSP 165562), 1 $ (IBSP 165563); Sauce Perez 163724); Cuesta Los Cristales ( 71.1388, 31.70447, Chanaral,~ Atacama (UFMG 15339), dorsal. F, same, ventral. G, syntype of Thomisoides rubripes Nicolet, 1849 (MNHN), À À [ 71.11667, 28.63333, 853 m], L.E. Pena,~ 26/X/ 659 m), 1 $ 8 imm. (IBSP 163744), 1 # (IBSP dorsal. H, subadult from Chanaral,~ Atacama (IBSP 162897), ventral. Abbreviation: VB, ventral branch. Scale À À 1980, 1 $ 8 imm. (AMNH); Huasco, Agua de Luna 165453), 1 $ (IBSP 165454), 1 # (UFMG 15350), 1 $ bars = 0.1 mm. ( 71.16181, 28.33556, 107 m), C.J. Grismado (UFMG 15351); Cuz-Cuz ( 71.29342, 31.67, À À À À et al., 18/VIII/2009, 1 $ 2 imm. (MACN-Ar 32336); 252 m), 1 # 7 $ 8 imm. (IBSP 163734), 1 # (IBSP 165548); same locality [ 70.93333, 27.33333, 140 m], N.I. Platnick et al., 10/X/1992, 1 $ (AMNH); Caleta Angosta ( 71.14792, 28.22308, 97 m), 1 # 2 165509), 1 # (IBSP 165510); Salamanca, El Tambo À À À À 14 m], L.E. Pena,~ 15–16/X/1992, 1 # 2 imm. Rodillo [ 70.83333, 27.08333, 26 m], 1989–1991, $ 6 imm. (MACN-Ar 32334); Carrizal Bajo [ 71.15, ( 71.0685, 31.77175, 453 m), 5 $ 5 imm. (IBSP À À À À À (AMNH), 1 # 2 imm. (AMNH); Quebrada de Leon 67 # (LEULS); [ 70.8, 27.06667, 26 m], J. Moreno, 28.08333, 39 m], J. Cepeda, X/2000, 2 # 1 $ 3 163747); Los Vilos, Caleta Chigualoco ( 71.51118, À À À À [ 70.66667, 26.98333, 799 m], 8–9/X/1990, 2 $ 8 13/X/1982, 1 $ (AMNH); 20/VII/1983, 1 # 3 $ 3 imm. imm. (LEULS); Freirina ( 71.16243, 28.48288, 31.75522, 20 m), 7/IV/2013, 3 imm. (IBSP 163757); À À À À À imm. (AMNH); same locality [ 70.66667, 26.98333, (AMNH); Copiapo,� 40 km S Copiapo[� 70.31667, 118 m), A.D. Brescovit et al., 9/XI/2013, 1 # (IBSP Near Puerto Punta Chungo ( 71.49122, 31.8767, À À À À À

79 m), 8/IV/2013, 13 $ 7 imm. (IBSP 163731), 1 # 15333); Chungungo, Playa Blanca [ 71.30283, Platnick et al., 9/XI/1993, 1 # (AMNH); Caleta El ( 70.93057, 32.49723, 415 m), A.D. Brescovit & À À À (IBSP 165507), 1 # (IBSP 165508), 1 # (UFMG 29.4446, 36 m], J. Pizarro-Araya, 27/II/2005, 2 # 1 Limar�ı[ 71.68333, 30.71667, 148 m], D. Valdivia, I.L.F. Magalhaes,~ 6/IV/2013, 1 $ 8 imm. (IBSP À À À 15361); [ 71.5, 31.91667, 189 m], J. Pizarro-Araya, $ 1 imm. (LEULS); El Tofo ( 71.25, 29.43333, 21/IX/2004, 1 $ 1 imm. (LEULS); Caleta La Cebada 163718); San Felipe, Putaendo ( 70.71002, À À À À À 28/X/2009, 1 $ (LEULS); Mincha, Caleta Oscuro 465 m), IX/1916, 1 # 1 imm. (AMNH); El Totoralillo ( 71.64278, 30.97275, 53 m), A.D. Brescovit & 32.56667, 1017 m), 1 $ 10 imm. (IBSP 163723), 1 À À À [ 71.6, 31.41667, 38 m], Mojano, XI/1974, 1 # ( 71.2615, 29.59264, 154 m), M. Izquierdo et al., I.L.F. Magalhaes,~ 14/XI/2013, 1 # (IBSP 165553), 1 $ (IBSP 165505), 1 $ (IBSP 165506); ( 70.73738, À À À À À (UCCC); R.C. Digue-Loja, 2/XI/1974, 1 $ (UCCC); 12/VII/2009, 1 $ 18 imm. (MACN-Ar 32337); Isla $ (IBSP 165554), 1 # (UFMG 15329); Camino a 32.706, 739 m), 4 imm. (IBSP 163737); Valparaıso, À Canela Baja, Los Pozos ( 71.20947, 31.30627, Choros [ 71.5394, 29.25539], J. Pizarro-Araya, X/ Fray Jorge ( 71.65365, 30.59622, 420 m), 12/XI/ Puchuncav�ı, La Laguna ( 71.424, 32.64933, 30 m), À À À À À À À À 912 m), A.D. Brescovit et al., 7/XI/2013, 1 $ (IBSP 2012, 2 $ 5 imm. (LEULS); L.E. Pena,~ 7/VIII/1980, 2013, 1 imm. (IBSP 165557), 1 # (UFMG 15330), 1 9/IV/2013, 2 # 8 $ (IBSP 163735), 1 # 6 $ 9 imm. 165558), 1 $ (IBSP 165559), 1 $ (IBSP 165560); 3 $ 1 imm. (AMNH); Llano de la Higuera $ (UFMG 15331); Huampulla [ 71.2, 30.6, 272 m], (IBSP 163736); La Laguna, road to Catapilco, km 16 À À Ermita Padre Hurtado [ 71.55, 31.46667, 256 m], [ 71.26667, 29.5, 645 m], 6/X/1990, 1 $ 1 imm. L.E. Pena,~ 30/IX/1993, 1 $ 11 imm. (AMNH); ( 71.3974, 32.6155, 105 m), 2 # 5 $ 5 imm. (IBSP À À À À À À A. Levican, 24/IX/2004, 1 # 1 imm. (LEULS); (AMNH); 17/X/1983, 1 $ 3 imm. (AMNH); Los Huilmo, 11 km SSW Ovalle, 4 km N Los Cristales 163730), 2 # 5 $ 1 imm. (IBSP 163732), 1 # 4 $ 4 Huentelauquen� [ 71.53333, 31.58333, 127 m], L.E. Choros [ 71.46047, 29.24025, 40 m], A. Gonzalez, [ 71.13333, 30.76667, 858 m], 22–25/IX/1982, 1 $ imm. (IBSP 163746), 1 # (IBSP 165455), 1 # (IBSP À À À À À À Pena,~ 26/IX/1980, 10 $ 10 imm. (AMNH); 3/IX/2007, 1 # 3 $ 3 imm. (LEULS);, 2007, 1 $ 9 5 imm. (AMNH); Lorenzo Peralta ( 71.58207, 165513), 1 # (IBSP 165514). Region� À Salamanca, Entre San Agust�ın y Zapallar imm. (LEULS); J. Pizarro-Araya, 27/V/2006, 3 $ 4 30.5741, 230 m), A.D. Brescovit & I.L.F. Metropolitana: Cachapoal, Mostazal, Huelquen� À ( 70.79339, 31.70568, 1430 m), M.J. Ram�ırez imm. (LEULS); R. Villalon,� 4/VI/2005, 1 # (LEULS); Magalhaes,~ 12/XI/2013, 1 # (IBSP 165585), 1 $ ( 70.67338, 33.8749, 385 m), A.D. Brescovit & À À À À et al., 1/XI/2011, 1 # 1 $ 22 imm. (MACN-Ar 32341), Punta Choros ( 71.46028, 29.249, 40 m), A.D. (IBSP 165586), 1 imm. (IBSP 165587); Los I.L.F. Magalhaes,~ 25/III/2013, 6 imm. (IBSP À À 1 # 1 $ (MACN-Ar 32343); ( 70.98692, 31.79583, Brescovit et al., 11/XI/2013, 1 # (IBSP 165590), 1 # Maitenes, Corral El Julio [ 71.68052, 30.35201], 163726); Chacabuco, La Dormida, Til-Til À À À À 771 m), A.D. Brescovit & I.L.F. Magalhaes,~ 8/IV/ (UFMG 15336), 1 $ (UFMG 15337); same locality S. Zunino, 6/VI/1975, 1 $ 4 imm. (UCCC); Los [ 70.91667, 33.08333, 2000 m], L. Irarrazaval, 16– À À 2013, 2 imm. (IBSP 163760); same locality [ 71.46028, 29.249, 40 m], V. Rodriguez, 4/VI/ Maitenes, N of Amolantas [ 70.91667, 30.36667, 18/XI/1984, 1 # (AMNH); Cordillera, Pirque, Cerro À À À À [ 70.96667, 31.78333, 563 m], J. Aros, 1/V/1961, 1 2005, 1 $ (LEULS); Quebrada Los Choros [ 71.066, 1384 m], L.E. Pena,~ 27/IX/1980, 1 # 1 $ 3 imm. Chena [ 70.75, 33.58333, 529 m], G. Arriagada, À À À À À $ 10 imm. (AMNH); El Temblador [ 71.31667, 29.35, 476 m], 1 # (LEULS); Totoralillo (AMNH); Socos [ 71.49294, 30.72781, 125 m], L. VIII/1985, 1 $ (MHNS 922); San Jose de Maipo, À À À À 29.46667, 83 m], 1/V/2004, 2 $ 2 imm. (LEULS); [ 71.31667, 29.48333, 83 m], D. Valdivia, 1/IX/ Espinoza, 27/IX/2003, 2 $ 5 imm. (LEULS); same Camino al Embalse de Yeso ( 70.20333, 33.79117, À À À À À Elqui, Coquimbo, 16 km S Cruz Grande [ 71.41667, 2006, 1 # (LEULS); La Serena, 19 km N La Serena locality [ 71.48333, 30.7, 151 m], T. Cekalovic, 8/ 1965 m), A.D. Brescovit et al., 22/III/2013, 2 $ 12 À À À 30.35, 140 m], N.I. Platnick et al., 7/X/1992, 2 $ 1 [ 71.33333, 29.76667, 182 m], N.I. Platnick & R.T. IX/1968, 1 $ 2 imm. (UCCC); same locality imm. (IBSP 163749), 1 # (IBSP 165517), 1 # (IBSP À À À imm. (AMNH); Andacollo [ 71.2804, 30.2355, Schuh, 4/XI/1981, 1 $ (AMNH); 20 km N La Serena, ( 71.49294, 30.72781, 125 m), 20/VII/2009, 1 # 2 $ 165518); Camino Central Hidreletrica Los Maitenes À À À À 260 m], J. Pizarro-Araya, VI–VII/2007, 1 # 3 $ 25 Rt. 5, km 491 [ 71.33333, 29.76667, 120 m], N.I. 8 imm. (MACN-Ar 32338); Petorca, La Ligua, ( 70.34133, 33.5705, 1329 m), 1 $ 4 imm. (IBSP À À À À imm. (LEULS); P. Augusto, 14/IX/2003, 1 $ 1 imm. Platnick et al., 6/X/1992, 1 # (AMNH); same locality Pichidangu�ı, Valle de Quillimar�ı(71.50652, 163710); El Canelo [ 70.38333, 33.61667, 1243 m], À À À (LEULS); Horcon� [ 71.215, 29.9243, 162 m], J. ( 71.33333, 29.76667, 120 m), 8/II/1994, 1 $ 32.10803, 25 m), A.D. Brescovit & I.L.F. L.E. Pena,~ XII/1984, 2 $ 10 imm. (AMNH); À À À À À Pizarro-Araya, 14/VIII/2004, 1 # 1 imm. (LEULS); (AMNH); Cuesta Porotitos [ 71.29217, 29.79447, Magalhaes,~ 7/IV/2013, 4 $ 5 imm. (IBSP 163728); Farellones, Plazoleta Los Pumas ( 70.29033, À À À La Serena, Cerro Grande [ 71.21667, 29.91667, 48 m], 5/VII/2004, 1 $ (LEULS); Marquesa Playa La Ballena ( 71.46095, 32.29212, 169 m), 3 33.35983, 2821 m), A.D. Brescovit et al., 21/III/ À À À À À 162 m], L. Espinosa, 16/VIII/2003, 1 # 2 imm. ( 70.95792, 29.94158, 626 m), 10/I/2006, 1 $ 2 $ 7 imm. (IBSP 163750); Unspeciﬁc province, El 2013, 1 # 1 imm. (IBSP 163727); Farellones, Puente À À (LEULS); Las Cardas [ 71.2613, 30.291, 360 m], imm. (LEULS); Monte Grande [ 71.2513, 29.881, Encanto, 9/I/1972, 1 # 1 $ (UCCC). Region� V de Nilhue~ [ 70.3, 33.35, 2207 m], M. Elgueta, 1982, 1 À À À À À À 13/XII/2003, 1 $ 2 imm. (LEULS); M. Moreno, I/ 64 m], L. Espinoza & J. Pizarro-Araya, 13/VIII/2004, Valpara�ıso: Los Andes, Calle Larga [ 70.58333, $ 1 imm. (MHNS 600); Penanolen~ ( 70.51667, À À 1979, 1 # (LEULS); Pan de Azucar� ( 71.2321, 3 $ 2 imm. (LEULS); Pelicana [ 71.0087, 29.9871, 32.83333, 878 m], H. Heatwole, 30/IX/1965, 1 # 33.48333, 892 m), C. Veloso, IV/2013, 4 imm. À À À À À 30.12235, 174 m), A.D. Brescovit et al., 8/XI/2013, 414 m], J. Pizarro-Araya, 1/XI/2006, 1 # (LEULS); (AMNH); Los Andes, Guarda Vieja ( 70.2737, (IBSP 163751); road to Farellones ( 70.36967, À À À 1 $ (IBSP 165588), 1 imm. (IBSP 165589); Ruta 5, Punta de Teatinos [ 71.28333, 29.81667, 48 m], L. 32.90292, 2094 m), A.D. Brescovit & I.L.F. 33.35483, 1738 m), A.D. Brescovit et al., 21/III/ À À À À km. 453 ( 71.37845, 30.02547, 212 m), A.D. Espinoza, 6/IX/2003, 1 $ 2 imm. (LEULS); R�ıo Magalhaes,~ 6/IV/2013, 1 # (IBSP 165515); 2013, 1 $ (IBSP 163740); Maipo, Calera de Tango, À À Brescovit & I.L.F. Magalhaes,~ 12/XI/2013, 1 # (IBSP Hurtado, El Panque, 20 km S Vicuna [ 70.68333, Hidroeletrica� Guarda Vieja, 1 $ 11 imm. (IBSP Quebrada de la Plata [ 70.88333, 33.5, 612 m], C. À À À 165573), 1 $ (IBSP 165574); Tongoy [ 71.48333, 30.21667, 1500 m], N.I. Platnick et al., 4/X/1992, 1 163748); same locality ( 70.5626, 32.83133, Veloso, 24/X/2010, 1 $ (IBSP 161103); San Felipe de À À À À 30.25, 31 m], 21/IX/1996, 1 # 2 $ 4 imm. (AMNH); # 1 $ 6 imm. (AMNH); Morrillos [ 70.7092, 878 m), 1 imm. (IBSP 163754); same locality Aconcagua, Llaillay, Puente Pedro Bustus M. À À La Higuera, 6 km E El Tofo [ 71.16667, 29.43333, 30.2894, 1507 m], 24–31/X/2008, 1 # (LEULS); ( 70.2737, 32.90292, 2094 m), 1 $ (IBSP 165516); ( 70.84783, 32.86867, 757 m), A.D. Brescovit À À À À À À À 764 m], N.I. Platnick & O.F. Francke, 7/I/1985, 1 $ Vicuna,~ Bahia Mausa [ 70.7, 29.71667, 1866 m], R�ıo Blanco ( 70.45452, 32.8555, 1226 m), 12 imm. et al., 23/III/2013, 1 imm. (IBSP 163705), 5 imm. À À À À 1 imm. (AMNH); 6 km S Cruz Grande [ 71.31667, L.E. Pena,~ 2–3/X/1993, 2 $ (AMNH); Diaguitas (IBSP 163706); Petorca, La Ligua, 7 km E La Ligua (IBSP 163753); Santiago, Santiago, Cuesta Lo Prado À 29.48333, 20 m], N.I. Platnick et al., 6/X/1992, 1 $ ( 70.62603, 30.00844, 1005 m), O. Alcagaya, 18/ [ 71.3, 32.46667, 185 m], N.I. Platnick & O.F. ( 70.8986, 33.45818, 562 m), 24/III/2013, 6 $ 14 À À À À À À À (AMNH); same locality ( 71.31667, 29.48333, IX/2006, 1 # (LEULS); Mayacun� [ 70.7, 29.71667, Francke, 9/I/1985, 1 # (AMNH); Pichicuy, Quebrada imm. (IBSP 163745), 1 # (IBSP 165511), 1 $ (IBSP À À À À 5 m), 11/XI/1993, 2 # 3 $ 1 imm. (AMNH); 79 km N 1866 m], R. Donoso?, 1/V/1961, 1 $ 6 imm. (AMNH); Huaquen [ 71.46667, 32.33333, 10 m], N.I. 165512); Quilicura [ 70.71667, 33.35, 489 m], L.E. À À À À La Serena, Rt. 5, km 553 [ 71.31667, 29.35, Huasco, Huasco, Llanos de Challe [ 71.06667, Platnick et al., 2/X/1992, 1 $ (AMNH); ( 71.32983, Pena,~ 25/V/1979, 1 $ 2 imm. (AMNH); V/1979, 1 # 3 À À À À 300 m], 15/X/1992, 1 # (AMNH); Caleta Hornos 28.21667, 291 m], 19–21/XII/1997, 1 # 2 imm. 32.44067, 53 m), A.D. Brescovit & I.L.F. imm. (AMNH); VIII–IX/1979, 1 # 3 imm. (AMNH); À À ( 71.28072, 29.62743, 449 m), A.D. Brescovit (LEULS); Limarı, Combarbala,� Illapel, Cuesta El Magalhaes,~ 7/IV/2013, 1 $ 4 imm. (IBSP 163725); Renca [ 70.75, 33.38333, 514 m], 11/X/1984, 1 # À À À À et al., 8/XI/2013, 1 # (IBSP 165582), 1 $ (IBSP Espino [ 71.06667, 31.31667, 2200 m], L.E. Pena,~ Zapallar, Papudo ( 71.46545, 32.50513, 76 m), 9/ (AMNH); Valle Lo Campino ( 70.72447, 33.38665, À À À À À À 165583); Choros Bajos [ 71.3, 29.28333, 300 m], 1/IX/1984, 4 $ 2 imm. (AMNH), 1 # 3 $ 3 imm. IV/2013, 1 # 3 $ 10 imm. (IBSP 163733); Quillota, 514 m), A.D. Brescovit et al., 24/III/2013, 3 imm. À À L.E. Pena,~ 10–12/XI/1981, 1 $ 2 imm. (AMNH); (AMNH), 3 # (AMNH); IX/1984, 2 # 3 $ 2 imm. Olmue, Palmas de Ocoa ( 71.08547, 32.93833, (IBSP 163707). Region� VI de O’Higgins: À À same locality [ 71.3, 29.28333, 79 m], 19/X/1983, 1 (AMNH); La Ligua Baja ( 71.04758, 31.0251, 697 m), A.D. Brescovit et al., 23/III/2013, 4 imm. Cachapoal, Machal�ı, Coya ( 70.61452, 34.1963, À À À À À À $ 8 imm. (AMNH); Chungungo ( 71.30283, 711 m), A.D. Brescovit et al., 7/XI/2013, 1 $ (IBSP (IBSP 163742); same locality ( 71.09033, 941 m), 5/IV/2013, 1 $ 4 imm. (IBSP 163708); À À 29.4446, 36 m), A.D. Brescovit et al., 11/XI/2013, 1 165570), 1 $ (IBSP 165571); Ovalle, 24 km S Socos, 32.91633, 377 m), 1 # 2 $ 7 imm. (IBSP 163743); Requinao, Donihue~ ( 70.84495, 34.18863, 483 m), À À À À $ (IBSP 165569), 1 # (UFMG 15332), 1 $ (UFMG Rt. 5, km 347 ( 71.61667, 30.88333, 360 m), N.I. San Felipe de Aconcagua, Catemu, El Guayacan� 25/III/2013, 3 imm. (IBSP 163755). No further À À

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016 © 20172016 The The Linnean Linnean Society Society of of London, London, ZoologicalZoological Journal Journal of of the the Linnean Linnean Society Society, 2017,, 2016 179, 767–864 81246 I. I. L. L. F. F. MAGALHAES MAGALHAESET ET AL. AL. SICARIID PHYLOGENY AND REVISION OF SICARIUS 47 data: Quebrada de Talca, 7/VIII/1997, 1 # (MHNS). Sicarius fumosus: Simon, 1893: 271. Sicarius defor- ARGENTINA. Buenos Aires: [ 58.36667, 34.6, mis: Simon, 1893: 271. À À 24 m], 15/IX/1996, 1 # 1 $ 2 imm. (AMNH) (certainly a mislabelling). Synonymy: This species was recognized based on the shape of femoral macrosetae, which are not medially widened as in fig. 7 of Nicolet’s paper (widened SICARIUS FUMOSUS (NICOLET, 1849) in S. crustosus and S. lanuginosus). Nicolet’s Figures 4A, B, 15E, 17B, 18B, 24B, 26C, 31A, 32B, S. deformis is an immature, but it fits with this 33D, E, 38, 39, 40B species, which is also the most common belonging to Thomisoides fumosus Nicolet, 1849: 354, plate 1, fig. his rugosas group. 7. Immature holotype from Chile deposited in MNHN 4160, examined, not found after the Instituto Diagnosis: Differs from S. thomisoides and Butantan fire in 2010. S. yurensis stat. nov. in the smaller size and by Thomisoides deformis Nicolet, 1849: 357. Immature possessing a truncate sternum, and white macrosetae holotype from Chile deposited in MNHN, not found. and black prolateral setae in the femora. Differs from SYN. NOV. the similar S. crustosus and S. lanuginosus by the

Figure 39. Sicarius fumosus (Nicolet, 1849) from Chile, female genitalia, digested. A, San Felipe, Valparaıso (IBSP 163738), dorsal. B, same, ventral. C, Belloto, Valparaıso (UCCC), dorsal. D, same, ventral. E, Quinahue, O’Higgins Figure 38. Sicarius fumosus (Nicolet, 1849) from Chile, male genitalia in prolateral view. A, Palmas de Ocoa, Val- (IBSP 163720), dorsal. F, same, ventral. G, subadult from El Radal, Maule (IBSP 163761), ventral. H, Los Vilos, paraıso (IBSP 165465). B, same, cymbium and bulb. C, Pichidanguı, Coquimbo (IBSP 163709). D, Cauquenes, Maule Coquimbo (AMNH), dorsal. I, same, ventral. J, Farellones, Santiago (IBSP 163722), dorsal. K, same, ventral. L, Cale- (AMNH). E, Farellones, Santiago (IBSP 163722). F, Huelquen, Santiago (IBSP 165461). Scale bars = 0.1 mm (except A, tones, O’Higgins (MHNS 678), dorsal. M, same, ventral. Scale bars = 0.1 mm. 1 mm).

Distribution: Southern Coquimbo region to Maule 33.5, 612 m), E.I. Schlinger, 5/IX/1966, 1 $ 2 imm. À region, Chile (Fig. 26C). (CAS 9052043). Region� VI de O’Higgins: Cachapoal, Machal�ı, Caletones, Embalse Sapos Additional material examined: CHILE. Region� IV [ 70.43333, 34.08333, 2233 m], M. Elgueta, 16/XII/ À À de Coquimbo: Choapa, Los Vilos, 19 km N Los Vilos, 1982, 1 # 2 $ (MHNS 678); Coya ( 70.50928, Rt. 5, km 244 ( 71.51667, 31.75, 5 m), N.I. Platnick 34.20235, 1043 m), A.D. BrescovitÀ & I.L.F. À À À et al., 9/XI/1993, 1 $ (AMNH); 6.5 km N Los Vilos Magalhaes,~ 5/IV/2013, 6 # 3 $ 3 imm. (IBSP 163714); [ 71.5, 31.85, 10 m], 16/X/1992, 1 $ (AMNH); same locality ( 70.61452, 34.1963, 941 m), 4 $ 4 À À À À Petorca, La Ligua, Pichidangu�ı, Valle de Quillimar�ı imm. (IBSP 163729); same locality ( 70.50928, À ( 71.50652, 32.10803, 25 m), A.D. Brescovit & 34.20235, 1043 m), 2 # (IBSP 163758), 1 # (UFMG À À À I.L.F. Magalhaes,~ 7/IV/2013, 1 # (IBSP 163709). 15352), 1 $ (UFMG 15353); Peumo, Pichidegua Region� V de Valpara�ıso: Petorca, Zapallar, Papudo ( 71.32548, 34.34533, 226 m), 26/III/2013, 1 $ 1 ( 71.46545, 32.50513, 76 m), A.D. Brescovit & imm.À (IBSP 163756);À Requinao, Donihue~ ( 70.84495, À À À I.L.F. Magalhaes,~ 9/IV/2013, 1 $ (IBSP 163712); 34.18863, 483 m), 25/III/2013, 2 $ (IBSP 163719), 1 À Quillota, Olmue, cerro La Campana [ 71.11667, $ (IBSP 165459); Los Nogales ( 70.65958, 34.22452, À À À 32.95, 818 m], R. Calderon,� 23/X/1971, 1 # (UCCC); 750 m), 5/IV/2013, 3 # 3 $ (IBSP 163711); San PalmasÀ de Ocoa ( 71.08547, 32.93833, 697 m), A.D. Vincente, between Coltauco and Telmo ( 71.15167, À À À Brescovit et al., 23/III/2013, 1 # (IBSP 163741), 1 $ 1 34.3798, 220 m), 26/III/2013, 1 # (IBSP 163763), 1 # À imm. (IBSP 165465); same locality [ 71.08547, (IBSP 165458); Colchaga, Chepica,� Quinahue, À 32.93833, 475 m], N.I. Platnick & O.F. Francke, 12/ Motocross road ( 71.32832, 34.69947, 198 m), 1 $ 2 Figure 40. Femur I, dorsal macrosetae, anterior view. A, Sicarius lanuginosus (Nicolet, 1849), male. B, Sicarius fumo- À À À I/1985, 1 # (AMNH); same locality [ 71.08547, imm. (IBSP 163720). Region� VII de Maule: sus (Nicolet, 1849), male. C, Sicarius crustosus (Nicolet, 1849), male. D, Sicarius crustosus (Nicolet, 1849), female. Note À 32.93833, 697 m], R. Calderon� G., 25/V/1984, 1 # Cauquenes, W of Cauquenes [ 72.31667, 35.96667, sexual dimorphism in S. crustosus, absent in other species. Scale bar, 0.1 mm. À À À (AMNH), 1 # (AMNH); Palmas de Ocoa, Parque 149 m], L. Irarrazaval, V/1984, 1 # (AMNH); same Nacional La Campana, 30/XI/1984, 1 # (AMNH), 1 # locality [ 72.31667, 35.96667, 350 m], 4/X/1983, 1 # À À (AMNH); 21/XII/1984, 1 # (AMNH); 29/I/1985, 1 $ (AMNH); Curico, Molina, El Radal, Parque Ingles� macrosetae in the femora having parallel sides interdistances: PME 0.13, ALE 0.15, PLE 0.18, (AMNH); San Felipe de Aconcagua, San Felipe ( 71.05915, 35.41338, 909 m), A.D. Brescovit & (Fig. 40B; widened in the other species, Fig. 40A, C, PME–PLE 0.36, ALE–PLE 0.18. Clypeus height À À ( 70.73738, 32.706, 739 m), A.D. Brescovit & I.L.F. I.L.F. Magalhaes,~ 3/IV/2013, 1 # 3 imm. (IBSP D), and by lacking brown markings at the border of 0.64. Sternum length 1.68, width 1.91, cordiform. À À Magalhaes,~ 6/IV/2013, 1 $ (IBSP 163738); Valparaıso, 163761); Teno, Los Quenes� [ 70.81667, 35, 880 m], the sternum. Males are further distinguished by the Femoral thorns: 4. Leg I: femur 3.84 long, patella À À Quilpue, Belloto [ 71.4, 33.03333, 197 m], A. R. Calderon� & J. Solenicens, 27/IV/1980, 1 $ (UCCC); long and curved embolus (Fig. 38). Females are 1.37, tibia 4, metatarsus 2.85, tarsus 1.68. II: fe À À Valenzuela, XI/1970, 1 $ (UCCC); Parque Nacional La Talca, R�ıo Claro, Los Maitenes ( 71.14145, further distinguished by the lack of ventral branches 4.01, ti 4. III: fe 3.92, ti 3.59. IV: fe 4.08, pa 1.3, ti Campana ( 71.12728, 32.98142, 947 m), A. 35.38833, 770 m), A.D. BrescovitÀ & I.L.F. in the spermathecae and by the median branches 3.92, mt 2.98, ta 1.68. Abdomen: length 3.27, width À À À Ojanguren et al., 22–23/VII/2010, 1 $ (MACN-Ar Magalhaes,~ 3/IV/2013, 1 $ (IBSP 163717); San being strongly coiled at the apex and much longer 3.55. Spermathecae branches: 10. Ventral branches 32326); Vina~ del Mar, Concon� ( 71.54267, 32.94467, Clemente, Las Garzas, La Suiza ( 70.99533, than the lateral branches (Fig. 39). absent (Fig. 39). À À À 91 m), A.D. Brescovit et al., 23/III/2013, 1 $ (IBSP 35.77283, 888 m), Hidreletrica� Pehuenche, 2/IV/ À 163713). Region� Metropolitana: Cachapoal, 2013, 2 $ (IBSP 163721), 1 # (IBSP 165457). No Description: Male (Palmas de Ocoa, Valparaıso, Variation: Males (N = 5): carapace length 2.95–4.36 Mostazal, Huelquen� ( 70.67338, 33.8749, 385 m), further data: 1 # (AMNH). Chile, IBSP 163741). Coloration: carapace, chelicerae, (3.74), femoral thorns 3–4, femur II length 3.96–6.07 À À 25/III/2013, 1 $ (IBSP 163762), 1 # (IBSP 165461); labium and endites reddish brown, sternum orange, (4.86). Females (N = 5): carapace length 3.63–4.92 Chacabuco, Cerro El Roble [ 71, 32.96667, 2200 m], SICARIUS CRUSTOSUS (NICOLET, 1849) legs orange, abdomen brownish grey. Total length (4.14), femoral thorns 3–4, femur II length 3.96–5.18 À À M. Elgueta, XI/1982, 1 $ (MHNS 608); Cordillera, San 6.68. Carapace length 3.59, anterior width 1.3, (4.42), spermathecae branches 4–10. Males (Fig. 38) Figures 4C, D, 10, 15C, D, 26D, 30D, 33L, 40C, D, 41 Jose de Maipo, El Canelo ( 70.45617, 33.58133, thoracic region width 3.36. Eye diameters and vary mainly in the length of the bulb body. Females À À Thomisoides crustosus Nicolet, 1849: 355, plate 1, 1350 m), A.D. Brescovit et al., 22/III/2013, 1 $ (IBSP interdistances: PME 0.13, ALE 0.15, PLE 0.15, PME– (Fig. 39) vary in number of spermathecae branches ﬁg. 8. Immature holotype from Chile deposited in 163759); same locality [ 70.38333, 33.61667, PLE 0.23, ALE–PLE 0.15. Clypeus height 0.43. Eye and in whether the lateral branches are strongly À À MNHN 4157, examined, not found after the Instituto 1000 m], L.E. Pena,~ 1980, 1 $ (AMNH); El Manzano tubercles present. Macrosetae in lateral border of the coiled. One female from Belloto, Valparaıso, lacks Butantan ﬁre in 2010. [ 70.4115, 33.5795, 1263 m], D. Jackson, VIII/1979, carapace in multiple rows. Femora with prolateral the rounded reservatory at the apex (Fig. 39C, D). À À 1 $ (MHNS 924); Farellones, Plazoleta Los Pumas black setae in a zig-zag pattern, white macrosetae Female subadults have pre-spermathecae with the Sicarius crustosus: Simon, 1893: 271. ( 70.29033, 33.35983, 2821 m), A.D. Brescovit present. Sternum length 1.63, width 1.88, cordiform. median branches digitiform, not coiled, much longer À À et al., 21/III/2013, 8 # 5 $ 1 imm. (IBSP 163716), 1 # 2 Note: This species has been recognized by the Femoral thorns: 3. Leg I: femur 4.36 long, patella than the laterals (Fig. 39G). $ (IBSP 163752), 1 # (IBSP 165460); road to strongly widened macrosetae in the femora. 1.37, tibia 4.67, metatarsus 3.39, tarsus 1.76. II: fe Farellones ( 70.36967, 33.35483, 1738 m), 2 $ 4.62, ti 4.67. III: fe 4.12, ti 4.08. IV: fe 4.31, pa 1.27, ti Natural history: This species has been collected (IBSP 163715);À same localityÀ ( 70.32933, 33.3525, Diagnosis: Differs from S. thomisoides and 4.26, mt 3.55, ta 1.86. Abdomen: length 3.22, width under rocks, logs and roots in xeric woodlands, high- À À 2207 m), 3 # 6 $ 1 imm. (IBSP 163722); same locality S. yurensis stat. nov. in the small size and by 2.95. Palp as in Figure 38A and B. altitude scrublands and beaches. Some of the places ( 70.36967, 33.35483, 1738 m), 1 # 1 $ (IBSP possessing a posteriorly truncate sternum, and white Female (Palmas de Ocoa, Valparaıso, Chile, IBSP where it has been collected (such as Farellones, À À 163739); Valle del R�ıo Mapocho entre El Arrayan� y macrosetae and black prolateral setae in the femora. 165466). As in male, except for the following. Total Santiago) are known to be covered by snow during Farellones [ 70.36967, 33.35483, 1738 m], W. Differs from S. fumosus by the widened macrosetae length 7.5. Carapace length 3.8, anterior width 1.4, the winter (adult specimens were collected in À À Noodt, X/1958–VI/1960, 1 # (MHNS 429); Maipo, in the femora (Fig. 40C, D; not widened in thoracic region width 3.71. Eye diameters and autumn). Eggcase as in Figure 17B. Calera de Tango, La Rinconada, Maipu(� 70.91667, S. fumosus, Fig. 40B), and by having brown À

markings at the border of the sternum (Fig. 30D). Natural history: Specimens have been collected The most similar species is S. lanuginosus: males under rocks in deserts, dry coastal vegetation and differ by the longer, slightly sinuous embolus of the xeric scrublands. This is a very cryptic species: male (Fig. 41A–C) (shorter and with a proximal individuals remain motionless when disturbed. hump in S. lanuginosus); females differ by lacking rounded reservatories at the apex of the Distribution: Dry coast of Chile, from Antofagasta to spermathecae branches, and by the gradual Coquimbo (Fig. 26D). reduction in length of the spermathecae branches (Fig. 41D–I) (rounded reservatories present and the Additional material examined: CHILE. Region� II two median pairs of branches clearly longer than the de Antofagasta: Antofagasta, Antofagasta, 4 km N laterals in S. lanuginosus). Paposo [ 70.38333, 23.51667, 20 m], N.I. Platnick À À et al., 11/X/1992, 1 $ (AMNH); 6 km E Paposo Description: Male (Paposo, Antofagasta, Chile, [ 70.33333, 23.56667, 894 m], 12/X/1992, 1 # 1 AMNH). Coloration: carapace and chelicerae reddish imm.À (AMNH);À Taltal, 24 km S Paposo [ 70.41667, À brown, labium and endites light brown, sternum light 25.2, 50 m], 11/X/1992, 1 # (AMNH). Region� III de brown with dark brown markings on margin, legs ÀAtacama: Chanaral,~ Las Lomitas, Parque Nacional light brown, abdomen brownish yellow. Total length Pan de Azucar� ( 70.60601, 26.00979, 614 m), M.J. À À 6.14. Carapace length 3.12, anterior width 1.39, Ram�ırez et al., 26/X/2011, 1 # 1 $ (MACN-Ar 32330); thoracic region width 2.92. Eye diameters and Copiapo,� Caldera, Rodillo [ 70.83333, 27.08333, À À interdistances: PME 0.15, ALE 0.18, PLE 0.15, PME– 26 m], XI/1991, 1 $ (LEULS); Copiapo,� 35 km S PLE 0.25, ALE–PLE 0.25. Clypeus height 0.56. Eye Copiapo[� 70.31667, 27.66667, 1326 m], L.E. Pena,~ À À tubercles present. Macrosetae in lateral border of the 16–17/X/1992, 1 $ (AMNH); 40 km S Copiapo� carapace in multiple rows. Femora with prolateral [ 70.31667, 27.73333, 700 m], 23–25/X/1983, 2 $ black setae in a zig-zag pattern, white macrosetae (AMNH);À RutaÀ 5, km. 758 ( 70.48717, 27.76693, À À present, widened at the base. Sternum length 1.35, 610 m), A.D. Brescovit et al., 9/XI/2013, 1 $ (IBSP width 1.63, cordiform. Femoral thorns: 5. Leg I: 165572); Huasco, Huasco, Caleta Angosta femur 3.88 long, patella 1.14, tibia 4.36, metatarsus ( 71.14792, 28.22308, 97 m), C.J. Grismado et al., À À 3.2, tarsus 1.63. II: fe 4.16, ti 4.2. III: fe 3.65, ti 3.63. 18/VIII/2009, 1 # (MACN-Ar 32331), 1 # (MACN-Ar IV: fe 3.96, pa 1.09, ti 3.8, mt 3.14, ta 1.6. Abdomen: 34224); Freirina ( 71.16243, 28.48288, 118 m), À À length 3.18, width 3.02. Palp as in Figure 41A and B. A.D. Brescovit et al., 9/XI/2013, 1 # (IBSP 165577), 1 Female (Paposo, Antofagasta, Chile, AMNH). As in $ (IBSP 165578); Parque Nacional Llanos de Challe male, except for the following. Total length 7.58. ( 71.11356, 28.18403, 304 m), C.J. Grismado et al., À À Carapace length 3.1, anterior width 1.27, thoracic 17–18/VIII/2009, 1 $ (MACN-Ar 32328); same locality region width 2.98. Eye diameters and interdistances: ( 71.0885, 28.19222, 304 m), 1 $ 1 imm. (MACN-Ar À À PME 0.13, ALE 0.13, PLE 0.14, PME–PLE 0.25, 32329). Region� IV de Coquimbo: Elqui, La ALE–PLE 0.2. Clypeus height 0.56. Femora macrose- Higuera, El Totoralillo ( 71.2615, 29.59264, À À tae widened at the apex. Sternum length 1.3, width 154 m), M. Izquierdo et al., 12/VII/2009, 1 $ (MACN- 1.5, cordiform. Femoral thorns: 4. Leg I: femur 3.08 Ar 30189), 1 $ 1 imm. (MACN-Ar 30205), 1 $ (MACN- long, patella 1.14, tibia 3.22, metatarsus 2.17, tarsus Ar 30206); Los Choros [ 71.46047, 29.24025, À À 1.19. II: fe 3.12, ti 3.15. III: fe 2.78, ti 2.75. IV: fe 40 m], A. Gonzalez, 2007, 1 $ (LEULS); La Serena, 3.12, pa 1.12, ti 2.21, mt 2.32, ta 1.37. Abdomen: Cuesta Porotitos ( 71.29217, 29.79447, 48 m), C.J. À À length 4.67, width 4.36. Spermathecae branches: 8. Grismado et al., 16/VIII/2009, 1 $ (MACN-Ar 32327); Ventral branches present, one or two pairs, straight. 1 # (MACN-Ar 30212); Limar�ı, Ovalle, Lorenzo Spermathecae as in Figure 41D and E. Peralta ( 71.58207, 30.5741, 230 m), A.D. À À Brescovit & I.L.F. Magalhaes,~ 12/XI/2013, 1 $ (IBSP Variation: Males (N = 3): carapace length 3.1–3.27 165584). (3.16), femoral thorns 4–5, femur II length 4.08–4.87 (4.37). Females (N = 5): carapace length 2.85–3.76 SICARIUS LANUGINOSUS (NICOLET, 1849) (3.15), femoral thorns 4–5, femur II length 3.08–4.11 (3.39), spermathecae branches 8–12. The number of Figures 15F, 26C, 40A, 42 ventral branches in the spermathecae of females Thomisoides lanuginosus Nicolet, 1849: 356. Imma- varies (one pair, one pair and one unpaired, two ture holotype from Chile deposited in MNHN 4155, Figure 41. Sicarius crustosus (Nicolet, 1849) from Chile. A–C, male genitalia in prolateral view; D–I, female genitalia, pairs). The femoral macrosetae morphology is examined, not found after the Instituto Butantan ﬁre digested. A, Paposo, Antofagasta (AMNH). B, same, cymbium and bulb. C, Paposo, Antofagasta (AMNH). D, Paposo, different in males and females, the ﬁrst having them in 2010. Antofagasta (AMNH), dorsal. E, same, ventral. F, Rodillo, Atacama (LEULS), dorsal. G, same, ventral. H, Copiapo, widened at the base (Fig. 40C) and the latter having Atacama (AMNH), dorsal. I, same, ventral. Abbreviation: VB, ventral branch. Scale bars = 0.1 mm (except A, 1 mm). them widened at the apex (Fig. 40D). Sicarius lanuginosus: Simon, 1893: 271.

macrosetae in the femora and black prolateral setae spermathecae in the females varies (Fig. 42F, H). in the femora. Differs from S. fumosus by the Female subadults have spermathecae with four medially widened macrosetae in the femora (Fig. 40A; short, medially grouped branches (Fig. 42D). not widened in S. fumosus, Fig. 40B), and by having brown markings at the border of the sternum (as in Natural history: This is a very elusive species; we Fig. 30D). The most similar species is S. crustosus: have visited some of the localities where museum males differ by the shorter embolus bearing a specimens were collected, but could not ﬁnd proximal hump (Fig. 42B, C) (longer and sinuous in individuals by active search. It is very rare in S. crustosus); females differ by having rounded collections, the only large series being that of long- reservatories at the apex of the spermathecae term ecological studies deposited in LEULS. branches, and by having the two median pairs of Specimens come from areas of xeric coastal vegetation. branches with about the same size, but clearly longer than the most lateral ones (Fig. 42E–H) (rounded Distribution: Apparently endemic to a small coastal reservatories absent and gradual reduction of length area in Southern Atacama region and Northern in the branches in S. crustosus). Coquimbo region, Chile (Fig. 26C).

Description: Male (Elqui, Coquimbo, Chile, AMNH). Additional material examined: CHILE. Region� III Coloration: carapace, chelicerae, labium and endites de Atacama: Huasco, Freirina, Caleta Chanaral~ de reddish brown, sternum light brown with dark Aceituno ( 71.48759, 29.07202, 41 m), M.J. À À brown markings on margin, legs light brown, Ram�ırez et al., 30/X/2011, 1 # (MACN-Ar 32332); abdomen brownish yellow. Total length 9.38. Isla de Chanaral~ [ 71.57649, 29.03346, 65 m], L.E. À À Carapace length 4.92, anterior width 1.67, thoracic Pena,~ 30/X/1980, 1 $ 1 imm. (AMNH). Region� IV region width 4.82. Eye diameters and interdistances: de Coquimbo: Elqui, Coquimbo, La Pampilla PME 0.2, ALE 0.2, PLE 0.2, PME–PLE 0.31, ALE– [ 71.33333, 29.95, 2 m], P. Vera, 11/VI/2011, 1 # PLE 0.23. Clypeus height 0.86. Eye tubercles (LEULS);À LaÀ Higuera, 9 km S Cruz Grande present. Macrosetae in lateral border of the carapace ( 71.31667, 29.5, 5 m), N.I. Platnick et al., 11/XI/ À À in multiple rows. Femora with prolateral black setae 1993, 1 # (AMNH); Los Choros [ 71.46047, À in a zig-zag pattern, white macrosetae present, 29.24025, 40 m], D. Valdivia, 7/X/2005, 1 $ medially widened. Sternum length 2.12, width 2.55, À(LEULS); D. Valdivia & J. Pizarro-Araya, 2007, 1 cordiform. Femoral thorns: 4. Leg I: femur 6.08 long, imm. (LEULS); F. Alfaro, 1 # (LEULS); J. Pizarro- patella 2.03, tibia 6.34, metatarsus 4.62, tarsus 2.43. Araya, 25/VIII/2005, 1 imm. (LEULS); J. Pizarro- II: fe 6.75, ti 6.34. III: fe 6.27, ti 5.39. IV: fe 6.2, pa Araya & D. Valdivia, 26/VIII/2005, 1 imm. (LEULS); 1.9, ti 5.39, mt 4.23, ta 2.32. Abdomen: length 4.43, L. Espinosa, 18/XI/2005, 1 # (LEULS); P. Gachon, width 4.02. Palp as in Figure 42A and B. 19/II/2005, 1 $ (LEULS); 17–20/XI/2005, 1 $ 1 imm. Female (Punta Choros, Coquimbo, Chile, LEULS). (LEULS); Punta Choros [ 71.46028, 29.249, 40 m], As in male, except for the following. Total length C. Farias, 7/X/2005, 1 imm.À (LEULS);À C. Flores, 4/ 10.88. Carapace length 4.46, anterior width 1.6, tho- VI/2005, 1 # (LEULS); J. Pizarro-Araya, 25/VIII/ racic region width 4.21. Eye diameters and interdis- 2005, 1 imm. (LEULS); L. Espinoza, 27/VIII/2005, 1 tances: PME 0.18, ALE 0.23, PLE 0.2, PME–PLE # (LEULS); 7/X/2005, 1 $ (LEULS); P. Gachon, 4/VI/ 0.31, ALE–PLE 0.13. Clypeus height 0.66. Sternum 2005, 1 imm. (LEULS); 8/X/2005, 1 imm. (LEULS); length 1.91, width 2.2, cordiform. Femoral thorns: 4. V. Rodriguez, 4/VI/2005, 1 $ (LEULS); 25–28/VIII/ Leg I: femur 4.77 long, patella 1.67, tibia 4.82, 2005, 1 # (LEULS), 4 imm. (LEULS); 7/X/2005, 1 # metatarsus 3.4, tarsus 2.08. II: fe 5.25, ti 4.67. III: fe 1 imm. (LEULS); 6–7/X/2005, 3 # 1 imm. (LEULS); 4.77, ti 4.67. IV: fe 4.92, pa 1.53, ti 4.98, mt 1.98, ta 6–9/X/2005, 1 # (LEULS), 2 # 3 imm. (LEULS); 17– 3.22. Abdomen: length 6.41, width 5.45. Spermathe- 20/XI/2005, 1 # 3 imm. (LEULS), 2 # 2 imm. cae branches: 8. Ventral branches present, one pair, (LEULS); Quebrada Los Choros [ 71.066, 29.35, À À straight. Spermathecae as in Figure 42E and F. 476 m], 1 (LEULS); La Serena, 20 km N La Figure 42. Sicarius lanuginosus (Nicolet, 1849) from Chile. A–C, male genitalia in prolateral view; D–H, female geni- # Serena, Rt. 5, km 491 [ 71.33333, 29.76667, talia, digested. A, Cruz Grande, Coquimbo (AMNH). B, same, cymbium and bulb. C, La Serena, Coquimbo (AMNH). D, À À subadult from Los Choros, Coquimbo (LEULS), ventral. E, Los Choros, Coquimbo (LEULS), dorsal. F, same, ventral. G, Variation: Males (N = 5): carapace length 3.09–5.66 120 m], N.I. Platnick et al., 6/X/1992, 1 # (AMNH). Isla Chanaral,~ Coquimbo (AMNH), dorsal. H, same, ventral. Abbreviation: VB, ventral branch. Scale bars = 0.1 mm (ex- (4.28), femoral thorns 4–5, femur II length 3.92–7.92 (5.73). Females (N = 3): carapace length 3.15–4.62 cept A, 1 mm). PERUVIAN SPECIES (4.08), femoral thorns 4, femur II length 3.76–5.73 (4.91), spermathecae branches 8–9. Some males Three species, one of them with two subspecies, have Note: This species was recognized on the basis of Diagnosis: Differs from S. thomisoides and have the proximal hump more subtle than others been cited from Peru: S. terrosus, S. peruensis (Key- the proportionally longer legs and morphology of the S. yurensis stat. nov. in the small size and by (Fig. 42B, C), making differentiation from serling, 1880), S. gracilis (Keyserling, 1880) and femoral macrosetae. possessing a posteriorly truncate sternum, white S. crustosus difﬁcult at times. The distance between S. terrosus yurensis Strand, 1908. The record of

Description: Male (Elqui, Coquimbo, Chile, AMNH). Additional material examined: CHILE. Region� III Coloration: carapace, chelicerae, labium and endites de Atacama: Huasco, Freirina, Caleta Chanaral~ de reddish brown, sternum light brown with dark Aceituno ( 71.48759, 29.07202, 41 m), M.J. À À brown markings on margin, legs light brown, Ram�ırez et al., 30/X/2011, 1 # (MACN-Ar 32332); abdomen brownish yellow. Total length 9.38. Isla de Chanaral~ [ 71.57649, 29.03346, 65 m], L.E. À À Carapace length 4.92, anterior width 1.67, thoracic Pena,~ 30/X/1980, 1 $ 1 imm. (AMNH). Region� IV region width 4.82. Eye diameters and interdistances: de Coquimbo: Elqui, Coquimbo, La Pampilla PME 0.2, ALE 0.2, PLE 0.2, PME–PLE 0.31, ALE– [ 71.33333, 29.95, 2 m], P. Vera, 11/VI/2011, 1 # PLE 0.23. Clypeus height 0.86. Eye tubercles (LEULS);À LaÀ Higuera, 9 km S Cruz Grande present. Macrosetae in lateral border of the carapace ( 71.31667, 29.5, 5 m), N.I. Platnick et al., 11/XI/ À À in multiple rows. Femora with prolateral black setae 1993, 1 # (AMNH); Los Choros [ 71.46047, À in a zig-zag pattern, white macrosetae present, 29.24025, 40 m], D. Valdivia, 7/X/2005, 1 $ medially widened. Sternum length 2.12, width 2.55, (LEULS);À D. Valdivia & J. Pizarro-Araya, 2007, 1 cordiform. Femoral thorns: 4. Leg I: femur 6.08 long, imm. (LEULS); F. Alfaro, 1 # (LEULS); J. Pizarro- patella 2.03, tibia 6.34, metatarsus 4.62, tarsus 2.43. Araya, 25/VIII/2005, 1 imm. (LEULS); J. Pizarro- II: fe 6.75, ti 6.34. III: fe 6.27, ti 5.39. IV: fe 6.2, pa Araya & D. Valdivia, 26/VIII/2005, 1 imm. (LEULS); 1.9, ti 5.39, mt 4.23, ta 2.32. Abdomen: length 4.43, L. Espinosa, 18/XI/2005, 1 # (LEULS); P. Gachon, width 4.02. Palp as in Figure 42A and B. 19/II/2005, 1 $ (LEULS); 17–20/XI/2005, 1 $ 1 imm. Female (Punta Choros, Coquimbo, Chile, LEULS). (LEULS); Punta Choros [ 71.46028, 29.249, 40 m], As in male, except for the following. Total length C. Farias, 7/X/2005, 1 imm.À (LEULS);À C. Flores, 4/ 10.88. Carapace length 4.46, anterior width 1.6, tho- VI/2005, 1 # (LEULS); J. Pizarro-Araya, 25/VIII/ racic region width 4.21. Eye diameters and interdis- 2005, 1 imm. (LEULS); L. Espinoza, 27/VIII/2005, 1 tances: PME 0.18, ALE 0.23, PLE 0.2, PME–PLE # (LEULS); 7/X/2005, 1 $ (LEULS); P. Gachon, 4/VI/ 0.31, ALE–PLE 0.13. Clypeus height 0.66. Sternum 2005, 1 imm. (LEULS); 8/X/2005, 1 imm. (LEULS); length 1.91, width 2.2, cordiform. Femoral thorns: 4. V. Rodriguez, 4/VI/2005, 1 $ (LEULS); 25–28/VIII/ Leg I: femur 4.77 long, patella 1.67, tibia 4.82, 2005, 1 # (LEULS), 4 imm. (LEULS); 7/X/2005, 1 # metatarsus 3.4, tarsus 2.08. II: fe 5.25, ti 4.67. III: fe 1 imm. (LEULS); 6–7/X/2005, 3 # 1 imm. (LEULS); 4.77, ti 4.67. IV: fe 4.92, pa 1.53, ti 4.98, mt 1.98, ta 6–9/X/2005, 1 # (LEULS), 2 # 3 imm. (LEULS); 17– 3.22. Abdomen: length 6.41, width 5.45. Spermathe- 20/XI/2005, 1 # 3 imm. (LEULS), 2 # 2 imm. cae branches: 8. Ventral branches present, one pair, (LEULS); Quebrada Los Choros [ 71.066, 29.35, À À straight. Spermathecae as in Figure 42E and F. 476 m], 1 (LEULS); La Serena, 20 km N La Figure 42. Sicarius lanuginosus (Nicolet, 1849) from Chile. A–C, male genitalia in prolateral view; D–H, female geni- # Serena, Rt. 5, km 491 [ 71.33333, 29.76667, talia, digested. A, Cruz Grande, Coquimbo (AMNH). B, same, cymbium and bulb. C, La Serena, Coquimbo (AMNH). D, À À subadult from Los Choros, Coquimbo (LEULS), ventral. E, Los Choros, Coquimbo (LEULS), dorsal. F, same, ventral. G, Variation: Males (N = 5): carapace length 3.09–5.66 120 m], N.I. Platnick et al., 6/X/1992, 1 # (AMNH). Isla Chanaral,~ Coquimbo (AMNH), dorsal. H, same, ventral. Abbreviation: VB, ventral branch. Scale bars = 0.1 mm (ex- (4.28), femoral thorns 4–5, femur II length 3.92–7.92 (5.73). Females (N = 3): carapace length 3.15–4.62 cept A, 1 mm). PERUVIAN SPECIES (4.08), femoral thorns 4, femur II length 3.76–5.73 (4.91), spermathecae branches 8–9. Some males Three species, one of them with two subspecies, have Note: This species was recognized on the basis of Diagnosis: Differs from S. thomisoides and have the proximal hump more subtle than others been cited from Peru: S. terrosus, S. peruensis (Key- the proportionally longer legs and morphology of the S. yurensis stat. nov. in the small size and by (Fig. 42B, C), making differentiation from serling, 1880), S. gracilis (Keyserling, 1880) and femoral macrosetae. possessing a posteriorly truncate sternum, white S. crustosus difﬁcult at times. The distance between S. terrosus yurensis Strand, 1908. The record of

S. terrosus by Chamberlin (1916) probably refers to differences were true or caused by the poor conserva- S. boliviensis sp. nov. (see below). Sicarius peruensis tion of his specimen, he stated that, if those differ- and S. gracilis are here revised and considered valid; ences proved to be meaningful, his ‘variety could be S. terrosus yurensis is given species status; and two named yurensis’ (Strand, 1908, p. 227). Nonetheless, new species are herein described. subsequent catalogues treated yurensis as a valid subspecies (Roewer, 1942; Bonnet, 1958; Brignoli, 1983; World Spider Catalog, 2015). We could not find SICARIUS YURENSIS STRAND, 1908 STAT. NOV. Strand’s specimen, but we have examined a male Figures 24E, 26B, 30B, 37B, 43, 44 from Yura and it agrees with his description, notably Sicarius terrosus yurensis Strand, 1908: 226–227. in a diagnostic character: the width of the cephalic Female from Yura, Peru, most probably deposited in region (4 mm in S. terrosus yurensis, 2.8 in Keyser- Wiesbaden Museum, not found (F. Geller-Grimm, ling’s description, both with 8 mm in carapace pers. comm.). length). Based upon this morphological remark and Sicarius peruvianus: Lehtinen, 1986: 152, fig. 3 the type locality, we could recognize this wide- (misidentified, invalid emendation). headed Sicarius, which we elevate to species status. Note: Strand (1908) was not explicit about his Diagnosis: Males, females and juveniles are easily intentions to describe this as a species or a sub- distinguished from all Sicarius by the wide cephalic species; he was merely recording the presence of region (~50% of the carapace length) (Fig. 37B). S. terrosus in Peru, and noted some differences Males are further distinguished by the long and between his specimen and the description by Keyser- slender palpal bulb, with very thin embolus ling (1880). Facing doubts about whether these

Figure 44. Sicarius yurensis Strand, 1908 stat. nov., female genitalia, digested. A, Las Yaras, Tacna, Peru (AMNH), dorsal. B, same, ventral. C, Quebrada Chipana, Tarapaca, Chile (IBSP 161783), dorsal. D, same, ventral. E, San Pedro de Atacama, Antofagasta, Chile (AMNH), dorsal. F, same, ventral. G, subadult from Ninabamba, Peru (AMNH), ventral. H, subadult from Chincha, Ica, Peru (UFMG 15341), ventral. Abbreviation: VB, ventral branch. Scale bars = 0.1 mm.

(Fig. 43). Females are further distinguished (except labium, endites, sternum and legs light brown, from S. thomisoides) by the spermathecae with 10– abdomen brownish yellow. Total length 14.7. 18 branches usually bent at the apex and without a Carapace length 8, anterior width 3.82, thoracic rounded reservatory (Fig. 44). We have been unable region width 7.42. Eye diameters and interdistances: to ﬁnd a genital feature in females to distinguish PME 0.25, ALE 0.28, PLE 0.23, PME–PLE 1.37, this from S. thomisoides. ALE–PLE 0.28. Clypeus height 1.45. Eye tubercles Figure 43. Sicarius yurensis Strand, 1908 stat. nov., male genitalia in prolateral view. A, male from Yura, Arequipa, absent. Macrosetae in lateral border of the carapace Peru (bulb in prolateral view, rest of the palp twisted laterally) (AMNH). B, same, palpal femur, prolateral. C, same, Description: Male (Yura, Arequipa, Peru, AMNH). in a single row. Femora without prolateral black cymbium and bulb. D, Tacna, Peru (UAP). Scale bars = 0.1 mm (except A and B, 1 mm). Coloration: carapace and chelicerae reddish brown, setae. Sternum length 3.59, width 3.18, oval.

Femoral thorns: 6. Leg I: femur 9.4 long, patella 3.3, (AMNH); L.E. Pena,~ 29–31/I/1992, 1 imm. (AMNH); tibia 9.9, metatarsus 7.8, tarsus 4.9. II: fe 10.5, ti Valle R�ıo Lluta, [ 70.3, 18.4, 48 m], 2/II/1992, 1 10.38. III: fe 9.3, ti 9.1. IV: fe missing. Abdomen: imm. (AMNH); [ À70.31667,À 18.48333, 44 m], J. À À length 6.2, width 4.98. Palp as in Figure 43A–C. Aros, VII/1961, 2 $ (AMNH). Region� I de Female (Las Yaras, Tacna, Peru, AMNH). As in Tarapaca:� Iquique, Pozo Almonte, Pampa del male, except for the following. Total length 18.45. Tamarugal, ( 69.54533, 20.43567, 997 m), A.D. À À Carapace length 7.67, anterior width 4.33, thoracic Brescovit et al., 10/VII/2012, 1 # (IBSP 165501); region width 6.92. Eye diameters and interdistances: Quebrada Chipana, ( 69.12861, 20.81556, 2540 m), À À PME 0.28, ALE 0.25, PLE 0.31, PME–PLE 1.45, A. Taucare-R�ıos, 23/VIII/2009, 1 $ (IBSP 161783); ALE–PLE 0.31. Clypeus height 1.47. Sternum length Quebrada Duplijsa, Mamina,~ [ 69.4, 20.01667, 3.65, width 3.3, oval. Femoral thorns: 4. Leg I: femur 1923 m], L.E. Pena,~ VI/1985,À 1 imm.À (AMNH); 6.83 long, patella 2.74, tibia 6.5, metatarsus 4.58, Poconchile [ 70.06648, 18.44994, 749 m], 23/VI/ À À tarsus 3.75. II: fe 7.92, ti 6.83. III: fe 7.5, ti 6.42. IV: 2010, 1 $ 1 imm. (UAP). Region� II de Antofagasta: fe 7.08, pa 2.58, ti 6, mt 4.5, ta 3.92. Abdomen: Antofagasta, Antofagasta, 61 km S Antofagasta, length 9.9, width 8.2. Spermathecae branches: 17. [ 70.41667, 24.21667, 1637 m], 21/VIII/1982, 2 Ventral branches present, one pair, straight. Sper- imm.À (AMNH);À El Loa, San Pedro de Atacama, mathecae as in Figure 44A and B. Guatin, [ 68.08333, 22.78333, 3300 m], 23–30/VIII/ À À 1982, 1 $ 8 imm. (AMNH); Peine, [ 68.05, À Variation: Males (N = 3): carapace length 6.34–8 23.66667, 2453 m], XI/1975, 1 $ (AMNH); Talabre, À (7.45), femoral thorns 5–6, femur II length 9.1–10.5 [ 67.91667, 23.25, 3200 m], 14/II/1992, 1 $ 5 imm. À À (9.99). Females (N = 5): carapace length 7.17–8.8 (AMNH); Tilomonte, [ 68.1, 23.78333, 2374 m], 23/ À À (7.68), femoral thorns 4–5, femur II length 7.92–9.8 I/1996, 1 $ 4 imm. (AMNH); [ 68.08333, 22.78333, À À (8.45), spermathecae branches 10–17. Genital mor- 3271 m], Guat�ın, 8/II/1997, 1 $ 1 imm. (UCCC); phology is slightly variable in both sexes (Figs 43, same locality, [ 68.1, 23.78333, 2374 m], L.E. À À 44). Subadult females have pre-spermathecae with Pena,~ 23–30/VIII/1982, 2 $ 4 imm. (AMNH); same four to several short, digitiform branches (Fig. 44G, H). locality, ( 68.06667, 23.68333, 2453 m), 10/IV/ À À 1990, 1 $ 5 imm. (AMNH). Region� III de Atacama: Natural history: This species has been collected Toconao, [ 68, 23.18333, 2684 m], E. Casanueva, À À from desert areas in coastal Peru and Chile; it seems 12/XI/1982, 1 $ (UCCC). more common in the higher altitudes of northern Chile than its sister species, S. thomisoides. SICARIUS PERUENSIS (KEYSERLING, 1880) Distribution: Desertic coast in Central to Southern Figures 16C, 17C, 18D, 27B, 31C, 32C, 33F–H, 45 Peru and Northern Chile (Fig. 26B). The Thomisoides peruensis Keyserling, 1880: 272, plate northernmost records from Peru are based on 8, fig. 150. Syntypes from several localities in Peru, immatures only; examining adults from those deposited in PAN, partially examined, only an imma- regions would be desirable to confirm co-specificity. ture syntype (PAN 10a) from Chorillos, Peru (see below). Additional material examined: PERU. Lima: Sicarius peruensis: Simon, 1893: 271; Aguilar & Canete,~ Lunahuana,� [ 76.11667, 12.96667, À À Mendez,� 1971: 143, plate 2. 1214 m], 1 imm. (MUSM 504943). Ica: Chincha, Grocio Prado, San Juan, ( 75.98883, 13.40575, Note: This species was described from several À À 768 m), A.D. Brescovit et al., 19/VI/2012, 3 imm. localities in Peru, including Lima, Monterico, San (IBSP 165447), 1 imm. (UFMG 15341). Ayacucho: Malu, Montana Nancho and Chorillos (Keyserling, Lamar, Ninabamba, R�ıo Pampas, [ 73.86667, 1880). Although Keyserling mentions at least a male 12.95, 1800 m], W. Weyrauch, 2/X/1947,À 1 imm. in the description, we could only find an immature À (AMNH). Arequipa: Arequipa, Yura, [ 71.66667, syntype in PAN. He did illustrate a palp, and À 16.25, 2829 m], K.P. Schmidt, 10/VIII/1939, 1 # 1 although his illustration is not very detailed, it À imm. (AMNH). Tacna: Tacna, Las Yaras, [ 70.55, shows the projecting cymbium and long embolus À 17.86667, 400 m], 3–5/XI/1983, 1 $ (AMNH); 28/X/ characteristic of this species. In addition, this spe- À Figure 45. Sicarius peruensis (Keyserling, 1880) from Peru. A–C, male genitalia in prolateral view; D–K, female geni- 2009, 1 # (UAP). CHILE. Region� XV de Arica y cies is very common in Lima and surrounding areas, Parinacota: Arica, Arica, 2.5 km W, 3.2 km S which are localities from where some of Keyserling’s talia, digested. A, Canta, Lima (IBSP 162924). B, same, cymbium and bulb. C, Isla Guanape Norte, La Libertad (AMNH). D, Ricardo Palma, Huarochiri, Lima (IBSP 162929), dorsal. E, same, ventral. F, same, anterior. G, Pisco, Ica Poconchile, Ruta A-143, [ 70.09382, 18.48273, material came. We decided to retain the name (IBSP 162915), dorsal. H, same, ventral. I, subadult female from Asia, Canete,~ Lima (IBSP 162931), ventral. J, Ciene- 932 m], R. Sage, 31/VII/2013,À 2 2 imm.À (MACN-Ar S. peruensis for the species common around Lima, $ guilla, Huaycan, Lima (MUSM 504947), dorsal. K, same, ventral. PP, pore plate; VB, ventral branch. Scale 30188), 1 (MACN-Ar 30210); Azapa Valley, which has been cited under this name in a few $ bars = 0.1 mm (except A, 1 mm). [ 70.18333, 18.51667, 800 m], X/1941, 1 imm. works (e.g. Aguilar & Mendez,� 1971; Alegre et al., À À

1977; Binford et al., 2008, 2009). The name S. gra- varies only slightly in both sexes. Subadult females A.D. Brescovit et al., 16/VI/2012, 6 # 9 $ (IBSP S. peruensis and S. utriformis by the more robust cilis is here retained for the rarer species, which is have pre-spermathecae with widely spaced, 162917); same locality [ 76.75, 12.06667, 917 m], branches of the spermathecae and by the ventral À À partially sympatric (see discussion below, under digitiform branches, and a small, slightly coiled D. Silva D., 19/I/1986, 1 $ (MUSM 504947); same branches of the spermathecae with a single, usually S. gracilis). ventral branch (Fig. 45I). locality ( 76.81667, 12.1, 475 m), P. Centeno, 23/ incomplete coil (Fig. 46D–I) (slender branches, À À VI/2007, 1 $ (MUSM 504934); E. Sta. Maria del Mar, double coil in S. peruensis and S. utriformis). Diagnosis: Distinguished from all Neotropical Natural history: Specimens have been collected in Puquio de Chilca, Quebrada de Chilca, 7 km species, except S. gracilis and S. utriformis, by coastal deserts, under rocks and cattle dung. [ 76.75, 12.4, 720 m], O.F. Francke, 24/XII/1973, 1 Description: Male (Chincha, Ica, Peru, UFMG À À having a single, large distal patch of prolateral black Eggcase as in Figure 17C. Aspects of the natural $ (AMNH); La Molina Vieja [ 76.95, 12.08333, 15342). Coloration: carapace and chelicerae reddish À À setae in the femora (Fig. 24D). Males differ from history of this species were given by Aguilar & 319 m], O. Meneses, 4/I/1968, 1 $ 2 imm. (IBSP brown, labium, endites, sternum and legs orange, S. gracilis and S. utriformis by the long embolus and Mendez (1971). 2578); Lomas de Atocongo [ 76.9, 12.2, 342 m], C. abdomen brownish grey. Total length 8.7. Carapace À À the abrupt transition between the base and body of Mudahn, 26/V/1984, 1 $ (MUSM 504942); Lomas length 4.92, anterior width 1.98, thoracic region width the bulb (Fig. 45A–C) (smoother transition in Distribution: Coastal deserts in central and Flor de Amancaes ( 77.11667, 11.98333, 36 m), C. 4.72. Eye diameters and interdistances: PME 0.2, À À S. gracilis and S. utriformis). Females are northern Peru (Fig. 27B). A record from Yuyapichis Castillo, 15–29/III/2005, 1 # (MUSM 501530); ALE 0.2, PLE 0.18, PME–PLE 0.58, ALE–PLE 0.18. distinguished from S. gracilis by the slender branches River (IBSP 2921), an area of rainforest in Huanuco, Zapallal [ 77.08333, 11.83333, 503 m], J. Utene, Clypeus height 0.84. Eye tubercles present. À À of the spermathecae and by the ventral branches with is probably a labelling error and has not been 21/VII/1986, 1 $ (MUSM 504941). Ica: Chincha, Macrosetae in lateral border of the carapace in a double coil (Fig. 45D–K) (robust branches, single included in the map. Grocio Prado, San Juan ( 75.98883, 13.40575, multiple rows. Femora with prolateral black setae in a À À coil in S. gracilis), and from S. utriformis by lacking 768 m), A.D. Brescovit et al., 19/VI/2012, 3 # 5 $ 1 large distal patch. Sternum length 2.21, width 2.34, digitiform projections in the spermathecae. Additional material examined: PERU. La imm. (IBSP 162912); ( 76.10361, 13.41483, 171 m), cordiform. Femoral thorns: 3. Leg I: femur 6.92 long, À À Libertad: Guanape, Isla Norte [ 78.95, 8.53333], 2 # 4 $ 1 imm. (IBSP 162930); Pisco, Humay patella 2.16, tibia 7.42, metatarsus 5.52, tarsus 3.12. À À Description: Male (Canta, Lima, Peru, UFMG D. Duffey, II/1979, 3 # 4 imm. (AMNH); Trujillo, ( 75.82083, 13.70058, 718 m), 1 # 5 $ 3 imm. II: fe 7.67, ti 7.75. III: fe 7.17, ti 6.92. IV: fe 7.17, pa À À 15348). Coloration: carapace and chelicerae reddish Hacienda Cartavio [ 79.01667, 8.1, 46 m], W. (IBSP 162915), 1 # (UFMG 15344), 1 $ (UFMG 2.08, ti 6.82, mt 5.86, ta 3.1. Abdomen: length 3.96, À À brown, labium and endites light brown, sternum Weyrauch, VI–VII/1939, 1 $ (AMNH). Lima: Canete,~ 15345). Huanuco: Panguana, Rio Yuyapichis width 3.88. Palp as in Figure 46A and B. and legs brownish yellow, abdomen brownish Asia, Via Coyallo ( 76.54303, 12.76814, 130 m), [ 74.93333, 9.6, 239 m], U. Meede, 7/IX/1977, 1 $ Female from Chincha, Ica, Peru (IBSP 165448). As À À À À yellow. Total length 9.6. Carapace length 5.59, A.D. Brescovit et al., 18/VI/2012, 1 # 2 $ 6 imm. (IBSP 2921) (this might be a labelling error). in male, except for the following. Total length 10.75. anterior width 2.18, thoracic region width 5.52. (IBSP 162931); Chilca ( 76.70336, 12.52661, Carapace length 5.73, anterior width 2.39, thoracic À À Eye diameters and interdistances: PME 0.2, ALE 246 m), 8 # 10 $ (IBSP 162916); Huarochıri, Chilca, region width 5.07. Eye diameters and interdistances: SICARIUS GRACILIS KEYSERLING, 1880 0.2, PLE 0.2, PME–PLE 0.66, ALE–PLE 0.2. Via Santo Domingo de los Olleros ( 76.62456, PME 0.2, ALE 0.2, PLE 0.2, PME–PLE 0.71, ALE– À Clypeus height 0.92. Eye tubercles present. 12.45039, 504 m), 1 # 1 $ 3 imm. (IBSP 162919); Figures 26B, 46 PLE 0.2. Clypeus height 1.12. Sternum length 2.49, À Macrosetae in lateral border of the carapace in Canta, Canta ( 76.77486, 11.62753, 1612 m), 13/ Thomisoides gracilis Keyserling, 1880: 275. Female width 2.44, cordiform. Femoral thorns: 4. Leg I: À À multiple rows. Femora with prolateral black setae VI/2012, 1 # 5 $ (IBSP 162924), 1 # (UFMG 15348), holotype from Monterico, Peru, deposited in PAN, femur 6.61 long, patella 2.29, tibia 6.92, metatarsus in a large distal patch. Sternum length 2.29, width 1 $ (UFMG 15349); Road to Arahuay ( 76.76869, not found. 4.77, tarsus 3.05. II: fe 7.08, ti 7.25. III: fe 7, ti 6.61. À 2.49, cordiform. Femoral thorns: 4. Leg I: femur 11.66231, 1612 m), 1 $ (IBSP 162928); Santa Rosa IV: fe 6.92, pa 2.16, ti 6.55, mt 5.18, ta 3.09. Abdo- À 8.17 long, patella 2.41, tibia 8.33, metatarsus 6.14, de Quives, Trapiche ( 76.81631, 11.69161, Sicarius gracilis: Simon, 1893: 271. men: length 5.02, width 4.82. Spermathecae À À tarsus 3.95. II: fe 9.1, ti 8.9. III: fe 8.33, ti 7.67. 1344 m), 1 # (IBSP 162914); Desembocadura R�ıo branches: 4. Ventral branches present, one pair, with IV: fe 7.67, pa 1.96, ti 7.25, mt 5.45, ta 3.68. Chillon, D. Silva D., 17/II/1984, 1 $ (MUSM 504939); Note: The recognition of this species was hampered a single coil. Spermathecae as in Figure 46D and E). Abdomen: length 4.16, width 4.01. Palp as in Huaral, Reserva Nacional de Lachay ( 77.36858, by the fact that we could not ﬁnd Keyserling’s speci- À Figure 45A and B. 11.41228, 157 m), A.D. Brescovit et al., 15/VI/2012, men in PAN and the original description is vague and Variation: Males (N = 5): carapace length 4.51–5.73 À Female (same locality, UFMG 15349). As in male, 1 $ 2 imm. (IBSP 162922); same locality ( 77.32353, could refer to several Sicarius species. Additionally, (5.13), femoral thorns 3–4, femur II length 6.61–8.7 À except for the following. Total length 11.38. Carapace 11.37772, 181 m), 3 # 3 $ 3 imm. (IBSP 162923), 1 Keyserling only recorded S. gracilis for Monterico, (7.83). Females (N = 5): carapace length 5.02–6.2 À length 6.34, anterior width 2.45, thoracic region # (UFMG 15346), 1 $ (UFMG 15347); same locality Peru, an unspeciﬁc location that could refer to many (5.48), femoral thorns 3–4, femur II length 6.07–7.5 width 6. Eye diameters and interdistances: PME [ 77.35, 11.36667, 417 m], P. Aguilar, X/1968, 1 $ places. However, he also recorded another species, (6.68), spermathecae branches 4–6. Genital À À 0.18, ALE 0.2, PLE 0.2, PME–PLE 0.74, ALE–PLE 1 imm. (MUSM 504932); Huarochiri, Nieve Nieve S. peruensis, for this locality. We found two species in morphology is slightly variable in both sexes; some 0.28. Clypeus height 1.14. Sternum length 2.75, ( 76.68642, 12.02531, 1102 m), A.D. Brescovit the coast of Peru with partially overlapping distribu- females, especially those from the northern range of À À width 2.78, cordiform. Femoral thorns: 5. Leg I: et al., 16/VI/2012, 1 # 1 $ 1 imm. (IBSP 162932); tions: one very common, especially around Lima, and the species, have a sclerotized pore plate in the femur 7.42 long, patella 2.58, tibia 7.42, metatarsus Ricardo Palma ( 76.65406, 11.92508, 1384 m), 14/ another one much rarer. Thus, we decided to retain spermathecae (Fig. 46I), while others do not (Fig. 46E, À À 5.02, tarsus 3.35. II: fe 8.08, ti 7.75. III: fe 7.67, ti VI/2012, 1 # 1 $ 2 imm. (IBSP 162929), 1 $ (IBSP the name S. peruensis for the more common species, G). Female subadults have pre-spermathecae with 6.83. IV: fe 7.58, pa 2.58, ti 6.92, mt 5.08, ta 4.25. 165445); Sisicaya ( 76.63144, 12.02775, 1263 m), and the name S. gracilis for the rarer species. short and blunt branches (Fig. 46J). À À Abdomen: length 5.18, width 4.5. Spermathecae 16/VI/2012, 1 $ 2 imm. (IBSP 162926); Verrugas branches: 8. Ventral branches present, one pair, with Canon [ 76.4875, 11.88944, 2329 m], R.C. Diagnosis: Distinguished from all Neotropical Natural history: This species has been collected À À a double coil (Fig. 45). Shannon, 20/VI/1928, 1 # (SINHM); Lima, Ancon species, except S. gracilis and S. utriformis, by from coastal deserts and dry forests. [ 77.11667, 11.7, 450 m], J.C. Pallister, 8/II/1947, having a single, large distal patch of prolateral black À À Variation: Males (N = 5): carapace length 5.39–6 1 $ (AMNH); Atocongo [ 76.9, 12.2, 342 m], setae in the femora (Fig. 24D). Males differ from Distribution: Southern Ecuador and coastal areas À À (5.69), femoral thorns 3–5, femur II length 7.92–9.5 Koepcke, 4/XI/1951, 1 $ 1 imm. (MUSM 504945); S. peruensis by the shorter embolus (Fig. 46A–C) from northern to central Peru (Fig. 26B). (8.84). Females (N = 5): carapace length 5.25–6.34 Chosica, Santa Marıa de Chosica [ 76.7, 11.93333, and from S. utriformis by the more abrupt transition À À (5.93), femoral thorns 3–5, femur II length 6.61–8.08 1124 m], O. Meneses, X/1968, 1 $ 1 imm. (IBSP between the base and body of the bulb (smooth in Additional material examined: ECUADOR. (7.5), spermathecae branches 6–10. Genital shape 2579); Cieneguilla ( 76.74375, 12.05431, 917 m), S. utriformis). Females are distinguished from Manabı: Jipijapa, Puerto Lopez, Parque Nacional À À

Machalilla ( 80.76389, 1.59083, 247 m), A. Chagas ( 75.86797, 13.37006, 997 m), A.D. Brescovit et al., À À À À et al., 2/IV/2011, 3 # 1 imm. (MNRJ 6259). PERU. 19/VI/2012, 4 # 6 imm. (IBSP 162921), 1 $ (IBSP Piura: Talara, Negritos [ 81.30472, 4.65389], H. 165448), 1 # (UFMG 15342); Nazca, Palpa, Dunas de À À Exline & W.B. Peck, 13/III/1939, 1 # 1 $ (CAS Usaca, Tambo de Perro ( 75.20594, 14.82369, À À 9052051). Lambayeque: Lambayeque, Las Pampas, 354 m), M. Vilchez & D. Silva D., 3/IV/2010, 1 $ km 885 old Pan-American Highway [ 79.76667, (MUSM 501200); [ 74.93333, 14.83333, 656 m], H. À À À 5.86667, 157 m], S.W. Cardiff, 8/IV/1981, 1 # Aslencios, IV/1990, 1 # (MUSM 504935). À (AMNH); Lobos de Tierra, Lobos de Tierra Island ( 80.88333, 6.45), K.P. Schmidt, VIII/1939, 2 # À À SICARIUS ANDINUS SP. NOV. (AMNH), 1 # 1 $ (AMNH). Lima: Canete,~ Lunahuana( 76.16278, 12.97691, 668 m), A.D. Figures 27B, 47 À À Brescovit et al., 18/VI/2012, 4 # 6 imm. (IBSP 162920); same locality ( 76.15811, 12.98589, Type material: Male holotype from PERU. À À 668 m), 3 # 4 $ 3 imm. (IBSP 162925), 3 $ 3 imm. Huanuco: Acomayo, Andes [ 76.05, 9.46, 1291 m], À À (IBSP 162927); same locality ( 76.16278, 12.97691, F. Woytkowski, VII/1946 (AMNH). Paratypes: À À 668 m), 1 # (IBSP 165452); same locality PERU. Huanuco: Ambo [ 76.2, 10.13333, À À [ 76.11667, 12.96667, 1214 m], 1 $ (MUSM 2100 m], W. Weyrauch, 2 $ 2 imm. (AMNH); À À 504946). Ica: Chincha, Laran, Luchumarca Acomayo [ 76.05, 9.46, 2100 m], F. Woytkowski, À À

Figure 46. Sicarius gracilis (Keyserling, 1880). A–C, male genitalia in prolateral view; D–J, female genitalia, digested. A, Chincha, Ica, Peru (UFMG 15342). B, same, cymbium and bulb. C, Puerto Lopez, Manab�ı, Ecuador (MNRJ 6259). D, Figure 47. Sicarius andinus sp. nov. from Peru. A–B, male genitalia in prolateral view; C–G, female genitalia, Chincha, Ica, Peru (IBSP 165448), dorsal. E, same, ventral. F, Lunahuana, Canete,~ Lima, Peru (IBSP 162925), dorsal. digested. A, male holotype from Acomayo, Huanuco (AMNH) (right palp, mirrored). B, same, cymbium and bulb. C, G, same, ventral. H, Negritos, Piura, Peru (CAS 9052051), dorsal. I, same, ventral. J, subadult female from Lunahuana, female paratype from Ambo, Huanuco (AMNH), dorsal. D, same, ventral. E, subadult from Huanuco (AMNH), dorsal. F, Canete,~ Lima, Peru (IBSP 162925), ventral. PP, pore plate; VB, ventral branch. Scale bars = 0.1 mm (except A, 1 mm). same, ventral. Scale bars = 0.1 mm (except A, 1 mm).

carapace in multiple rows. Femora without prolateral and orange (Fig. 49) (other species, except S. levii black setae. Sternum length 2.86, width 3.06, cordi- sp. nov., with light brown pores; see also colour form. Femoral thorns: 7. Leg I: femur 6.75 long, photos in the interactive key provided as Supporting patella 2.55, tibia 7, metatarsus 5,39, tarsus 2.53. II: Information Appendix S1). fe 7.17, ti 7.08. III: fe 6.61, ti 6.27. IV: fe 6.75, pa 2.27, ti 6.27, mt 4.97, ta 2.55. Abdomen: length 6.41, width Description: Male holotype (Lagunillas, Santa Cruz, 6.34. Spermathecae branches: 5. Ventral branches Bolivia, MZSP 48990). Coloration: carapace and absent. Spermathecae as in Figure 47C and D. chelicerae reddish brown, labium brown, endites reddish brown, sternum and legs orange brown, Variation: Males (N = 3): carapace length 5.07–5.93 abdomen light brown. Total length 13.35. Carapace (5.44), femoral thorns 4–6, femur II length 7.92–9.3 length 6.75, anterior width 2.47, thoracic region (8.49). Females (N = 2): carapace length 5.93–6 width 7. Eye diameters and interdistances: PME (5.97), femoral thorns 5–7, femur II length 7.17–7.33 0.23, ALE 0.23, PLE 0.25, PME–PLE 0.58, ALE–PLE (7.25), spermathecae branches 5–6. Genital 0.23. Clypeus height 1.17. Eye tubercles present. morphology does not vary in the small series we Macrosetae in lateral border of the carapace in have had the chance to examine. Subadult females multiple rows. Femora without prolateral black have pre-spermathecae with four branches grouped setae. Sternum length 2.98, width 3.32, cordiform. in the middle of the bursa (Fig. 47E, F). Femoral thorns: 4. Leg I: femur 9.4 long, patella 2.8, tibia 9.7, metatarsus 7.5, tarsus 4. II: fe 11.42, ti Distribution: Known only from the Peruvian Andes, 11.42. III: fe 10.25, ti 9.5. IV: fe 9.7, pa 2.7, ti 8.9, mt in the province of Huanuco (Fig. 27B). 7.8, ta 3.9. Abdomen: length 6.2, width 5.8. Palp as in Figure 48A and B. Additional material examined: PERU. Huanuco: Female (San Antonio de Parapetı, Santa Cruz, Acomayo [ 76.05, 9.46, 2100 m], W. Weyrauch, III/ Bolivia, AMNH). As in male, except for the following. 1940, 1 imm.À (AMNH);À same data as holotype, 6 Total length 16.32. Carapace length 8.2, anterior imm. (AMNH). width 3.29, thoracic region width 8.29. Eye diameters and interdistances: PME 0.23, ALE 0.23, PLE 0.25, PME–PLE 0.74, ALE–PLE 0.36. Clypeus height SICARIUS BOLIVIENSIS SP. NOV. 1.58. Sternum length 3.4, width 3.5, cordiform. Figures 18E, 26B, 31E, 32E, 33J, 48, 49 Femoral thorns: 4. Leg I: femur 9.6 long, patella 3.1, Figure 48. Sicarius boliviensis sp. nov., male genitalia in prolateral view. A, holotype from Lagunillas, Santa Cruz, Bolivia (MZSP 48990). B, same, cymbium and bulb. C, Corumba, Mato Grosso do Sul, Brazil (MZSP 8261). D, Quispi- Sicarius terrosus: Chamberlin, 1916: 214, plates 9–10 tibia 9.8, metatarsus 7.1, tarsus 4.1. II: fe 10.63, ti canchi, Cusco, Peru (UNSAAC). Abbreviation: SA, sclerotized area. Scale bars = 0.1 mm (except A, 1 mm). (misidentified, probably this species). 10.38. III: fe 10, ti 9.3. IV: fe 9.5, pa 2.6, ti 8.9, mt 7.5, ta 4. Abdomen: length 7.86, width 7.95. Sper- Note: We could not re-examine Chamberlin’s speci- mathecae branches: 8. Ventral branches absent. VII/1946, 1 # 1 imm. (AMNH); J.C. Pallister, 27/I/ sternum orange, legs brownish orange, abdomen men (MCZ 20753) before it was lost at the fire at Spermathecae as in Figure 49A and B. 1947, 1 # (AMNH). brownish yellow. Total length 9.8. Carapace length Instituto Butantan in 2010, but the locality data 5.32, anterior width 1.98, thoracic region width 5.07. (Santa Ana, Peru) and sternum morphology indicate Variation: Males (N = 5): carapace length 6.48–7.69 Etymology: The name is an adjective referring to the Eye diameters and interdistances: PME 0.18, ALE it most probably refers to this species. (7.01), femoral thorns 4–5, femur II length 9.8– Andean distribution of this species. 0.18, PLE 0.18, PME–PLE 0.51, ALE–PLE 0.18. 12.16 (11.08). Females (N = 5): carapace length Clypeus height 0.86. Eye tubercles present. Type material: Male holotype from BOLIVIA. 7.17–8.2 (7.58), femoral thorns 3–5, femur II length Diagnosis: Males can be distinguished by the long Macrosetae in lateral border of the carapace in Santa Cruz: Cordillera, Lagunillas ( 63.68583, 8.5–10.63 (9.09), spermathecae branches 4–8. Males À and robust embolus set off at approximately 90° from multiple rows. Femora without prolateral black 19.64858, 1132 m), R. Pinto da Rocha et al., 4/XII/ vary in the length of the embolus (Fig. 48); females À the rest of the bulb (Fig. 47A, B); they are somewhat setae. Sternum length 2.49, width 2.52, cordiform. 2010 (MZSP 48990). from Peru have a very subtle reservatory at the similar to those of S. diadorim and S. jequitinhonha Femoral thorns: 4. Leg I: femur 6.95 long, patella apex of the spermathecae. It might be that sp. nov. (Fig. 58), but differ by the more robust 2.19, tibia 7.67, metatarsus 6, tarsus 2.64. II: fe 8.25, Etymology: The name is an adjective referring to the populations from Cusco represent a distinct species, embolus and by lacking prolateral black setae in the ti 8.5. III: fe 6.83, ti 6.61. IV: fe 82, pa 2.03, ti 6.68, country from where most of the records of this but additional material is necessary to elucidate femora. Females can be distinguished by the mt 5.86, ta 2.49. Abdomen: length 4.67, width 4.46. species come. that. Subadult females have pre-spermathecae with spermathecae with six finger-like branches slightly Palp as in Figure 47A and B. short, triangular branches (asymmetric in some convoluted and having a small reservatory at the Female paratype from Ambo, Huanuco, Peru Diagnosis: Males, females and immatures can be cases), with a tiny reservatory at the apex apex; and by the the most ectal branches well (AMNH). Coloration as in male, except for reddish recognized by the strong sclerotization around the (Fig. 49I, J). separated from the medians and pointing laterad brown carapace and light brown legs. Total length most proximal femoral thorn (Fig. 48A, sa). Males (Fig. 47C, D). 12.38. Carapace length 5.93, anterior width 2.41, tho- are further distinguished by the robust bulb with Distribution: Dry forests in Bolivia, western Brazil, racic region width 5.86. Eye diameters and interdis- very short embolus (Fig. 48). Females are northern Paraguay and Cusco, Peru, (Fig. 26B). This Description: Male holotype (Acomayo, Huanuco, tances: PME 0.2, ALE 0.23, PLE 0.23, PME–PLE distinguished by the finger-like, robust branches in species occurs next to the Argentinean northern Peru, AMNH). Coloration: carapace reddish, 0.58, ALE–PLE 0.31. Clypeus height 1.17. Eye tuber- the spermathecae, with rounded reservatories absent border, and thus its presence in this latter country is chelicerae, labium and endites reddish brown, cles present. Macrosetae in lateral border of the or very subtle, and with the glandular pores large not unlikely.

Additional material examined: PERU. Cusco: females: we revised female specimens identified by Quispicanchis, [ 71.95075, 13.5311, 3419 m], C. them as ‘S. patagonicus’ and the vial contained À À Mart�ınez, 1973, 1 # (MACN-Ar 30216); same locality, both S. rupestris and S. mapuche sp. nov. As a [ 71.93333, 13.51667, 3419 m], Huacarpay, 15/III/ consequence, the female they illustrated as À À 2012, 1 # 1 $ 10 imm. (UNSAAC). BOLIVIA. S. patagonicus is not co-specific with the males Cochabamba: Campero, Marquilla, E. Aiquile they considered to be the same species (see their [ 65.16667, 18.18333, 2200 m], 10/II/1976, 1 # 3 $ plate 2). Some subsequent authors apparently used (AMNH);À PeÀna~ Colorado [ 64.86667, 18.16667, their work to identify Argentinean specimens (e.g. À À 1600 m], 13/II/1976, 3 $ (AMNH); Charamoco Binford et al., 2008, 2009), spreading misidentifica- [ 66.283, 17.6, 2438 m], J.A. Vellard, VII/1940, 1 $ tions. À À (MACN-Ar 3697) (spermathecae in very bad state of We here describe two new species, Sicarius levii conservation). Santa Cruz: Cordillera, Lagunillas sp. nov. (Gerschman & Schiapelli’s ‘terrosus’) and ( 63.6775, 19.65167, 1132 m), R. Pinto-da-Rocha Sicarius mapuche sp. nov. (Gerschman & Schiapelli’s À À et al., 4/XII/2010, 1 imm. (MZSP 48991); San Antonio ‘patagonicus’, in part). Sicarius rupestris is rede- de Parapeti [ 63.21667, 20.01667, 842 m], B. scribed and material from the type locality is illus- À À Malkin, 1991, 2 # 2 $ (AMNH); Nuflo de Chavez, La trated, but its resolution here is far from satisfactory Bola [ 61.58333, 17.38333, 264 m], P.A. Goloboff as it seems clear that it might have to be split into À À et al., 8/I/1991, 1 $ (AMNH). Chuquisaca: at least two or three species in the future. However, Hernando Siles, E. Monteagudo [ 63.95, 19.8, we do not count on enough material to resolve this À À 1600 m], L.E. Pena,~ 21–24/XII/1984, 1 $ (AMNH). matter at present, as a very large collection of Sicar- BRAZIL. Mato Grosso do Sul: Corumba, Serra do ius from Argentina was lost in the fire at Instituto Urucum [ 57.60047, 19.19924, 662 m], C. Lenko, Butantan in 2010 before we could examine them, À À 25–30/XI/1960, 1 # (MZSP 8261). PARAGUAY. leaving us with scarce material. Boqueron:� Filadelfia, 19 km N Filadelfia [ 60.05, 22.16667, 138 m], K.L. Anderson, 27/IX/1978,À 1 À SICARIUS RUPESTRIS (HOLMBERG, 1881) imm. (AMNH); 5/X/1978, 1 imm. (AMNH), 1 # 1 $ (AMNH); F. Jaggli, 1 # 1 $ (IBSP 166121). Figures 15H, 26C, 50, 51, 52 Thomisoides rupestris Holmberg, 1881: 153, plate 4, fig. 9; Mello-Leitao,~ 1940: 8, figs 6, 7. Syntypes from ARGENTINEAN SPECIES Sierra Pichy-Mahuida, Argentina, lost (see Ger- Sicarius taxonomy in Argentina is rather confus- schman & Pikelin, 1979). ing, mainly because previous authors studying its Sicarius rupestris: Tullgren, 1901: 187; Gerschman species did not examine types and misidentified de Pikelin & Schiapelli, 1979: 88, fig. 1c; Labarque & species. Mello-Leitao~ (1941) erroneously identified Ram�ırez, 2012: 3, figs 2F, 5C–D, 8C–D, 9F, 10F, 11F, S. levii sp. nov. as S. terrosus, and was followed by 12F, 13F, 14F, 15F, 16F, 17F, 18F, 19F, 20F, 26A–F, subsequent authors (Gerschman de Pikelin & Schi- 27A, B, 30E. apelli, 1979; Binford et al., 2008, 2009). To make Thomisoides patagonicus Simon, in Mello Leitao,~ things worse, Gerschman de Pikelin & Schiapelli 1919: 475, fig. 1. Syntypes from Patagonia, Argen- (1979) made a regional revision of the genus that tina, allegedly in MZSP (Mello Leitao,~ 1919), not only added to the confusion. They recognized the found, lost (R. Pinto da Rocha, pers. comm.); Mello- same three species we do, but, despite their efforts, Leitao,~ 1940: 8, fig. 8; Gerschman de Pikelin & could not examine any types apart from that of Schiapelli, 1979: 88, fig. 1a. SYN. NOV. S. irregularis (Mello-Leitao,~ 1941). Thus, they erro- Thomisoides irregularis Mello-Leitao,~ 1940: 9, fig. 9. neously considered S. levii sp. nov. to be S. terro- Female holotype from Arroyito, Neuquen,� Argentina, sus (as did Mello-Leitao,~ see S. terrosus in their deposited in MLP 14270, examined. Synonymized fig. 1) and S. mapuche sp. nov. to be S. patagoni- under S. patagonicus by Gerschman de Pikelin & cus (see palp SEM image in their plate 2); on the Schiapelli, 1979. SYN. NOV. other hand, they correctly recognized S. rupestris. Sicarius patagonicus: Gerschman de Pikelin & However, they clearly had trouble identifying Schiapelli, 1979: 88, fig. 1a.

Figure 49. Sicarius boliviensis sp. nov., female genitalia, digested. A, San Antonio del Parapeti, Santa Cruz, Bolivia (AMNH), dorsal. B, same, ventral. C, Filadelﬁa, Boqueron,� Paraguay (AMNH), dorsal. D, same, ventral. E, Chuquisaca, Bolivia (AMNH), dorsal. F, same, ventral. G, Quispicanchi, Cusco, Peru (AMNH), dorsal. H, same, ventral. I, subadult from Pena~ Colorado, Cochabamba, Bolivia (AMNH), ventral. J, subadult female from La Bola, Santa Cruz, Bolivia (AMNH), ventral. Scale bars = 0.1 mm.

Figure 50. Sicarius rupestris (Holmberg, 1881) from Argentina, male genitalia in prolateral view. A, Caleta Olivia, Santa Cruz (UCCC). B, same, cymbium and bulb. C, Languineo, Chubut (AMNH). D, Uspallata, Mendoza (CAI 3047). E, Pichi Mahuida, R�ıo Negro (type locality) (MACN 33863). F, Pampa de Achala, Cordoba� (IBSP 36170). G, Santa Rosa, Mendoza (CAI 3062). H, Sicarius patagonicus Simon in Mello-Leitao,~ 1919, reproduction of original drawing by Mello- Leitao~ (1919). Scale bars = 0.1 mm (except A, 1 mm).

Synonymy: The types of S. patagonicus could not be holotype (Fig. 51C, D) and it falls within the found and the type locality is unspeciﬁc (‘Patagonia’). variation we recognize for S. rupestris. However, Mello-Leitao’s~ illustrations of a male with a long, S. irregularis may eventualy be considered valid thin, curved embolus (Fig. 50H) do not really ﬁt any when the limits of S. rupestris are re-evaluated (see of the specimens we examined, except perhaps a variation below); we refrain from revalidating it at deviant male from Mendoza (Fig. 50G), but some of this time as we presently cannot diagnose it from his illustrations are known to be innacurate. S. rupestris. One male (Fig. 50E) and one female we Figure 51. Sicarius rupestris (Holmberg, 1881) from Argentina, female genitalia, digested. A, Pichi Mahuida, R�ıo Negro Regardless, his illustration is certainly more similar illustrated (Fig. 51A, B) come from the type locality (type locality) (MACN 33864), dorsal. B, same, ventral. C, holotype of Thomisoides irregularis Mello-Leitao,~ 1940 from to males of S. rupestris than to any of the other two of S. rupestris. Arroyito, Neuquen� (MLP 14270), dorsal. D, same, ventral. E, Uspallata, Mendoza (CAI 3048), dorsal. F, same, ventral. Argentinean species. Gerschman de Pikelin & G, Malarque,€ Mendoza (CAI 3045), dorsal. H, same, ventral. I, Punta de Vacas, Mendoza (CAI 3129), dorsal. J, same, Schiapelli (1979) had placed S. irregularis as a Diagnosis: Males differ from other Argentinean ventral. K, Languineo, Chubut (AMNH), dorsal. L, same, ventral. Scale bars = 0.1 mm. synonym of S. patagonicus. We have examined its species, especially the similar S. mapuche sp. nov. by

and interdistances: PME 0.18, ALE 0.18, PLE 0.18, Gigantes ( 64.59257, 31.37536, 899 m), A.V. À À PME–PLE 0.33, ALE–PLE 0.15. Clypeus height 0.69. Peretti, VI/2013, 1 # 4 imm. (UFMG 15823), 1 # Sternum length 2.01, width 2.11, cordiform. Femoral (UFMG 15824); VIII/2013, 1 # 3 imm. (IBSP thorns: 3. Leg I: femur 4.16 long, patella 1.75, tibia 166389), 2 $ (IBSP 166390); VII–VIII/2013, 2 $ 4.04, metatarsus 2.92, tarsus 2.32. II: fe 4.46, ti 3.91. (UFMG 15822); R�ıo Primero, Camino del Cuadrado, III: fe 4.36, ti 3.59. IV: fe 4.7, pa 1.58, ti 3.71, mt La Falda ( 63.57367, 30.8765, 1295 m), G.D. Rubio À À 2.26, ta 2.29. Abdomen: length 4.36, width 4.06. Sper- et al., 17/II/2013, 1 # 4 imm. (MACN-Ar 30187*), 1 $ mathecae branches: 16. Ventral branches absent. (MACN-Ar 30207*), 1 # (MACN-Ar 30208*); San Spermathecae as in Figure 51C and D. Alberto, Pampa de Achala ( 64.86667, 31.6, 2184 m), C.A.Rheims, A.J.SantosÀ & M.O.Gonzaga,À 2/ Variation: Males (N = 7): carapace length 4.46–6.14 III/2002, 1 # (IBSP 36168*), 1 # 1 $ (IBSP 36170*); (5.12), femur II length 5.59–8.5 (6.81). Females Santa Mar�ıa, Villa Carlos Paz, 2 km NE Villa Carlos (N = 5): carapace length 4.16–6.14 (5.47), femoral Paz [ 64.43333, 31.38333, 810 m], F.A. Coyle À À thorns 3–4, femur II length 4.46–6.75 (5.96), et al., 15/III/1988, 1 $ (AMNH); P.A. Goloboff, 16/VII/ spermathecae branches 8–16. Genital morphology 1979, 1 # (MACN-Ar 30230), 1 # (MACN-Ar 32321*). and total length are painfully variable (Figs 50, 51), San Juan: Calingasta, 8 km S Tocota [ 69.41667, À suggesting this species might possibly be a complex 30.66667, 2490 m], 1 $ 2 imm. (CAI 3132-4); Sierra of 2–3 independent lineages. This is a problem we Àdel Tontal, Camino a la antena, 16.91 km E de cannot resolve with the material at hand, and that Barreal ( 69.27496, 31.62592, 2686 m), C. À À will be probably be better addressed using an Grismado et al.,4–6/XI/2013, 1 $ (MACN-Ar 30214), integrative approach in the future. Specimens 1 # (MACN-Ar 30215), 1 # 2 $ 2 imm. (MACN-Ar marked with an asterisk (*) in the list of material 30955), 1 # 5 $ 3 imm. (MACN-Ar 31010). Mendoza: examined are some of those found by us to be the Las Heras, Quebrada del Toro ( 69.064, 32.48381, À À most similar to the population from the type locality. 2735 m), C.J. Grismado et al., 30/VIII/2008, 1 $ (MACN-Ar 30232); Uspallata, 14 km NW Tambillos Natural history: Specimens have been collected [ 68.86667, 32.86667, 760 m], S. Roig & S. Lagos, À À from monte (a xeric scrubland typical of Argentina) 13/I/1999, 1 $ (CAI 3154); Puma - Proy. Potrerillos and grasslands, usually under rocks. We have [ 69.33333, 32.58333, 2087 m], S. Lagos, 20/VI/ À À observed courtship and mating behaviour of a pair 2001, 1 # 1 $ (CAI 3047-8*); Lujan,� Las Heras, from Sierra del Tontal, Argentina (Fig. 52). It is Punta de Vacas [ 69.75, 32.85, 2408 m], G. Flores À À quite simple, with the male approaching the female, & A. Scollo, 13–23/II/2006, 1 # (CAI 3091); same touching her with the first two pairs of legs; the locality ( 69.7663, 32.85588, 2452 m), S. Claver & À À female responds by bending her abdomen upwards. A. Scollo, 4–14/XII/2004, 1 # (CAI 3065), 1 $ (CAI The male then embraces her and inserts both 3129); Malarque,€ Agua Escondida [ 68.30307, Figure 52. Sicarius rupestris (Holmberg, 1881), pair from Sierral del Tontal, San Juan, Argentina, courtship and cop- À pedipalps simultaneously, as described by Levi & 36.15024, 1129 m], A. Roig, 29/I/1979, 1 $ (MACN- ula (male to the left/top, female to the right/bottom). A, male approaches female. B, male starts touching female with his À Levi (1969) for Sicarius levii sp. nov. and by Ar 32323); Nevado Medio ( 68.51308, 35.13562, first legs. C, female assumes a vertical position and starts bending her abdomen towards the male. Male grasps female’s À À Magalhaes~ et al. (2013) for S. saci sp. nov. 2940 m), G. Debandi & E. Ruiz, 8–17/II/2005, 1 legs with his first legs, and starts touching her with his second legs. D, female bends her abdomen so as the venter is $ (misidentified in the latter work as S. cariri). This (CAI 3045); Santa Rosa, Nacu~ n~an� [ 67.96667, facing upside, while the cephalothorax is in a completely vertical position. The male grasps the female with all his legs, À mating was apparently not successful, however, as 34.03333, 589 m], S. Lagos, 7/II/1998, 1 # (CAI except the fourth. The male introduces both pedipalpal bulbs at the female’s genital opening at once (inset in E). À the female did not produced an eggcase even 3062). San Luis: Jun�ın, Merlo, Mirador del Sol several months after mating. [ 65.03112, 32.3494, 839 m], A. Porta, 1 $ 1 imm. À À the curved and thin embolus (robust and shorter in PLE 0.2, PME–PLE 0.43, ALE–PLE 0.15. Clypeus (MACN-Ar 30211*); M.E. Galiano, 22/XI/1983, 1 $ 3 S. mapuche) and by the body of the bulb not so height 0.69. Eye tubercles absent. Macrosetae in Distribution: Argentinean arid lands from imm. (MACN-Ar 30229); Libertador General San strongly curved. Females can be distinguished from lateral border of the carapace in a single row. Catamarca to Chubut (Fig. 26C). Mart�ın, San Felipe [ 65.46328, 32.80287, 844 m], À À other Argentinean species by the strongly coiled Femora without prolateral black setae. Sternum Luchini, 9/X/1960, 1 $ 1 imm. (MACN-Ar 30223); 23/ branches of the spermathecae with a small length 2.47, width 2.21, cordiform. Femoral thorns: Additional material examined: ARGENTINA. X/1960, 1 $ (MACN-Ar 30224); 28/X/1960, 1 $ 1 imm. reservatory at the apex (straight branches, large 4. Leg I: femur 6.2 long, patella 2.21, tibia 5.93, Catamarca: Andalgala,� Cuesta Minas, 5 km S (MACN-Ar 30222); 1 $ (MACN-Ar 30225). La reservatory in S. mapuche sp. nov.). metatarsus 5.02, tarsus 3.39. II: fe 6.82, ti 6.07. III: Capillitas [ 66.37802, 27.37044, 2337 m], E. Pampa: Curaco,� RN 152 ( 66.48925, 38.14441, À À À À fe 6.41, ti 5.52. IV: fe 6.34, pa 1.71, ti 5.59, mt 4.91, Maury, 1/I/1981, 1 $ (MACN-Ar 32324). Cordoba:� 283 m), A.D. Brescovit et al., 11/X/2014, 1 # 1 imm. Description: Male (Caleta Olivia, Santa Cruz, ta 3.3. Abdomen: length 4.57, width 4.11. Palp as in no further data [ 64.18248, 31.40337, 415 m], J. (MACN-Ar 34247); Lihuel Calel, Estacion� de servicio À À Argentina, UCCC 362). Coloration: carapace Figure 50A and B. Bretes, 1924, 1 # (MACN-Ar 6791); Pocho, El sobre la ruta 152, cercana al PN Lihue Calel chelicerae, labium and endites reddish brown, Female (Arroyito, Neuquen, Argentina, MLP Potrero, 35 km W Salsacate [ 65.31479, 31.31492, ( 65.58408, 38.01265, 346 m), 1 $ 1 imm. (MACN- À À À À sternum and legs orange, abdomen brownish grey. 14270, holotype of Thomisoides irregularis). As in 1184 m], Maury, 5/II/1981, 1 $ 3 imm. (MACN-Ar Ar 34092), 1 $ 1 imm. (IBSP 167028*), 1 $ 1 imm. Total length 8.5. Carapace length 5.11, anterior male, except for the following. Abdomen light brown. 32325*); Punilla, Observatorio Astronomico� Bosque (UFMG 19265*); Parque Nacional Lihue Calel, cerro width 1.93, thoracic region width 5.07. Eye Total length 8.25. Carapace length 4.16, anterior Alegre [ 64.56893, 31.60613, 1043 m], P.A. Fortaleza ( 65.59225, 38.02075, 384 m), 1 # À À À À diameters and interdistances: PME 0.18, ALE 0.2, width 1.58, thoracic region width 3.92. Eye diameters Goloboff, IX/1987, 1 $ (MACN-Ar 32322); Tanti, Los (MACN-Ar 35932*), 1 $ (MACN-Ar 35933*), 1 $

(MACN-Ar 35934*), 1 $ (MACN-Ar 35935*), 1 # Leitao’s~ (1919) illustration does not match its palpal (IBSP 167029*), 1 $ (IBSP 167030*), 1 # (UFMG morphology. 19266*), 1 $ 1 imm. (UFMG 19267*), 1 # (MACN-Ar 34248*); RN 152 km 30 ( 65.45084, 37.86591, Type material: Male holotype from ARGENTINA. À À 238 m), 1 # 1 imm. (IBSP 167031*), 1 # 1 imm. R�ıo Negro: Los Lagos, Cerca del Embalse Alicura� (UFMG 19268*); Lihuel Calel, 1 1 imm. (MACN-Ar (LT3S7E5) ( 70.88539, 40.58961, 772 m), P. # À À 34246*). Buenos Aires: Patagones, Carmen de Sackmann, I/2005 (MACN-Ar 33860). Paratypes: Patagones ( 63.02, 40.78, 37 m), H.A. Iuri, 1 $ (no same collecting data, 1 # 1 imm. (MACN-Ar 33861), À À voucher); Puan, Arroyo Chasico[� 63.04635, 1 $ (MACN-Ar 33862); Mendoza: Papagallo, no À 38.56103, 5 m], Spivak & Goloboff, 13/VI/1980, 1 # collector, no date, 1 # (MLP 15584). À (MACN-Ar 30217*), 1 # (MACN-Ar 30218*), 1 # (MACN-Ar 30220*), 1 # (MACN-Ar 30221*); 14/VI/ Etymology: The specific name honours the 1980, 1 $ (MACN-Ar 30219*); Tornquist, Sierra de la Mapuches, a group of Native American people Ventana [ 61.79403, 38.13943, 279 m], A. inhabiting portions of southern Chile and Argentina. À À Ruggiaro, 22–27/X/1981, 1 $ (MACN-Ar 32320). The name is to be treated as a noun. Neuquen:� Confluencia, Arroyito [ 68.58278, À 39.07889, 302 m], M. Biraben,� 1 $ (MLP 14270). Diagnosis: Males are similar to S. rupestris by the À R�ıo Negro: El Cuy, Paso Cordoba� [ 67.62524, smooth transition between the base and the body of À 39.12836, 403 m], E. Maury, I/1974, 1 # (MACN-Ar the bulb, but differ by the curved, robust and short À 6914); Pichi Mahuida, Pichi Mahuida, RP57 embolus (thinner and longer in S. rupestris) and ( 64.95407, 38.8408, 158 m), A.D. Brescovit et al., body of the bulb very curved (Fig. 53A–D) (straighter À À 15/X/2014, 1 # (MACN-Ar 33863), 1 $ 1 imm. (IBSP in S. rupestris). Females can be distinguished from 167033*), 1 $ (MACN-Ar 33864*); San Antonio, other Argentinean species by the long and straight Playas Doradas ( 65.02467, 41.64305, 16 m), 14/X/ stalks of the spermathecae with a large, oval À À 2014, 1 # (MACN-Ar 35929); Valcheta, RP 58 reservatory at the apex (strongly coiled, small ( 65.81116, 40.92843, 258 m), 13/X/2014, 1 # reservatory in S. rupestris; stalks absent in S. levii À À (MACN-Ar 35930), 1 # (IBSP 167032), 1 # (UFMG sp. nov., Figure 53F–K). 19269); same locality ( 65.79502, 40.88402, À À 239 m), 1 $ (MACN-Ar 34093); same locality Description: Male (holotype from Embalse Alicura,� ( 65.81116, 40.92843, 258 m), 1 $ (MACN-Ar Neuquen,� Argentina, MACN-Ar 33860). Coloration: À À 34244); Valcheta ( 66.1457, 40.6884, 183 m), H.A. carapace chelicerae, labium and endites reddish À À Iuri, 2013, 1 $ (MACN-Ar 30186). Chubut: brown, sternum, legs and abdomen brownish yellow. Languineo, E. Manantiales [ 70.13333, 43.4, Carapace length 3.86, anterior width 1.47, thoracic À À 1118 m], L.E. Pena,~ 6–10/XI/1985, 1 # 3 $ (AMNH). region width 3.72. Eye diameters and interdistances: Santa Cruz: Deseado, Caleta Olivia, 20 km N PME 0.13, ALE 0.15, PLE 0.13, PME–PLE 0.3, ALE– Caleta Olivia [ 67.60583, 46.26306, 22 m], T. PLE 0.1. Clypeus height 0.43. Eye tubercles absent. À À Cekalovic, 13/II/1962, 1 # 1 imm. (UCCC); Cerro Pico Macrosetae in lateral border of the carapace in a Truncado ( 68.13733, 46.84568, 163 m), A. Porta, single row. Femora without prolateral black setae À À 3/XI/2014, 1 $ (MACN-Ar 34250). Sternum length 1.73, width 1.8, cordiform. Femoral thorns: 4. Leg I: femur 3.99 long, patella 1.4, tibia 3.86, metatarsus 2.86, tarsus 2.26. II: fe 4.52, ti 3.99. SICARIUS MAPUCHE SP. NOV. III: fe 3.99, ti 3.72. IV: fe 4.39, pa 1.3, ti 3.72, mt 2.66, ta 2.39. Abdomen: length 3.46, width 3.19. Palp Figures 18C, 26D, 31D, 32D, 33I, 53 as in Figure 53C. Sicarius patagonicus: Gerschman de Pikelin & Schi- Female (Valle de Altares, Chubut, Argentina, apelli, 1979: 88, plate 2, figs 7, 8, 11 (misidentified). AMNH). As in male, except for the following. Cara- Note: Gerschman de Pikelin & Schiapelli (1979) pace light reddish brown. Total length 11.38. Cara- identified this species as S. patagonicus, but Mello- pace length 4.26, anterior width 1.98, thoracic region

Figure 53. Sicarius mapuche sp. nov. from Argentina. A–C, male genitalia in prolateral view; D–H, female genitalia, digested. A, Valle de los Altares, Chubut (AMNH). B, same, cymbium and bulb. C, male holotype from Embalse Alicura,� Neuquen� (MACN 33860). D, San Carlos, Mendoza (CAI 3085). E, subadult from Valle de los Altares, Chubut (AMNH), ventral. F, Valle de los Altares, Chubut (AMNH), dorsal. G, same, ventral. H, San Carlos, Mendoza (CAI 3078), dorsal. I, same, ventral. J, female paratype from Embalse Alicura,� Neuquen� (MACN 33861), dorsal. K, same, ventral. Scale bars = 0.1 mm (except A, 1 mm).

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016 © 20172016 The Linnean Society Society of of London, London, ZoologicalZoological Journal Journal of of the the Linnean Linnean Society Society, 2017,, 2016 179, 767–864 83872 I. I. L. L. F. F. MAGALHAES MAGALHAESET ET AL. AL. SICARIID PHYLOGENY AND REVISION OF SICARIUS 73 width 4.11. Eye diameters and interdistances: PME et al., 16–31/I/2015, 1 $ (MACN-Ar 34089), 1 $ 0.13, ALE 0.13, PLE 0.15, PME–PLE 0.48, ALE–PLE (MACN-Ar 34090). R�ıo Negro: El Cuy, Paso 0.15. Clypeus height 0.66. Sternum length 1.91, Cordoba� [ 67.62524, 39.12836, 403 m], R. À À width 2.19, cordiform. Femoral thorns: 4. Leg I: Bachmann, III/1959, 1 $ (MACN-Ar 6912); femur 4.04 long, patella 1.58, tibia 3.75, metatarsus Pilcaniyeu, Cerro Alto [ 70.85832, 40.86466, À À 2.64, tarsus 2.34. II: fe 4.41, ti 3.8. III: fe 4.36, ti 1276 m], E. Maury & Toth, I/1975, 1 # 2 $ (MACN- 3.51. IV: fe 4.31, pa 1.6, ti 3.63, mt 2.64, ta 2.37. Ar 6783). Chubut: Martines, Rocky Tripp, 70 km Abdomen: length 6.83, width 6.14. Spermathecae Las Plumas [ 67.25742, 43.70216, 282 m], C. À À branches: 11.Ventral branches absent. Spermathecae Castro, 12/IV/1970, 1 # (MACN-Ar 98); Paso de as in Figure 53J and K. Indios, Valle de los Altares [ 68.4, 43.88333, À À 319 m], L.E. Pena,~ 5/XI/1990, 2 # 1 $ (AMNH); 3–5/ Variation: Males (N = 6): carapace length 3.47–5.76 XI/1990, 1 $ 3 imm. (AMNH); Puerto Madryn (4.12), femoral thorns 3–4, femur II length 4.21–4.82 [ 65.05828, 42.76138, 37 m], Cosztonyi, III/1969, 1 À À (4.48). Females (N = 5): carapace length 3.8–4.26 # (MACN-Ar 6918); E. Maury, I/1975, 1 # (MACN-Ar (4.14), femoral thorns 3–4, femur II length 3.92–4.41 6913); Puerto Piramides� [ 64.27207, 42.57173, À À (4.23), spermathecae branches 7–11. Genital 67 m], P.A. Goloboff, 5/II/1980, 2 # 1 $ (MACN-Ar morphology is slightly variable in both sexes (53). 30233). Female subadults have pre-spermathecae with 4–5 reduced branches, not globose at tip (Fig. 53E). SICARIUS LEVII SP. NOV. Distribution: Arid areas in northern Patagonia, in Figures 2, 5, 9, 16A, B, 18F, 26D, 30C, 31F, 32F, the provinces of Chubut, Neuquen,� R�ıo Negro and 33K, 54 Mendoza, Argentina (Fig. 26D). Thomisoides terrosus: Mello-Leitao,~ 1941: 108, fig. 4 (misidentified). Additional material examined: ARGENTINA. Sicarius sp. 1: Levi, 1967: 322; figs 1–3; Levi & Levi, Mendoza: Malargue,€ Fort�ın Malargue€ [ 69.5798, 1969: 31, figs 1–15. À 35.47211, 1408 m], A. Roig A., 3/IV/1976, 1 $ 4 Sicarius sp. 1: Reiskind, 1966: 195. À imm. (MACN-Ar 30234); Malarque,€ Sierra Palauco Sicarius terrosus: Gerschman de Pikelin & Schia- ( 69.50578, 36.1298, 2457 m), F. Campon,� 27/I/ pelli, 1979: fig. 1b, plate 1, 4-5 (misidentified). À À 2010, 1 $ (CAI 3131); San Carlos, Reserva Laguna del Diamante, Alvarado ( 69.37978, 34.24617, Note: Reiskind (1966), Levi (1967) and Levi & 2337 m), A. Marvaldi & G. Debandi,À 4–14/XII/2004,À 1 Levi’s (1969) illustrations and locality data indicate # (CAI 3093); G. Debandi & A. Marvaldi, 3 # (CAI that the species they studied is this one; their vouch- 3074-6), 1 $ (CAI 3092), 1 # (CAI 3130); G. Debandi ers were deposited in the MCZ and were destroyed & S. Claver, 8–17/II/2005, 1 # 1 imm. (CAI 3070-1); in the fire at the Instituto Butantan before we could same locality ( 69.37789, 34.24407, 2362 m), 1 # examine them. This species is probably the largest (CAI 3098); sameÀ localityÀ ( 69.37978, 34.24617, known haplogyne spider in terms of body length and À À 2337 m), G. Flores et al., 17/XII/2012, 1 # (CAI body mass, although some Central American Scy- 3173); S. Claver & G. Debandi, 8–17/II/2005, 1 # todes might have a larger leg span. (CAI 3077), 1 # (CAI 3079), 3 # (CAI 3088-90); S. Claver & R. Carrara, 13–23/II/2006, 1 # (CAI 3094); Type material: Male holotype from ARGENTINA. S. Roig & G. Debandi, 1–5/I/2006, 1 # (CAI 3095), 1 R�ıo Negro: El Cuy, RP 6 ( 67.79195, 39.25879, À À # (CAI 3096); 5–8/I/2006, 1 # 2 imm. (CAI 3066-8), 1 309 m), A.D. Brescovit et al., 12/X/2014 (MACN-Ar $ (CAI 3069), 2 # 1 imm. (CAI 3080-2), 2 # (CAI 33859). Paratypes: same province, El Cuy, Paso 3085-6); 23–25/II/2006, 1 $ (CAI 3078); 5/VIII/2006, 2 Cordoba� ( 67.61586, 39.14034, 403 m), H.A. Iuri, À À $ (CAI 3072-3); same locality ( 69.37789, 34.24407, 20–21/I/2015, 1 # (MACN-Ar 34200), 1 # (MACN-Ar À À 2362 m), 1 # (CAI 3087); same locality ( 69.37978, 34201), 1 # (MACN-Ar 34202), 1 $ 5 imm. (MACN- À 34.24617, 2337 m), 26–31/XII/2006, 1 # (CAI, Ar 34203); General Roca, Cinco Saltos, camino a 2097);À same locality ( 69.37978, 34.24617, Lago Pellegrini ( 68.0519, 38.81157, 311 m), A.D. À À À À 2709 m), XII/2004, 2 # 1 $ (CAI 3101-3); San Rafael, Brescovit et al., 16/X/2014, 1 # 1 $ 3 imm. (MACN- Sosneado [ 68.34203, 34.61649, 716 m], E. Maury, Ar 33865); Santiago del Estero: Campo Gallo Figure 54. Sicarius levii sp. nov. from Argentina. A–D, male genitalia in prolateral view; E–I female genitalia, À À I/1975, 1 $ 1 imm. (MACN-Ar 6917). Neuquen:� ( 62.85, 26.5833), J. Abalos,� no date, 3 # (MLP digested. A, male holotype from El Cuy, Rıo Negro (MACN 33859). B, same, cymbium and bulb. C, Rıo Salı, Tucuman À À Anelo,~ Punta Carranza [ 68.78251, 38.35056, 13736). (AMNH). D, Lavalle, Mendoza (CAI 3049). E, subadult female from Mendoza (MHNS), ventral. F, La Curtiembre, Salta À À 440 m], O. De Ferrariis, 23/XII/1976, 1 $ (MACN-Ar (AMNH), dorsal. G, same, ventral. H, Lavalle, Mendoza (CAI 3050), dorsal. I, same, ventral. J, paratype from Cinco Sal- 30226); Chos Malal, Parque Provincial Tromen Etymology: This species, the largest sicariid, is tos, Rıo Negro (MACN 33865), dorsal. K, same, ventral. DHT, dorsal setae tuft; VB, ventral branch; VHT, ventral setae ( 70.12253, 37.08607, 2184 m), P.D.P. Pinheiro named in honour of the recently deceased Dr tuft. Scale bars = 0.1 mm (except A, 1 mm). À À

Herbert W. Levi, a giant among arachnologists. This morphology is slightly variable in both sexes (MACN-Ar 6909); Santa Marıa, Puenta Balasta Ar 34245); same locality ( 67.66825, 39.12691, À À is in recognition of his huge contributions to (Fig. 54). Female subadults with spermathecal (Punta de Balastro?), 15 km N of Puenta Balasta 380 m), 15/X/2014, 1 imm. (IBSP), 1 imm. (IBSP), 1 arachnology in the form of his beautiful, excellent branches unformed, with two triangular outgrowths [ 66.28333, 26.93333, 2000 m], L.E. Pena,~ 13/II/ imm. (UFMG), 1 imm. (MACN-Ar 34249); RP 6 À À taxonomic revisions and his ever inspiring example in the place of the future spermathecal branches 1993, 1 # 2 $ (AMNH). Santiago del Estero: Ojo de ( 67.79195, 39.25879, 309 m), A.D. Brescovit et al., À À to new generations of students. (Fig. 54E). Agua [ 64.06415, 29.31096, 276 m], Rumboll, X/ 12/X/2014, 1 imm. (MACN-Ar); General Roca, Coronel À À 1969, 1 imm. (MACN-Ar 6042). La Rioja: Arauco, Juan JoseG� omez� [ 67.63624, 39.01893, 246 m], À À Diagnosis: This species can be easily recognized by Natural history: Levi (1967) and Levi & Levi (1969) Ascha, Aimogasta [ 66.81783, 28.55518, 863 m], Sosa, 12/XI/1946, 1 # (MACN-Ar 30236); V/1946, 1 $ À À its large size and cephalic region slightly wider than studied the predatory and sexual behaviour of this J.C. Freyre, III/1947, 1 # 1 imm. (MACN-Ar 2207); (MACN-Ar 30235); General Roca, Belsa [ 67.575, À that of S. rupestris and S. mapuche sp. nov. (compare species in detail; and Reiskind (1966) described its General Belgrano, Iliar [ 66.18539, 30.67851, 39.02682, 248 m], R. Bachmann, I/1962, 1 # 2 imm. À À À Figs 15H, 16B). Males are further distinguished by self-burying behaviour. We have collected it from 438 m], P. Gomez, 1940, 1 # (MACN-Ar 6795); (MACN-Ar 5483); Chichinales [ 66.94212, À having a ventral tuft of setae in the palpal femur monte xeric scrub, usually under large rocks or in General Sarmiento, Vinchina [ 68.16675, 28.77002, 39.11329, 226 m], Schigone, XI/1965, 1 # (MACN-Ar À À À (Fig. 54A, VHT) and by the straight bulb with a the sediment accumulated next to cliffs. 1477 m], J.M. Viana, XI/1970, 1 $ (MACN-Ar 6911); 6902), 1 # (MACN-Ar 6906); [ 67.575, 39.02682, À À distinct base and a short embolus (Fig. 54B–D) Independencia, Patqu�ıa, Guayapa [ 66.87937, 248 m], R. Bachmann, X/1963, 1 # (MACN-Ar 5029), 1 À (smooth transition between base and body, and long Distribution: Western Argentina, from Salta to R�ıo 30.04386, 423 m], L. Yivoff, 24/XI/1961, 1 # 2 imm. # (MACN-Ar 6910). No further data: F.A. Enders, 1 À embolus in S. rupestris and S. mapuche sp. nov.). Negro provinces (Fig. 26D). A record from Buenos (MACN-Ar 30238); 20/III/1962, 1 # 1 $ (MACN-Ar $ (SINHM). CHILE. Region� V de Valpara�ıso: Cerro Females are easily distinguished from all Sicarius by Aires (IBSP 4057) is doubtful; it could refer to 30239); La Rioja [ 66.85722, 29.4178, 508 m], San Cristobal [ 70.63389, 33.29611, 643 m], Fritz, À À À À the very short spermathecae branches, barely visible somewhere in Buenos Aires province (where some Bowman & Pendola, 1914, 1 # (MACN-Ar 15925); VIII/1963, 1 # (MACN-Ar 6804) (doubtful record). in dorsal view (Fig. 54F–K). records of S. rupestris are known), but the species Manciniraudix, 1941, 1 # (MACN-Ar 767); Viana, has never been collected so far east. A vial in the J.M., 1970, 1 $ (MACN-Ar 6903); Sanagasta, Sierra BRAZILIAN SPECIES Description: Male (holotype from El Cuy, R�ıo Negro, MACN says ‘Chile, Cerro San Cristobal’.� We have de Velazco [ 66.88749, 29.18932, 2277 m], À À Argentina, MACN-Ar 33859). Coloration: carapace, examined hundreds of Chilean Sicarius specimens, Cranwell J., III/1940, 1 $ (MACN-Ar 6907). The sole representantive of Sicarius in Brazil had chelicerae, labium and endites reddish brown, and never found this species there. The only Cerro Cordoba:� Cruz del Eje, Pocho, Chancan�ı( 65.45311, been Sicarius tropicus (Mello-Leitao,~ 1936) until we À sternum orange, legs brownish orange, abdomen light San Cristobal� we know of is inside Santiago, capital 31.41616, 819 m), M.A. Izquierdo, 15/IX/2005, 1 # revised the genus for that country, describing three À brown. Carapace length 9.04, anterior width 4.39, of Chile; Gerschman de Pikelin & Schiapelli (1979) (MACN-Ar 34088); Punilla, La Falda [ 64.48368, more species (Magalhaes~ et al., 2013). These four spe- À thoracic region width 9.04. Eye diameters and placed this record in the Valpara�ıso region, in the 31.08835, 958 m], Cesari, IV/1975, 1 $ 1 imm. cies will not be treated again here, so readers must À interdistances: PME 0.23, ALE 0.32, PLE 0.22, PME– Andes range, next to the border with Mendoza, (MACN-Ar 6904). Mendoza: Las Heras, Hornito del refer to our previous publication for a more complete PLE 1.03, ALE–PLE 0.27. Clypeus height 1.73. Eye Argentina. We followed them and included this Gringo [ 68.81459, 32.84853, 714 m], L. Yapur, 19/ picture of Brazilian Sicarius. Here, we present the À À tubercles absent. Macrosetae in lateral border of the record in the distribution map, but it would be better IX/1976, 1 # (MACN-Ar 30227), 2 $ 1 imm. (MACN- descriptions of two new species which we had previ- carapace in a single row. Femora without prolateral regarded as a mislabelling. Ar 30228); Maipu, Rodeo del Medio [ 68.66667, ously considered as morphological variants of S. cariri black setae. Sternum length 3.86, width 3.72, 32.98333, 679 m], 13/III/1954, 1 imm.À (MHNS); and S. diadorim, but that have been shown to be par- À cordiform. Femoral thorns: 5. Leg I: femur 10.24 long, Additional material examined: ARGENTINA. Santa Rosa, Lavalle, Reserva Natural Provincial ticularly divergent based on DNA sequence data (see patella 3.99, tibia 9.98, metatarsus 7.32, tarsus 4.52. Salta: Cafayate [ 65.97448, 26.07291, 1648 m], Telteca ( 68.046333, 32.375583, 558 m), R. Botero- Magalhaes et al., 2014). Upon a closer look at their À À À À II: fe 11.44, ti 11.04. III: fe 10.51, ti 9.44. IV: fe 10.37, Viana, J.M., IX/1963, 2 # (MACN-Ar 6793); General Trujillo & A.L. Carbajal, 13–19/III/2015, 1$ 6 imm. genital morphology, we could identify some features pa 3.33, ti 9.31, mt 7.32, ta 4.39. Abdomen: length Jose de San Martın, La Curtiembre [ 63.21667, (MACN-Ar 34845); G. Flores, 25/II/1994, 1 imm. (CAI to distinguish them from known species. À 7.98, width 6.18. Palp as in Figure 54A and B. 22.91667, 1200 m], L.E. Pena,~ 15/II/1993, 1 $ 3142), 1 # 1 imm. (CAI 528); 1/XI/1994, 3 $ 2 imm. À Female (La Curtiembre, Salta, Argentina, AMNH). (AMNH); R�ıo Seco, 4 km S Cafayate [ 64.1627, (CAI 3135-9); 2–30/XI/1994, 1 imm. (CAI 3153); 15/ À SICARIUS SACI SP. NOV. As in male, except for the following. Labium, endites, 22.8722, 754 m], C.J. Grismado & A. Compagnucci, XII/1995, 1 # 1 imm. (CAI 3140-1); G. Flores & S. À sternum and legs light brown, abdomen brownish 2/XI/2004, 1 imm. (MACN-Ar 30213); Molinos, Cerro Roig, 14/III/1995, 1 # 3 imm. (CAI 3145); G. Flores Figures 16D, 27A, 55, 56 yellow. Total length 24. Carapace length 10.25, ante- Colorado [ 66.49966, 25.80662, 3590 m], 1 # 3 et al.,1–18/XII/2012, 1 imm. (CAI 3174); S. Roig, 24/ Sicarius cariri: Magalhaes~ et al., 2013: figs 44, 70, À À rior width 5.18, thoracic region width 10.25. Eye imm. (MACN-Ar 30237); Laguna Brealito, 15 km W XII/1995, 1 # 1 $ (CAI 3049-50); S. Roig & G. Flores, 71, 82, 84 (misidentified, in part, see note below). diameters and interdistances: PME 0.23, ALE 0.28, Seclantas� [ 66.35743, 25.29545, 2603 m], E. 31/X/1995, 3 imm. (CAI 3149-51); 5/XI/1996, 1 # 1 Sicarius cariri clade 2: Magalhaes et al., 2014: 5328, À À PLE 0.25, PME–PLE 1.6, ALE–PLE 0.31. Clypeus Maury, 29/I/1981, 1 # 3 imm. (MACN-Ar 30231). imm. (CAI 3143-4); Nacu~ n~an� [ 67.96667, 34.03333, figs 1, 2. À À height 2.31. Sternum length 2.01, width 4.51, cordi- Tucuman:� Tafı del Valle, Amaicha� del Valle, 9.8 km 589 m], S. Lagos, 7/II/1998, 1 # 1 imm. (CAI 3059- form. Femoral thorns: 6. Leg I: femur 9.25 long, E Amaicha� del Valle [ 65.91667, 26.58333, 3061). San Luis: Ayacucho, R�ıo Lujan� [ 65.94764, Note: All the records of Sicarius cariri from Barra, À À À patella 4.01, tibia 8.6, metatarsus 6.55, tarsus 4.15. 2033 m], P. Devries et al., 28/III/1992, 1 $ (AMNH); 32.31985, 555 m], Viana, J.M., 1947, 1 # (MACN-Ar Lençois,� Morro do Chapeu� and Xique-Xique cited in À II: fe 10, ti 9.38. III: fe 9.75, ti 8.38. IV: fe 9.3, pa Los Zazos, C.J. Grismado & A. Compagnucci, 1/XI/ 2195); Belgrano, Parque Nacional Sierra de las our previous contribution (Magalhaes~ et al., 2013) 3.96, ti 8.5, mt 6.75, ta 4.09. Abdomen: length 12.9, 2004, 1 $ 2 imm. (MACN-Ar 30209); Trancos, R�ıo Quijadas, administrative area ( 66.96085, 32.469, are of this species instead. This species has been À À width 12. Spermathecae branches: 8. Ventral Sali, below rt. 9 bridge, 6.3 km S border with Salta 686 m), C. Grismado et al.,3–4/XI/2013, 1 imm. demonstrated to be an independently evolving lin- branches present, one pair, straight. Spermathecae province [ 65.25, 26.23333, 786 m], F.A. Coyle (MACN-Ar 30994). Buenos Aires: Buenos Aires eage by Magalhaes et al. (2014). À À as in Figure 54J and K. et al., 17/III/1988, 1 # (AMNH). Catamarca: [ 58.36667, 34.6, 24 m], L. Coronel, 24/VIII/1988, 1 À À Andalgala, Villavil, 8 km E Andalgala [ 66.31623, # 3 $ 1 imm. (IBSP 4057) (almost certainly a wrong Type material: Male holotype from BRAZIL. Bahia: À Variation: Males (N = 6): carapace length 7.83–9.6 27.6004, 1010 m], F.A. Enders, 22/II/1973, 1 # label). Neuquen:� Confluencia, Senillosa (Senilopa?) Morro do Chapeu, Parque Estadual Morro do À (8.93), femoral thorns 5–6, femur II length 9.9–12.5 (SINHM); 11/II/1974, 1 # (SINHM); 30/XII/1974, 1 # [ 68.41667, 39, 300 m], O. de Ferrariis, I–II/1973, 1 Chapeu� ( 41.33036, 11.49756, 1001 m), I.L.F. À À À À (11.42). Females (N = 5): carapace length 8.8–10.5 (SINHM); 28/VII/1972, 1 # (SINHM); 23/XI/1972, 1 $ $ (AMNH). R�ıo Negro: El Cuy, Paso Cordoba� Magalhaes~ et al., 20/I/2012 (UFMG 11021). (9.69), femoral thorns 5–7, femur II length 8.9–11.63 (SINHM); 30/XII/1973, 1 # 1 $ (SINHM); no further ( 67.63204, 39.12521, 380 m), A.D. Brescovit et al., Paratypes: same data, 1 # (IBSP 162262), 1 imm. À À (10.08), spermathecae branches 6–12. Genital data [ 65.76947, 28.48545, 514 m], 1971, 1 $ 12/X/2014, 6 imm. (MACN-Ar 35931), 1 imm. (MACN- (IBSP 162264), 1 $ (IBSP 162265), 1 $ (IBSP À À

Figure 56. Sicarius saci sp. nov. from Brazil, female genitalia, digested. A, female paratype from Morro do Chapeu,� Bahia (UFMG 11686), dorsal. B, same, ventral. C, Barra, Bahia (IBSP 162691), dorsal. D, same, ventral. E, Xique-Xique, Bahia (IBSP 162288), dorsal. F, same, ventral. E and F reproduced from Magalhaes~ et al. (2013) with permission from Figure 55. Sicarius saci sp. nov. from Brazil, male genitalia in prolateral view. A, male holotype from Morro do Magnolia Press. Scale bars = 0.1 mm. Chapeu,� Bahia (UFMG 11021). B, same, cymbium and bulb. C, same, apical. D, Barra, Bahia (UFMG 11017). E, Lençois,� Bahia (MNRJ 6264). F, Xique-Xique, Bahia (IBSP 162285). E and F reproduced from Magalhaes~ et al. (2013) with permission from Magnolia Press. Scale bars = 0.1 mm (except A, 1 mm). have few (10–15), coiled spermathecae branches Femora with prolateral black setae in several with a small reservatory at the apex (Fig. 56); at discrete patches. Sternum length 2.58, width 3.08, 162266), 1 $ (IBSP 162267), 1 $ (IBSP 162268), 1 $ nothing more than a red hood. It is said to find great present we have not been able to find a diagnostic cordiform. Femoral thorns: 6. Leg I: femur 7.75 long, (IBSP 162269), 1 $ (UFMG 11022), 1 $ (UFMG pleasure in acting mischievously and tricking people. feature for females based on morphology alone. patella 2.64, tibia 8.17, metatarsus 6.48, tarsus 3.39. 11023), 1 $ (UFMG 11686); same locality Likewise, the new species tricked us into thinking it Identification of females without accompanying II: fe 8.8, ti 8.6. III: fe 8, ti 7.42. IV: fe 7.83, pa 2.33, ( 41.25144, 11.49914, 1095 m), 21/I/2012, 1 $ was only a variety of S. cariri before, and all the males can be done using COI or H3 sequences (see ti 7.17, mt 5.5, ta 2.83. Abdomen: length 5.66, width À À (IBSP 162272), 1 # (IBSP 162273), 1 # (IBSP individuals listed above lack at least one leg, which Magalhaes et al., 2014). 5.25. Palp as in Figure 55A and B. 162274), 1 $ (IBSP 162276), 1 $ (IBSP 162277), 1 we used for DNA extraction. The name is to be Female paratype from Morro do Chapeu,� Bahia, imm. (IBSP 162278), 1 $ (IBSP 162279), 1 $ (IBSP treated as a noun in apposition. Description: Male holotype from Morro do Chapeu,� Brasil (UFMG 11022). As in male, except for the fol- 162280), 1 $ (IBSP 162281), 1 $ (IBSP 162282), 1 # Bahia, Brasil (UFMG 11021). Coloration: carapace, lowing. Total length 13.2. Carapace length 7, anterior (UFMG 11024), 1 $ (UFMG 11025), 1 $ (UFMG Diagnosis: Males are similar to those of Sicarius chelicerae, labium, endites and sternum reddish width 2.67, thoracic region width 6.83. Eye diameters 11026); ( 41.36886, 11.47756, 892 m), 19/I/2012, 1 cariri by the strongly marked transition between brown, legs orange brown, abdomen brownish yellow. and interdistances: PME 0.23, ALE 0.25, PLE 0.28, À À # (IBSP 162258), 1 imm. (IBSP 162259), 1 imm. the base and body of the bulb, and by the short Total length 11.13. Carapace length 6, anterior PME–PLE 0.66, ALE–PLE 0.23. Clypeus height 1.02. (IBSP 162260), 1 # (UFMG 11685); embolus. They can be distinguished by the embolus width 2.32, thoracic region width 6.07. Eye Sternum length 2.82, width 3.22, cordiform. Femoral being straight in apical view (Fig. 55C) (with a diameters and interdistances: PME 0.2, ALE 0.25, thorns: 6. Leg I: femur 7.5 long, patella 2.75, tibia Etymology: The Saci is a creature from Brazilian retrolateral concavity in S. cariri, Magalhaes~ et al., PLE 0.23, PME–PLE 0.51, ALE–PLE 0.18. Clypeus 7.08, metatarsus 5.52, tarsus 3.14. II: fe 8.17, ti 7.5. folklore; it has the form of a one-legged, black boy 2013, fig. 66) and by the somewhat shorter bulb. height 0.84. Eye tubercles present. Macrosetae in III: fe 7.25, ti 6.67. IV: fe 7.5, pa 2.48, ti 6.61, mt who hops around smoking a pipe and wearing Females are very similar to those of S. cariri; they lateral border of the carapace in multiple rows. 5.07, ta 2.69. Abdomen: length 6.08, width 6.

Spermathecae branches: 10. Ventral branches absent. Etymology: The name is a noun in apposition to the Spermathecae as in Figure 56A and B. type locality. Itaobim is located in the Jequitinhonha River valley. Variation: Males (N = 5): carapace length 5.02–6 (5.56), femoral thorns 5–6, femur II length 7.58–8.8 Diagnosis: Males are similar to those of (8.02). Females (N = 5): carapace length 4.3–7 (5.59), S. diadorim by the long, thin embolus; they can be femoral thorns 6–7, femur II length 4.98–8.17 (6.65), distinguished by the embolus being more curved in spermathecae branches 10–15. Genital morphology is lateral view and by the more sinuous bulb body slightly variable in both sexes (Fig. 55); the male (Fig. 58F, G) and by the straight embolus in apical from Lençois� is somewhat deviant, but it fits with view (Fig. 58E) (S. diadorim with embolus not so this species. curved in lateral view, bulb body straight, and embolus curved in apical view; Fig. 58A–D). Natural history: Specimens have been collected in Females can be distinguished from those of sandy Caatinga, usually buried in the sediment near S. diadorim by the presence of ventral branches in cliffs, trees or under logs. A pair has been observed the spermathecae, and from those of S. ornatus by mating (Magalhaes~ et al., 2013, fig. 44). having only one pair of ventral branches (usually two or more in S. ornatus, especially in southern Distribution: Central Bahia, Brazil (Fig. 27A). populations) and by the longer spermathecae branches, with larger reservatories in the apex Additional material examined: BRAZIL. Bahia: (Fig. 57C–F). Barra, Brejo Olhos D’Agua� ( 43.35269, 10.80744, À À 481 m), I.L.F. Magalhaes~ et al., 26/I/2012, 1 # (IBSP Description: Male (holotype, Itaobim, Minas Gerais, 162247), 1 $ (IBSP 162688), 1 # (IBSP 162690), 1 $ Brazil, UFMG 10157). Coloration: carapace, (IBSP 162691), 1 # (UFMG 11017), 1 $ (UFMG chelicerae, labium and endites reddish brown, 11684); Lençois� [ 41.38333, 12.55, 512 m], A.P.L. sternum and legs light brown, abdomen brownish À À Giupponi & R.L.C. Baptista, 13/XI/2002, 1 # (MNRJ yellow. Total length 9.3. Carapace length 5.1, 6264); Xique-Xique ( 42.72953, 11.08025, 475 m), anterior width 2.2, thoracic region width 5.39. Eye À À 27/I/2012, 1 # (IBSP 162285), 1 $ (IBSP 162286), 1 $ diameters and interdistances: PME 0.25, ALE 0.25, (IBSP 162287), 1 $ (IBSP 162288), 1 # (IBSP 162695), PLE 0.25, PME–PLE 0.51, ALE–PLE 0.18. Clypeus 1 # (IBSP 166323), 1 # (IBSP 166324), 1 $ (UFMG height 0.84. Eye tubercles present. Macrosetae in 11027), 1 # (UFMG 11687). lateral border of the carapace in multiple rows. Femora with prolateral black setae in several discrete patches. Sternum length 2.08, width 2.67, SICARIUS JEQUITINHONHA SP. NOV. cordiform. Femoral thorns: 5. Leg I: femur 6.08 long, patella 2.16, tibia 6.82, metatarsus 5.18, tarsus 2.14. Figures 16E, 27A, 57, 58E–G II: fe 6.92, ti 6.75. III: fe 6.25, ti 6.2. IV: fe missing. Sicarius diadorim: Magalhaes~ et al., 2013: 91, 98, Abdomen: length 4.31, width 4.41. Palp as in 99, 146, 147 (misidentified, in part, see note below). Figs 57A, B, 58F. Sicarius diadorim clade 2: Magalhaes et al., 2014: Female (paratype, Itaobim, Minas Gerais, Brazil, 5328, figs 1, 2. UFMG 10156). As in male, except for the following. Note: All the records of Sicarius diadorim from Sternum reddish brown and legs orange brown. Total Espinosa and Itaobim cited in our previous contribu- length 13.5. Carapace length 6.14, anterior width tion (Magalhaes~ et al., 2013) are of this species 2.49, thoracic region width 6.27. Eye diameters and instead. This species has been demonstrated to be an interdistances: PME 0.23, ALE 0.25, PLE 0.23, Figure 57. Sicarius jequitinhonha sp. nov. from Brazil. A–B, male genitalia in prolateral view; C–F, female genital- independently evolving lineage by Magalhaes et al. PME–PLE 0.58, ALE–PLE 0.28. Clypeus height 1.25. ia, digested. A, male holotype from Itaobim, Minas Gerais (UFMG 10157). B, same, cymbium and bulb. C, female paratype from Itaobim, Minas Gerais (UFMG 10156), dorsal. D, same, ventral. E, Espinosa, Minas Gerais (IBSP 162292), (2014). Sternum length 2.61, width 2.98, cordiform. Femoral dorsal. F, same, ventral (redrawn from Magalhaes~ et al., 2013, figs 98–99). Scale bars = 0.1 mm (except A, 1 mm). thorns: 4. Leg I: femur 6.07 long, patella 2.52, tibia Type material: Male holotype from BRAZIL. Minas 6.41, metatarsus 4.98, tarsus 2.25. II: fe 6.42, ti 6.5. Gerais: Itaobim ( 41.57925, 16.59553, 367 m), III: fe 6.25, ti 5.58. IV: fe missing. Abdomen: length (5.78), femoral thorns 4–5, femur II length 5.93–6.58 (copulation was not observed). The male was À À I.L.F. Magalhaes~ et al., 25/XI/2011 (UFMG 10157). 7.5, width 6.92. Spermathecae branches: 15. Ventral (6.34), spermathecae branches 14–16. Genital removed from the container after the female laid an Paratypes: same locality and collector ( 41.51042, branches present, one pair, straight. Spermathecae morphology is variable (Fig. 57). eggcase; after that, she laid three more eggcases in a À 16.53294, 434 m), 25/XI/2011, 1 $ (UFMG 10156); as in Figure 57C and D. period of 4 months. The only viable eggcase gave rise À Espinosa ( 42.83497, 14.94664, 580 m), I.L.F. Natural history: Specimens have been collected to five young and the remaining were attacked by À À Magalhaes~ et al., 10/I/2012, 1# (IBSP 163683), 1 $ Variation: Males (N = 2): carapace length 5.1–5.39 buried near large rocks or under logs in Caatinga mites. The spiderlings took between 21 and (IBSP 162290), 1 $ (IBSP 162291), 1 $ (IBSP (5.25), femoral thorns 4–5, femur II length 6.92–7.17 areas. One virgin pair from Espinosa, raised in the 26 months to reach maturity (N = 3); one of them 162292), 1 imm. (IBSP 162294); (7.05). Females (N = 5): carapace length 5.52–6.14 laboratory until maturity, was allowed to mate had not yet become an adult after 29 months.

Figure 58. Male palpal bulbs in prolateral view (except D–E, apical). A–D, Sicarius diadorim Magalhaes,~ Brescovit & Santos, 2013. A, holotype from Itacarambi, Minas Gerais, Brazil (UFMG 10789). B, male from Sao~ Felix� do Coribe, Bahia, Brazil (DZUB 6229). C, male from Cocos, Bahia, Brazil (DZUB 5000). D, holotype from Itacarambi, Minas Gerais, Brazil (UFMG 10789), apical. E–G, Sicarius jequitinhonha sp. nov. E, holotype from Itaobim, Minas Gerais, Brazil (UFMG 10157). F, same, apical. F, male from Espinosa, Minas Gerais, Brazil (IBSP 163683). Scale bars = 0.1 mm.

Distribution: Northern Minas Gerais and south- island other than the type locality. Note that Baert eastern Bahia, Brasil (Fig. 27A). et al. (2008) examined specimens from several islands and considered all of them to be S. utriformis. Additional material examined: BRAZIL. Bahia: Itapetinga [ 40.24991, 15.25145, 259 m], C.M.P. Diagnosis: Differs from all species, except À À Leite, 2013, 1 # (IBSP 166469), 2 $ (IBSP 166470), 3 # S. peruensis and S. gracilis, by having a single, large 1 $ (IBSP 166471), 1 $ (IBSP 166472). Minas Gerais: distal patch of prolateral black setae in the femora Itaobim ( 41.51042, 16.53294, 434 m), 1 imm. (Fig. 24D, arrow). The male is distinguished from (UFMG 10147);À same localityÀ ( 41.48086, 16.56875, these two species by the gently curved bulb, with a 316 m), 27/XI/2011, 1 imm. (UFMGÀ 10155).À smooth transition between the base and the body of the bulb (Fig. 59A, B) (well-marked transition in S. peruensis and S. gracilis). Females are very similar NORTHERN SPECIES to those of S. peruensis: they have short, widely Besides the greater diversity found in Chile, Argen- spaced spermathecae branches without reservatory in tina, Peru and Brazil, Sicarius are known from iso- the apex, a sclerotized pore plate and ventral lated species in the Galapagos islands (S. utriformis) branches with a double coil (Fig. 19B). They can be and Central America (S. rugosus). distinguished by having several tiny digitiform projections arising from the branches (Fig. 59C, D, E). Figure 59. Sicarius utriformis (Butler, 1877) from the Galapagos Islands (Ecuador). A–B, male genitalia in prolateral SICARIUS UTRIFORMIS (BUTLER, 1877) Description: Male (Espanola,~ Galapagos Islands, view; C–F, female genitalia, digested. A, Espanola~ Island (RBINS). B, same, cymbium and bulb. C, Espanola~ Island RBINS). Coloration: carapace and chelicerae reddish (RBINS), dorsal. D, same, ventral. E, same, anterior. F, subadult female from Fernandina (AMNH), not to scale, ventral. Figures 17E, 19B, 24D, 27D, 59 brown, labium, endites and sternum light brown, PP, pore plate; VB, ventral branch. Scale bars = 0.1 mm (except A, 1 mm). Thomisoides utriformis Butler, 1877: 77, plate 13, legs yellow, abdomen brownish yellow. Total length ﬁg. 4. Immature holotype from Charles Island (also 12. Carapace length 6.92, anterior width 2.64, Female (Espanola,~ Galapagos Islands, RBINS). As Variation: Subadult females have pre-spermathecae known as Floreana or Santa Mar�ıa), Galapagos thoracic region width 6.61. Eye diameters and in male, except for the following. Total length 15.75. with short and blunt branches, with a primordial, Islands, Ecuador, deposited in BMNH, examined. interdistances: PME 0.23, ALE 0.33, PLE 0.28, Carapace length 7.25, anterior width 2.78, thoracic very slightly coiled pair of ventral branches Sicarius utriformis: Simon, 1893: 270;. Baert et al., PME–PLE 0.58, ALE–PLE 0.23. Clypeus height 1.22. region width 6.68. Eye diameters and interdistances: (Fig. 59F). 2008: 63, 102. Eye tubercles present. Macrosetae in lateral border PME 0.23, ALE 0.33, PLE 0.28, PME–PLE 0.71, Sicaroides ultriformis: Banks, 1902: 55, 71 (lapsus). of the carapace in multiple rows. Femora with ALE–PLE 0.33. Clypeus height 1.25. Sternum length Natural history: Baert et al. (2008) report that the Note: Butler’s (1877) immature specimen comes prolateral black setae in a large distal patch. 3.08, width 3.32, cordiform. Femoral thorns: 3. Leg I: species occurs in littoral zones, low arid zones and from Floreana Island; we have also examined speci- Sternum length 3.19, width 3.15, cordiform. Femoral femur 9 long, patella 2.94, tibia 9.4, metatarsus 6.92, transition forest. Banks (1902) collected specimens mens from Espanola~ (an adult pair and some imma- thorns: 3. Leg I: femur 10.63 long, patella 2.81, tibia tarsus 3.2. II: fe 9.8, ti 10. III: fe 8.7, ti 8.3. IV: fe 8.7, under rocks in Espanola,~ and described the eggcase tures) and Fernandina (a subadult female). Until it is 11.5, metatarsus 8.9, tarsus 3.88. II: fe 12, ti 12.63. pa 2.69, ti 8.42, mt 6.75, ta 3.31. Abdomen: length (his illustration is reproduced in Fig. 17E). determined whether a single species occurs in all III: fe 10.63, ti 10.5. IV: fe 10.75, pa 3.05, ti 9.7, mt 8.7, width 8.3. Spermathecae branches: 6. Ventral islands of the archipelago, our diagnosis must be 8, ta 3.8. Abdomen: length 5.73, width 5.73. Palp as branches present, one pair, with a double coil. Sper- Distribution: Endemic to the Galapagos Islands. We taken with caution, as it is based on adults from an in Figure 59A and B. mathecae as in Figure 59C, D and E. examined specimens from Espanola,~ Fernandina and

Floreana (Fig. 27D); Baert et al. (2008) provide patches, white macrosetae present. Sternum length additional records from Eden, Gardner (near 1.86, width 2.44, cordiform. Femoral thorns: 4. Leg I: Espanola),~ Rabida,� Santiago, Santa Cruz and Santa femur 4.97 long, patella 1.83, tibia 6.07, metatarsus Fe� islands, not examined by us. 4.92, tarsus 2.34. II: fe 6.2, ti 6.14. III: fe 5.32, ti 5.11. IV: fe 5.45, pa 1.6, ti 5.07, mt 4.72, ta 2.34. Abdomen: Additional material examined: ECUADOR. length 3.65, width 4.23. Palp as in Figure 60A and B. Galapagos Islands: Espanola,~ Espanola~ Female from Santa Ana, Sonsonate, El Salvador [ 89.63664, 1.40092, 68 m], L. Baert, 1991, 1 # (AMNH). As in male, except for the following. À À (RBINS 27720); L. Baert et al., 16/IV/1991, 1 $ Labium, endites, sternum and legs light brown. Total (RBINS); Punto Cevallos, L. Baert, 1 imm. (RBINS); length 8.8. Carapace length 4.51, anterior width Fernandina, Tagus Cove [ 91.36667, 0.25, 90 m], 2.08, thoracic region width 4.51. Eye diameters and À À Arcturus, 1929, 1 imm. (AMNH); 1 imm. (AMNH). interdistances: PME 0.23, ALE 0.2, PLE 0.23, PME– PLE 0.58, ALE–PLE 0.2. Clypeus height 0.81. Ster- num length 2.17, width 2.38, cordiform. Femoral SICARIUS RUGOSUS thorns: 4. Leg I: femur 4.77 long, patella 1.78, tibia (F. O. PICKARD-CAMBRIDGE, 1899) 5.45, metatarsus 4.31, tarsus 2.01. II: fe 5.18, ti 5.05. Figures 12B, 15G, 24C, 27C, 60 III: fe 4.98, ti 4.67. IV: fe 5.18, pa 1.75, ti 4,67, mt Sicarioides rugosus F.O. Pickard-Cambridge, 1899b: 4.31, ta 2.11. Abdomen: length 4.57, width 5.25. 49, plate 3, fig. 12. Immature holotype from Bebe- Spermathecae branches: 6. Ventral branches absent. dero, Guanacaste, Costa Rica, deposited in BMNH, Spermathecae as in Figure 60E and F. examined. Kraus, 1955: 11, fig. 16. Sicarioides rugosus: Lehtinen, 1986: 156, fig. 7. Variation: Males (N = 3): carapace length 4.11–5.25 (4.56), femoral thorns 4–5, femur II length 6.08–7.58 Note: We have not examined adult specimens from (6.62). Females (N = 3): carapace length 4.46–4.83 the exact type locality, but this species is very com- (4.6), femoral thorns 4–5, femur II length 4.92–5.32 mon in nearby areas, and thus we have no doubts of (5.14), spermathecae branches 4–6. Genital its identity. morphology variable from small branches without reservatory to branches a bit longer, with Diagnosis: This species can be distinguished by reservatory on the median ones (Fig. 60E–J). having white macrosetae and several discrete Subadult females with only two digitiform, elongate patches of prolateral black setae in the leg femora spermathecal branches (Fig. 60D). (Fig. 24C). Males are further distinguished by the very short bulb with a long, bent embolus (Fig. 60A, Natural history: This species inhabits dry forests in B, C). Females are further distinguished by the Central America; it is the Sicarius species recorded spermathecae with few short and robust branches, from the most humid habitats (1300–1900 mm of and without ventral branches (Fig. 60E–J). annual precipitation, but with a 3-month dry season receiving less than 20 mm of rain). Description: Male (Santa Ana, Sonsonate, El Salvador, AMNH). Coloration: carapace and Distribution: Dry west coast of Central America, chelicerae reddish brown, labium light brown, endites, from southern El Salvador to north-western Costa sternum and legs light brown, abdomen brownish Rica (Fig. 27C). yellow. Total length 8.42. Carapace length 4.11, anterior width 1.75, thoracic region width 4.06. Eye Additional material examined: EL SALVADOR. diameters and interdistances: PME 0.23, ALE 0.23, Sonsonate: Santa Ana, Laguna de Coatepeque, PLE 0.2, PME–PLE 0.48, ALE–PLE 0.18. Clypeus [ 89.53333, 13.83333, 730 m], Zilck, 20/IX/1957, 1 # height 0.66. Eye tubercles present. Macrosetae in À 1 $ (AMNH). NICARAGUA. Leon:� La Paz Centro, lateral border of the carapace in multiple rows. Volcan� Momotombo, El Cardon,� [ 86.59734, 12.4332, Femora with prolateral black setae in several discrete À

Figure 60. Sicarius rugosus (F. O. Pickard-Cambridge, 1899). A–C, male genitalia in prolateral view; D–J, female genitalia, digested. A, Santa Ana, Sonsonate, El Salvador (AMNH) (right palp, mirrored). B, same, bulb. C, Parque Nacional Santa Rosa, Guanacaste, Costa Rica (INBIO 47116) (right palp, mirrored). D, subadult female from Liberia, Guanacaste, Costa Rica (AMNH), ventral. E, Santa Ana, Sonsonate, El Salvador (AMNH), dorsal. F, same, ventral. G, Leon,� Nicara- gua (INBIO), dorsal. H, same, ventral. I, Parque Nacional Santa Rosa, Guanacaste, Costa Rica (INBIO 47116), dorsal. J, same, ventral. Scale bars = 0.1 mm (except A, 1 mm).

83 m], C. Vıquez & J. Mata, 12/II/2007, 1 # 1 $ than long and posteriorly truncate. Palp with simple, (INBIO NIC17). COSTA RICA. Guanacaste: La not sculptured femoral thorns (Fig. 18G). Leg femora Cruz, Parque Nacional Santa Rosa [ 85.76667, without prolateral black setae, with white macrosetae À 10.88333, 162 m], Goodnight, 24/VII/1978, 1 imm. (and brown macrosetae in H. spatulata comb. nov.), (AMNH); Liberia, [ 85.53333, 10.6, 90 m], F.S. which might be medially widened (Fig. 24A). À Truxal, 14/VII/1962, 1 $ (AMNH); 5 km S Liberia Trichobothria with a distal smooth surface (Fig. 31H). [ 85.43333, 10.58333, 126 m], S. Peck, 10/VII/1966, ALS with black setae short, not covering the spinning À 1 imm. (AMNH); Estacion Biologica Maritza, field, without major ampullate gland spigots. PMS Sendero a La Toma de Agua ( 85.49524, 10.96254, absent, sometimes replaced by setae (Fig. 11C). Male À 849 m), R. Gutierrez, 22/IV/2004, 1 $ (INBIO 76804); palpal bulb with parallel sides, with embolus long and La Cruz, Parque Nacional Santa Rosa, Area sinuous (Figs 20A, C and 21A, C) or broad and blunt, Administrativa ( 85.61549, 10.83641, 272 m), D. with a keel (Figs 20E, 22C, D, E). Inner spermathecae À Jansen, 1—31/I/1991, 2 # 3 $ 6 imm. (INBIO 47096); branched, with branches arising from a single stalk same locality ( 85.62648, 10.83998, 272 m), I/1991, 1 (Figs 19C, 20B, D). Outer spermathecae present, not À imm. (INBIO 47095); same locality ( 85.95077, branched, reduced (Figs 19C, 20B, D) (branched in H. À 10.85433, 300 m), J. Jacobs, 24/XI/2001, 1 $ (INBIO spatulata comb. nov.). 72236); same locality ( 85.62648, 10.83998, 272 m), À R.R. Dastreich, 2/III/1995, 1 imm. (INBIO 47164); Distribution: Dry regions of southern Africa. Costa Playa Naranjo ( 85.63742, 10.8327, 172 m), C. À Cano, 6/II/1998, 1 $ (INBIO 49656); same locality Species included: Hexophthalma hahni (Karsch) ( 85.63283, 10.82639, 172 m), 10/II/1999, 2 $ 2 imm. comb. rest., H. testacea (Purcell) comb. nov., À (INBIO 49655); D. Garcia, 10/II/1998, 1 # 1 $ H. albospinosa (Purcell) comb. nov., H. damarensis (INBIO 49647); Estacion Murcielago, 8 km SW (Lawrence) comb. nov., H. dolichocephala (Lawrence) Cuaginiquil ( 85.73102, 10.90193, 180 m), P. Rios, comb. nov., H. spatulata (Pocock) comb. nov. À 28/II/1991, 3 # 3 $ 3 imm. (INBIO 47116); Estacion Santa Rosa ( 85.61549, 10.83641, 272 m), D. HEXOPHTHALMA HAHNI (KARSCH, 1878), Jansen, I/1991, 3À imm. (INBIO 45948); Isla San Jose COMB. REST. ( 85.92289, 10.85628), J. Jacobs, 9/IX/2001, 1 $ À (INBIO 72223); 21–23/XI/2001, 1 # (INBIO 72196); Figures 13B, 14B–C, 18G, 19C, 23A–C, 24A, 61 Sect. Palo Verde ( 85.35144, 10.35048, 20 m), Hexomma hahnii Karsch, 1878: 325, plate 9, fig. 2. Mantillo, 1—24/IV/1992,À 1 imm. (INBIO 47466); Male holotype from ‘Herrero Land’ (currently in Setor San Josecito, Rio Tempisquito ( 85.54888, Namibia), deposited in ZMB 722, examined (Fig. 61). 10.87555, 219 m), D. Garcia, 25/III/1998,À 3 imm. Hexophthalma hahni Karsch, 1879: 109. (INBIO 49629); Sitios de Pocosol, Sitios de Pocosol Sicarius hahni Simon, 1893: 269; figs 225–231; Pur- ( 85.51206, 10.93264, 364 m), D. Jansen, 4/III/1990, cell, 1908: 225; Lotz, 2012: 8, figs 2C, 3C, 4D. Figure 61. Hexophthalma hahni (Karsch, 1878) comb. rest., male holotype (MfN 722). A, habitus, dorsal. B, right À palp, prolateral. C, right bulb, prolateral. Notice broken tip of embolus. D, femur II, macrosetae, anterodorsal. E, repro- 2 # 2 imm. (INBIO 47097). duction of the original illustration by Karsch (1878). Notice complete embolus. Scale bars = A, B, 1 mm; C, D, 0.1 mm. Additional material examined: NAMIBIA. Khomas: Windhoek, Eros Mountains [17.1, 22.56667], E. HEXOPHTHALMA KARSCH, 1879, REVALIDATED À Younga, 20/X/1974, 1 # (TM 22904); Windhoek Kenhardt, towards Upingotn [21.15, 29.35], 1 $ (TM (Fig. 62A–C). Lawrence, 1938: 215, fig. 1; Lotz, 2012: À Hexomma Karsch, 1878: 325. Type species by mono- [17.08361, 22.57], P.G. Olivier, 19/IV/1972, 1 # (TM 19872); Namaqualand, Springbok [17.88342, 3, figs 2A, 3A, 4A. À typy Hexomma hahnii Karsch, 1878. exSMN 35427). Narras: 10 km W Gobabeb (15.05, 29.66655], H. Snyman, 18/IX/1970, 1 # (TM 9904); À Note: Lotz (2012) redescribed this species, but his Hexophthalma Karsch, 1879: 109. Replacement 23.56667, 1432 m), C.E. Griswold et al.,2–5/X/1984, Twee Rivigren, Kalahari Gemsbok National Park À illustrations of genitalia (see his figs 3A, 4A) do not name for Hexomma Karsch, 1878; preoccupied by 1 $ (KNM 21144). Hardap: Stampriet [18.40189, [20.6117, 26.47241], Leriche Family, 1960–1970, 1 # À fit with the morphology of the paralectotype exam- Hexomma Thorell, 1876 (Opiliones: Sclerosomatidae). 24.34369], C. Owen, 9/III/1934, 1 $ (TM 18605); 1 $ 1 imm. (KNM 24870). NO DATA. 1 # 2 $ (TM À ined by us (Fig. 62A–C), nor with that of a pair from Tsondab Valley, Tsondab Valley [15.83499, 23.87], 6511). À the type locality (Figs 21A, B, 20A, B). Our speci- Diagnosis: Species of Hexophthalma can be G. Newlands, 2/VII/1970, 1 imm. (TM 6986). Karas: mens have an embolus almost as long as the bulb distinguished from Sicarius by the lack of posterior Fish River Canyon, Ai-Ais [17.61444, 27.58889], 2/ À HEXOPHTHALMA ALBOSPINOSA (PURCELL, 1908) body in males, and spermathecal branches long and median spinnerets (Fig. 11A, C); palpal bulb with a III/1981, 1 $ 1 imm. (TM 19869). SOUTH AFRICA. COMB. NOV. finger-like in females. We refrain from re-diagnosing long body with parallel sides (Fig. 23C) (subconical Limpopo: Louistrichardt, 8 km W of Bluegunspoort the species as we have not examined the lectotype in Sicarius); and lack of a setae tuft anterior to the road [29.90371, 23.04693], C.R. Owen, 24/XII/1984, Figures 20A and B, 21A and B, 62A–C À designated by Lotz (2012), but the identity of the spinnerets in females. 1 $ (TM 19543); no specific data, Crimea farm, N. Sicarius albospinosus Purcell, 1908: 224. Female lec- species should be clarified in the future. Jacobson, II/1979, 1 # (TM 19545). Northern Cape: totype from Namibia, Great Namaqualand, Luderitz€ Description: As for the subfamily, except for the 127.4 km W Pofadder [19.39472, 29.12861], 2/XII/ Bucht, L. Schultze (SAM-ENW-X 150521), desig- À Additional material examined: NAMIBIA. Karas: following. Chelicerae lamina with weakly sclerotized 1984, 1 $ (TM 22763); 45 km S of Vioolsdrift nated by Lotz (2012), not examined; female paralec- 6 km N Luderitz€ , dunes behind Agate beach, apex (Fig. 13B) (except H. spatulata comb. nov., with [17.84563, 29.10823], C.R. Owen, 28/IX/1988, 1 $ totype from Namibia, Great Namaqualand, Angra À (15.16667, 26.58333), C.E. Griswold & T. Meikle strongly sclerotized apex). Sternum cordiform, wider (TM 19549), 1 # (TM 19550), 1 $ (TM 19551); Pequena, L. Schultze (ZMB 10468), examined À

31.61596], 9/IV/1984, 1 # (TM 22764); Niedwondt’s from known populations of H. spatulata comb. nov. À Pass (19.00622, 32.34892), E. Nortje & S. They might represent an undescribed species. Kritzinger-Klopper,À X/2005, 1 imm. (AcAT 2012/ 2865); Nieuwoudtlville, Van Rhyns Pass [19.01606, Material examined: SOUTH AFRICA. Eastern 31.37482], C.R. Owen, 26/IX/1987, 2 # (TM 22757). Cape: Port Elizabeth, Van der Kemps Kloof Nature À Reserve [25.49741, 33.88725], B. Milne, II/2011, 1 $ (AcAT 2013/1514) (typeÀ locality of the species; see HEXOPHTHALMA SP.1 Pocock, 1900). North West: Silikaatsnek, Brits, Figure 11 2527Db, A. Roberts, VII/1921, 2 $ (TM 13600). Western Cape: 21.8 km N of Mosselbay [22.05575, 34.01042], C.R. Owen, 25/I/1988, 1 # 1 $ 2 imm. Material examined: NAMIBIA. Karas: Fish River À Canyon [17.61444, 27.58889], N.A. Leroy, VII/1981, (TM 22762). À 1 imm. (TM 19547), 1 $ (TM 19548), 1 $ (TM 19552), 1 imm. (TM 19553), 1 imm. (TM 19554), 1 # (TM LOXOSCELINAE SIMON, 1893 19555), 1 $ (TM 19556), 1 imm. (TM 19557). Loxoscelinae: Simon, 1893: 271.

HEXOPHTHALMA SP.2 Diagnosis: Easily distinguished from Sicariinae by several characters, the most notable being: absence of Figures 20C, D, 21C, D soil-adhering setae; a single femoral thorn in the Note: This probably represents an undescribed spe- palpal femur (several in Sicariinae); a large colulus; cies of Hexophthalma gen. reval., not described here absence of spigots in the posterior median spinnerets. because the fine taxonomy of African Sicariinae is beyond the scope of this revision. Their genitalia is Description: Araneomorphae, ecribellate, haplogyne nevertheless illustrated (Fig. 20C, D). spiders. Six eyes (the AME absent), arranged in three diads. Carapace about as wide as long, with Material examined: NAMIBIA. Erongo: 70 km E one row of macrosetae in the lateral borders, with a Walvis Bay, (15.253, 22.962), D. Jacobs & P. Webb, À median, violin-shaped dark band. Labium longer 10–14/II/2012, 1 $ (AcAT 2013/4240); 73 km E Walvis than wide, pointed, partially fused to sternum. Bay, (15.221, 22.921), D. Jacobs, 10–14/II/2013, 1 $ À Endites with a white apex, touching in front of (AcAT 2013/4239); Arandis, Karub, [14.28264, labium. Sternum oval and longer than wide, with 22.11898], 2/VII/1937, 1 $ (TM 7775); Rooikop, three outer rows of longer macrosetae near the À10 km E Walvisbaai, [14.65746, 22.98356], J. À border, and shorter macrosetae in the middle. Londt, 1 $ (KNM 21151); Swakopmund, 40 km ESSE Chelicerae fused at the base, without teeth, with a Figure 62. Hexophthalma spp., syntypes. A–C, Hexophthalma albospinosa (Purcell, 1908) comb. nov., female (MfN Swakopmund, [14.89926, 22.80942], D. Jacobs, 14/ À chelicera lamina along the promargin ending in a 10468). A, habitus, dorsal. B, spermathecae, digested, dorsal. C, femur II, macrosetae, anterodorsal. D, Hexophthalma II–7/VI/2013, 1 # (AcAT 2013/4241); Vogelfederberg, pointed, bifid apex, with fang serrula (Figs 13C, testacea (Purcell, 1908) comb. nov., immature (MfN 48699), habitus, dorsal. Inset: femur II macrosetae, anterodorsal. 55 km E Walvis Bay, (14.91667, 23.1, 500 m), C.E. À 14A). Palp without claw, with one small prolateral Scale bars = A, D, 1 mm; B, C, 0.1 mm. Griswold & T. Meikle Griswold, 5/X/1984, 1 # 2 imm. thorn (Fig. 18I), without a distal crack in the ventral (KNM 21148); Hentiesbaai, [14.96667, 22.41667], C. face of the femur. Femora with normal, pointed Owen, 24/III/1994, 1 (TM 18604). À Griswold, 9/X/1984, 1 # 1 $ 2 imm. (KNM 21143); $ HEXOPHTHALMA TESTACEA (PURCELL, 1908) macrosetae, with a dorsal, longitudinal line without Fish River Canyon, Ai-Ais, [17.61444, 27.58889], setae. Tibia with eight rows of pointed macrosetae: À COMB. NOV. N.A. Leroy, VII/1981, 1 $ (TM 19868). SOUTH HEXOPHTHALMA SPATULATA (POCOCK, 1900) two dorsal, two prolateral, two retrolateral and two AFRICA. Northern Cape: Bloeddrift, West of Figure 62D COMB. NOV. ventral, with all macrosetae with approximately the Arnisfont, Richtersveld, [17.09935, 29.00169], Sicarius testaceus Purcell, 1908: 224. Immature syn- same length. Tarsus with onychium, with two claws, À Figures 6, 12C, 14D, 18H, 19D, 20E, F, 22, 23D–F, W.D. Maacke, 25/II/1962, 1 $ 1 imm. (TM 22784). type from Namibia, Kl. Namaqualand, Kamaggas, with tips of normal length (not longer than the 31H VII/1904 (ZMB 48700), examined (Fig. 62D). Lotz, teeth). Trichobothria with a wide socket, a proximal Sicarius spatulatus Pocock, 1900: 321 [male lecto- 2012: 12, figs 2E, 3E, 4F. hood and a distal ridged or smooth surface. Tarsal HEXOPHTHALMA DAMARENSIS (LAWRENCE, 1928) type, designated by Lotz (2012), from Port Elizabeth, organ capsulate. Abdomen oval, longer than wide. COMB. NOV. South Africa, probably in the BMNH, not examined]; Material examined: NAMIBIA. Karas: Nakeis, Epiandrium composed of a single, dense group of Lotz, 2012: 10, figs 2D, 3D, 4E. Sicarius damarensis Lawrence, 1928: 221; Lotz, Karasburg [18.75, 28.01667], P.A. Methuen, 5/XII/ spigots. Posterior trachea composed of a long and À 2012: 4, figs 2B, 3B, 4B. Holotype female from from 2012, 1 imm. (TM 13949). NO DATA. 1 $ (TM 6357). Note: We examined a female from the type locality broad 3rd fused entapophysis; lateral tracheae Namibia: Outjo, deposited in Iziko Museum, Cape SOUTH AFRICA. Mpumalanga: Lydenburg, (Port Elizabeth) and our specimens fit the descrip- present, with numerous branches. Colulus present Town (SAM 7057), not examined. Lydenburg Dish [30.44593, 25.09574], P.A. Kranti, tion by Pocock (1900) and the re-description by Lotz (Figs 7A, 8A). ALS with normal setae, with two À Material examined. NAMIBIA. Erongo: Karibib, 1897, 1 # (TM 16354). Western Cape: Cederberg (2012). The two females from Brits (TM 13600, geni- segments, numerous piriform gland spigots, and one Ervango Mountains [15.85439, 21.93808], V. Mountains, A. Heeres, 12/III/1993, 1# (DNSM talia pictured in Fig. 23D, E, F) have an aberrant major ampullate gland spigot with a slit-like opening À Fitzsimmans, 8/V/1959, 1 # (TM 16290). ARA174); 2 km W of Van Rhyns Dorp [18.73387, genital morphology and are geographically isolated (Figs 7B, 8C). PMS present, without spigots (Figs 7C,

8D). PLS with aciniform gland spigots and a setal pit Loxoscella Strand, 1906: 668. Synonymized by Material examined. ARGENTINA. Cordoba� : LOXOSCELES SIMILLIMA LAWRENCE with a group of modified setae (Figs 7D, 8E). Palpal Gertsch & Ennik, 1983. Cordoba� ( 64.19192, 31.41975), A. Vexenat, 07/VI/ � � bulb simple, piriform (Fig. 63A, B). Inner Calheirosia Mello-Leitao,~ 1917: 80. Synonymized by Figures 18I, 63A, 8A–E 1975, 2 # 10 $ (MACN-Ar 21483). spermathecae usually simple (Fig. 63C), branched in Brignoli, 1978. the rufescens group. Outer spermathecae absent or Material examined: SOUTH AFRICA. Limpopo: LOXOSCELES CARIBBAEA GERTSCH present. Spermathecae pores small, with a single Diagnosis: As that of the subfamily. Soutpansberg, Farm Amsterdam 116, A. Schoeman, flagellum arising from them (Fig. 63D). Subadult 1 # 2 $ 2 imm. (TM 10611); 23/XI/1970, 1 # 5 $ 1 females without pre-spermatheca. Description: As that of the subfamily. imm. (TM 10602); Farm Weipe 47, W.D. Haacke, Material examined: CUBA. Santiago de Cuba: 23/II/1970, 1 # (TM 10738). Kwazulu-Natal: Reserva Ecologica� Siboney-Jutic�ı, Cueva La V�ırgen Distribution: Native of Africa, America and the Mzuke, Ndumo Game Reserve (32.15933, ( 75.7144, 19.96083), A. Perez� Gonzalez,� 04/V/2010, LOXOSCELES HEINECKEN & LOWE, 1832 � Mediterranean region. Some species (notably 26.87089), C. Haddad, 21/I/2006, 1 imm. (MACN- 1 # 1 $ (MACN-Ar 32734); 1 # 3 $ 3 imm. (MACN-Ar � Loxosceles Heinecken & Lowe in Lowe, 1832: 321. L. rufescens and L. laeta Nicolet) have been Ar 11316), 1 $ (MACN-Ar 11317); 17/I/2006, 1 # 32736). Type species Loxosceles citigrada Heinecken & Lowe, introduced to other regions (Gertsch & Ennik, 1983). (MACN-Ar 11319), 1 imm. (MACN-Ar 11320), 1 $ 1832 (= Loxosceles rufescens Dufour, 1820). (MACN-Ar 11321). LOXOSCELES AMAZONICA GERTSCH Omosites Walckenaer, 1833: 438. Synonymized by Species included: 107 known species (World Spider Simon, 1893. Catalog, 2015). LOXOSCELES SIMILIS MOENKHAUS Material examined: BRAZIL. Bahia: Central, Toca de Piloes~ (11°010–11°190S, 41°470–42°100W), E.F. Material examined: BRAZIL. Minas Gerais: Ramos & F.S. Cunha, III/2002, 4$ (IBSP 89007), 1# Simonesia� , RPPN Estacao~ Biologica� da Mata do 5$ (IBSP 89016), 1# 1 $ (IBSP 89071). Ceara:� Sossego ( 20.07361, 42.07028, 648 m), M.T.T. Quixada�, Fazenda Mage,� I.L.F. Magalhaes~ et al., 6/ � � Santos, 10-12/XII/2009, 1 $ (UFMG 12070), 1 $ IV/2012, 1 $ (UFMG 12044). (UFMG 12072), 1 # (UFMG 12073); Caratinga, RPPN Feliciano Miguel Abdala ( 19.77279, � LOXOSCELES DESERTA GERTSCH 42.16764, 742 m), A.A. Carosini Vera, XI/2011, 1 # (UFMG� 12075); same locality ( 20.07361, � 42.07028, 648 m), 1 $ (UFMG 12074); Imbe� de Material examined: USA. Arizona: Tucson, F.E. � Minas ( 19.86833, 43.96583, 818 m), C.S. Souza, 6/ Russell, 1 # (MACN-Ar 21497), 1 $ (MACN-Ar 21498). � � VIII/2011, 1 # (UFMG 12127). ACKOWLEDGEMENTS LOXOSCELES LAETA (NICOLET, 1849) We are deeply indebted to the curators of the institu- Figure 2 tions housing the material studied during the course of this work. We express our gratitude to all of them Material examined: BRAZIL. Santa Catarina: and to P. Marais (AcAT), N. I. Platnick and L. Sor- Curitibanos (27°16058″S, 50°35004″W), Taipas, S. kin (AMNH), A. Carmichael (CAS), F. Campon� Suzuki, 22/IV/1950, 1# (IBSP 34953). Sao~ Paulo: (CAI), N. Govender (DNSM), C. Stoffels (KNM), C. Osasco (23°31057″S, 46°47030″W), J.R. Bettinazzi, 26/ Grismado (MACN), L. Leibensperger (MCZ), A. Frie- VII/1982, 3# 3 $ (IBSP 34948). CHILE. Valpara�ıso: derichs and J. Kapp (ZMB), C. Barros (MNRJ), V. S. Ocoa, Border of Parque Nacional La Campana, alt. Costa (UFMG) and D. DeRoche (USNM), who kindly 355 m (32°54098″S, 71°05042″W), A.D. Brescovit made material from their collections available, pack- et al., 23/III/2013 (IBSP 163704) aged and sent specimens, and also offered their help and hospitality during visits to their institutions. We also thank several people who sent material under LOXOSCELES INTERMEDIA MELLO-LEITAO~ their care or specifically kept an eye for Sicarius in the field: A. V. Peretti, A. O. Porta, A. Taucare R�ıos, Material examined: BRAZIL. Santa Catarina: C. Calitra, H. A. Iuri, J. Ochoa, L. S. Carvalho, L. E. Sao~ Bento do Sul, Centro de Estudos e Pesquisas C. Schmidt and R. Botero-Trujillo. Many colleagues Ambientais Rugendas, I.L.F. Magalhaes~ et al., also accompanied us in the field trips: A. Taucare 13-16/XII/2011, 1 # (UFMG 12097), 1 # (UFMG R�ıos, B. T. Faleiro, C. Veloso, F. M. Hughes, G. F. B. 12098), 2 # 3 $ (UFMG 12099), 1 $ (UFMG Pereira, J. Cabra, L. Piacentini and T. J. Porto. Col- Figure 63. Scanning electronic microscopy, Loxosceles genitalia. A, Loxosceles simillima Lawrence, 1927, male (TM 12101). lecting permits were kindly issued by SEMA-Bahia 10611), left palpal bulb, prolateral. B, Loxosceles hirsuta Mello-Leitao,~ 1931, male (MACN 21483), right palpal bulb, pro- and ICMBio (Brazil), Subsecretar�ıa de Ecolog�ıa de lateral. Inset showing bulb microteeth. C, Loxosceles hirsuta, female (MACN 21483), left spermathecae, digested. D, La Pampa and Secretar�ıa de Ambiente y Desarrollo LOXOSCELES HIRSUTA MELLO-LEITAO~ same, close-up of pores. IS, inner spermatheca; OS, outer spermatheca; UE, uterus externus. Scale bars = A, 0.5 mm; B, Sustentable de R�ıo Negro and Administracion� de 0.1 mm; C, 0.05 mm; D, 0.02 mm. Figures 7, 14A, 63B–D Parques Nacionales (Argentina), Direccion� General

8D). PLS with aciniform gland spigots and a setal pit Loxoscella Strand, 1906: 668. Synonymized by Material examined. ARGENTINA. Cordoba� : LOXOSCELES SIMILLIMA LAWRENCE with a group of modified setae (Figs 7D, 8E). Palpal Gertsch & Ennik, 1983. Cordoba� ( 64.19192, 31.41975), A. Vexenat, 07/VI/ � � bulb simple, piriform (Fig. 63A, B). Inner Calheirosia Mello-Leitao,~ 1917: 80. Synonymized by Figures 18I, 63A, 8A–E 1975, 2 # 10 $ (MACN-Ar 21483). spermathecae usually simple (Fig. 63C), branched in Brignoli, 1978. the rufescens group. Outer spermathecae absent or Material examined: SOUTH AFRICA. Limpopo: LOXOSCELES CARIBBAEA GERTSCH present. Spermathecae pores small, with a single Diagnosis: As that of the subfamily. Soutpansberg, Farm Amsterdam 116, A. Schoeman, flagellum arising from them (Fig. 63D). Subadult 1 # 2 $ 2 imm. (TM 10611); 23/XI/1970, 1 # 5 $ 1 females without pre-spermatheca. Description: As that of the subfamily. imm. (TM 10602); Farm Weipe 47, W.D. Haacke, Material examined: CUBA. Santiago de Cuba: 23/II/1970, 1 # (TM 10738). Kwazulu-Natal: Reserva Ecologica� Siboney-Jutic�ı, Cueva La V�ırgen Distribution: Native of Africa, America and the Mzuke, Ndumo Game Reserve (32.15933, ( 75.7144, 19.96083), A. Perez� Gonzalez,� 04/V/2010, LOXOSCELES HEINECKEN & LOWE, 1832 � Mediterranean region. Some species (notably 26.87089), C. Haddad, 21/I/2006, 1 imm. (MACN- 1 # 1 $ (MACN-Ar 32734); 1 # 3 $ 3 imm. (MACN-Ar � Loxosceles Heinecken & Lowe in Lowe, 1832: 321. L. rufescens and L. laeta Nicolet) have been Ar 11316), 1 $ (MACN-Ar 11317); 17/I/2006, 1 # 32736). Type species Loxosceles citigrada Heinecken & Lowe, introduced to other regions (Gertsch & Ennik, 1983). (MACN-Ar 11319), 1 imm. (MACN-Ar 11320), 1 $ 1832 (= Loxosceles rufescens Dufour, 1820). (MACN-Ar 11321). LOXOSCELES AMAZONICA GERTSCH Omosites Walckenaer, 1833: 438. Synonymized by Species included: 107 known species (World Spider Simon, 1893. Catalog, 2015). LOXOSCELES SIMILIS MOENKHAUS Material examined: BRAZIL. Bahia: Central, Toca de Piloes~ (11°010–11°190S, 41°470–42°100W), E.F. Material examined: BRAZIL. Minas Gerais: Ramos & F.S. Cunha, III/2002, 4$ (IBSP 89007), 1# Simonesia� , RPPN Estacao~ Biologica� da Mata do 5$ (IBSP 89016), 1# 1 $ (IBSP 89071). Ceara:� Sossego ( 20.07361, 42.07028, 648 m), M.T.T. Quixada�, Fazenda Mage,� I.L.F. Magalhaes~ et al., 6/ � � Santos, 10-12/XII/2009, 1 $ (UFMG 12070), 1 $ IV/2012, 1 $ (UFMG 12044). (UFMG 12072), 1 # (UFMG 12073); Caratinga, RPPN Feliciano Miguel Abdala ( 19.77279, � LOXOSCELES DESERTA GERTSCH 42.16764, 742 m), A.A. Carosini Vera, XI/2011, 1 # �(UFMG 12075); same locality ( 20.07361, � 42.07028, 648 m), 1 $ (UFMG 12074); Imbe� de Material examined: USA. Arizona: Tucson, F.E. � Minas ( 19.86833, 43.96583, 818 m), C.S. Souza, 6/ Russell, 1 # (MACN-Ar 21497), 1 $ (MACN-Ar 21498). � � VIII/2011, 1 # (UFMG 12127). ACKOWLEDGEMENTS LOXOSCELES LAETA (NICOLET, 1849) We are deeply indebted to the curators of the institu- Figure 2 tions housing the material studied during the course of this work. We express our gratitude to all of them Material examined: BRAZIL. Santa Catarina: and to P. Marais (AcAT), N. I. Platnick and L. Sor- Curitibanos (27°16058″S, 50°35004″W), Taipas, S. kin (AMNH), A. Carmichael (CAS), F. Campon� Suzuki, 22/IV/1950, 1# (IBSP 34953). Sao~ Paulo: (CAI), N. Govender (DNSM), C. Stoffels (KNM), C. Osasco (23°31057″S, 46°47030″W), J.R. Bettinazzi, 26/ Grismado (MACN), L. Leibensperger (MCZ), A. Frie- VII/1982, 3# 3 $ (IBSP 34948). CHILE. Valpara�ıso: derichs and J. Kapp (ZMB), C. Barros (MNRJ), V. S. Ocoa, Border of Parque Nacional La Campana, alt. Costa (UFMG) and D. DeRoche (USNM), who kindly 355 m (32°54098″S, 71°05042″W), A.D. Brescovit made material from their collections available, pack- et al., 23/III/2013 (IBSP 163704) aged and sent specimens, and also offered their help and hospitality during visits to their institutions. We also thank several people who sent material under LOXOSCELES INTERMEDIA MELLO-LEITAO~ their care or specifically kept an eye for Sicarius in the field: A. V. Peretti, A. O. Porta, A. Taucare R�ıos, Material examined: BRAZIL. Santa Catarina: C. Calitra, H. A. Iuri, J. Ochoa, L. S. Carvalho, L. E. Sao~ Bento do Sul, Centro de Estudos e Pesquisas C. Schmidt and R. Botero-Trujillo. Many colleagues Ambientais Rugendas, I.L.F. Magalhaes~ et al., also accompanied us in the field trips: A. Taucare 13-16/XII/2011, 1 # (UFMG 12097), 1 # (UFMG R�ıos, B. T. Faleiro, C. Veloso, F. M. Hughes, G. F. B. 12098), 2 # 3 $ (UFMG 12099), 1 $ (UFMG Pereira, J. Cabra, L. Piacentini and T. J. Porto. Col- Figure 63. Scanning electronic microscopy, Loxosceles genitalia. A, Loxosceles simillima Lawrence, 1927, male (TM 12101). lecting permits were kindly issued by SEMA-Bahia 10611), left palpal bulb, prolateral. B, Loxosceles hirsuta Mello-Leitao,~ 1931, male (MACN 21483), right palpal bulb, pro- and ICMBio (Brazil), Subsecretar�ıa de Ecolog�ıa de lateral. Inset showing bulb microteeth. C, Loxosceles hirsuta, female (MACN 21483), left spermathecae, digested. D, La Pampa and Secretar�ıa de Ambiente y Desarrollo LOXOSCELES HIRSUTA MELLO-LEITAO~ same, close-up of pores. IS, inner spermatheca; OS, outer spermatheca; UE, uterus externus. Scale bars = A, 0.5 mm; B, Sustentable de R�ıo Negro and Administracion� de 0.1 mm; C, 0.05 mm; D, 0.02 mm. Figures 7, 14A, 63B–D Parques Nacionales (Argentina), Direccion� General

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Petrunkevitch A. 1911. A synonymic index-catalogue of spi- Sarkinen€ T, Iganci JRV, Linares-Palomino R, Simon APPENDIX 1 latter, the sternum has an anteriorly pointing ders of North, Central and South America with all adjacent MF, Prado DE. 2011. Forgotten forests–issues and pro- notch (CI = 0.333, RI = 0.818). List of characters used for inferring the trees. The islands, Greenland, Bermuda, West Indies, Tierra del spects in biome mapping using Seasonally Dry Tropical 12. Sternum, coloration. 0- uniform, 1- with dark first three characters have not been used in the Fuego, Galapagos, etc. Bulletin of the American Museum of Forests as a case study. BMC Ecology 11: 1–15. brown markings on border (CI = 0.5, RI = 0.5). Bayesian analysis or in the cladistic analysis with Natural History 29: 1–791. Simon E. 1893. Histoire naturelle des araignees� . Librairie 13. Chelicerae, fang serrula: 0- present (Fig. 14A), equal weights. Many characters, especially those of Pickard-Cambridge FO. 1899a. On some spiders from Encyclopedique� de Roret, Paris 1: 257–488. 1- absent (Fig. 14B) (CI = 1, RI = 1). outgroups, are only autapomorphic in the context of Chili and Peru collected by Dr. Plate of Berlin. Journal of Simon E. 1903. Histoire naturelle des araignees� . Librairie 14. Chelicerae, apophysis on paturon (male): 0- our dataset. Characters are unordered unless stated the Linnean Society of London 27: 15–22. Encyclopedique� de Roret, Paris 2: 669–1080. absent, 1- present. Autapomorphic for P. globo- Pickard-Cambridge FO. 1899b. Arachnida – Araneida and Strand E. 1906. Diagnosen nordafrikanischer, hauptsachlich€ otherwise. Consistency index and retention index sus. Opiliones. In: Godman FDC, Salvin O, eds. Biologia Cen- von Carlo Freiherr von Erlanger gesammelter Spinnen. refer to the tree in Figure 2. Characters used by 15. Chelicerae, stridulatory apparatus: 0- present, trali-Americana, Zoology. London: R.H. Porter, 41–88. Zoologischer Anzeiger 30: 604637, 655–690. Platnick et al. (1991) and Labarque & Ramırez 1- absent. Autapomorphic for D. spelunca Bon- Pizarro CM. 1944. El itinerario de Don Claudio Gay. Bolet�ın Strand E. 1908. Exotisch araneologisches.-I. Amerikanische (2012) are labelled respectively as P91: X and LR12: aldo, Rheims & Brescovit. del Museo Nacional de Historia Natural 22: 22–44. hauptsachlich€ in Peru, Bolivien und Josemitetal in Cali- X, with X denoting the character number in those 16. Chelicerae, teeth: 0- present, 1- absent. Autapo- Platnick NI, Coddington JA, Forster RR, Griswold CE. fornien gesammelte Spinnen. -II. Spinnen aus Kamerun. - studies. The character matrices analyzed in our work morphic for D. spelunca. 1991. Spinneret morphology and the phylogeny of haplog- III. Ubersicht€ der bekanten Hysterocrates-Arten. -IV. Zur are available in different formats as electronic sup- 17. Chelicerae, promarginal lobe, size: 0- small, 1- yne spiders (Araneae, Araneomorphae). 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Chelicerae, promarginal lobe, position in rela- (CI = 0.561, RI = 0.887). urwissenschaftlichen Gesellschaft zu Jena, Vol. 13. Jena: Tullgren A. 1901. Contribution to the knowledge of the spi- tion to the apex of cheliceral lamina: 0-lateral 2. Spermathecae, branches, number (female) Gustav Fischer, 203–246. der fauna of the Magellan Territories. Svenska Expeditio- (Fig. 14A, pl), 1- frontal (Fig. 14B, pl). In Rambaut A, Suchard MA, Xie D, Drummond AJ. 2014. nen till Magellanslanderna€ 2: 181–263. (CI = 0.394, RI = 0.649). Sicariinae, the promarginal lobe covers the Tracer v1.6. Online at http://beast.bio.ed.ac.uk/Tracer, Ung V, Dubus G, Zaragueta-Bagils€ R, Vignes-Lebbe R. apex of the chelicera lamina entirely in ante- accessed 31 August 2015. 2010. Xper²: introducing e-taxonomy. Bioinformatics 26: Cephalothorax rior view (see Labarque & Ramırez, 2012, fig. Ram�ırez MJ. 1989. Lista de los typos de Araneae descriptos 703–704. 3. Cephalic region, shape: 0- narrow (around 30% 15) (CI = 1, RI = 1). por Nicolet depositados en el MNHN. Bulletin du Centre Walckenaer CA. 1833. Memoire� sur une nouvelle classifica- of thoracic region width), 1- enlarged (around 19. Chelicerae, promarginal lobe setae, shape International de Documentation Arachnologique 6: 7–11. tion des araneides.� Annales de la Societ� e� Entomologique de 50% of thoracic region width) (Fig. 37). Autapo- (LR12: 29): 0- long and slender, 1- short and Ram�ırez MJ. 2000. Respiratory system morphology and the France 2: 414–446. morphic for S. yurensis stat. nov. conical (see Labarque & Ramırez, 2012, fig. 15) phylogeny of haplogyne spiders (Araneae, Araneomorphae). Walckenaer CA. 1847. Dernier Supplement.� In: Walckenaer 4. Anterior median eyes (P91: 0, LR12: 36): 0- pre- (CI = 1, RI = 1). 149–157. CA, Gervais P, eds. � . Journal of Arachnology 28: Histoire Naturelle des Insectes: Apteres sent, 1- absent. Autapomorphic for P. globosus 20. Chelicerae, lamina, apex, shape: 0- simple Ram�ırez MJ. 2014. The morphology and phylogeny of diony- Paris: Librairie Encyclopedique� de Roret, 365–564. (Taczanowski). (Fig. 13A, B) 1- bifid (Figs 13C, 14A, cl). In chan spiders (Araneae, Araneomorphae). Bulletin of the Werneck FP. 2011. The diversification of eastern South 5. Lateral eyes, position in relation to each other: addition to this, we considered a character American Museum of Natural History 390: 1–375. American open vegetation biomes: historical biogeography 0- juxtaposed, 1- apart by at least 1/2 diameter on the sclerotization of the lamina apex; it Reiskind J. 1966. Stereotyped burying behavior in Sicarius. and perspectives. Quaternary Science Reviews 30: 1630– (CI = 0.5, RI = 0.923). appears to be very sclerotized in Sicarius American Zoologist 9: 195–200. 1648. 6. Lateral eyes, position: 0- sitting directly on the Roewer CF. 1942. Katalog der Araneae von 1758 bis, 1940, World Spider Catalog. 2015. World Spider Catalog. Natu- and H. spatulata comb. nov., and weakly so carapace (Fig. 12A), 1- on the top of a tubercle in other species of Hexophthalma gen. reval. vol. 1. Bremen: Kommissions-Verlag von ‘Natura’. ral History Museum Bern, online at http://wsc.nmbe.ch, (Fig. 12B) (CI = 0.167, RI = 0.706). Ronquist F, Teslenko M, Van der Mark P, Ayres DL, version 16.5, accessed on August 27, 2015. However, we observed too many intermediate 7. Carapace, shape (LR12: 19). 0- moderately Darling A, Hohna€ S, Larget B, Liu L, Suchard MA, Zobel-Thropp PA, Bodner MR, Binford GJ. 2010. Com- conditions, and did not include a character arched, 1- strongly arched. Autapomorphic for Huelsenbeck JP. 2012. MrBayes 3.2: efficient Bayesian parative analyses of venoms from American and African on sclerotization in our analysis (CI = 1, S. fusca. phylogenetic inference and model choice across a large Sicarius spiders that differ in sphingomyelinase D activity. RI = 1) 8. Carapace, coloration pattern: 0- uniform, 1- model space. Systematic Biology 61: 539–542. Toxicon 55: 1274–1282. 21. Chelicerae, lamina, apex, membranous sheath: with dark lateral bands, 2- with a median vio- 0- absent, 1- present (Figs 13A, B, 14C, ms) lin-shaped band (CI = 1, RI = 1). (CI = 1, RI = 1). 9. Carapace, setae, texture: 0- smooth. 1- with SUPPORTING INFORMATION barbs. Setae with barbs adhere to soil particles Additional supporting information may be found online in the supporting information tab for this article: and are typical of Sicariinae (see Duncan et al., Male palp 2007) (CI = 1, RI = 1). 22. Palp, endite, serrula (LR12: 38): 0- present, 1- Appendix S1. Interactive key for identifying Sicarius using the Xper2 software. 10. Carapace, lateral macrosetae, number of rows: 0- absent (CI = 1, RI = 1). Appendix S2. Input file (.tnt) and trees from the discrete-only, unweighted parsimony analysis. a single row (Fig. 12A), 1- more than one row 23. Palp, femur, distal crack: 0- absent, 1- present. Appendix S3. Input file (.tnt) and trees from the discrete-only, weighted parsimony analysis. (Fig. 12B). Inapplicable for taxa without macrose- In Sicariinae, the palpal femur has a distinct Appendix S4. Input file (.tnt) and trees from the discrete and continuous, weighted parsimony analysis. tae in the carapace (CI = 0.167, RI = 0.688). sclerotized pseudosegment near the patella Appendix S5. Input file (.nex for MrBayes) and consensus tree from the discrete-only Bayesian inference 11. Sternum, posterior end, shape: 0- rounded to (Figs 22C, 43A, B) (CI = 1, RI = 1). analysis. pointed, 1- posteriorly notched. In the first 24. Palp, femur, thorns, position. 0- next to the Appendix S6. Character matrix in .nex format for Mesquite. state, the end of the sternum points to the femur base, 1- apart from the femur base (by space between the fourth coxae, while in the at least 4–5 diameters). This and the next

character is inapplicable for taxa without 40. Palp, bulb, embolus, shape (male): 0- straight prolateral, one retrolateral and one ventral. The Scytodoid families. We here report an addi- stridulatory apparatus (CI = 0.5, RI = 0.875). to slightly curved (Figs 41, 50), 1- distally bent four ventral ones may have longer macrosetae, tional state (short and bifid/trifid) which occurs 25. Palp, femur, proximal thorn, size and shape. 0- (Figs 46, 60), 2- sigmoid (Fig. 20A, B) (CI = 0.4, giving the leg a somewhat raptorial appearance in most specimens of Brazilian species small and spiniform (Fig. 18B, D, G, I), 1- large RI = 0.571). (Fig. 4D). In our dataset, none of the non-sicar- (although some specimens exhibit state 1). Con- and sculptured, with a median crest (Fig. 18A, 41. Palp, bulb, embolus, curvature in apical view iid outgroups has tibial macrosetae but we sidered unordered, although state 2 is almost C, E) (CI = 0.5, RI = 0.9). (male): 0- straight (Fig. 58E), 1- retrolaterally believe that this might be an artefact due to certainly derived from state 1. There is consid- 26. Palp, femur, dorsal proximal tuft of setae curved (Fig. 58D) (CI = 0.333, RI = 0.333). biased outgroup choice (see character 45), and erable variation in 3rd entapohysis shapes (male): 0- absent, 1- present (Fig. 54A, dht) 42. Palp, bub, embolus, keels (male). 0- absent, 1- that the synapomorphic condition for sicariid in coded as state 1: Loxosceles has it distally (CI = 1, RI = 1). Male Sicariinae have a con- present (Fig. 20C, E, arrows; 63B) (CI = 0,25, this case is not the presence of macrosetae per broadened; Sicarius has it distally tapering spicuous dorsal patch of long setae near the RI = 0.25). se, but its particular arrangement in eight rows (Fig. 5F); Hexophthalma gen. reval. has it base of the palpal femur (CI = 1, RI = 1). (CI = 1, RI = 1). broader than in Sicarius; and H. spatulata 27. Palp, femur, ventral proximal tuft of setae Legs 50. Legs, tibiae, macrosetae, length: 0- all macrose- comb. nov. has it very broad, short and squar- (male): 0- absent, 1- present (Fig. 54A, vht). 43. Legs, femora, prolateral black setae: 0- absent tae with the same length, 1- long macrosetae ish (Fig. 6D). These conditions are somewhat Autapomorphic for S. levii sp. nov. (Fig. 24A, E), 1- present (Fig. 24B–D). Some intercalated with short macrosetae (Fig. 4D). continuous and we preferred not to subdivide 28. Palp, femur, region near proximal thorn (male): Sicarius have groups of black setae in the Sicariinae have tibial macrosetae arranged in a this state (CI = 1, RI = 1). 0- not sclerotized, 1- sclerotized (Fig. 48, sa). femora; these setae differ structurally from the way reminiscent of the Mimetidae, as already 59. Abdomen, posterior respiratory system, fused Autapomorphic for S. boliviensis sp. nov. surrounding setae, resembling miniaturized noted by some previous authors (e.g. Mello- 3rd opisthosomal entapophyses, lateral 29. Palp, femur, length (male): 0- normal, 1- greatly versions of the dorsal macrosetae (Magalhaes~ Leitao,~ 1936) (CI = 1, RI = 1). branches (female) (LR12: 59): 0- present, 1- elongated. In most Loxosceles, the palpal femur is et al., 2013, figs 19, 20). They do not adhere to 51. Legs, tarsi, distal end, segmentation (LR12: absent. A Sicariinae synapomorphy, convergent longer than 55% the length of the carapace, and soil particles and can be seen even in dirty live 10): 0- entire (Fig. 4A), 1- distal articulation with P. globosus in our dataset (CI = 0.5, very slender (Gertsch, 1967: figs 1, 5, 8); in other specimens as black patches in the otherwise delimiting an onychium (Labarque & Ram�ırez, RI = 0.875). groups, the femur is shorter than up to 45% of soil-covered cuticle (Fig. 16C, E) (CI = 1, 2012, fig. 5A) (CI = 1, RI = 1). 60. Abdomen, posterior respiratory system, fused carapace length (CI = 1, RI = 1). RI = 1). 52. Leg I, tarsus, claw, tip, length: 0- same as 3rd opisthosomal entapophyses, lateral 30. Palp, tibia, length (male): 0- normal (up to 30% 44. Legs, femora, prolateral black setae, arrange- teeth, 1- greatly elongated (Fig. 4A). The sec- branches, shape (female) (LR12: 60): 0- simple, of carapace length), 1- greatly elongated ment: 0- large distal patch (Fig. 24D), 1- zig- ond condition is more common, and more 1- branched. Recovered as an ambiguous Lox- (around 60% of carapace length). Autapomor- zag pattern (Fig. 24B), 2- several discrete extremely pronounced, in desert species, espe- osceles synapomorphy in our analysis (CI = 1, phic for L. laeta. patches (Fig. 24C). Inapplicable for taxa with- cially S. mapuche sp. nov. and S. rupestris RI = 1). 31. Palp, cymbium, shape (male): 0- cup-shaped out black setae (CI = 1, RI = 1). (see Labarque & Ram�ırez, 2012, fig. 5C). Is 61. Abdomen, setae tuft near spinnerets (female): (Figs 34A, 59A, 60A), 1- distally projected (Figs 45. Legs, femora, macrosetae: 0- absent, 1- present. this an adaptation to psammophilous habitats? 0- absent, 1- present (Fig. 9A; Magalhaes~ et al., 41A, 48A, 20A) (CI = 0.2, RI = 0.429). The presence of femoral macrosetae is recov- (CI = 0.250, RI = 0.688). 2013; fig. 31, ht; Levi & Levi, 1969; figs 7–8). 32. Palp, cymbium, procursus (male): 0- absent, 1- ered as a sicariid synapomorphy in our analy- 53. Leg I, tarsus, prolateral claw, teeth rows Adult female Sicarius have this patch of very present. Autapomorphic for P. globosus. sis, although this is probably an artefact (LR12: 12): 0- one, 1- two. A classical character long and plumose setae (Fig. 9D) which they 33. Palp, base, spermophor (male): 0- not enlarged, caused by the choice of non-sicariid outgroups uniting Scytodidae and Drymusidae (Labarque apparently use for building the eggcase (see 1- enlarged, forming a bump against the wall (CI = 1, RI = 1). & Ram�ırez, 2012, and references therein) Levi & Levi, 1969: 32–33, 35). Please note that of the bulb (Fig. 21A, C, arrows) (CI = 1, 46. Legs, femora, macrosetae, colour: 0- brown, 1- (CI = 1, RI = 1). at least some Hexophthalma gen. reval. also RI = 1). white, 2- brown and white. This and the next 54. Legs, tarsi, inferior tarsal claw (P91: 36, LR12: build modified eggcases despite lacking this 34. Palp, bulb, transition between bulb base and two characters are inapplicable for taxa with- 11): 0- present, 1- absent. Two-clawed legs are setae tuft (see character 92), so characters are body (male): 0- smooth (Figs 50, 59), 1- steep out femora macrosetae (CI = 0.667, a classic synapomorphy uniting Sicariinae and not correlated (CI = 1, RI = 1). (Figs 48, 60) (CI = 0.25, RI = 0.25). RI = 0.875). Loxoscelinae (see Lehtinen, 1986; Platnick 62. Abdomen, spinnerets, colulus (female): 0- pre- 35. Palp, bulb, body, position (male): 0- inserted in 47. Legs, femora, dorsal macrosetae, shape: 0- et al., 1991) (CI = 1, RI = 1). sent, 1- absent. A classical Sicariinae synapo- the ventral portion of the base (Figs 50, 48), 1- with parallel sides, 1- medially widened 55. Leg I, metatarsus, prolateral group of curved morphy (CI = 0.5, RI = 0.9). inserted in the middle of the base (Fig. 20A, C; (Fig. 40A, C; Lotz, 2012; fig. 2B, E), 2- club- spines (male): 0- absent, 1- present. Autapo- 63. Abdomen, spinnerets, anterior lateral spin- Lotz, 2012, fig. 3A–C) (CI = 0.143, RI = 0.143). shaped (Figs 40D, 14D, 22A; Lotz, 2012, fig. morphic for L. laeta. nerets, shape (female). 0- short and conical, 1- 36. Palp, bulb, body, shape (male): 0- subconical 2D). The macrosetae are sexually dimorphic long and tubular. Loxosceles have ALS with (Figs 50, 48), 1- with parallel sides and distally in S. crustosus (medially widened in males, basal articles which are around three times narrowing (Fig. 20A, C; Lotz, 2012, fig. 3A–C) club-shaped in females); we coded this taxon Female abdomen longer than wide, and with a reduced spinning (CI = 1, RI = 1). as 1 because our observations were made pri- 56. Abdomen, coloration pattern: 0- uniform, 1- field with few piriform spigots (Fig. 8B) 37. Palp, bulb, embolus, embolic sclerite (male): 0- marily from males (CI = 0.667, RI = 0.667). striped or spotted (CI = 0.5, RI = 0.667). (CI = 1, RI = 1). absent, 1- present. Autapomorphic for P. globo- 48. Leg I, femora, anterior face, macrosetae, 57. Abdomen, macrosetae, shape: 0- with parallel 64. Abdomen, spinnerets, anterior lateral spin- sus. length: 0- shorter than dorsal macrosetae (less sides, 1- club-shaped (Fig. 22A) (CI = 0.5, nerets, setae, colour and shape (female): 0- 38. Palp, bulb, embolus base, shape (male): 0- slen- than 50% of the length) (Fig. 24A, D), 1- as RI = 0). brown and simple, 1- black and plumose (Figs der, 1- flat, ribbon-shaped (Fig. 61C) (CI = 1, long as the dorsal macrosetae (Fig. 24B, E) 58. Abdomen, posterior respiratory system, fused 9A–C, 12C, Magalhaes~ et al., 2013, fig. 32) RI = 1). (CI = 0.333, RI = 0.867). 3rd opisthosomal entapophyses, shape (female) (CI = 1, RI = 1). 39. Palp, bulb, embolus, shape: 0- slender, 1- broad 49. Legs, tibiae, macrosetae: 0- absent, 1- present, (LR12: 55): 0- short and slender, 1- elongated 65. Abdomen, spinnerets, anterior lateral spin- and blunt (Fig. 22D, E). Autapomorphic for in eight rows. Sicariids have tibial macrosetae and broad, 2- short and bifid/trifid (Fig. 12D). A nerets, setae, length (female): 0- shorter than H. spatulata comb. nov. arranged in four pairs of rows, one dorsal, one character differentiating Sicariidae from other spigots (Fig. 12C), 1- longer than spigots

© 2017 The Linnean© 2016 TheSociety Linnean of London, Society Zoological of London, JournalZoological of the JournalLinnean of Society the Linnean, 2017, 179 Society, 767–864, 2016 © 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016 94 I. L. F. MAGALHAES ET AL. SICARIIDSICARIID PHYLOGENY PHYLOGENY AND AND REVISION REVISION OF OF SICARIUSSICARIUS 86195 character is inapplicable for taxa without 40. Palp, bulb, embolus, shape (male): 0- straight prolateral, one retrolateral and one ventral. The Scytodoid families. We here report an addi- stridulatory apparatus (CI = 0.5, RI = 0.875). to slightly curved (Figs 41, 50), 1- distally bent four ventral ones may have longer macrosetae, tional state (short and bifid/trifid) which occurs 25. Palp, femur, proximal thorn, size and shape. 0- (Figs 46, 60), 2- sigmoid (Fig. 20A, B) (CI = 0.4, giving the leg a somewhat raptorial appearance in most specimens of Brazilian species small and spiniform (Fig. 18B, D, G, I), 1- large RI = 0.571). (Fig. 4D). In our dataset, none of the non-sicar- (although some specimens exhibit state 1). Con- and sculptured, with a median crest (Fig. 18A, 41. Palp, bulb, embolus, curvature in apical view iid outgroups has tibial macrosetae but we sidered unordered, although state 2 is almost C, E) (CI = 0.5, RI = 0.9). (male): 0- straight (Fig. 58E), 1- retrolaterally believe that this might be an artefact due to certainly derived from state 1. There is consid- 26. Palp, femur, dorsal proximal tuft of setae curved (Fig. 58D) (CI = 0.333, RI = 0.333). biased outgroup choice (see character 45), and erable variation in 3rd entapohysis shapes (male): 0- absent, 1- present (Fig. 54A, dht) 42. Palp, bub, embolus, keels (male). 0- absent, 1- that the synapomorphic condition for sicariid in coded as state 1: Loxosceles has it distally (CI = 1, RI = 1). Male Sicariinae have a con- present (Fig. 20C, E, arrows; 63B) (CI = 0,25, this case is not the presence of macrosetae per broadened; Sicarius has it distally tapering spicuous dorsal patch of long setae near the RI = 0.25). se, but its particular arrangement in eight rows (Fig. 5F); Hexophthalma gen. reval. has it base of the palpal femur (CI = 1, RI = 1). (CI = 1, RI = 1). broader than in Sicarius; and H. spatulata 27. Palp, femur, ventral proximal tuft of setae Legs 50. Legs, tibiae, macrosetae, length: 0- all macrose- comb. nov. has it very broad, short and squar- (male): 0- absent, 1- present (Fig. 54A, vht). 43. Legs, femora, prolateral black setae: 0- absent tae with the same length, 1- long macrosetae ish (Fig. 6D). These conditions are somewhat Autapomorphic for S. levii sp. nov. (Fig. 24A, E), 1- present (Fig. 24B–D). Some intercalated with short macrosetae (Fig. 4D). continuous and we preferred not to subdivide 28. Palp, femur, region near proximal thorn (male): Sicarius have groups of black setae in the Sicariinae have tibial macrosetae arranged in a this state (CI = 1, RI = 1). 0- not sclerotized, 1- sclerotized (Fig. 48, sa). femora; these setae differ structurally from the way reminiscent of the Mimetidae, as already 59. Abdomen, posterior respiratory system, fused Autapomorphic for S. boliviensis sp. nov. surrounding setae, resembling miniaturized noted by some previous authors (e.g. Mello- 3rd opisthosomal entapophyses, lateral 29. Palp, femur, length (male): 0- normal, 1- greatly versions of the dorsal macrosetae (Magalhaes~ Leitao,~ 1936) (CI = 1, RI = 1). branches (female) (LR12: 59): 0- present, 1- elongated. In most Loxosceles, the palpal femur is et al., 2013, figs 19, 20). They do not adhere to 51. Legs, tarsi, distal end, segmentation (LR12: absent. A Sicariinae synapomorphy, convergent longer than 55% the length of the carapace, and soil particles and can be seen even in dirty live 10): 0- entire (Fig. 4A), 1- distal articulation with P. globosus in our dataset (CI = 0.5, very slender (Gertsch, 1967: figs 1, 5, 8); in other specimens as black patches in the otherwise delimiting an onychium (Labarque & Ram�ırez, RI = 0.875). groups, the femur is shorter than up to 45% of soil-covered cuticle (Fig. 16C, E) (CI = 1, 2012, fig. 5A) (CI = 1, RI = 1). 60. Abdomen, posterior respiratory system, fused carapace length (CI = 1, RI = 1). RI = 1). 52. Leg I, tarsus, claw, tip, length: 0- same as 3rd opisthosomal entapophyses, lateral 30. Palp, tibia, length (male): 0- normal (up to 30% 44. Legs, femora, prolateral black setae, arrange- teeth, 1- greatly elongated (Fig. 4A). The sec- branches, shape (female) (LR12: 60): 0- simple, of carapace length), 1- greatly elongated ment: 0- large distal patch (Fig. 24D), 1- zig- ond condition is more common, and more 1- branched. Recovered as an ambiguous Lox- (around 60% of carapace length). Autapomor- zag pattern (Fig. 24B), 2- several discrete extremely pronounced, in desert species, espe- osceles synapomorphy in our analysis (CI = 1, phic for L. laeta. patches (Fig. 24C). Inapplicable for taxa with- cially S. mapuche sp. nov. and S. rupestris RI = 1). 31. Palp, cymbium, shape (male): 0- cup-shaped out black setae (CI = 1, RI = 1). (see Labarque & Ram�ırez, 2012, fig. 5C). Is 61. Abdomen, setae tuft near spinnerets (female): (Figs 34A, 59A, 60A), 1- distally projected (Figs 45. Legs, femora, macrosetae: 0- absent, 1- present. this an adaptation to psammophilous habitats? 0- absent, 1- present (Fig. 9A; Magalhaes~ et al., 41A, 48A, 20A) (CI = 0.2, RI = 0.429). The presence of femoral macrosetae is recov- (CI = 0.250, RI = 0.688). 2013; fig. 31, ht; Levi & Levi, 1969; figs 7–8). 32. Palp, cymbium, procursus (male): 0- absent, 1- ered as a sicariid synapomorphy in our analy- 53. Leg I, tarsus, prolateral claw, teeth rows Adult female Sicarius have this patch of very present. Autapomorphic for P. globosus. sis, although this is probably an artefact (LR12: 12): 0- one, 1- two. A classical character long and plumose setae (Fig. 9D) which they 33. Palp, base, spermophor (male): 0- not enlarged, caused by the choice of non-sicariid outgroups uniting Scytodidae and Drymusidae (Labarque apparently use for building the eggcase (see 1- enlarged, forming a bump against the wall (CI = 1, RI = 1). & Ram�ırez, 2012, and references therein) Levi & Levi, 1969: 32–33, 35). Please note that of the bulb (Fig. 21A, C, arrows) (CI = 1, 46. Legs, femora, macrosetae, colour: 0- brown, 1- (CI = 1, RI = 1). at least some Hexophthalma gen. reval. also RI = 1). white, 2- brown and white. This and the next 54. Legs, tarsi, inferior tarsal claw (P91: 36, LR12: build modified eggcases despite lacking this 34. Palp, bulb, transition between bulb base and two characters are inapplicable for taxa with- 11): 0- present, 1- absent. Two-clawed legs are setae tuft (see character 92), so characters are body (male): 0- smooth (Figs 50, 59), 1- steep out femora macrosetae (CI = 0.667, a classic synapomorphy uniting Sicariinae and not correlated (CI = 1, RI = 1). (Figs 48, 60) (CI = 0.25, RI = 0.25). RI = 0.875). Loxoscelinae (see Lehtinen, 1986; Platnick 62. Abdomen, spinnerets, colulus (female): 0- pre- 35. Palp, bulb, body, position (male): 0- inserted in 47. Legs, femora, dorsal macrosetae, shape: 0- et al., 1991) (CI = 1, RI = 1). sent, 1- absent. A classical Sicariinae synapo- the ventral portion of the base (Figs 50, 48), 1- with parallel sides, 1- medially widened 55. Leg I, metatarsus, prolateral group of curved morphy (CI = 0.5, RI = 0.9). inserted in the middle of the base (Fig. 20A, C; (Fig. 40A, C; Lotz, 2012; fig. 2B, E), 2- club- spines (male): 0- absent, 1- present. Autapo- 63. Abdomen, spinnerets, anterior lateral spin- Lotz, 2012, fig. 3A–C) (CI = 0.143, RI = 0.143). shaped (Figs 40D, 14D, 22A; Lotz, 2012, fig. morphic for L. laeta. nerets, shape (female). 0- short and conical, 1- 36. Palp, bulb, body, shape (male): 0- subconical 2D). The macrosetae are sexually dimorphic long and tubular. Loxosceles have ALS with (Figs 50, 48), 1- with parallel sides and distally in S. crustosus (medially widened in males, basal articles which are around three times narrowing (Fig. 20A, C; Lotz, 2012, fig. 3A–C) club-shaped in females); we coded this taxon Female abdomen longer than wide, and with a reduced spinning (CI = 1, RI = 1). as 1 because our observations were made pri- 56. Abdomen, coloration pattern: 0- uniform, 1- field with few piriform spigots (Fig. 8B) 37. Palp, bulb, embolus, embolic sclerite (male): 0- marily from males (CI = 0.667, RI = 0.667). striped or spotted (CI = 0.5, RI = 0.667). (CI = 1, RI = 1). absent, 1- present. Autapomorphic for P. globo- 48. Leg I, femora, anterior face, macrosetae, 57. Abdomen, macrosetae, shape: 0- with parallel 64. Abdomen, spinnerets, anterior lateral spin- sus. length: 0- shorter than dorsal macrosetae (less sides, 1- club-shaped (Fig. 22A) (CI = 0.5, nerets, setae, colour and shape (female): 0- 38. Palp, bulb, embolus base, shape (male): 0- slen- than 50% of the length) (Fig. 24A, D), 1- as RI = 0). brown and simple, 1- black and plumose (Figs der, 1- flat, ribbon-shaped (Fig. 61C) (CI = 1, long as the dorsal macrosetae (Fig. 24B, E) 58. Abdomen, posterior respiratory system, fused 9A–C, 12C, Magalhaes~ et al., 2013, fig. 32) RI = 1). (CI = 0.333, RI = 0.867). 3rd opisthosomal entapophyses, shape (female) (CI = 1, RI = 1). 39. Palp, bulb, embolus, shape: 0- slender, 1- broad 49. Legs, tibiae, macrosetae: 0- absent, 1- present, (LR12: 55): 0- short and slender, 1- elongated 65. Abdomen, spinnerets, anterior lateral spin- and blunt (Fig. 22D, E). Autapomorphic for in eight rows. Sicariids have tibial macrosetae and broad, 2- short and bifid/trifid (Fig. 12D). A nerets, setae, length (female): 0- shorter than H. spatulata comb. nov. arranged in four pairs of rows, one dorsal, one character differentiating Sicariidae from other spigots (Fig. 12C), 1- longer than spigots

(Fig. 9A–C; Magalhaes~ et al., 2013; fig. 32). As H. spatulata comb. nov. has just bare cuticle. different coding schemes. Thus, we prefer to 91. Abdomen, spermathecae, ventral branches, noted by Platnick et al. (1991), Neotropical Autapomorphic for H. spatulata comb. nov. take the first approach, mostly because in Hex- shape (female): 0- straight, 1- with a single coil Sicarius have very long and dense ALS setae, 72. Abdomen, posterior median spinnerets, spigots ophthalma gen. reval. and Loxosceles, the (Fig. 46), 2- with a double coil (Figs 19B, 45, which hamper examination of the spigots; other (female): 0- present, 1- absent (Figs 7C, 8D). A outer spermathecae is commonly reduced (or 59, vb). Considered ordered (CI = 1, RI = 1). Sicariinae have normal setae (CI = 1, RI = 1). synapomorphy of Loxosceles (CI = 1, RI = 1). absent) (except for L. hirsuta, which has a 92. Abdomen, spermathecae, sclerotized pore plates 66. Abdomen, anterior lateral spinnerets, number 73. Abdomen, posterior lateral spinnerets, modified reduced inner spermathecae) (CI = 0.25, (female): 0- absent, 1- present (Figs 19B, 45, of articles (female) (P91: 40, LR12: 67): 0- setae (female) (see P91: 57, LR12: 82): 0- RI = 0.75). 46I, 59, pp) (CI = 1, RI = 1). three, 1- two (CI = 1, RI = 1). absent, 1- present (Figs 7D, 8F). These setae 79. Abdomen, spermathecae, outer spermathecae, 67. Abdomen, anterior lateral spinnerets, major arise from a prolateral pit in the PLS of Lox- shape (female): 0- well developed, with a stalk Behaviour ampullate gland spigot, number (female) (P91: osceles (CI = 1, RI = 1). (Fig. 20F, 63C), 1- reduced, without stalk For Sicarius, observations are from our own field 33, LR12: 70): 0- two, 1- one, 2- none (Figs 74. Abdomen, posterior lateral spinnerets, modified (Fig. 20B, D). This and the next character are work. Data for Hexophthalma have been coded from 6A, 11B). Considered ordered. Species with setae, apex shape (female): 0-spatulate inapplicable for taxa without outer spermathe- photos of unidentified specimens from Africa, which one MAP spigot plus one MAP spigot nubbin (Fig. 8F), 1- ragged (Fig. 7D). Inapplicable for cae (CI = 0.5, RI = 0.8). have been coded under H. hahni comb. rest. have been coded as 0. Sicarius have been taxa without modified setae. Although in our 80. Abdomen, spermathecae, outer spermathecae, 93. Eggcase, position: 0- carried in the chelicerae, coded as 0 or 1 (contra Platnick et al., 1991; tree this character has ambiguous optimization, shape (female): 0- simple (Fig. 20B, D), 1- 1- attached to the substrate. Drymusa spelunca see comment in next character) (CI = 0.4, we believe the ragged state to be apomorphic branched (Fig. 20F). Autapomorphic for has been coded after other species of Drymusa RI = 0.5). because a spatulate morphology is more similar H. spatulata comb. nov. (D. serrana Goloboff & Ram�ırez does not attach 68. Abdomen, anterior lateral spinnerets, major to that of ordinary setae. This would lend fur- 81. Abdomen, spermathecae, inner spermathecae, the eggcase to the substrate, but rather to its ampullate gland spigots, shape (female): 0- nor- ther support to the position of Loxosceles simil- branches, insertion (female): 0- arising from a web; D. dinora Valerio and D. spectata Alayon-� mal, 1- one spigot modified into a nubbin lima (coded 0) as sister to all other Loxosceles single stalk (Fig. 19C), 1- inserted throughout Garc�ıa carry it in the chelicerae; see Labarque (Labarque & Ram�ırez, 2012; fig. 24B, C), 2- (coded 1) in our analysis. the bursa (Fig. 19A, B, D) (CI = 1, RI = 1). & Ram�ırez, 2007) (CI = 1, RI = 1). long and filiform (Figs 9E, 10B, map). This and 82. Abdomen, spermathecae, ventral pores, size 94. Eggcase, shape: 0- rounded to oval, 1- pot- the next character are inapplicable for taxa Female genitalia and colour (female): 0- small, inconspicuous shaped and covered with dirt (Fig. 17). A classi- without MAP spigots. Platnick et al. (1991) 75. Abdomen, epigynum (female): 0- absent, 1- pre- and light brown, 1- large and orange (Figs 49, cal synapomorphy of Sicariinae (CI = 1, RI = 1). coded Sicarius as lacking MAP spigots, sent. Autapomorphic for P. globosus. 54; also see photos in the interactive key, Sup- 95. Eggcase, collar: 0- absent (Fig. 17D), 1- present although they comment that ‘one spigot with a 76. Abdomen, post epigastric fovea (female) (LR12: porting Information, Appendix S1) (CI = 1, (Fig. 17A, B, C, E). In our analysis the collar is straighter shaft may serve the major ampullate 95): 0- absent, 1- present. Autapomorphic for RI = 1). a synapomorphy of all Sicarius but gland’ (p. 56). We prefer to follow this latter S. fusca. 83. Abdomen, spermathecae, length (female): 0- S. thomisoides and S. yurensis stat. nov. Unfor- interpretation, as those spigots have a different 77. Abdomen, spermathecae, pre-spermathecae longer than uterus externus, 1- shorter than tunately we could not observe eggcases of base morphology than the surrounding piri- (subadult females): 0- absent, 1- present. In uterus externus (Fig. 54). Autapomorphic for S. yurensis stat. nov., which would provide fur- forms and lie in the posterior margin of the most Sicarius and at least two Hexophthalma S. levii sp. nov. ther data to optimize this character (CI = 1, spinning field, well apart from other spigots gen. reval., we could find subadult females bear- 84. Abdomen, spermathecae, spacing (female): 0- RI = 1). (CI = 1, RI = 1). ing a pre-spermathecae. This pre-spermathecae closely spaced (Fig. 33A, D, I, J, L), 1- widely 96. Self-burying behaviour: 0- absent, 1- present. 69. Abdomen, anterior lateral spinnerets, major has a shape reminiscent of that of adults, but spaced (Figs 20B, D, 23A, 33G). Spermathecae Sicariinae have a stereotyped behaviour for ampullate gland spigot, aperture (female) not yet totally formed (it seems to lack glandu- were considered widely spaced when their burying themselves in fine sediment (Reiskind, (LR12: 73): 0- rounded, 1- slit-like (Figs 7B, lar pores and usually has shortened propor- bases are apart by a distance greater than 1966) (CI = 1, RI = 1). 8C). A character traditionally recognized as a tions). At least Neotropical Loxosceles subadult their length (CI = 0.2, RI = 0.692). 97. Web-building behaviour: 0- present, 1- absent. synapomorphy of Loxosceles. We have coded females lack such pre-spermathecae (A. D. 85. Abdomen, spermathecae, rounded reservatory None of the species of Sicarius which we Physocyclus Simon, Drymusa Simon and Scy- Brescovit, pers. observ.), although Brignoli in the apex (female): 0- absent (Fig. 46), 1- pre- have had the opportunity to observe in the todes Latreille based on co-familiars imaged (1969, figs 26, 40, 46) reported variations in sent (Fig. 51) (CI = 0.125, RI = 0.5). field spins webs or drag-lines, or uses silk for and analysed by Labarque & Ram�ırez (2012): L. rufescens spermathecae that might be consid- 86. Abdomen, spermathecae, apex, shape (female): any purpose other than reproductive activities L. laeta, L. rufescens and L. deserta (a close ered pre-spermathecae. We have not examined 0- not bent (Figs 46, 51), 1- bent (Figs 35, 39) (eggcases and sperm webs) (CI = 0.5, relative of L. reclusa) after Platnick et al. enough material of other taxa in our dataset, so (CI = 0.5, RI = 0.5). RI = 0.857). (1991); L. hirsuta, L. simillima, S. crustosus they have been coded ‘?’ (CI = 1, RI = 1). 87. Abdomen, spermathecae, stalk, shape (female): and S. levii after our own SEM images; and 78. Abdomen, spermathecae, outer spermathecae 0- straight (Fig. 53), 1- coiled (Fig. 51) the remaining Sicarius species by examining (female): 0- absent, 1- present (Fig. 20, os). (CI = 0.167, RI = 0.545). Venom their spigots under the stereomicroscope Neotropical Sicarius have branched spermath- 88. Abdomen, spermathecae, digitiform projections 98. Venom, SicTox alpha protein: 0- absent, 1- pre- (CI = 1, RI = 1). ecae which are directly inserted in the bursa. (female): 0- absent, 1- present (Figs 19B, 59). sent. The SicTox gene family includes several 70. Abdomen, posterior median spinnerets (fe- It seems impossible to determine whether Autapomorphic for S. utriformis. paralogues of genes expressing proteins similar male): 0- present, 1- absent (Figs 6A, B, 11A, these spermathecae are homologous to the 89. Abdomen, spermathecae, ventral branches (fe- to sphingomyelinase D. These paralogues have C, 12C). African Sicariinae have lost their PMS inner spermathecae, to the outer one, or both. male): 0- absent (Figs 60, 49), 1- present (Figs been found to belong in two main clades, ter- (CI = 1, RI = 1). We have tried coding Neotropical Sicarius as 19A, B, 42, 54, vb) (CI = 0.2, RI = 0.556). med a and b (Binford et al., 2009). This charac- 71. Abdomen, posterior median spinnerets setae lacking outer spermathecae (and considering 90. Abdomen, spermathecae, ventral branches, ter refers to the presence of proteins derived (female): 0- present (Fig. 11A, C), 1- absent their spermathecae as homologous to the inner number (female): 0- one, 1- two or more from paralogues of the SicTox a clade in the (Figs 12C, 6A, B). Most Hexophthalma gen. spermathecae), or the inverse pattern. Topol- (Fig. 41; Magalhaes~ et al., 2013, fig. 119) spider’s venom. This and the next two charac- reval. retains setae in the place of PMS, while ogy and tree length were unaffected by these (CI = 0.5, RI = 0). ters have been coded after Binford et al. (2009).

Loxosceles simillima has been coded after APPENDIX 2 L. spinulosa, L. similis after L. variegata. This and the next character have been coded as 0 MATERIAL EXAMINED FOR SCORING NON-SICARIID for non-sicariid outgroups because, in spiders, OUTGROUPS. proteins of the SicTox family have been detected only in sicariids so far. In our study, Physocyclus globobus (Taczanowski) (Pholcidae): the possession of SicTox alpha protein par- BRAZIL. Minas Gerais: Marlieria� , Parque alogues emerges as the synapomorphy of all Estadual do Rio Doce ( 19.71628, 42.7338, 597 m), Loxosceles but L. simillima (CI = 1, RI = 1). À À T. Rodrigues et al., 2/IX/2003, 3 # 4 $ (UFMG 99. Venom, SicTox beta protein: 0- absent, 1- pre- 11457); Prudente de Morais, Fazenda do Sape,� sent. This character refers to the presence of rodovia MG-424 ( 19.46838, 44.24161, 850 m), proteins derived from paralogues of the SicTox À À E.S.S. Alvares,� 28/V/2000, 2 # 2$ (UFMG 384). b clade in the spider’s venom (see discussion under character 98). Recovered in our study as Drymusa spelunca Bonaldo, Rheims & Brescovit a Sicariidae synapomorphy (CI = 1, RI = 1). (Drymusidae): BRAZIL. Para�: Parauapebas, 100. Venom, sphingomyelinase D activity: 0- absent, Floresta Nacional de Carajas,� Pereira, Gruta N1-170, 1- present. Scoring for S. ornatus from Lopes Equipe Carste col., 28/IX–03/X/2007, 2# 16$ (IBSP et al. (2013). Optimization for this character is 97678), 2# 5$ (IBSP 97680). ambiguous, as both Loxosceles and Hexoph- thalma gen. reval. venoms show Smase-D Scytodes fusca Walckenaer (Scytodidae): BRAZIL. activity in vitro or in vivo, while most Sicarius Acre: Senador Guiomard, Estacao~ Experimental screened lack this activity (but see Lopes et al., Catuaba ( 10.07333, 67.62389, 231 m), G.H.F. 2013). It must be noted that our current under- À À Azevedo & A.J. Santos, XI/2010, 1 $ (UFMG 10579). standing on SMase-D activity across different Minas Gerais: Belo Horizonte ( 19.86139, Sicariidae species is still incomplete, so its scor- À 43.96333), A.J. Santos et al., VI–VIII/1993, 1 # 4 $ ing is not straightforward. Thus, the optimiza- (UFMGÀ 232); Bairro Horto Florestal ( 19.91889, tion of this character must be viewed with À 43.93861, 845 m), E.S.S. Alvares,� I/2001, 1 # À caution. Hexophthalma spatulata comb. nov. (UFMG 230); Tres^ Marias ( 19.86667, 43.96667), has not been subject to studies of this kind and À À N.G. Fonseca, 5-12/XII/2007, 1 $ 1 i mm. (UFMG, thus its venom activity remains unknown 1972). (CI = 0.333, RI = 0.667).