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1 August 2003 THE SPIDER SUBFAMILY AMAUROBIOIDINAE (ARANEAE, ANYPHAENIDAE): A PHYLOGENETIC REVISION AT THE GENERIC LEVEL
MARTÍN J. RAMÍREZ
Author Affiliations +
Abstract

A cladistic phylogenetic analysis at generic level of the subfamily Amaurobioidinae is presented. The analysis is based on a dataset of 93 representative species scored for one behavioral and 199 morphological characters. Tree searches were made under equal and implied weights according to homoplasy, and the results were compared in terms of sensitivity to jackknife resampling. Mildest weighting functions produced trees more robust to resampling, and those results were selected as the working phylogenetic hypotheses. Groups of weak support as identified by jackknifing and Bremer indices are in general those that vary in resolution with different character-weighting schemes.

Seven outgroup representatives were included (Malenella nana Ramírez, from Malenellinae, and six Anyphaeninae species). In this analysis Anyphaeninae, previously identified as sister group of Amaurobioidinae, is paraphyletic, but forcing its monophyly does not alter the groupings within Amaurobioidinae. The monophyly of the genera is in general well supported, but some particularly conflicting groups are discussed. In contrast, the relationships among genera are in general problematic.

Amaurobioidinae is diagnosed by a pronounced indentation at the base of male palpal tegulum, and by a particular male copulatory bulb conformation, with a paramedian apophysis. The subfamily is classified in two tribes (Gayennini and Amaurobioidini); the genus Josa Keyserling, probably sister group to Gayennini, is not assigned to either tribe.

The tribe Amaurobioidini is mainly diagnosed by an apical loop of the sperm duct in the male copulatory bulb. It includes 10 genera: Amaurobioides O.P.-Cambridge is restricted to seashores of southern continents. Clubiona chilensis Nicolet, transferred to Amaurobioides, is the first true record of the genus for South America. The male of Axyracrus elegans Simon, three species of Aysenia Tullgren, and three of Coptoprepes Simon are newly described. Four new genera are proposed in Amaurobioidini: Gamakia, Selknamia (described for one new species each), Aysenoides (for three new species), and Negayan (type species Gayenna tridentata Simon, including also Axyracrus coccineus Mello-Leitão, Clubiona paduana Karsch, Gayenna excepta Tullgren, Gayenna exigua Mello-Leitão, and Tomopisthes lebruni Simon). The previously revised genera Acanthoceto Mello-Leitão and Ferrieria Tullgren are also included in the tribe. The basal branch and most intergeneric branches of the tribe have low support values. Amaurobioides and Negayan, however, are relatively well supported.

The tribe Gayennini is well defined by a homogeneous conformation of male and female genitalia, with a distinctive secondary conductor and spherical spermathecae. It includes 11 genera: Gayenna Nicolet includes only G. americana Nicolet from Chile and adjacent Argentina. Arachosia O.P.-Cambridge comprises many species previously assigned to Oxysoma. Abuzaida striata

INTRODUCTION

The Anyphaenidae is a homogeneous family of small to medium-sized, wandering hunter spiders. The group is relatively uniform and well defined, both morphologically and geographically. They have characteristic claw tufts composed of flattened setae, and a particularly well-developed tracheal system, externally evident by the wide, advanced tracheal spiracle. Most diversity of the family occurs in the New World, especially South America, with 29 endemic genera out of 54.

The extended tracheal system of Anyphaena Sundevall motivated Bertkau (1878) to erect the family Anyphaenidae. In subsequent years, anyphaenids were considered either a separate family or members of the families Clubionidae and the old Drassidae (today mostly Gnaphosidae), depending on the inclination of the authors to cherish or decry characters from internal anatomy (see Platnick, 1974; Brescovit, 1997; Ramírez, 1995a). However, the group in itself, beyond its hierarchic position, was never disputed.

A small cladistic analysis of the family (Ramírez, 1995a) resulted in a classification of Anyphaenidae in three subfamilies. The most basal group, Malenellinae, includes only the Chilean Malenella nana Ramírez. The Anyphaeninae was revised at the generic level by Brescovit (1997), and it is probably the sister group of Amaurobioidinae. These last two subfamilies mostly correspond to the main divisions in traditional keys to genera (e.g., Simon, 1897a). The monophyly of Anyphaeninae is not well documented, but Amaurobioidinae may be easily recognized by a very characteristic male copulatory palp, with a deep basal indentation in the tegulum, occupied by a membranous area (Platnick, 1977; Ramírez, 1995a).

Most species of Amaurobioidinae were described in Gayenna Nicolet, Tomopisthes Simon, Oxysoma Nicolet, or Tasata Simon. These four genera were ambiguously defined in the classic literature (see Tullgren, 1901; Ramírez and Kochalka, 1993; Ramírez, 1995b), being mostly diagnosed by details in the position of the eyes. As shown here, these characters are among the most homoplasious of all the analysis. A nomenclatorial twist in the early history of the group further complicates this unfortunate fact. The most popular genera of Amaurobioidinae seemed to be Gayenna and Oxysoma, in the sense that specialists were inclined to describe new species in those genera. Both genera were proposed by Nicolet (1849) for two species of remarkable appearance and morphology, common in temperate forests of Chile. Nicolet's descriptions are of little use for identification, but the illustrations are good enough to distinguish these two species. This work by Nicolet is included in the “Historia Física y Política de Chile”, compiled by Claudio Gay in Spanish, but printed in Paris. The collection of spiders studied by Nicolet was deposited in the Museum National d'Histoire Naturelle in Paris, where Eugène Simon worked. Most of Nicolet's types, however, were thought to be lost, until about half of them were found mixed with general collections (Ramírez, 1989). It seems that Simon (1889, 1904) only examined a few of those types. That, together with the fact that the library of the museum in Paris apparently lacked the atlas with the illustrations from Nicolet, may explain how things unfolded. In 1897 Simon described two monotypic genera, Aporatea and Mezenia (Simon, 1987a), precisely for the same species for which Nicolet created Oxysoma and Gayenna, while ascribing Nicolet names to very different spiders. Simon placed in Oxysoma those amaurobioidines with a markedly procurved posterior eye row (here Arachosia and Tasata), but he left Gayenna loosely defined by characters also attributed to other genera (notably Tomopisthes; see Tullgren, 1901; Ramírez and Kochalka, 1993). Subsequent authors followed the characterization of Gayenna and Oxysoma given by Simon, often using Gayenna as a dump group for species that lacked the specific characters of better defined genera.

Modern revisionary works in Amaurobioidinae cover only a few genera. Forster (1970) revised Amaurobioides Hewitt, and Ramírez (1993, 1995b, 1997, 1999) presented cladistic revisions of Liparotoma (here part of Philisca), Monapia, Acanthoceto, and Ferrieria. The relationships among genera were preliminary studied for the closer relatives of Monapia and Acanthoceto (Ramírez, 1995b, 1997, 1999). Kochalka (1980) revised Josa Keyserling in an unpublished Master's thesis, which also includes considerations of the systematics of Amaurobioidinae, most of them already discussed in my previous papers.

The 22 genera of Amaurobioidinae that result from this revision include about 140 known species. In addition, the collections that I examined hold a similar number of undescribed species. The aim of this contribution is to produce a phylogenetic classification of the subfamily, settling the limits of the genera. Besides the representatives selected for the cladistic analysis, I assign revised generic placement to other species that I examined in the preliminary stages of this work. As explained in Cladistic Analysis, these species fit well in the genera as defined here.

MATERIALS AND METHODS

Specimens examined for this study are deposited in the following institutions:

  1. AMNH   American Museum of Natural History, New York, Norman Platnick

  2. BMNH   The Natural History Museum, London, Janet Beccaloni

  3. BPBM   Bishop Museum, Honolulu, David Preston

  4. CAS   California Academy of Sciences, San Francisco, Charles Griswold

  5. IBNP   Inventario Biológico Nacional de Paraguay, Asunción, John Kochalka

  6. IBSP   Instituto Butantan, São Paulo, Antonio Brescovit

  7. IML   Fundación Miguel Lillo, Tucumán, Pablo Goloboff

  8. IRSN   Institut Royal des Sciences Naturelles de Belgique, Brussels, Léon Baert

  9. MACN-Ar   Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires, Cristina Scioscia

  10. MBUV   Museo de Biología, Universidad Central de Venezuela, Caracas, R. Candia

  11. MCN   Fundação Zoobotânica do Rio Grande do Sul, Erica Buckup

  12. MCTP   Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Arno Lise

  13. MCZ   Museum of Comparative Zoology, Cambridge, Massachusets, Herbert Levi

  14. MHNP   Muséum National d'Histoire Naturelle, Paris, Christine Rollard

  15. MHNS   Museo Nacional de Historia Natural, Santiago, Ariel Camousseight

  16. MLP   Museo de La Plata, Luis Pereira

  17. MMLS   Museo Municipal de Ciencias Naturales Lorenzo Scaglia, Mar del Plata, Juan Farina

  18. MNRJ   Museu Nacional, Universidade Federal do Rio de Janeiro, Adriano Kury

  19. MZUSP   Museu de Zoologia, Universidade de São Paulo, Ricardo Pinto da Rocha

  20. NRS   Naturhistoriska Riksmuseet, Stockholm, Torbjörn Kronestedt

  21. SMF   Senkenberg Museum, Frankfurt, Manfred Grasshoff

  22. UC   Universidad de Concepción, Viviane Jerez

  23. UCB   University of California, Berkley, Elizabeth Arias

  24. UNESP   Instituto de Biociências, Universidade de São Paulo, Isabela Rinaldi

  25. UPBS   University of Plymouth, UK, Department of Biological Sciences, Peter Smithers

  26. USNM   National Museum of Natural History, Smithsonian Institution, Washington, D.C., Jonathan Coddington

  27. ZMB   Musem für Naturkunde, Institut für Systematische Zoologie, Berlin, Jason Dunlop

  28. ZMH   Zoologisches Museum, Hamburg, Hieronymus Dastych

  29. ZMK   Zoologisk Museum, Copenhagen, Nikolaj Scharff

The lists of material examined (other than the types) include over 6400 specimens. Localities are listed mostly as they appear in the labels, but were checked with maps and gazetteers. Distances and measures from labels are transcribed without conversion to the metric system. Annotations on the original labels are included (e.g., “under stones”, Malaise trap), but not taxonomic identifications.

Format of Descriptions

Measurements are in millimeters, taken with an ocular micrometer on a Leitz stereomicroscope. Measurements are given with two decimals, but accuracy between 1% and 2% is as follows: ±0.017 mm (for measurements <1.67 mm), 0.033 mm (1.70–3.33 mm), 0.133 (>3.33 mm). Carapace length is the maximum in dorsal view, not including chelicerae; total length is without chelicerae or spinnerets. Tibiae are measured between condyla, metatarsi from basal condyle to dorsal apical end. Measurements of total length, abdomen dimensions, and position of tracheal spiracle vary with physiological condition or preservation artifacts and are only intended to give an idea of the size and aspect of the specimen. Other measurements are difficult to take and may have considerable error: the distance between tracheal spiracle and spinnerets is measured up to the posterior margin of abdominal cuticle (because spinnerets are telescopable) and is quite inaccurate for small specimens with few setae. Length of female tarsal palp is problematic if telescoped into the tibia. Some types were examined without access to an ocular micrometer, and thus measurements are lacking for some species known only from the types.

Spine (macroseta) pattern is described in standard format, with slight intuitive modifications (ap = apical, bas = basal, d = dorsal, p = prolateral, r = retrolateral, v = ventral). In case a spine is not paired, it is indicated whether it is placed on a particular side: p 2-d1–1 means four spines on the prolateral side (two basals, one median dorsally displaced, and one apical on the median line). In case a segment bears only basal or apical spines, notation is abbreviated: 2ap is equivalent to 0–0–2, 1bas to 1–0–0, and p1ap to 0–0-p1. Only surfaces bearing spines are listed. Occasionally (mostly on femora) two spines are not strictly paired but are close to each other; these are associated in parentheses: 0-d1-(1-d1) would be equivalent to 0-0-0-0-d1-0-0-0-1-d1. If bristles (thin, reduced macrosetae) occur in place of spines, it is indicated: 1–1 bristles (two bristles in a median line) or 1-(1 bristle) (one spine and one bristle). When referring to individual spines, other positions of the generalized pattern (for instance, in characters 129–199) are replaced by “x”: v x-p1-x is the ventral median prolateral spine, regardless of whether the specimen bears v 2–2–2 or v 0-p1–2. Spine positions are approximate, with reference to a generalized pattern, and the notation is not strictly literal: v 2ap spines are close to the apical margin on tibia, but not so close on the metatarsus. In many cases notations like p d1-x-x and d p1-x-x may be equally appropriate, and they were arbitrarily (but consistently) settled according to the generalized pattern.

Abbreviations Used in Text and Tables

  1. AB   accessory bulb

  2. ALE   anterior lateral eye

  3. ALS   anterior lateral spinneret

  4. AME   anterior median eye

  5. APmf   anterior pouch on median field

  6. C1   primary conductor

  7. C2   secondary conductor

  8. C2p   prolateral portion of C2

  9. C2r   retrolateral portion of C2

  10. CD   copulatory duct

  11. CO   copulatory opening

  12. E   embolus

  13. FD   fertilization duct

  14. LL   lateral lobe

  15. LT   lateral trachea

  16. MA   median apophysis

  17. MT   median trachea

  18. PLE   posterior lateral eye

  19. PMA   paramedian apophysis

  20. PME   posterior median eye

  21. PMS   posterior median spinneret

  22. RTA   retrolateral tibial apophysis

  23. SCG   index of supported/contradicted groupings

  24. SD   sperm duct

All drawings were made with a camera lucida on a Leitz stereo or compound microscope. Spermathecae were cleared in clove oil and are illustrated with a compound microscope. The tracheal system was examined after digestion in KOH 10–20% in a double boiler. Spinnerets were critical-point dried for the scanning electorn microscope; all other structures were air dried. Primordia of epigyne were dissected from penultimate females close to ecdysis from regular collections. Spermathecae were prepared for scanning after digestion with contact lens cleaner overnight (Sierwald, 1990).

CLADISTIC ANALYSIS

Representatives

In this analysis, terminals are exemplar species, instead of hypothetical constructs or bauplans. A first step for terminal selection was the examination of all available type specimens and the larger collections of Amaurobioidinae over the world. From all these specimens I made a preliminary list of species, with at least a few sketches and notes for each one. These files comprise about three times as many species as included here, a number far beyond the scope of this paper. I decided then to base the analysis on representative species. More than one representative is included for each genus (except monotypic ones) and for groups of species that seemed reasonable candidates to deserve genus rank. For the sake of nomenclatorial stability, type species of genera are also included, when available. Because the main purpose of this study is to settle the limits and relationships of genera, I also included species bearing particularly conflicting combinations of characters. In turn, no further species was added if a very similar one had been included, or if it seemed evident that such species would not imply a problematic generic assignation. Finally, I included all species treated in previous cladistic analyses (genera Monapia, Acanthoceto, Liparotoma), because they do not add complications to the project, and it seemed interesting to test those previous hypotheses. Taxon sampling was increased after some preliminary analysis, but still some problems are not adequately resolved (e.g., the limits of Sanogasta). These weak points of the analysis are noted and will be addressed in the future.

Outgroups

The monophyly of Anyphaenidae is reasonably well founded, and the grouping of Anyphaeninae plus Amaurobioidinae is very well supported and has never been disputed (Ramírez, 1995a). This is so even though the closest relatives of Anyphaenidae are not yet determined, as is true for most dionychan families (Coddington and Levi, 1991), because anyphaenids are quite uniform in some characters not commonly found on other spiders (the details of the claw tufts and the tracheal system). Malenella nana was used to root the analysis. The Anyphaeninae, recently revised at the generic level by Brescovit (1997), includes 33 genera whose relationships are practically unknown. The monophyly of Anyphaeninae is at best weakly supported. I included six representative species, selected by having characters potentially conflicting with the monophyly and internal resolution of Amaurobioidinae. That is, they share conditions that are diagnostic for genera or groups of genera within Amaurobioidinae. This selection biases the analysis against monophyly of Anyphaeninae, and hence solutions with Anyphaeninae constrained to be monophyletic were also explored. Admittedly, some statements of homology among amaurobioidines and anyphaenines are questionable (e.g., those of male palpal conductors—see the taxonomic section Anyphaeninae for details).

Characters

Of the 200 characters listed below, 189 are informative for phylogeny. Most characters are from male (59) and female (28) sexual structures and from spines (71). The general somatic morphology, coloration, and spinnerets are coded in 38 characters. The tracheal system is coded in three characters that are considered logically related. Only one character of sexual behavior is included. Table 3 lists character statistics, and figure 4 shows the average fit that the character systems had after the analysis.

Coding

Multistate characters are considered additive when the states were interpreted as internested homologies; this is not intended to express assumptions on the evolution of characters, but merely reflects degrees of similarity (Lipscomb, 1992; Goloboff, 1997a). Morphoclines were interpreted as internested homologies. Because some authors have criticized this approach as an unjustified assumption, additional analyses were made with all characters set as nonadditive. These analyses produced identical trees (but only one of the two trees for concavity K = 6; see also note under character 11).

Weight for each character is a function of the homoplasy it implies on the tree. However, there is an amount of a priori homoplasy, as determined by intraspecific variability (e.g., Negayan tridentata may have two or three teeth on the cheliceral retromargen, and the scoring [12] adds one internal step to character 20). Those cells of the matrix were coded as polymorphisms (letters a–j in the datamatrix), and the appropriate internal steps were added manually with command ccode= of Pee-Wee. Polymorphic entries were also used to express ambiguous homology, because of intermediate conditions (e.g., the intermediate shapes of the paramedian apophysis of some Gaynennini, character 68). These entries do not count for internal steps. The complete data matrix is listed in table 1.

Character Description and Optimizations

Except as noted, states are (0) absent, (1) present, and “synapomorphy” represent a gain (transformation 0 → 1). To make reading easier, after definition of each character and states, a synopsis of its evolution is given, as optimized on the preferred trees of figure 3.

Color and Body Pattern

  1. Body pattern: (0) absent, color uniform; (1) present, with contrasting patches, spots or dots. Color uniform is a synapomorphy (reversion) of Sanogasta puma and S. tenuis, both being pale brown. Coptoprepes species have a weakly contrasting pattern; only C. flavopilosus is uniformly dark brown (fig. 35A). Malenella nana is uniformly pale green (see character 3). In previous contributions (Ramírez, 1995b, 1999), a pattern of dark small dots on a pale background appeared as a synapomorphy of Monapia, Tasata, and Oxysoma. With a wider selection of representatives, there are many intermediate or ambiguous conditions, and the character is not used here (but see characters 4 and 5).

  2. Ventral longitudinal dark stripe on abdomen. It may be dark and homogeneous, or formed by aligned spots. It appears independently many times.

  3. Red lateral bands on carapace. A synapomorphy of Araiya.

  4. Color green. A potential synapomorphy of Wulfila and Malenella nana; appears as a plesiomorphy in this analysis.

  5. Anterior dorsal dark dot on abdomen (fig. 125). Potential synapomorphy of some Tasata, Oxysoma, and Monapia species, arises independently in all three genera.

  6. Two pairs of dark dots on the anterior half of abdomen. Similar distribution as character 3, ariable in Oxysoma punctatum and Tasata unipunctata.

Carapace

  1. Carapace very narrow: (0) normal, female carapace width/length > 0.65; (1) narrow, width/length < 0.55 (figs. 22A, 88A). Estimated from the male for a few species with unknown females. Because measurements were taken from one specimen only, statistical analysis of continuous characters derived from measurements is impossible. Only a clear gap across all terminals was considered as the limit between states (fig. 1A), and any intermediate group is coded as ambiguous. A potential synapomorphy of Aysenia, but ambiguous at node 117 because Axyracrus and Aysenoides parvus are intermediate and were coded [01].

  2. Carapace and chelicerae Amaurobioides-like. In this genus and Aysenia the carapace is wide in front, the chelicerae are very strong, and the ocular area is relatively small. It appears as a convergence in the optimal trees; however, in Axyracrus and Aysenoides the carapace is intermediate between the generalized shape and that of Amaurobioides.

  3. Thoracic groove. Absent in Malenella, shallow in Wulfila argentina (coded [01]), absent in clade 126.

Eyes

  1. Ocular area black. Appears independently in several groups.

  2. Ocular area protruded (fig. 22A). Synapomorphy of clade 118, but also of clades 141, 152, and of Tasata taim.

  3. Anterior eye row: (0) procurved; (1) straight; (2) recurved. States are ordered. Very homoplasious. Characters 11–16 are perhaps the most traditional characters of spider systematics. They are continuously variable, with boundaries between states arbitrarily defined. A small variation in the position of AME can change the row from recurved to straight or procurved, while a wide change to the other side will still be considered recurved. Eye rows are seen from standard views (anterior, dorsal), which may not be comparable when the shape of the cephalic area varies. The ambiguity of definition is reflected in their high levels of homoplasy. Eliminating these characters only affects local resolution in clades 95 and 128, involving groups with tiny support.

  4. Posterior eye row: (0) procurved or straight; (1) recurved. Synapomorphy of clade 119, with reversals in clade 100 and Amaurobioides africana; ambiguous at nodes 95 and 96 of Josa.

  5. Posterior eye row strongly procurved (fig. 129A, B). Traditionally considered synapomorphy of Arachosia (what Simon referred as Oxysoma), appears also in some Tasata, in Gayenna americana, and in Oxysoma itambezinho.

  6. Separation between lateral eyes: (0) up to one diameter; (1) more than one diameter. Appears independently in Arachosia bergi, Tomopisthes horrendus, and clade 93 of Josa.

  7. Ratio AME/ALE: (0) AME minute; (1) AME < ALE; (2) AME = ALE; (3) AME > ALE. States are ordered. Extremely homoplasious.

  8. Ratio PME/PLE: (0) PME < PLE; (1) PME = PLE; (2) PME > PLE. States are ordered. Very homoplasious. State 2 is an autapomorphy of Josa calilegua.

Chelicerae and Endites

  1. Male chelicerae: (0) strong, as in female or larger; (1) smaller than in female. Appears and reverts in several groups.

  2. Male retromarginal distal tooth: (0) similar to the basal; (1) much larger than the basal (Ramírez, 1997: figs. 50, 56). Synapomorphy of clade 99.

  3. Male distal pro- and retromarginal teeth: (0) separate; (1) contiguous, on a common protuberance (Ramírez, 1997: figs. 50, 56). Synapomorphy of clade 99, absent in Acanthoceto riogrande (contra Ramírez, 1997).

  4. Number of retromarginal teeth: (0) one; (1) two (fig. 12A); (2) three; (3) four or more (fig. 11B, C). States are unordered, because the homology among individual teeth is unclear (apical, basal, or intermediate teeth might be added or lost). Very homoplasious. Having two teeth is a synapomorphy of Gayennini and Josa, with subsequent reversals in several groups, also variable within Amaurobioidini. See character 23.

  5. Size of retromarginal teeth: (0) small denticles; (1) regular teeth. Quite homoplasious, ambiguous in Malenella and some Anyphaeninae, because they have both regular and small teeth. State 1 arises independently in clades 118 and 176, and there are several other convergences and reversals.

  6. Male median promarginal tooth: (0) unmodified, slightly larger than the laterals; (1) thick, elevated (fig. 96B). Synapomorphy of clade 132 of Philisca.

  7. Number of promarginal teeth: (0) three; (1) four; (2) five or more. States are unordered for the same reason as character 20. The general number is three. Two gains of promarginal teeth are associated with a gain of one or more retromarginals, two other gains are not; all other changes are ambiguously optimized.

  8. Male chelicerae modified (fig. 97D). Similar to those of Dictyninae males, it is a synapomorphy of a group of Philisca species, here represented by clade 132.

  9. Male endites modified (fig. 96B). In this analysis the protuberances at the external sides of the endites are an autapomorphy of Philisca ornata, also present in other related species.

Female Legs and Palp

  1. Leg III orientation: (0) backward; (1) forward (fig. 24A). Synapomorphy of Aysenia and Aysenoides. The habits of these spiders are almost unknown; the body shape and position of third legs suggest that they might live in narrow tubes.

  2. Leg III much shorter that IV: (0) unmodified, female tibia + metatarsus III equal or longer to tibia IV; (1) shorter than 80% of tibia IV (fig. 1B). Estimated from the male for the few species with unknown females. Synapomorphy of clade 151, convergent in Sanogasta tenuis. All three species live on grasses and have similar cryptic habits.

  3. Tibia I sinuous: (0) straight (fig. 112C); (1) slightly sinuous (fig. 112A, B). Synapomorphy of Araiya (with a convergence in Aysenia elongata, fig. 22B). In other genera only males have a sinuous tibia I.

  4. Patch of blunt hairs on palp. Autapomorphy of Malenella nana (fig. 10E, F).

  5. Thick female palp. Autapomorphy of Malenella nana (Ramírez, 1995a; this paper, compare figs. 10F and 12C).

  6. Palpal claw blunt, compressed. Synapomorphy of clade 134 (Ramírez, 1993: fig. 4; this paper, fig. 101).

Claw tufts and Scopulae

  1. Orientation of claw tuft setae: (0) flat sides up and down, the generalized condition in Dionycha (Platnick and Lau, 1975; this paper, fig. 10C, D); (1) ridged sides directed outward (fig. 13B, C, G). A synapomorphy of Anyphaeninae plus Amaurobioidinae (Ramírez, 1995a). Each claw tuft seta has a vertical widened surface, with the petiolus extending in an external rib (fig. 13G). Some dionychans like Lygromma (Prodidomidae) or Scotinella (Liocranidae) have superficially similar setae (Platnick and Lau, 1975).

  2. Scopulae on anterior tibiae: (0) absent; (1) present. Independently lost in several groups.

  3. Scopulae on posterior tibiae. Synapomorphy of Arachosia, with convergences in clade 93 of Josa, and in Tomopisthes horrendus.

Abdomen

  1. PMS with many aciniform gland spigots (fig. 15C). Synapomorphy of Amaurobioides, probably used to build waterproof retreats.

  2. Male abdomen projecting over anal tubercle (Ramírez, 1997: fig. 16). Synapomorphy of Acanthoceto.

  3. Thick setae on ALS base (fig. 61D–F) of males, females and immatures. Synapomorphy of Arachosia. The males of P. huapi, and at least the adult males and females of other undescribed Philisca species also have similar setae on ALS.

Tracheae

  1. Tracheal spiracle position: (0) closer to spinnerets, or midway to epigastric furrow; (1) closer to epigastric furrow. A potential synapomorphy of Anyphaeninae (Ramírez, 1995a, but here nonmonophyletic), with convergences in some Arachosia and Acanthoceto acupicta. Because the next two characters are most probably a consequence of the advancement of the tracheal spiracle, the weight of these three characters is 1, with all others being 3; equal weights produce the same results, including the nonmonophyly of Anyphaeninae.

  2. Length of lateral tracheae: (0) short; (1) long. Same as preceding.

  3. Position of first bifurcation of median tracheae: (0) separate from lateral tracheae; (1) contiguous. Potential synapomorphy of Anyphaeninae, see character 38.

Male Palp

  1. Femoral apophysis (fig. 60E). Synapomorphy of Josa.

  2. Retrolateral tibial apophysis (RTA): (0) absent; (1) present. The RTA is primitive for Anyphaenidae, but is independently lost in Wulfila argentina (where a basal apophysis is present instead, see character 46), Josa + Gayennini, and Coptoprepes campanensis; Gamakia hirsuta has a tiny relict of RTA (fig. 46).

  3. Shape of RTA: (0) thick or spatulate; (1) thin, narrow, spine-shaped (fig. 18C); (2) Negayan type, elongate and distally hooked (fig. 50C). State 2 is a synapomorphy of Negayan; node 119 and descendants optimize state 1, node 108 optimizes state 0, all other Amaurobioidinae [01]. The relict of RTA of Gamakia hirsuta (fig. 46) is coded uncertain; if coded 1, only the assignments of clades 106, 120, and 121 change from [01] to 1.

  4. RTA extremely thin (Ramírez, 1997: fig. 9). Synapomorphy of clade 101 of Acanthoceto.

  5. Retrolateral or dorsal apophysis, additional to the RTA. Only in Malenella nana and Aysha prospera. Otoniela is coded ambiguous because it has a low ridge. Xiruana hirsuta has only one RTA, but X. gracilipes has two; if Xiruana is coded as present, it becomes sister to Aysha, a more reasonable placement according to Brescovit (1997). The optimizations for the basal node of Amaurobioidinae do not change much with that alternative resolution (but see character 79).

  6. Basal retrolateral tibial apophysis. A potential synapomorphy of some Anyphaeninae, unclear in Xiruana and Otoniela.

  7. Cymbial conductor (figs. 34A, 61). Synapomorphy of Anyphaeninae plus Amaurobioidinae (Coddington, 1990; Ramírez, 1995a).

  8. Cymbial conductor subapical: (0) apical (figs. 34A, 61); (1) subapical (fig. 53B). Synapomorphy of clade 106, with convergence in Aysenoides parvus and Coptoprepes valdiviensis.

  9. Cymbial conductor width: (0) narrow (fig. 61); (1) wide (fig. 34A). The narrow condition is a synapomorphy of Gayennini, convergent in some Anyphaenines. There is a reversal to wide in clade 158 of Monapia (Ramírez, 1995b: figs. 41, 55, 62, 68, 74).

  10. Retrolateral apical notch on cymbium. The notch is usually fitted to the median apophysis (fig. 42A, B), even also to the secondary conductor (fig. 38A). Convergent in Coptoprepes, Negayan, and Amaurobioides maritima. Sanogasta approximata and Monapia dilaticollis have similar notches, but they are not clearly associated with the median apophysis.

  11. Apical patch of thick, bent setae on cymbium (fig. 119B, C; Gerschman and Schiapelli, 1970: fig. 23). In this analysis an autapomorphy of Oxysoma longiventre, present also in some closely related, undescribed species.

  12. Retrolateral basal notch on cymbium (figs. 66D, 118E). Appears convergently in Gayennoides, clade 149 of Monapia, and Oxysoma itambezinho.

Copulatory Bulb

  1. Apical loop of the sperm duct (SD), dorsal to secondary conductor (figs. 21H, 26C). Synapomorphy of Amaurobioidini, with a reversal in Acanthoceto acupicta group (clade 100), and a convergence in Monapia angusta. Sanogasta approximata and several Anyphaeninae have a weak curve, coded as uncertain.

  2. Loop of the SD dorsal to median apophysis. Appears convergently in clade 114 of Aysenoides (fig. 26C) (they have both loops, see preceding character) and in clade 134 of Philisca (fig. 102A).

  3. Apical dorsal margin of tegulum extended over the secondary conductor (figs. 82A, 84D). Synapomorphy of clade 174, reverts in clade 167 of Sanogasta, probably masked by the hypertrophy of secondary conductor (fig. 78B). Convergent in Phidyle punctipes. The bulbs of Amaurobioidini, Josa, and several Anyphaeninae are very different; hence, the character is scored as missing.

  4. Ventral loop of the sperm duct reaching the apical margin of tegulum. Quite notorious in Tasata and close relatives (figs. 128A, 137C), but ambiguous in many other species. Synapomorphy of clade 165, with a few reversals, and probable convergences in Anyphaeninae (here in Xiruana hirsuta).

  5. Sperm duct suddenly narrowed before reaching the embolus (figs. 18G, 50D). Convergent in Amaurobioides and Negayan paduana.

  6. Spiral loop of the sperm duct, before reaching the embolus: (0) absent (fig. 121); (1) present, slightly coiled (fig. 129E); (2) well coiled in spiral (fig. 128D). States are ordered. Synapomorphy of clades 147 (state 1) and 145 (state 2) of Tasata.

  7. Tegulum displaced basally (figs. 25B, 36F). Caused by the hypertrophy of the distal sclerites, it is a synapomorphy of Coptoprepes, also of clade 112 of Aysenia (but the male of Aysenia elongata is still unknown).

  8. Basal notch on tegulum. Synapomorphy of Amaurobioidinae (Platnick, 1977; Ramírez, 1995a).

  9. Basal notch of tegulum displaced prolaterally (fig. 58A). Convergence of Josa and clade 156 of Monapia.

  10. Ventral cusp on tegulum (fig. 47A). Synapomorphy of clade 104 of Negayan.

  11. Triangular sclerotized area from the sperm duct to the base of median apophysis (fig. 107E). Extremely homoplasious, present in Tomopisthes, but variable in Philisca. In several Monapia all the area is sclerotized (coded uncertain), variable in Monapia alupuran.

  12. Median apophysis (MA): (0) present; (1) reduced; (2) absent. States are ordered. Reduced independently in Ferrieria, Acanthoceto acupicta group, Negayan, Axyracrus, and Sanogasta maculatipes group. Totally absent in Negayan coccinea, Malenella, and the anyphaenine Italaman santamaria.

  13. Shape of MA: (0) relatively thick (fig. 18H); (1) slender and elongate (fig. 66E). The slender MA is a synapomorphy of Gayennini, with convergence in several Amaurobioidini. Only Gamakia and some Aysenoides have similar S-shaped MA.

  14. MA with thin branches (fig. 93A–C). Synapomorphy of Philisca, lost in P. tripunctata; convergent in Tasata centralis, with very short splinters (fig. 133).

  15. Paramedian apophysis (PMA; see Morphological Remarks): (0) absent; (1) present. Absent in Malenella and Anyphaenines, but coded uncertain in Otoniela adisi. This species has a palpal sclerite (the “ventral tegular projection”, Brescovit, 1997), which might be homologous to either PMA or the primary conductor (C1). In Josa the PMA is fused to the tegulum (character 74); Josa nigrifrons and Coptoprepes campanensis have no traces of PMA.

  16. Shape of PMA: (0) one short cusp (figs. 42C, 37A); (1) two or more short cusps (figs. 26C, D, 52A); (2) thick, simple, and elongate, type Philisca (fig. 98A); (3) slender, type Monapia or Sanogasta (fig. 82A); (4) bifid (fig. 107D, E). States are unordered; coding details within Gayennini sacrificed the presumed homology of their conspicuous, projecting PMA. State 1 is a synapomorphy of clade 121 with subsequent reversal in some Acanthoceto. State 2 appears in Philisca except P. puconensis and some Tasata, and state 3 appears in Sanogasta (including Arachosia) and Monapia. State 4 seems to be the primitive condition for Gayennini. Several Gayennini have intermediate shapes and were coded as polymorphic.

  17. One cusp of the PMA on the primary conductor (fig. 26). The cusp is clearly visible in some amaurobioidines, but is dubious in others; scored missing in Gayennini, which has reduced C1.

  18. PMA slender, associated with MA (figs. 82A, 84D). Synapomorphy of clade 172.

  19. Membranous area in the base of PMA, as seen in the unexpanded bulb: (0) absent; (1) present (fig. 85A); (2) completely surrounding the base of PMA (fig. 81B, D, F). This is a part of the distal hematodocha, which is visible in the unexpanded bulb; the main inflatable portion is mostly folded between PMA and tegulum. States are ordered. Synapomorphy of clades 172 (state 1) and 167 (state 2).

  20. PMA Oxysoma type. Forming a hollow under the tegulum, with a cusp close to the base, and an elongate, recurved tip (fig. 116B). Synapomorphy of clade 141.

  21. Globose lobe on C1, at the origin of PMA (fig. 47C). Convergence of Negayan coccinea with clade 108 of Coptoprepes.

  22. PMA fused to tegulum (fig. 56A, B). Present in Josa, but the PMA is missing in J. nigrifrons.

  23. Primary conductor (C1; see Morphological Remarks): (0) absent; (1) present, without canal (fig. 93B); (2) with a canal where the embolus fits (fig. 44B); (3) massive, with canal (fig. 47D). States are ordered. Potential homologs of C1 occur in many dionychans, Malenella (a hyaline conductor, but not a separate sclerite), Wulfila (the “ventral tegular projection”, Brescovit, 1997), and Otoniela (see character 67); Anyphaena accentuata (fig. 11A) has a potential homolog of the C1 quite similar to the massive conductor found in some Amaurobioidinae (state 3; “median apophysis” of Huber, 1995; “ventral tegular projection” of Brescovit, 1997), including a shallow canal. In this analysis with restricted outgroups, the C1 is ambiguous [01] through outgroup internal nodes and is present without canal at base of Amaurobioidinae, Josa, and Gayennini. The canal (state 2) arises at the base of Amaurobioidini; the C1 becomes massive (state 3) at clades 100 and 106 (and in Anyphaena accentuata). Josa has a C1 fused to the tegulum (fig. 60B), but otherwise is similar in shape and position to that of Amaurobioidini. In Gayennini the C1 is small, not associated with the embolus, and is even lost independently in several groups; it optimizes as a regain in Arachosia bergi, but the putative C1 is quite different, being fused to the PMA.

  24. Translucent vertical lamina on C1 (figs. 27C, 28C). Synapomorphy of clade 114.

  25. Prolateral process on C1, crossing the canal (figs. 47C, 52B). A potential synapomorphy of Selknamia minima and some Negayan.

  26. Apex of C1 displaced close to the median apophysis (Ramírez, 1997: fig. 44; this paper, figs. 32B, 33D). Convergent to state 1 in clade 100, Selknamia and in Aysenoides parvus. All these species have small bulbs, with crowded apical sclerites. The morphology in Axyracrus elegans is quite different, and is coded as missing.

  27. Secondary conductor (C2; see Morphological Remarks). (0) absent; (1) fused to anterior dorsal margin of tegulum; (2) free. States are unordered. Among anyphaenid outgroups, potential homologs of the C2 occur in Anyphaeninae. As discussed by Brescovit (1997), the “anyphaenine conductor” is a good candidate, here present in Xiruana. In Anyphaena accentuata, the prolateral tegular projection (Brescovit, 1997: fig. 4) is also a putative homolog of a free C2, according to its apical position, arising close to the anterior dorsal margin of tegulum (fig. 11A). The “prolateral tegular projection” of Wulfila argentina (Brescovit, 1997: fig. 19) arises between the base of the embolus and the (presumably) primary conductor, and may be a tegular or embolar apophysis instead of a C2. With this outgroup selection, a fused C2 arises in clade 178. If Anyphaeninae is constrained to be monophyletic, the assignment of the basal node of Amaurobioidinae is [012] instead of 1. According to Brescovit, Xiruana, Aysha, and several other genera without putative homologs of secondary conductor form a monophyletic group with Xiruana, and hence their conductors should have appeared independently. If Xiruana is forced to be sister to Aysha (see character 45), the optimization of C2 becomes ambiguous and thus compatible with Brescovit's interpretation. The optimization on most parsimonious trees implies multiple separations and losses of C2. Among the Amaurobioidini, the C2 is free only in some Coptoprepes (clade 108) and is independently lost several times; it becomes free also in node 176, fused at node 172, and separate again at node 169! The C2 of Josa is different and could not be scored for several of the following details.

  28. Wide membranous area separating C2 from tegulum: (0) membranous area narrow retrolaterally, just a suture, or C2 fused to tegulum (fig. 78B); (1) wide (figs. 105A, 93D). A potential synapomorphy of Tomopisthes and Philisca, also present in Gayenna americana, Oxysoma punctatum, and Tasata centralis.

  29. Prolateral process on C2: (0) absent; (1) elongate, rounded lobe, Arachosia type (fig. 70); (2) flattened lobe, directed basally, Tasata type (figs. 116C, 126D). States are unordered. State 1 is a synapomorphy of clade 124 of Arachosia. State 2 is a synapomorphy of clade 163. In most Monapia and Philisca the morphology is confusing, but definitely not of the Arachosia type (scored [02]).

  30. Dentate prolateral ridge or lobe on C2 (figs. 102B, 126D). Appears convergently in Tasata and clade 136 of Philisca.

  31. Apex of C2 (the piece where the canal ends): (0) apical (fig. 82A); (1) median or basal, the C2 extended as anterior border (fig. 84E). Synapomorphy of clade 165, with several reversals; convergent in Sanogasta approximata and S. x-signata; confusing in several Gayennini with modified C2.

  32. Canal on C2: (0) absent; (1) present, short; (2) deep, long, arising under the PMA, Gayenna type (fig. 82B). The canal appears at node 178 (with same provisions as character 79) and is lost several times in both tribes. State 2 is a synapomorphy of clade 174, being subsequently lost in Sanogasta pehuenche and S. approximata.

  33. C2 divided by a membranous area: (0) undivided; (1) totally divided by a membranous area, retrolateral to the canal (Ramírez, 1995b: fig. 40; this paper, figs. 133A, 135D). Convergent in clades 161 and 145.

  34. C2 with membranous area prolateral to the canal. Similar to preceding, but at the other side of the canal (figs. 118H, 124E). A potential synapomorphy of Oxysoma saccatum and O. itambezinho.

  35. Membranous lobe on C2. Synapomorphy of Monapia, it is an outgrowth of the unsclerotized area dividing the C2 (Ramírez, 1995b: fig. 40); lost in M. angusta.

  36. Denticles on prolateral portion of C2. Extremely homoplasious in clade 165, with many independent gains and losses. Weakly sclerotized areas of the bulb seem especially susceptible to develop such sculptures.

  37. Teeth on C2 apex, regularly disposed, pointing backward (fig. 111A, C). A potential synapomorphy of Araiya and Tomopisthes (with somewhat similar disposition in Philisca amoena, coded ambiguous).

  38. Denticles on retrolateral portion of C2 (fig. 116E). Convergent in Oxysoma and clade 157 of Monapia, lost in M. pichinahuel.

  39. Base of C2r: (0) thick; (1) wide, thin, translucent (Ramírez, 1995b: figs. 25, 63). Synapomorphy of clade 159 of Monapia, lost in M. carolina and clade 155; also present in some Philisca, with ambiguous optimizations.

  40. C2 Josa type (figs. 56B, 60B). Hypertrophied and complex, a clear synapomorphy for the genus.

  41. Shape of the relictual C1 in Gayennini: (0) conical (figs. 63C, 105C); (1) acute (Ramírez, 1999: figs. 27, 39); (2) thin, rounded (fig. 126B). States are unordered, only scored in Gayennini. Very homoplasious, it optimizes state 2 in most internal nodes.

  42. Articulation of embolus: (0) fixed, fused to tegulum; (1) movable, membranous in part. In this analysis the embolus became movable at node 179; I think that the embolus of Aysha is articulate (contra Brescovit, 1997).

  43. Embolus very long. Arises independently in Wulfila argentina, Josa, Negayan, Coptoprepes campanensis, and clades 111 or 112 of Aysenia and 96 of Monapia.

  44. Basal process on embolus. For the anyphaenines, I scored this character present for both the “apophysis of embolic process” and the “basal process of embolus” as distinguished by Brescovit (1997). It optimizes as present at clade 180, and is lost in Xiruana and Selknamia. In the Gayennini the shape of the process is homogeneous, but it becomes very small and of dubious homology in Monapia, where it optimizes as three independent regains.

  45. Shape of basal process of embolus: (0) flattened (fig. 44B); (1) thin, hyaline (fig. 33B, C); (2) membranous, expansible (Acanthoceto acupicta group, fig. 33E); (3) complex (fig. 54B); (4) spinelike (Aysenoides, fig. 26B); (5) thick, conical (Negayan, fig. 50D); (6) Gayennini type, an extension of a narrow sclerotized stripe, partly bordered by a membranous area which may confer variable shape (figs. 93F, 116C). States are unordered. Because of the outgroup selection, the primitive state is determined by that of Aysha and Josa (complex, state 3). In Amaurobioidini the ancestral state is 0. All Gayennini have state 6 or absent. Axyracrus was coded [36] because the complex process arises from a membranous area similar to that of Gayennini. State 1 is a synapomorphy of clade 103 (parallel in Amaurobioides africana), changing to state 2 in the Acanthoceto acupicta group.

  46. Base of embolus flattened: (0) unmodified, approximately cylindrical; (1) flattened (Ramírez, 1995b: fig. 62). Synapomorphy of clade 156 of Monapia, convergent (not very similar, though) in M. angusta, Josa, and Xiruana.

  47. Base of embolus with anterior longitudinal projecting ridge (fig. 99F). Totally homoplasious, appears independently in some Philisca and Phidyle.

Epigyne

  1. Epigastrium partially sclerotized (fig. 59C). Arises independently four times in species of Josa, Sanogasta, Tasata, and Wulfila, which have dark sclerotized cuticle around the epigynum, instead of soft, clear cuticle.

  2. Insertions of epigastric muscles at sides of epigyne: (0) superficial; (1) depressed (fig. 69B). Synapomorphy of clade 171, reverts in Sanogasta tenuis.

  3. Epigyne projecting posteriorly. Convergent in Josa nigrifrons and Sanogasta x-signata, also occurs in species of both genera not included here.

  4. Semicircular ridges anterior to epigyne. All convergences in species of Tomopisthes, Tasata, Oxysoma, and Araiya.

  5. Anterior pouch on median field (APmf): (0) absent; (1) opening forward (fig. 94E); (2) opening backward (fig. 80). States are unordered. Anyphaeninae has a wide diversity of pouches; here, Anyphaena accentuata has state 2. State 1 is a synapomorphy of Gayennini, state 2 of clade 168. Coded uncertain for a few Amaurobioidini with irregular folding on median field.

  6. Position of APmf: (0) advanced; (1) close to epigastric furrow. Quite homoplasious, some interferences with next character. For Sanogasta, the optimal reconstruction implies that the pouch moved close to epigastrium in node 170, then advanced again at node 167 (see preceding character).

  7. Shape of APmf: (0) opening approximately circular; (1) opening transverse (fig. 69); (2) pouch widely distended (fig. 107F, G). States are unordered. State 1 is convergent in Monapia and Arachosia, state 2 in Tomopisthes, Tasata, Philisca amoena, and Araiya coccinea.

  8. Lumen of APmf: (0) simple; (1) double. A double cavity appears independently in Arachosia and clade 142 and is ambiguous through several nodes of Monapia.

  9. Median depression on epigyne: (0) absent; (1) present; (2) vestigial. States are unordered. States 1 and 2 are synapomorphies of groups in Monapia (see discussion in Ramírez, 1995b); among the outgroups, state 1 is present in Malenella and Anyphaena accentuata.

  10. Pouch on median depression (not the APmf). Synapomorphy of clade 153, reverts in Monapia angusta (Ramírez, 1999).

  11. Lateral lobes (LL): (0) separate; (1) contiguous; (2) fused with suture; (3) fused without suture. States are ordered. Synapomorphy of some clades in Philisca and Monapia (Ramírez, 1993, 1995b). State 2 appears convergently in Oxysoma punctatum.

  12. Posterior notch between LL. The LL are close together over the epigastric fold, limiting notch. Appears independently in Josa and clade 128 of Sanogasta (figs. 56H, 84A).

  13. Posterior depressions on LL. Appear independently in some Josa (fig. 55A), Ferrieria (Ramírez, 1997: fig. 67), and clade 104 of Negayan (shallow in N. tridentata).

  14. LL projecting posteriorly. Arises independently in Josa, clade 104, and Sanogasta x-signata.

Spermathecae and Ducts

  1. Fusion of proximal copulatory ducts (CD): (0) separate; (1) fused walls; (2) totally fused, with common lumen. States are ordered. States 1 and 2 are synapomorphy of clades 155 and 156 of Monapia, respectively (Ramírez, 1995b); state 1 is also a synapomorphy of clade 129 of Sanogasta.

  2. Copulatory openings on epigastric furrow. Convergent in several groups.

  3. CD slender. Often associated with the spherical spermathecae (character 124), appears convergently in Gayennini, Selknamia, and Aysenoides. Through outgroups, appears at clade 181 to revert in 178.

  4. CD coiled along longitudinal axes. Present in Josa, Ferrieria, Acanthoceto pichi (ambiguous at nodes 102 and 103), Negayan, and clade 111 of Aysenia. Because most entelegyne families have representatives with similarly coiled ducts, the convergences here are not surprising. However, those of Negayan, Ferrieria, Acanthoceto pichi, and Aysenia araucana are remarkably similar, yet mostly homoplasious.

  5. CD extremely coiled. Convergence in Aysenia segestrioides and clade 95 of Josa (figs. 23E and 57F, respectively).

  6. Lumen of proximal CD: (0) thin; (1) ample. Synapomorphy of clade 156 of Monapia, convergent in Xiruana. In this anyphaenine genus, the epigyne is extremely modified, folded on itself, with the copulatory openings at the end of an invagination, resembling those of some Monapia. In Xiruana, the median, sclerotized, elevated plate is in fact the lateral lobes fused to each other (personal obs.).

  7. Walls of proximal CD thin, flexible. Synapomorphy of clade 155 of Monapia, convergent in Xiruana.

  8. CD trajectory Oxysoma type (figs. 120E, 123B). Synapomorphy of Oxysoma (see diagnosis).

  9. Accessory bulbs (AB; see Morphological Remarks). Present in most Anyphaenidae (and entelegynes), absent in Malenella, Wulfila argentina, and Otoniela.

  10. Duct of the AB: (0) short (fig. 80E); (1) long (fig. 73B). Highly homoplasious, perhaps because of the somewhat dubious limits between states. In Amaurobioidini the duct is short.

  11. Spermathecae shape: (0) irregular; (1) approximately spherical. Synapomorphy of Gayennini and also of Aysenoides, convergent in Selknamia and the anyphaenine Otoniela.

  12. Spermathecae contiguous: (0) separate; (1) contiguous. Synapomorphy of Negayan and potential synapomorphy of Coptoprepes (ambiguous because of anyphaenine outgroups), also convergent in Aysenia elongata.

  13. Fertilization duct (FD) coiled along with the CD (fig. 57F). Potential synapomorphy of some Josa.

  14. FD distant from epigastric furrow (fig. 23E). Very homoplasious.

Sexual Behavior

  1. Copulatory plug. Among close outgroups, only seen in Malenella. Absent in Anyphaena accentuata (Huber, 1995), to my knowledge not reported for any other anyphaenine. Synapomorphy of Monapia (with reversal at clade 155), also convergent in Axyracrus elegans and Gamakia hirsuta.

Spines

  1. Spines on chelicerae. A thick spine on anterior face of paturon (Ramírez, 1999: fig. 14) is a synapomorphy of clade 151, convergent in Oxysoma itambezinho.

  2. Female palpal femur with a line of ventral spines. Synapomorphy of clade 152, also present in Aysha prospera (weak spines) and Wulfila argentina.

  3. A series of prolateral-ventral spines on femur I. Synapomorphy of clade 151.

Tibia I

  1. Supplementary ventral spines on tibia I: (0) 2–2–2 or less; (1) 2-2-2-2 or more. Synapomorphy of clade 153, also present in Otoniela. Ferrieria has v 2–2–2 (contra Ramírez, 1999), plus p and r 0-v1, resembling v 2-2-2-2.

  2. v p1-x-x.

  3. v r1-x-x.

  4. v x-p1-x.

  5. v x-p1-x displaced prolaterally (fig. 24E). Synapomorphy of Aysenia plus Aysenoides.

  6. v x-r1-x.

  7. v xap: (0) 2ap; (1) p1ap; (2) 0ap. States are ordered because individual spines are homologous; merged into one character because the r1ap is never present alone.

Metatarsus I

  1. v 2bas.

  2. v x-p1-x.

  3. v x-r1-x.

  4. p 1-x.

  5. r 1-x.

  6. d p1-x.

  7. d 2ap.

Tibia II

  1. v p1-x-x.

  2. v r1-x-x.

  3. v x-p1-x. Coded ambiguous in Monapia guenoana, because they have v r1-r1-r1-2-0, and the homology of individual spines is unclear.

  4. v x-r1-x.

  5. v p1ap.

  6. v r1ap.

  7. p x-1.

Metatarsus II

  1. p d1-x-x.

  2. p x-1-x.

  3. d p1-x.

  4. d p1ap.

  5. d r1ap.

Patella III

  1. r d1.

Tibia III

  1. Prolateral spines on tibiae III and IV displaced ventrally. Synapomorphy of clade 126.

  2. v p1-x-x.

  3. v r1-x-x.

  4. v x-p1-x.

  5. v x-r1-x.

  6. v x-x-p1.

  7. v x-x-r1.

Metatarsus III

  1. v 2-x-x.

  2. v x-p1-x.

  3. v x-r1-x.

  4. v ap: (0) 2; (1) p1; (2) 1; (3) 0. States are unordered, because the unpaired median spine (state 2) is not homologous with either of the laterals. Legs III and IV vary coordinately, and thus only one character is scored. One median spine is present in early immatures of at least Tomopisthes horrendus.

  5. Preening comb on metatarsi III and IV. In some cases poorly defined, only a group of thick setae (fig. 40). Some genera (e.g., Oxysoma) have a bunch of hairs in that position.

  6. p d1-x-x.

  7. p x-1-x.

  8. p x-x-1.

  9. r d1-x-x.

  10. r x-1-x.

  11. r x-x-1.

  12. d x-p1-x.

  13. d x-x-p1.

  14. d x-x-r1.

Patella IV

  1. r d1.

Tibia IV

  1. v p1-x-x.

  2. v r1-x-x.

  3. v x-p1-x.

  4. v x-r1-x.

  5. v x-x-p1.

  6. v x-x-r1.

Metatarsus IV

  1. v p1-x-x.

  2. v r1-x-x.

  3. v x-p1-x.

  4. v x-r1-x.

  5. p d1-x-x.

  6. p x-1-x.

  7. p x-x-1.

  8. r d1-x-x.

  9. r x-1-x.

  10. r x-x-1.

  11. d x-p1-x.

  12. d x-x-p1.

  13. d x-x-r1.

Characters Not Included

Several characters considered in preliminary data matrices were excluded from the analysis, and are discussed below. Some may be informative for less inclusive analyses.

Posterior slope of carapace, steep or attenuate (Ramírez, 1997: character 21). Posterior slope begining close behind the fovea (Ramírez, 1999: character 2). Posterior notch in carapace (Ramírez, 1993: character 2). Many intermediate conditions exist in this dataset.

Carapace globose in males. Seemingly correlated with the small chelicerae, already scored in character 17.

Sternum very narrow. Abdomen elongate. At least in some groups (e.g., Aysenia and close relatives), both characters are seemingly correlated with the elongate body in general (scored in character 6).

Proportions of male palpal tibia (length/width) are very variable and are somewhat phylogenetically conservative. Measures of these dimensions are coarsely approximate (e.g., the anterior limit of the tibia is poorly defined in the presence of an apophysis) and of dubious homology, since the general shape of the tibia is variable as well.

Wide diastema between copulatory bulb and tarsus (fig. 98A). Many intermediate conditions.

Retrolateral basal lamina on cymbium (figs. 34B, 42B). Many intermediate conditions.

Basal tegular notch short or deep (fig. 91A vs. 83B). Many intermediate conditions and interactions with other characters (e.g., character 59).

Median epigynal field elevated (Ramírez, 1997: character 42). Many intermediate conditions exist in this dataset.

Ducts of the accessory bulbs, converging or diverging. Many intermediate conditions exist, and there are ambiguities if the ducts are short (character 123).

Preference for habitats close to water (Ramírez, 1997: character 43). It is not convincing that the several conditions are homologs (e.g., at stony seashores or grasses on wetlands).

Preference for living on grasses, under stones, etc. Several details of habitat seemed dependent on biogeography (e.g., in Chilean temperate forests there are no grasslands, and there are no superficial rocks through most of the Pampas).

Evaluation of Cladistic Hypothesis

This dataset was analyzed with parsimony under equal weights with the program NONA, and under implied weights with Pee-Wee (Goloboff, 1994). A congruence-based test was made to select between these methods of analysis and among different strengths of the weighting function.

Tree Searches

For a dataset of this size, only heuristic solutions are available. However, because the same optimal trees were hit many times using different search strategies and under different conditions, they are almost certainly the exhaustive optimal sets. For instance, the parsimony ratchet (Nixon, 1999) found the two optimal trees in Pee-Wee (concavity K = 6) in 100 of 100 runs of tree bisection-reconnection (TBR) multiratchet (nixwts*12 100).

Character Weighting

A proposed refinement of parsimony is character weighting according to homoplasy. The best known and more widely used ways to perform this are successive weighting (Farris, 1969) and implied weights (Goloboff, 1993, 1995a). The aim of these procedures is to reach a classification that better explains those characters with a better fit to the cladistic hypothesis, at the expense of the more homoplasious ones. In successive weighting, a first run is made with all weights equal, and the weights are calculated from the set of most parsimonious trees, using a homoplasy index (e.g., consistency index, or rescaled consistency index; Farris, 1989). The data are then analyzed made under these weights, and the process is repeated until a stable result is reached. As noted by Goloboff (1993), successive weighting may produce inconsistent trees, which are not optimal under the weights they imply. Calculating the weights from a set of trees is also problematic, because the characters will have different homoplasy values on different trees (best scores are typically used). In this dataset, succesive weighting with the consistency index starting with only one of the most parsimonious trees under equal weights produces at least five different results, each from a different starting tree. Successive weighting is not an optimality criterion; hence, there is no way to select among these different results. Because the general idea of successive weighting is the same as in implied weights, and the problems of the former method may interact in complex ways with the resampling procedures used below to compare methods, only implied weighting and equal weights will be considered here.

Implied weighting as implemented in Pee-Wee assigns higher weight to those characters with less homoplasy, and the sum of weights over all characters is maximized during tree searches. Each tree is evaluated according the homoplasy it implies. The value to be maximized is Fit = Σ(fiti), where fiti = K/(K + Hi), Hi is the homoplasy of character i in the tree under evaluation, and K is a constant that defines the concavity of the function. The function decreases as H grows, so the more extra steps, the lower the weight. The concave shape means that the decrease in weight from 0 to 1 extra step is greater than from 1 to 2, and so on. The effect is that trees will be preferred that save steps in less homoplasious characters, at the expense of allocating some extra homoplasy in the more homoplasious ones. In Pee-Wee, K may be an integer between 1 and 6; the lower the K, the steeper the descent of the weight function. Compared to equal weights, implied weighting (as well as any fractional weighting strategy) typically produces a more resolved consensus. Trees from K = 6 are usually more similar to those from equal weights, but those from K = 1 are more different, often with bizarre clades. One might be inclined, on philosophical grounds, to a classification that better reflects the more reliable characters, but still a decision has to be made as to how strongly to weight against homoplasy.

There have been several approaches to decide among different weighting strategies in cladistic analyses, all based on some measure of congruence. Congruence may be measured comparing tree topologies or comparing how the dataset adjusts to trees (“taxonomic congruence” and “character congruence”, respectively; e.g., Wheeler, 1995). Measures based on character congruence express a more direct relationship between trees and datasets, because they are based on the same measures that are used to evaluate trees (e.g., the length of the characters optimized over the tree). A well-known index of this kind is the incongruence length difference (e.g., Farris et al., 1995). Comparing character congruence indices when they come from different ways of measuring the fit of a character to a tree may have its caveats; however, this approach has not yet been adequately studied. For the problem addressed here, the indices proposed so far are not comparable, because the weights are not a linear function of homoplasy. For example, the rescaled fit (Goloboff, 1994; analogous to the retention index) grows for greater values of K when calculated for random trees.

I used topological measures in experiments in line with those of Penny and Hendy (1985), and especially those of Goloboff (1997a). The idea of my experiments is to estimate the predictive power of the weighting procedures. An algorithm is deemed more predictive if it produces trees that better explain data not yet examined; that is, if it is superior in finding the correct tree without part of the data. Because the correct tree and all the future data are unknown, we can only rely on some kind of estimation. Here an estimation of predictivity was obtained by measuring how well an algorithm can retrieve the results from the complete dataset, based on the partial evidence of datasets where a portion of the characters was eliminated. A more efficient algorithm will be able to recover more groups, even without part of the data. This strategy has two properties that deserve mention. First, it does not rely on fixed partitions of the dataset, for example, systems of characters. Over a long series of replications, it is possible to estimate if the variation in the indices may be attributed to the random generation of resampled datasets. Second, the reference against which the results are compared is the optimal set of trees for the total evidence.

Some indices were constructed for these comparisons. Let T be the number of groups in the consensus from the complete dataset. J is the number of groups in the consensus from the pseudoreplicate (jackknifed) dataset. The value JT is the number of groups in common between the consensus from pseudoreplicate and the consensus of the complete datasets (that is, the ‘correct’ clades recovered in the pseudoreplicate). JT varies between 0 and T. Because T will be different for each concavity in general, JT alone (or divided by a constant value, as in the fork index; see Swofford, 1991) would favor weighting strategies producing a more resolved consensus from the complete dataset, irrespective of the accuracy. Fractional weighting is well known for producing fewer trees and a more resolved consensus. A normalized value between 0 and 1 is obtained as PC = JT/T, which is the proportion of correct groups recovered by the pseudoreplicate.

Greater resolution may also produce higher J values, thus increasing JT. It is interesting to quantify an error E = JJT, the number of incorrect groups in the consensus from the pseudoreplicate. E can take values between 0 and J, and thus it can be normalized as PE = (J − JT)/J, the proportion of wrong groups from the jackknifed dataset (similar to the error rate of Goloboff and Farris, 2001).

It may be interesting to examine whether these indices based on consensus are biasing the analysis in some direction, for example, against methods that produce more ambiguous, but perhaps robust, results. Equal weights for this dataset (and also in general) produce many more trees for both complete and pseudoreplicate datasets than do any of the fractional weightings examined. The individual trees of complete and jackknifed datasets may agree or conflict in ways that are not detected by comparing just their consensus. Hence, the same results were analyzed calculating max JT, the maximum number of groups in common between pairs of trees of the jackknifed and complete datasets. Because these trees are mostly resolved, no normalization or measure of error is necessary.

One hundred jackknifed pseudoreplicate datasets were generated with probability of elimination of 0.36 (approximately e−1, the probability used for jackknifing to give values comparable to those of bootstrapping; Farris et al., 1996). The pseudoreplicate datasets were analyzed under the six available values of the constant of concavity in Pee-Wee (K = 1 to 6) and equal weights (in NONA), with the commands hold 30 hold/1 nixwts*15 3 find* (three series of 15 runs of TBR parsimony ratchet each, keeping one shorter tree each run, then TBR swapping up to a maximum of 30 trees). In preliminary runs these commands sufficed to find the optimal trees in most of the replicates, collapsing the appropriate groups in the consensus. Optimal trees for the complete dataset were hit many thousands of times in aggressive searches, and are deemed exhaustive. Searches under equal weights produce at least many thousands of trees. Only 21 of these trees were sufficient to produce the same consensus. For the pairwise comparisons of trees, this set of 21 trees was extended to 50. All these calculations were made with simple macro files in Pee-Wee.

The results are shown in figure 2 and table 2. The number of correct groups is larger for the weighting concavities K = 4 to 6, both if analyzed as a proportion of the groups in the consensus (fig. 2A) or between pairs of trees (fig. 2C). The proportion of wrong groups in the consensus from the pseudoreplicate datasets is smaller for the concavities K = 3 to 6. All these point out to a better performance of the mildest weighting functions. The low performance of equal weights in recovering correct groups is somewhat compensated through conservative results, but the error rate is still greater. The two virtues of accuracy and robustness seem to be absent with stronger concavities (K = 1 or 2), which produce well-resolved but inaccurate trees. The results presented here are based on the two optimal trees for K = 6 (strict consensus in fig. 3, character indices in fig. 4 and table 3). One of these trees (with Josa personata sister of clade 93) is also the one optimal with K = 5 and 4, and it is also the only one optimal under K = 6 if the Fit is calculated with floating point precision in X-Pee-Wee (Goloboff, 1997b; both trees differ only by ≈0.04 units of fit). With K = 3, Coptoprepes is paraphyletic (clade 107 sister of 121). Under K = 2, major rearrangements are produced in the Amaurobioidini, and with K = 1, also in Gayennini. Searches under equal weights produce a much less resolved consensus (fig. 5), with most groups as in the jackknife majority rule consensus.

In this estimation process, predictivity is somewhat equated with stability, and that is why it uses a procedure often applied to estimate clade support (jackknifing). Some authors have used the sum of jackknifing or bootstrap proportions as a measure of the accuracy of weighting strategies (e.g., Penny and Hendy, 1986; Källersjö et al., 1999). The approach used here is slightly different, besides the attention paid to the problem of resolution. First, the reference set of trees for counting agreement or disagreement of groups is calculated from the complete dataset. Spurious groups produced in the majority-rule consensus from the pseudoreplicate datasets are counted against, not in favor of a method. Second, all groups are counted, even if their frequency is below 50%. Counting only those groups that would appear in the majority consensus produces results more in favor of mild weighting functions (results not shown), but the interpretation is less clear, because jackknifing is used to generate pseudoreplicate datasets, but also as a measure of support to decide which groups of the pseudoreplicate trees are to be counted. That seems like overuse of jackknifing.

Lists of Synapomorphies

Following the description of each group, a table summarizes its synapomorphies and those of the clades included. Only unambiguous synapomorphies are listed (e.g., 0 → 1, but not 01 → 1; 01 → 2, but not 01 → 12; option ambiguous- of Pee-Wee). Diagnosing polytomies (and groups close to polytomies) has its drawbacks (Maddison, 1989; Goloboff, 1994). For the diagnosis of groups, the six most parsimonious dichotomous trees for concavities K = 4 to 6 (two resolutions of clade 95, three of clade 167) were considered. The synapomorphies common to all six trees are reported in the first place; those of only some resolutions are listed separately (calculated by command apo/).

Indices of Support

Three indices for support of individual groups were explored in this analysis. The Bremer support (Bremer, 1988, 1994; Källersjö et al., 1992) was calculated heuristically searching trees suboptimal by 1.8, 2.2, 2.6, …, 4.0, and then 5, 6, 7, …, 17 units of fit. In each search the optimal trees were TBR swapped until 4000 trees were found. Lower values of Bremer support for all these searches are reported in figure 6. TBR swapping trees suboptimal by 1.8 produced only 1767 trees without overflow of tree space, hence the values of BS ≤ 1.8 are most probably exact.

Jackknifing frequencies were calculated with 1000 pseudoreplicate datasets, eliminating characters with p = 0.36 (Farris et al., 1996). Each pseudoreplicate was analyzed with three runs of TBR parsimony ratchet (Nixon, 1999) with 25 iterations each, saving only one tree for each iteration (with the default of 20% of characters reweighted; commands h/1 nixwts*25 3). This search is very aggressive for a dataset this size, and will most likely obtain the shortest trees (a pilot test of 15 replicates with h/2 nixwts*50 8 produced the same scores). For each pseudoreplicate, the strict consensus was saved to a file and submitted to the program FQ (for majority-rule consensus, distributed P. Goloboff). The majority rule-consensus from the jackknifed pseudoreplicate datasets is presented in figure 7.

The simple majority-rule consensus from the jackknifing is perhaps too conservative a measure of support. A more refined way to analyze frequencies of jackknifed groups is the SCG Index (“supported/contradicted groupings; Goloboff et al., in prep.). SCG = SC, where S is the proportion of the pseudoreplicate datasets that produce the group and C is the proportion that contains the most frequent incompatible group. Compared to the jackknifing index, this one has the advantage of preserving groups with low frequencies when they are mostly undisputed. Because the resampling algorithm is the same, it suffers from the same bias as the jackknifing. In this analysis, the SCG Index (fig. 6) was calculated from the same tree file from 1000 pseudoreplicate datasets as the jackknifing index.

As expected, all three measures are mostly correlated (fig. 8). However, some noticeable conflicts exist among the resampling measures and the Bremer support (fig. 8A, B). Several groups with low to very low Bremer support have moderate to very high SCG or jackknifing frequencies (e.g., clades 93, 95, 98, 150, 154; fig. 8). They have in common support by only one to a few very homoplastic synapomorphies, but their placement in the tree is nevertheless undisputed (similarly as in Goloboff and Farris, 2001: S30). For instance, the three unambiguous synapomorphies for clade 93 (Josa lutea + J. riveti) are extremely homoplastic, which is reflected in a very low Bremer support (BS = 0.2). These two species are extremely similar, they are nested in one of the better supported groups (the genus Josa), and there are no many conflicting characters suggesting alternative relationships within the genus.

The resampling indices of support examined here seem more appropriate than does the Bremer support to express robustness or stability of groups. This is so because they are sensitive both to the absolute evidence in favor of groups and to the conflicting characters favoring other resolutions. These indices are not free of bias, though. In the specific case of unequal character weighting, the support for groups is heterogeneously distributed among characters, thus biasing the resampling. This effect is patent when unsupported (or plainly contradicted) groups appear as supported. In this analysis, there are two equally parsimonious resolutions in Josa, A = (Josa personata + J. calilegua) and B = (J. personata (J. riveti + J. lutea)). Resolution A appears as supported in the jackknifing analyses (figs. 6 and 7; the support is low, though). This is so because comparing both trees, tree A has better fit for two characters (difference in fit = 2.5 + 3.2), while tree B has the same difference distributed over five characters (2.5 + 0.2 + 2.5 + 0.3 + 0.2), making it more probable that a jackknifed pseudoreplicate will have resolution A rather than B. Resampling with p = 0.36, pA ≈ 0.504, and pB ≈ 0.418, then pSCG = pApB ≈ 0.09%. Spurious groups are easily detected, but the same bias is surely affecting values for supported groups in an undetected way. Bootstrap frequencies have a comparable bias with unequal weights, but in the opposite direction (Goloboff et al., in prep.). Jackknifing is preferred here over bootstrapping, because in the first the frequency of a group is not influenced by characters irrelevant to that group (Farris et al., 1996). The support metrics used here should be interpreted only as approximate measures of stability of clades.

Weighting Functions and Support

Part of the problem of settling among alternative weighting functions may be subsumed under the estimation of support for groups. It is not surprising that the clades that have different resolutions under different weighting functions are also those with low support. Support indices are constructed looking at the effect that small alterations in the dataset have on the groups (resampling methods), or the effect that alterations in the groups have on the optimality measures (Bremer support). Looking at the sensitivity of groups to small changes in the way that the data are analyzed is yet another way of evaluating the stability of groups (Wheeler, 1995). The search for the best methods of analysis is legitimate, but the arguments and strategies used so far to compare methods are not totally convincing and will no doubt be debated for a long time. The fact that the better supported clades are robust to changes in the weighting functions may help circumvent part of the problem from a practical point of view.

DISCUSSION

Some critical considerations about the groups proposed here can be derived rather directly from the approximate measures of support obtained in the cladistic analysis. Additional points can be made taking into account the way in which characters and species were selected and processed for this monograph. These will be by force subjective, but of the most value for orienting future research.

The monophyly of Amaurobioidinae is mostly undisputed and will most probably remain undisturbed should additional outgroups be included. The peculiar conformation of the male copulatory bulb, especially of the basal tegular notch, is unparalleled in any other etelegyne known so far. Similarly, all members of the tribe Gayennini proposed here have a very conservative pattern of male and female genitalia. Even though some homologies of details of the male copulatory bulb might be disputed, alternative homology relations would still have the Gayennini as a very homogeneous, most probably monophyletic group, sharing many character states (although perhaps differently defined).

Most genera in Gayennini are reasonably well supported, but others deserve comment. The paraphyly of Sanogasta (which has Arachosia as an internal clade) may need several additional representatives to be solved. Arachosia is no doubt a monophyletic group, but some species not included are similar enough to A. praesignis to be potential candidates for basal clades in the genus. It may be expected that the basal resolution of Arachosia species will affect their relationships with groups of Sanogasta. The placement of Sanogasta approximata is not well supported; analysis of the several other species not included here may help elucidate problems in clade 172. Stability in this group may hopefully suffice to raise the support for clade 173 and Gayennoides. The intraspecific variation in Arachosia bergi, Sanogasta maculosa, and clade 167 will most likely have only local effects.

Philisca is a well-defined group, except for P. puconensis, which might be related to Tomopisthes instead. There are several species not included here (with a remarkable diversity in the Chilean Pacific Juan Fernández Islands), all of them with either the male modified chelicerae (as in clade 132) or the reduced leg spination of clade 135. Araiya and Tomopisthes are here represented by all known members. Their sister group relationship is weakly supported. Clade 163 is mostly supported by the spinose anterior metatarsi of females, a set of homoplasious characters. Clades 162 and 161 have weak support, mainly because their basal species are problematic: Oxysoma saccatum shares many characters with Monapia, Tasata chiloensis with Oxysoma, and Phidyle punctipes may conceivably be a Tasata instead. Of these, Phidyle is monotypic, all species of Monapia are included, and some undescribed Chilean species of Oxysoma are very closely related to O. longiventre (if not intraspecific variants). Additional representatives that may help resolve this clade may be found in Tasata and in the undescribed females of Oxysoma itambezinho and Monapia tandil.

The genus Josa is undisputedly monophyletic, as concluded by Kochalka (1980). It is very likely that further knowledge of the many species of the genus will challenge the relationships among species obtained here, but it is unlikely that the support for the genus could be considerably eroded. The Amaurobioidini is more problematic, as shown by the low support values for the tribe and several of its internal clades, mainly on the branches among genera. Negayan and Amaurobioides are very homogeneous, and it is not likely that additional representatives would improve or challenge their monophyly. There are several species of Aysenia and Aysenoides not included here, some with noticeable differences in the male copulatory bulb. Superficial examination of their morphology suggested that they are clearly assignable to either of these genera, but the analysis under equal weights splits clade 118 in two. It is conceivable that additional species of both genera may help stabilize the group. Perhaps more promising would be the inclusion of more Coptoprepes species, a genus that is also paraphyletic under equal weights, because of its basal position in the tribe and the quite variable male copulatory bulb and tibial apophysis. The remaining genera are either monotypic or all their species were included, except a probably undescribed species close to Acanthoceto pichi, which would most likely have only a local effect on the trees.

The results of previous analyses are tested with this one, including a wider range of representatives and characters. My analyses of Liparotoma (here subsumed into Philisca), Monapia, and Acanthoceto (Ramírez, 1993, 1995b, 1997, 1999) had the same outcome in the relationships among species, except for the further resolution obtained here in the placement of Monapia alupuran. In the revision of Acanthoceto, however, the outgroups belonging to Amaurobioidini had quite different relationships, because of the limited sampling of terminals and characters. Some of the groupings proposed by Kochalka (1980) in his unpublished work are confirmed here. His Gayenna-Oxysoma group is in fact Gayennini, supported by the conformation of epigyne and spermathecae as stated by him (characters 104, 116, 124, but also others). His Amaurobioides group (Amaurobioides, Axyracrus, and Aysenia) is also corroborated, with the addition of Aysenoides, but is supported by different characters. Kochalka diagnosed the group by the pointed retrolateral tibial apophysis in the male palp (character 43, state 1), a character that is more broadly distributed and ambiguously optimized. Kochalka gave some importance to the coiling of the female copulatory duct on a longitudinal axis (characters 117, 118). This condition is shown here to be very homoplasious.

TAXONOMY

ANYPHAENIDAE BERTKAU

  • Anyphaenidae Bertkau, 1878: 358, 379 (type genus Anyphaena Sundevall, 1833). Petrunkevitch, 1939: 187. Mello-Leitão, 1947: 289–292. Lehtinen, 1967: 384. Forster, 1970: 12–15, 18. Platnick, 1974: 205–211. Platnick and Lau, 1975: 4–6. Ramírez, 1995a: 373.

  • Diagnosis:

    Spiders with two tarsal claws. Claw tuft formed by 2–8 rows of spatulate setae (figs. 10D, 13G). Nonadhesive side of spatulate setae with thick, aligned microvilli; adhesive side facing inward or obliquely to ventral/inward. Tracheal system well developed, in characteristic pattern (Ramírez, 1995a: figs. 111), tracheal spiracle moderately to widely separate from spinnerets. Posterior spinnerets without cylindrical gland spigots.

    Description:

    Recently redescribed in Ramírez (1995a).

    Composition:

    Three subfamilies: Malenellinae, Anyphaeninae, and Amaurobioidinae. The limits and relationships among the subfamilies are discussed in Ramírez (1995a), but see comments under Anyphaeninae.

    MALENELLINAE RAMÍREZ

  • Malenellinae Ramírez, 1995a: 376 (type genus Malenella Ramírez, 1995), 1997: 178.

  • Diagnosis:

    Resembles some Anyphaeninae in having minute AME and pale green color, but distinguished by having the tracheal spiracle closer to spinnerets. Females and immatures are also distinguished by having a thickened palpal tarsus with an apical dorsal patch of blunt setae (fig. 10E, F), and males by the copulatory bulb with a short, thick embolus, without median apophysis.

    Description:

    See Ramírez (1995a).

    Distribution and Composition:

    Only Malenella nana Ramírez from southern Chile.

    MALENELLA NANA RAMÍREZ Figures 9A, 10

  • Malenella nana Ramírez, 1995a: 376.

  • New Records:

    Región Metropolitana (Santiago): Santiago: Bucalemi, San Antonio, 23–24.X.1994, L. Peña (AMNH). Región IX (Araucanía): Malleco: 16.5 km NE Pucón, 12.I.1951, Ross and Michelbacher, 1♀ (CAS). Cautín: Flor del Lago Ranch, Villarrica, Polo Field, 39°12.300′S, 72°08.367′W, 282 m, canopy fogging GT Nothofagus obliqua roble, 13.XII.2001, Arias et al., 1♀ (UCB). Región X (Los Lagos): Chiloé: Isla de Chiloé: El Pozuelo, 1.5 km NE Butalcura, 2.II.2001, T. Cekalovic, 1♀ (AMNH).

    ANYPHAENINAE BERTKAU Table 4, Figure 9B, C

  • Anyphaenidae Bertkau, 1878: 358 (type genus Anyphaena Sundevall, 1833).

  • Anyphaeninae Bertkau, 1878: 379. Reviewed at generic level by Brescovit, 1997.

  • Diagnosis:

    Tracheal spiracle advanced, at midpoint or closer to epigastrium than to spinnerets. Tegulum without basal notch.

    Note:

    In this analysis Anyphaeninae is paraphyletic in terms of Amaurobioidinae, but see comments below.

    Description:

    Recently redescribed by Brescovit (1997).

    Distribution and Composition:

    Thirty-three genera, mostly from the New World. Anyphaena Sundevall has representatives in Palearctic and Oriental regions; Australaena Berland is known only from Polynesia.

    Six representative species of Anyphaeninae are included as outgroups. They were selected for having putatively homologous conditions of important characters diagnostic of higher groups of Amaurobioidinae. Species of Xiruana Brescovit have a structure (“conductor of Anyphaeninae”, Brescovit, 1997) very similar to the secondary conductor found in amaurobioidines. The prolateral and retrolateral tegular projections in Anyphaena accentuata (Walckenaer) (Brescovit, 1997: figs. 3, 4) were both tentatively homologized with the secondary conductor. Species of Wulfila O.P.-Cambridge and Anyphaena accentuata have a putative homolog of a primary conductor (named “ventral tegular projection” by Brescovit, 1997). In A. accentuata, this structure has a shallow canal where the embolus fits, favoring homology with a conductor (fig. 11A). Species of Aysha Keyserling have a complex process at the embolar base comparable to that of Josa. Italaman santamaria Brescovit has an acute retrolateral tibial apophysis similar to that found in Amaurobioides and close relatives. Several conditions present in Malenella nana are also found in Wulfila (green color, minute AME, absence of accessory bulbs in the spermathecae), Italaman (simple male copulatory bulb), and Otoniela Brescovit (many anterior leg spines, absence of accessory bulbs). This selection of representative outgroup characters is clearly insufficient for a sound hypothesis of relationships among anyphaenines. Because only a few characters specific for that subfamily are here included, and because the rather unusual Malenella was used as outer outgroup, it is clear that a more complete analysis is needed to appropriately assess anyphaenine relationships. In this dataset, only three correlated characters would support the monophyly of Anyphaeninae (the advanced tracheal spiracle and associated modifications of the tracheal system, characters 38–40). The effect of the suspicious resolution of Anyphaeninae over the relationships among amaurobioidines, however, is limited: if Anyphaeninae is constrained to be monophyletic, all groups in Amaurobioidinae are the same in the optimal trees, except for the collapsing of clade 95 of Josa. Character optimizations at the basal node of Amaurobioidinae are the same except for characters 15 (state 1 instead of 2), 79 ([012] instead of 1), 123 ([01] instead of 1), 125 (0 instead of [01]), 138 ([01] instead of 0), and 163 ([01] instead of 1).

    Brescovit (1997) suggested a monophyletic group of nine genera including Aysha, united by having a process on the male embolar base. I cannot test his hypothesis here, not only because of my modest selection of representatives and specific characters for Anyphaeninae, but because there is not yet a coherent scheme of homologies for the many details of embolar morphology, embracing both anyphaenines and amaurobioidines.

    AMAUROBIOIDINAE HICKMAN Table 5

  • Amaurobioididae Hickman, 1949: 31 (type genus Amaurobioides O.P.-Cambridge, 1883). Forster, 1970: 165. Synonymized with Anyphaenidae by Platnick, 1974: 211.

  • Amaurobioidinae: Lehtinen, 1967: 211, 316, 320–321. Kochalka, 1980: 65. Ramírez, 1995a: 381.

  • Note:

    Simon (1897a: 99–100, 104, 1903a: 1032, 1903c: 29) used informal names for some Amaurobioidinae that might be confused with family-level names (Tomopisthini, Oxysomini, Oxysomate), which clearly referred to the members of each genus, and are not available at familylevel (ICZN, 1999: 11[f]2).

    Diagnosis:

    Distinguished from Anyphaeninae by the male tegulum with a deep notch occupied by the median hematodocha, visible in ventral view as a membranous area at base of the copulatory bulb. Most species have the tracheal spiracle closer to the spinnerets than to the epigastrium.

    Description:

    Body size small (2.50) to medium (22.00). Chelicerae (fig. 12A) with three teeth on promargin (exceptionally four or five), two to seven teeth on retromargin (exceptionally one). Labial apex rounded or slightly notched. Male palp with only one tibial apophysis in retrolateral apical position, or lacking apophysis. Tip of cymbium with ventral canal devoid of setae, often associated with embolus (cymbial conductor). Subtegulum prolateral, visible in unexpanded palp. Tegulum with deep basal notch occupied by median hematodocha. Sperm reservoir in subtegulum, sperm duct running to tegulum through apical margin of bulb (clockwise, left palp ventral view), then parallel through retrolateral border of cymbium, then bordering tegular notch and entering into embolar base. Embolus articulated prolaterally to tegulum. Embolar base usually with one process of diverse shape; terminal portion of embolus filiform or moderately thick, usually fitting into canal on conductor (primary or secondary) and into cymbial conductor. Apical portion of tegulum housing distal hematodocha, where rest of sclerites are inserted. Median apophysis retrolateral, usually articulated on weakly sclerotized area, commonly hook-shaped, sometimes bifid, reduced, even absent in some species. Paramedian apophysis arising from sclerotized plate on distal hematodocha, between median apophysis and retrolateral ventral border of tegulum, with one to several sclerotized cusps. Primary conductor arising between embolus and prolateral ventral border of tegulum (relictual in Gayennini), often with canal where embolus fits. Secondary conductor arising distally, associated with apical dorsal tegular stripe where sperm duct runs. Epigyne with paired lateral lobes and median field, copulatory openings usually associated with furrows between median field and lateral lobes. Copulatory duct weakly sclerotized from opening to joint with duct of accessory bulb; from that point duct well sclerotized, coming into spermatheca. “Dictynoid” pore conspicuous, close to union of copulatory duct with spermatheca and fertilization duct. Tracheal spiracle variable from close to spinnerets to midpoint between spinnerets and epigastrium (except in some Arachosia and Acanthoceto, slightly closer to epigastrium). Length of lateral tracheae variable according to spiracle position, reaching spinnerets. Trichobothriae in one row on metatarsi, two rows on tarsi (fig. 13A). Anterior lateral spinnerets with two major ampullate gland spigots (figs. 15B, 16B, 17B, 117B), or one plus nubbin (Ramírez, 1995a: fig. 39), and several unmodified piriform gland spigots. Posterior median spinnerets with two minor ampullate gland spigots (figs. 16C, D, 17C) and several aciniform gland spigots. One PMS minor ampullate detected in Amaurobioides (fig. 15C, contra Ramírez, 1995a), but it is so similar to aciniforms that second one may have been overlooked. Posterior lateral spinnerets only with aciniform gland spigots (figs. 15D, 16E, 17D). Gland spigots not sexually dimorphic, cylindrical gland spigots absent.

    Distribution:

    Mainly South America, but Josa extending also to Central America, Arachosia to Central and North America, and the coastal genus Amaurobioides in the seashores of Chile, South Africa, Australia, Tasmania, and New Zealand.

    Composition:

    Twenty-two genera grouped in two tribes, Amaurobioidini and Gayennini, here defined, plus the genus Josa, sister group of Gayennini, for which I declined to erect a tribe.

    Types Not Examined:

    The following species have been assigned to various amaurobioidine genera, but their placement cannot be clarified without examination of the types: Anyphaena trivittata Bertkau, 1880; Anyphaena furcata Keyserling, 1880; and Anyphaena vittata Keyserling, 1881.

    Nomina Dubia:

    The types of the following Chilean species have not been found in MHNP, and the original descriptions are very ambiguous: Clubiona lineata Nicolet, 1849; Clubiona limbata Nicolet, 1849; Clubiona nubes Nicolet, 1849; and Clubiona versicolor Nicolet, 1849.

    Morphological Remarks

    Male Copulatory Bulb

    The copulatory bulb is united to the cymbial alveolus by the basal hematodocha and a triangular petiole. Because neither the subtegulum nor the tegulum forms a complete ring, the three hematodochae (basal, median, and distal) are continuous (fig. 33E). The subtegulum is compact, partially visible in prolateral view in the unexpanded bulb (figs. 83A, 129E), but mostly distally in Amaurobioidini (fig. 47B). Part of the median hematodocha is visible in ventral view in the unexpanded palp, occupying a basal notch on the tegulum, which is the most conspicuous synapomorphy of Amaurobioidinae. The anterior part of the bulb is occupied by the distal hematodocha, from where the conductors, the median apophysis, and the paramedian apophysis arise.

    Conductors

    The homology of the extremely diverse structures accompanying the embolus of entelegyne spiders is a contentious field. Here I tried to establish the homology of the structures across the subfamily, with variable success. I call the primary conductor (C1) a structure arising between the base of the embolus and the prolateral margin of the tegulum. The C1 is most evident in Amaurobioidini, often bearing a long canal where the embolus fits. The apical portion of C1, where the canal ends, usually forms a heavily sclerotized, partially coiled beak (e.g., fig. 50D). The apical portion of C1 may look like a separate sclerite if the basal portion is protruding as well (fig. 33B). In the Gayennini, there is a small sclerite mostly hidden by the other structures arising from the distal hematodocha (figs. 63C, 105C), connected by a sclerotized stripe to the articulation between the embolus and prolateral margin of tegulum, which is a presumed homolog of the C1. In Josa the basal portion of the C1 is fused with the tegulum; the apical portion is no longer identifiable (fig. 60A, B).

    I call the secondary conductor (C2) a sclerite also associated with the embolus, arising from the apical-dorsal region of the distal hematodocha. It is closely related, often fused, to a sclerotized stripe of the tegulum where the sperm duct runs. In Amaurobioidini, the main structure leading the embolus is the C1, and the C2 is small, with a poorly defined canal (fig. 26A). In Gayennini this situation is reversed, and the C2 is a conspicuous structure. In Josa, the C2 is hypertrophied and very complex (figs. 56B, 60B).

    Paramedian Apophysis

    Hidden behind the retrolateral margin of the tegulum there is a tightly folded section of the distal hematodocha, which becomes inflated during expansion (fig. 50D; see also character 71). A sclerotized plate attached to this section of hematodocha is called the paramedian apophysis (PMA), which, in Amaurobioidini, is distally articulated to the C1 (fig. 50D). The PMA may have one (in Gayennini) to several cusps (in Amaurobioidini). The PMA is also connected, often more or less fused, with the median apophysis. A protruding PMA appears in Gayennini (see character 68), together with the anterior pouch on the median epigynal field (APmf, character 104). Because the shape of the PMA seems also to be somewhat correlated with the shape of the APmf (e.g., fig. 124D, H), it is possible that the PMA fits into the APmf during copulation, at least in some species.

    Median Apophysis

    The median apophysis (MA) is much more conservative than are the conductors. In most Amaurobioidinae the MA is very simple, small, and hooked, at some extent articulated or fused with the plate of the PMA. Species of Coptoprepes have a larger, complex MA. Davies (1998) doubted the homology of the MA in Amaurobioides, because it is not articulated, but is firmly fused to a complex of paramedian apophysis and part of the conductor. However, when close relatives of Amaurobioides are examined (Axyracrus, Aysenia, Aysenoides), it is clear that the distal hematodocha where C1, PMA, and AM are placed underwent variable degrees of sclerotization. Because the morphology of the area is otherwise conservative, the homology of the MA seems clear. The homology of the MA as identified in Josa is somewhat more problematic, because the entire distal region of the bulb is extremely modified. The MA becomes reduced independently in several clades (character 64), but in these cases the general morphology of the area is similar to that of close relatives.

    Embolus

    Coddington (1990) noted that the embolus of Amaurobioides runs counterclockwise, differing from most entelegynes. Other Amaurobioidini, and to a lesser degree to a Gayennini, also have that embolar course. The change from clockwise to counterclockwise involved only minor modifications of the copulatory bulb, compared to a more generalized conformation. The region of the basal articulation of the embolus has an acute to straight angle, thus changing the course, while the basal part of the sperm duct runs clockwise, as in other entelegynes. Amaurobioidines with a less pronounced angle (e.g., Selknamia, figs. 52, 53A, B) are ambiguous as to the curvature, while in others the curvature looks different from prolateral and ventral views (e.g., Negayan, figs. 47D, 50A, B). In Anyphaeninae there is great variability in embolus morphology, and sometimes it is not clear in what direction a contorted embolus is running.

    Alternative Interpretations of Male Palp Structures

    The identification of C1, C2, and cusps of PMA is disputable. The system of homologies adopted here is what seemed more congruent after detailed study of the anatomy and informal examination of alternative codings during the construction of the dataset. Those alternative codings did not produce very different results from those shown here, though. This is because alternative homologies have a coordinated effect on groups of terminals with similar morphology. The C1 in Gayennini, as interpreted here, is a small piece, compared to the structure found in Amaurobioidini. An alternative coding might consider this small piece the second cusp of the PMA (which is lacking in Gayennini, compared with most Amaurobioidini). The C2 of Gayennini might be considered a C1 (or conversely, the apical portion of C1 in some Amaurobioidini, a C2), because in Negayan and Selknamia the apical portion of C1 is similar to the C2 of Gayennini. This coding was used in preliminary datasets, but was later abandoned, because the homologies would be supported by intermediate morphologies that were not intermediate in phylogeny! Settling these problems of unclear homologies may involve the use of alternative codings, keeping those homologies (and the corresponding trees) that imply most parsimonious results (Rieppel, 1996; Wheeler, 1996).

    Spermathecae and Associated Ducts

    There is some disagreement on the terminology for these structures. I mostly followed Sierwald (1989), with modifications. The main difference is interpretation of the copulatory duct. In Sierwald's view, this duct runs from the copulatory opening to the connection with a duct from the “head of spermatheca”, here accessory bulb (AB) after Carico and Holt (1964). The segment from that point to the “base of spermatheca” (here spermatheca) is considered by Sierwald as part of the spermatheca itself, the “peduncle of spermatheca”, uniting the head and base. This terminology seems adequate for Lycosoidea (Griswold, 1993; Diana Silva, personal commun.), but not so much for Amaurobioidinae, at least in functional terms. In Amaurobioidinae, the accessory bulb joins the copulatory duct relatively close to the copulatory opening, and from this point to the origin of the fertilization duct there runs a tube that evidently functions as a duct rather than as a reservoir. In fact, the male embolus runs through that segment (Ramírez and Kochalka, 1993: fig. 4), and I could not find much justification for considering it part of the spermatheca (a name that indicates storage function). Comparative anatomy does not help much to settle the question, because the evolutionary transformations that lead to the entelegyne female genitalia are not well understood. I am inclined to accept the homology of the accessory bulb with one of the paired receptacles, as found in most Mygalomorphae and many Haplogynae. In this case, the fertilization ducts are homologous to the ducts leading to one pair of receptacles, the entelegyne copulatory openings are homologous to the duct leading to the second pair of receptacles, and the “peduncles” are invaginations of the body wall connecting the two receptacles of each side (see Sierwald, 1989).

    The development in Tomopisthes horrendus is in agreement with this hypothesis (fig. 106). The accessory bulb arises anteriorly on a cuticular fold separating the median field from the lateral lobes, the primordium of the copulatory opening leads to the primordium of the accessory bulb, and both primordia are connected by a deep folding corresponding to the copulatory duct. For the sake of descriptive power, I refer as “spermatheca” to the ample chamber immediately connected to the fertilization duct, “copulatory duct” to the tube running from the copulatory opening to spermatheca, and “accessory bulb” to the blind sac bearing conspicuous pores, connected to the copulatory duct by a tube of variable length. The accessory bulb was recently referred to as “seminal receptacle” (Brescovit, 1997) and “diverticulum” (Huber, 1995). Bonaldo (2000) called it “secondary spermatheca” on account of its function as a sperm reservoir in Corinninae.

    Leg Spines

    Goloboff (1995b) used several characters from patterns of spines in Nemesiidae. Bosselaers and Jocqué (2002) used both patterns and individual spines in an analysis of some Corinnidae and Liocranidae. They found those characters informative, with similar levels of homoplasy as for other somatic ones. Here many of the leg spines were coded as independent characters, homologized according to their position. This is possible since the spination in Anyphaenidae follows a rather conservative pattern.

    In most genera the spines on leg I are similarly distributed to those on leg II. Legs III and IV are also similar in spines, which are more numerous than on forelegs. Through the four pairs of legs, most spine positions are conserved, because they are serially homologous. A common pattern is:

    Legs I and II, femur d 1–1–1, p 0–1-(1-d1), r d1ap; tibia v 2–2–2; metatarsus v 2bas. III, femur d 1–1–1, p and r 0-d1-d1; patella r d1; tibia v 2–2–2, p and r d1–1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella, tibia, and metatarsus = III.

    In some groups the anterior legs are almost as spinose as the posterior legs. A common pattern of this type is:

    Leg I and II, femur d 1–1–1, p and r 0-d1-(1-d1); tibia v 2–2–2, p and r d1–1, d r1-0-1-0; metatarsus v 2bas, p and r d1–1–1, d 0-p1–2. III, femur = I; patella r d1; tibia = I; metatarsus = I, but v 2–2–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella, tibia, and metatarsus = III.

    Most spine patterns vary between these two examples. In the spinose pattern, spines on anterior and posterior legs differ mostly by the ventrals on metatarsi. There are only a few species with more than two ventral spines on metatarsus I or II, they are not especially spinose on other surfaces, and these spines are not usually sexually dimorphic. Some species have more than three pairs of ventral spines on tibiae I and II, conferring a raptorial appearance (e.g., some Monapia).

    Males are often more spinose than are females. The additional male spines appear after the last ecdysis. Spines of penultimates of both sexes are similar to those of the female. In some rare specimens (but commonly in Sanogasta backhauseni) there are supernumerary spines, for example, two or three spines where one is expected. Such an anomaly is often asymmetrical.

    Bristles (similar to spines but thinner and shorter) seem to be homologous to spines, because some specimens have a bristle where a spine is normally found. Frequent positions for replacement of spines by bristles are the prolaterals and retrolaterals on femora, and the v p1-x-x of tibia II. In species with spinose males, it is common that the male has a spine where the female has a bristle; common positions are the dorsals of tibiae (r1-0-1-0) and patellae (1–0–1).

    Inter- and intrasexual variabilities in spines were coded without distinction as polymorphisms in the data matrix, with internal steps added accordingly. It is expected that variability is underestimated in species known from a few exemplars, and more drastically if the males are unknown. To estimate the effect of variations in the internal steps for the spine characters, several replications were examined with the internal steps for the spine characters assigned randomly between 0 and 29. These produced virtually the same trees, except for some clades of little support, suggesting that underestimation of variability in spines is not decisive in this analysis.

    TRIBE AMAUROBIOIDINI HICKMAN Table 6

  • Amaurobioididae Hickman, 1949: 31 (type genus Amaurobioides O.P.-Cambridge, 1883).

  • Diagnosis:

    Distinguished from Gayennini and Josa by the male copulatory bulb with an apical dorsal loop on the sperm duct, visible in apical view (figs. 21H, 26C, white arrow; absent in Acanthoceto acupicta group, see character 53), and by the well-developed primary conductor with canal where the embolus fits (fig. 47).

    Description:

    Chelicerae commonly with three or more teeth on retromargin, sometimes only two. Male palp with one retrolateral tibial apophysis (reduced in Gamakia, absent in Coptoprepes campanensis). Cymbial conductor wide. Sperm duct with loop on apical side of copulatory bulb. Primary conductor with canal where embolus fits; basal portion often weakly sclerotized, close to base of embolus; apical portion, where canal ends, sclerotized, of varied shape, sometimes diverging from basal portion (fig. 33B). Secondary conductor generally small, partially or totally fused to apical tegular stripe where basal part of sperm duct runs, well developed only in some Coptoprepes. Paramedian apophysis not projecting, usually with two conical cusps. Epigyne without anterior pouch, lateral lobes separated from each other. Spermathecae usually of irregular shape, but spherical in Selknamia and Aysenoides.

    Distribution:

    South America, except the coastal genus Amaurobioides, which occur in the sea-shores of Chile, South Africa, Australia, Tasmania, and New Zealand, and Sanogasta maculatipes, probably introduced in Eastern Island.

    Composition:

    Ten genera, four of them newly proposed here: Acanthoceto Mello-Leitão, Amaurobioides O.P.-Cambridge, Axyracrus Simon, Aysenia Tullgren, Aysenoides, n. gen., Coptoprepes Simon, Ferrieria Tullgren, Gamakia, n. gen., Negayan, n. gen., and Selknamia, n. gen.

    Amaurobioides O.P.-Cambridge Table 7

  • Amaurobioides O.P.-Cambridge, 1883: 356 (type species by monotypy Amaurobioides maritima O.P.-Cambridge, 1883). Simon, 1897a: 89, 1903a: 1034. Hogg, 1909: 163. Hewitt, 1917: 704. Hickman, 1949: 31, 1951: 1. Forster, 1955: 184. Lehtinen, 1967: 211 (removed from synonymy of Uliodon L. Koch, 1873). Coddington, 1990: 10. Ramírez, 1995a: 366, 1997: 178. Davies, 1998: 212.

  • Cluilius Simon, 1889: 220 (type species by monotypy Clubiona chilensis Nicolet, 1849; see Simon, 1904: 100, and Synonymy in Philisca). new synonymy.

  • Synonymy:

    The type species of Cluilius is here considered a typical Amaurobioides. See also Synonymy in Philisca.

    Diagnosis:

    Distinguished from other genera of Anyphaenidae by having many aciniform gland spigots on posterior median spinnerets (Ramírez, 1995a: fig. 43).

    Description:

    Redescribed by Forster (1970).

    Morphological Remarks:

    The posterior median spinnerets densely covered by spigots might have a correlation with their aquatic habitats, because the retreat silk is much denser than that of other Anyphaenidae. In support of this association, the PMS are similarly enlarged and covered by many aciniform gland spigots in the European aquatic spider Argyroneta aquatica (Clerck) and in species of the intertidal spider genus Desis, from Pacific and South African coasts (Lehtinen, 1967). Forster (1970) reported three processes on the male palpal conductor of Amaurobioides maritima. One of them is here interpreted as the tip of the primary conductor, bearing the canal, the other two as cusps of the paramedian apophysis. Davies (1998) suggested that the structure that is here identified as secondary conductor may be a primary conductor (C1) instead, because in Amaurobioides isolata Hirst it is a well-defined sclerite, entirely bordered by a membranous area; however, the same is true for the structure here identified as C1.

    Distribution:

    Shores of Austral regions: one species from Chile, one from South Africa, all others from Australia, New Zealand, and Tasmania.

    Composition:

    In addition to the three species detailed below: A. isolata Hirst, 1993, A. litoralis Hickman, 1949, A. major Forster, 1970, A. minor Forster, 1970, A. pallida Forster, 1970, A. picuna Forster, 1970, A. piscator Hogg, 1909, A. pleta Forster, 1970, and A. pohara Forster, 1970.

    Amaurobioides maritima O.P.-Cambridge Figure 18A–D

  • A. maritima O.P.-Cambridge, 1883: 356 (male holotype from New Zealand, Otago, Allday Bay, in BMNH, not examined).

  • A. maritimus: Forster, 1970: 168. Ramírez, 1995a: 28.

  • Description and Diagnosis:

    See Forster (1970). Additional data are provided below.

    Female (Otago):

    Total length 15.45. Carapace length 5.45, width 4.26, wider on legs II–III. Leg III, length of tibia 2.50, metatarsus 2.67; leg IV, length of tibia 3.23. Sternum length 3.50, width 2.07. Abdomen length 10.00, width 6.25, spiracle–epigastrium 4.40, spiracle–spinnerets 1.10. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–2; metatarsus v 2bas. II, femur = I; tibia v r1–2–2 or 2–2–2, p 1–1; metatarsus v 2-(p1-r1)-0 or 2-r1–0, p 1–0. III, femur d 1–1–1, p and r 0-d1-d1; tibia v 0–2–2, p and r 1-0-1-0; metatarsus v 2–2–2, p d1–1–1, r 0–1–1, d 2ap. IV, femur d 1–1–1, p and r d1ap; tibia v p1–2–2 (plus r1 supernumerary), r d1ap or 0-d1-d1; metatarsus v 2-p1–2 or 2-p1-r1, p and r 0–1–1, d 2ap. Epigyne: lateral lobes separate, anterior margin elevated. Median field slightly sclerotized. Copulatory ducts irregular, contorted before reaching spermathecae, ducts of accessory bulbs long, thick, diverging (fig. 18A).

    Male (Otago):

    Spines as in female, except: leg I, femur d 1–1–1, p 0-d1-d1, r d1ap; tibia v 2-2-2-2 or 6 spines on each side, irregularly paired, p 1–2–1, r 1–0–1; metatarsus v 2bas, p d1–1–0, r 1, d p1. II, femur = I; tibia v 2–2–2, p and r 1–1–1; metatarsus v 2-r1–0, p d1–1–0, r 1. III, tibia v 2–2–2, p and r 1-d1–1. IV, femur = I; tibia = III, but p 0-d1–1 or 1ap; metatarsus v 2–2–2, p 0–1–1, r d1–1–1. Palp (fig. 18B–D): tibia short, width/legth 0.85, RTA very long, sharp, slightly concave ventrally. Cymbium with retrolateral notch where median apophysis fits, and short basal projection, opposing tibia. Sperm duct thick, suddenly narrowed in front of tegular notch. Embolus with basal process flat, well developed. Median apophysis small, thin, apical. Paramedian apophysis with three separate cusps, apical cusp concave, placed under median apophysis, median cusp conical, heavily sclerotized, ventral cusp flattened, weakly sclerotized. Primary conductor well developed, with conspicuous canal. Secondary conductor compressed, partly fused to anterior dorsal margin of tegulum.

    Variability:

    Spines, according to Forster (1970). Female: II, tibia p 0–1; metatarsus v 2bas. III, tibia p 1–1–1; metatarsus d p1ap. IV, tibia v p1-p1–2, p 0; metatarsus r 0–1–1. Male: I, tibia v 2–2–2, p and r 1–1–1; metatarsus v 2–2–0, p 1–1–0. II, metatarsus v 2–2–0, p and r 1–1–0. III, metatarsus r d1–1–1, d 0-p1–2. IV, tibia p 0–1–1, r 1–1–1.

    Material Examined:

    NEW ZEALAND: Otago, St. Clair Beach, on cliff face, 28.IV.1979, J. Carico, 1♂ 1♀ (MACN-Ar, identified by R. Forster).

    Amaurobioides africana Hewitt Figures 13A–D, 15, 18E–I

  • Amaurobioides africanus Hewitt, 1917: 704 (female holotype from South Africa, East London, in BMNH, examined). Lehtinen, 1967: 212. Forster, 1970: 176 (removed from synonymy of A. maritima O.P.-Cambridge). Lopez, 1974: 902.

  • Diagnosis:

    Distinguished from other Amaurobioides by the wide, sclerotized portion of paramedian apophysis with three separate cusps (fig. 18G–I), and by the shape of the elevated margins of epigyne (fig. 18F).

    Female (Namibia):

    Total length 14.00. Carapace length 4.92, width 3.60, wider on leg II. Length of tibia/metatarsus: I, 2.83/2.50; II, 2.67/2.10; III, 2.37/2.10; IV, 2.60/2.33. Palpal tarsus length 1.50. Chelicerae very strong, with three teeth on retromargin. Sternum length 2.83, width 1.77. Spines, all short: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–2; metatarsus v 2-r1–0. II, femur = I; tibia v r1–2–2, p 0–1; metatarsus v 2-r1–0, p 1. III, femur = I; tibia v p1-p1–2, p 1-d1–1 or 0–1, r d1–1; metatarsus v 2–0–1, p and r 0–1–1, d 2ap. IV, femur d 1–1–1 or d 1–0–1; tibia v p1-p1–2, p 0, r d1–1; metatarsus v 2–0–1 or 2-r1–1, p and r 0–1–1, d 2ap. Abdomen length 9.44, width 5.32. Spiracle–epigastrium 5.19, spiracle–spinnerets 1.07. Epigyne (fig. 18E, F): lateral lobes separate, depressed on anterior margin, prolonged in V-shaped marks on elevated median field. Copulatory ducts irregular, contorted before reaching spermathecae, ducts of accessory bulbs long, thick, converging; some of their gland ducts discharging before expansion of bulb.

    Male (Namibia):

    Total length 8.25. Carapace length 3.60, width 2.67. Length of tibia/metatarsus: I, 2.97/2.83; II, 2.93/2.73; III, 2.47/2.23; IV, 2.50/2.30. Chelicerae smaller than those of female, teeth evenly spaced. Spines as in female, except: leg I, tibia p 1–0–1. II, tibia v 2–2–2, p 1-d1–1; metatarsus v 2–2–0. III, tibia v 2–2–2, p and r 1-d1–1; metatarsus v 2-r1–1 or 2–2–1, p and r d1–1–1. IV, femur d 1–1–1, p d1ap; metatarsus v 2-r1–1, p and r d1–1–1. Palp: tibia short, width/length 1.07, RTA long, sharp, forming subtle angle at base. Cymbium with weak apical notch where median apophysis fits, and short, rounded basal projection opposing tibia. Copulatory bulb (fig. 18G–I): sperm duct thick, suddenly narrowed in front of tegular notch. Embolus with basal process flat, thin, striated longitudinally. Median apophysis long, apical. Paramedian apophysis with flattened portion close to median apophysis, wide, sclerotized, bearing three separate cusps, and weakly sclerotized portion close to primary conductor, projecting as flattened, triangular cusp. Primary conductor with open canal, bifurcating in elongate tips; canal ending on dorsal tip. Secondary conductor compressed, partly fused to anterior dorsal margin of tegulum, its membranous apex with vaguely defined canal.

    Natural History:

    Lamoral (1968) made a detailed ecological and physiological study, comparing A. africana with the sympatrid and also intertidal Desis formidabilis (Desidae). He found that A. maritima builds retreats using molusc shells, or only silk, in rock crevices, which endure daily periods of immersion as the tides rise. They resist long periods of immersion, taking oxygen from the water through an air film retained by hydrophobic hairs. They resist at least 12 hours of immersion after the air film has disappeared. The spiders are nocturnal and seem to prey mostly on isopod and amphipod crustaceans. The rhythm of silk nest building remained coordinated with the tides up to a week in the laboratory.

    Distribution:

    South Africa.

    Other Material Examined:

    NAMIBIA: Luderitzbucht, intertidal rocks (26°35′S, 15°10′E), 8–10.X.1984, C. Griswold and T. Meikle Griswold, 1♂ 3♀ 2 immatures (CAS). SOUTH AFRICAN REPUBLIC: Western Cape: Cape Peninsula (34°08′S, 18°20′E), intertidial rocks, 1966, B. Lamoral, 1♂ 1♀ (CAS); Cape of Good Hope, in rock crevices though white silken tubes, upper edge of average high tide, 6.II.1991, V. and B. Roth, 2♀ (CAS); Kommetjie, 34°9′S, 18°20′E, 30 air km S of Cape Town, intertidial zone, under rocks, 13.III.2001, L. Prendini, D. Ubick, 3♀ 3♀ penultimates, 1♀ (CAS); 3 mi S Port Nolloth, 1 m, 5.I.1967, E.S. Ross and K. Lorenzen, 1♀ (CAS).

    Amaurobioides chilensis (Nicolet), new combination Figures 19, 20

  • Clubiona chilensis Nicolet, 1849: 419 (female presumably holotype from Chile, no specific locality, in MHNP 4231, examined).

  • Cluilius chilensis: Simon, 1889: 220.

  • Philisca chilensis: Simon, 1897a: 86, 1904: 48.

  • Amaurobioides cf. boydi: Ramírez, 1995a: 366 (misidentification).

  • Note:

    The specimens examined in Ramírez (1995a), misidentified as Amaurobioides cf. boydi (here synonymyzed with Axyracrus elegans), are provisionally identified here as A. chilensis. The species was not included in the cladistic analysis, because the females are almost identical to those of A. maritima, and the males are unknown. There are some differences in the spermathecae between the type of A. chilensis and the specimens I collected in Chile, and it is possible that more than one species is involved. Opell (American Arachnological Society Annual Meeting, Keene, 2001) reported problems in species delimitation for New Zealand Amaurobioides.

    Diagnosis:

    Very similar to A. maritima, distinguished by the less curved epigynal lateral lobes (fig. 20B).

    Female (holotype, fig. 20A, spines from Cruz Grande MACN-Ar 9848):

    Total length ca. 8.68 (abdomen deteriorated). Carapace length 3.48, width 2.22, wider at leg II. Length of tibia/metatarsus: I, 1.75/1.70; II, 1.70/1.62; III, 1.30/1.40; IV, 1.65/1.65. Chelicerae strong, with three teeth on retromargin. Sternum length 1.91, width 1.11. Spines (six females): femora I–IV d 1–1–1, p d1ap or p 0. Leg I tibia v 2–2–2; metatarsus v 2bas. II, tibia v 2–2–2, p 0–1; metatarsus v 2bas, p 1–0. III, tibia v 0-p1–2 or p1–2–2, p 1-d1–1 or d1–1 or 0–1, r d1–1 or 0–1; metatarsus v 2–0–1 or 2-p1–1, p d1–1–1 or 0-d1–1, r 0-d1–1, d 0-p1–2. IV, tibia v p1-p1–2, r d1–1 or 0; metatarsus v p1-p1–1 or 2-p1–1, p 0-d1–1 or 1ap, r d1–0–1 or 1ap, d r1ap. Abdomen badly preserved, spiracle–epigastrium 2.77, spiracle–spinnerets 0.43. Color: type with abdomen totally faded. Fresh specimens are very similar to those of A. maritima (Forster, 1970: fig. 463). Epigyne (fig. 20B, C): lateral lobes separate, elevated at anterior margin. Median field weakly sclerotized. Copulatory ducts irregular, contorted before reaching spermathecae, ducts of accessory bulbs short, thick, diverging.

    Male:

    Unknown.

    Natural History:

    Collected from retreats made of white, very dense silk in rock crevices at the seashore in the spray zone (fig. 19).

    Distribution:

    The type lacks a precise locality. Collected in two widely separate localities at Chilean seashore, probably with a much more extensive distribution.

    Other Material Examined:

    CHILE: Región IV: Elqui: coast 6 km S Cruz Grande, 11.XI.1993, 29°29′S, 71°19′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 8 immatures (MACN-Ar), 6 immatures 2nd stage (MACN-Ar), 1♀ (MACN-Ar 9848), 2♀ (MACN-Ar 9849, photos MJR 1324–1326), 1♀ (MHNS), 2♀ (AMNH); same, 9 km S Cruz Grande, 5 immatures (MACN-Ar). Choapa: 12 km S Los Vilos, Rt. 5, km 213, elev. 5 m, 33°00′S, 71°31′W, 13.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1 immature (MACN-Ar). Región X: Llanquihue: 30 km E Puerto Montt, 41°36′S, 72°42′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH).

    Axyracrus Simon Table 8

  • Axyracrus Simon, 1884: 140 (type species by original designation Axyracrus elegans Simon, 1884), 1887: E23, 1897a: 90, 96–98. Keyserling, 1891: 83. Ramírez, 1995a: 381, 1997: 178.

  • Schiapellia Mello-Leitão, 1938: 115 (type species by original designation Schiapellia gerschmanni Mello-Leitão, 1938). new synonymy.

  • Synonymy:

    The type species of Schiapellia is here considered a junior synonym of Axyracrus elegans.

    Diagnosis:

    Resembles Amaurobioides and Aysenia in having a recurved posterior eye row, distinguished by the complex embolar process, reduced median apophysis, reduced basal portion of the primary conductor (without canal), and by the epigynal median field slightly elevated, frequently with copulatory plugs.

    Description:

    Carapace wide in front, posterior eye row recurved, ocular area projecting (fig. 21A). Chelicerae strong, slightly smaller in males, with three teeth on promargin, two on retromargin. Spines on anterior legs unmodified. Male palpal tibia slightly longer than wide, RTA long, pointed (fig. 21D, E). Tegulum with sort basal notch (fig. 21F). Sperm duct with pronounced loop on anterior dorsal margin (fig. 21H). Embolus not associated with canal on primary conductor, with complex basal process. Median apophysis reduced. Basal portion of primary conductor reduced to weakly sclerotized lobe, without canal. Apical portion thick, heavily sclerotized, with wide, shallow canal. Paramedian apophysis with two blunt, heavily sclerotized cusps (fig. 21G). Secondary conductor absent. Epigyne (fig. 21B, C) with median field slightly elevated, weakly sclerotized. Lateral lobes curved, narrow, widely separated. Copulatory openings often obstructed by copulatory plugs, occasionally entire median field covered by massive plug. Spermathecae elongate, copulatory ducts irregular, not coiled. Accessory bulbs voluminous.

    Composition:

    Only the type species.

    Axyracrus elegans Simon Figure 21

  • Axyracrus elegans Simon, 1884: 140 (several females, presumed syntypes, from Chile, Cabo de Hornos, Horn, in MHNP, examined), 1887: E23, 1897a: 98. Ramírez, 1995a: 366.

  • Schiapellia gerschmanni Mello Leitão, 1938: 116 (female holotype from Argentina, Isla de los Estados, in MACN-Ar 35425, examined). new synonymy.

  • Amaurobioides boydi Forster, 1970: 175 (female holotype from Chile, Magallanes, Isla Navarino, Pto. Willams, 30 m, 14.I.1964, J.C. Boyd no. 128 F. coll., BPBM 10678, right appendages mounted on separate slide, examined). new synonymy.

  • Synonymy:

    The presumed female syntypes are not labeled as types, but correspond well with the description and collection data. They were compared with the holotype of Schiapellia gerschmanni and Amaurobioides boydi, and with numerous specimens collected in the same area; no relevant differences were found. The epigyne of the holotype of A. boydi is lost, but the figures provided by Forster (1970) are enough for the identification.

    Diagnosis:

    See generic diagnosis.

    Female (Bahía San Antonio MACN-Ar 9807):

    Total length 6.25. Carapace length 2.93, width 1.77, wider on leg II. Length of tibia/metatarsus: I, 1.43/1.27; II, 1.33/1.23; III, 0.99/1.02; IV, 1.60/1.40. Palpal tarsus length 0.69. Chelicerae strong, with two teeth on retromargin, basal one largest. Sternum length 1.40, width 0.90. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–2; metatarsus v 2bas. II, femur = I; tibia v r1–2–2, p 0 or 0–1. Metatarsus = I. III, femur d 1–1–1, p and r d1ap; tibia v 0-p1–2, p 0–1, r d1–1; metatarsus v 2–0–2, p and r 0-d1–1, d 2ap or 0-p1–2. IV, femur d 1–1–1, r d1ap; tibia v p1-p1–2, r d1–1; metatarsus v 2-p1–2, p 1ap or 0-d1–1, r 1ap, d r1ap or 2ap. Abdomen length 3.60, width 1.90, spiracle–epigastrium 1.87, spiracle–spinnerets 0.18. Color: brown with dark brown pattern (fig. 21A); ocular area much darker, legs I and II darker from tibia to tarsus; sternum, endites, and labium dark brown, venter brown. Epigyne: see generic description.

    Male (Bahía San Antonio MACN-Ar 9807):

    Total length 5.45. Carapace length 2.67, width 1.70. Length of tibia/metatarsus: I, 2.20/1.93; II, 2.03/1.83; III, 1.47/1.50; IV, 1.97/1.83. Chelicerae smaller than those of female, vertical. Sternum length 1.37, width 0.80. Spines: leg I, femur d 1–1–1, p d1ap, r d1; tibia v 2–2–2, p 1-d1–1, r 0–1; metatarsus v 2–2–0, p d1-d1-d1, r d1. II, femur = I or p 0-d1-d1. Tibia and metatarsus = I. III, femur d 1–1–1, p and r 0-d1-d1; tibia v 2–2–2 or p1–2–2, p and r 1-d1-1-0; metatarsus v 2–2–2, p d1-d1–1, r 0-d1–1, d 0-p1–2. IV, femur d 1–1–1, p and r d1ap or p 0; tibia v 2–2–2, p 1-0-1-0, r 1-d1-1-0; metatarsus v 2–2–2, p 0-d1–1, r d1–0–1, d 0-p1–2. Abdomen length 2.67, width 1.50, spiracle–epigastrium 1.50, spiracle–spinnerets 0.15. Color as in female but more heavily contrasting, anterior legs much darker from tibia to tarsus. Palp: see generic description.

    Variability:

    The female holotype of Amaurobioides boydi has an additional, small apical tooth on the left cheliceral promargin, retromargin; spines also differ by: tibia II, v r1-r1–2; metatarsus III, IV, v 2–0–2. These differences are interpreted as anomalies and not included in the dataset.

    Natural History:

    According to the labels, many specimens were collected under stones or logs at seashore.

    Distribution:

    Tierra del Fuego and Magallanes.

    Other Material Examined:

    ARGENTINA: Tierra del Fuego: Bahía Aguirre, 13.II.1949, S. Núñez, 1♂ (MACN-Ar 2804); Bahía Buen Suceso, 16–31.I.1986, E. Maury, 1♂ 1♀ 1 immature (MACN-Ar 9806); Península Hardy, Isla Hoste, Bahía Orange, 2–3.I.1963, P.J. Darlington, 1♂ 2♀ (MCZ); Isla de los Estados, I–II.1935, J.A. Dagerre and A. Carcelles, 1♀ (MACN-Ar); XII.1967, A. Bachmann, 1♀ 2 immatures (MACN-Ar); Bahía Crosby, 18.X.1941, 6♀ 1 immature (MACN-Ar); Bahía San Antonio, 6–13.II.1982, J.C. Chébez, 1♂ 1♀ (MACN-Ar 9807); Puerto Perry, I.1982, J.C. Chébez, 1♀ (MACN-Ar); Puerto San Juan, en troncos podridos, A. Carcelles, I.1934, 2♀ (MACN-Ar 35425); Ushuaia, 25.II.1959, J. Vellard, 1 immature (MACN-Ar); 22.II.1959, J. Vellard, 1♀ (MACN-Ar). CHILE: Región XII (Magallanes y Antártica): Magallanes: Guarello, loc. 92, 18.XII.1978, S. Jacquemard, 1♀ 4 immatures (IRSN IG 25934), loc. 91, 1♀ (IRSN IG 25934); Isla Alacalufes, loc. 98, 20.XII.1978, S. Jacquemard, 1♂ (IRSN IG 25934); Isla Hermit, 13–14.III.1961, B. Malkin, 4♀ 4 immatures (CAS); Isla Hermit, St. Martin's Cove, 13–14.III.1961, B. Malkin, 4♀ 4 immatures (AMNH); Isla Hoste, Bahía Orange, 2–3.I.1963, P.J. Darlington, 1♂ 2♀ (MCZ); María Virginia, J. Vellard, 2♀ 1 immature (MACN-Ar); Rusffin, 9.III.1957, J. Vellard, 2 immatures (MACN-Ar); 9.II.1959, J. Vellard, 1♀ (MACN-Ar). No Locality: (presumably from Chile) 1♂ (IRSN IG 23934).

    Aysenia Tullgren Table 9

  • Aysenia Tullgren, 1902: 54 (type species by monotypy Aysenia elongata Tullgren, 1902). Simon, 1903a: 1031. Ramírez, 1995a: 381 (transferred from Clubionidae), 1997: 178.

  • Diagnosis:

    Resembles Aysenoides in having an elongate body, a recurved posterior eye row, and the third leg directed forward, but can be distinguished by having the carapace wider in front (fig. 22A; as in Amaurobioides), irregular spermathecae (instead of spherical), and an unmodified, flattened embolar process (instead of spinelike).

    Description:

    Carapace very narrow, wider anteriorly, posterior eye row recurved, ocular area projecting (fig. 22A). Chelicerae strong, with three teeth on promargin, 2–4 regular teeth on retromargin. Legs generally short, leg III directed forward (figs. 23F, 24A). Tibia I with ventral x-p1-x spine displaced prolaterally (fig. 24E). Male palp with long, pointed RTA. Copulatory bulb variable in size, median apophysis unmodified, paramedian apophysis with two cusps. Primary conductor well developed, with canal. Secondary conductor small, fused to anterior margin of tegulum, without canal. Embolus short, with basal process flattened (figs. 23B, 24B). Epigyne with median field not elevated, copulatory ducts contorted or coiled, spermathecae with irregular lumen (figs. 22D, 23E).

    Natural History:

    Almost nothing is known of the behavior and habitat of these spiders, and most species are rare in collections. The elongate body and the anteriorly oriented third legs suggest that they may live in tubes. Some immatures similar to A. cylindrica were collected by beating vegetation in Malleco, Contulmo.

    Distribution:

    Southern forests of Chile and Argentina.

    Composition:

    A. elongata Tullgren, three species here newly described, and several still undescribed species.

    Aysenia elongata Tullgren Figure 22

  • Aysenia elongata Tullgren, 1902: 54 (female lectotype and female paralectotype [only a carapace, broken] here designated, from Chile, Río Aisén Valley, in NRS, examined). Simon, 1903a: 1031. Merian, 1913: 12. Ramírez, 1995a: 366.

  • Diagnosis:

    Distinguished from other Aysenia species by having large female accessory bulbs.

    Female (lectotype):

    Total length 2.67. Carapace length 1.00, width 0.50, slightly wider on leg III (fig. 22A). Length of tibia/metatarsus: I, 0.42/0.32; II, 0.35/0.28; III, 0.23/0.18; IV, 0.37/0.25. Palpal tarsus length 0.20. Chelicerae strong, with thick anterior bristles, three contiguous teeth on retromargin, promargin not seen, three very small teeth according to Tullgren (1902). Sternum length 0.57, width 0.33. Spines (those on femora and legs III–IV hardly visible, tentative): femora all d 1–1–1. Leg I, tibia v 2–2–0 (x-p1-x displaced prolaterally, fig. 22B); metatarsus v 2bas. II, tibia r1-r1–0 or r1; metatarsus v r1-r1–0, p 1. III, tibia apparently 0 (all scored as missing entries); metatarsus v 1ap and apical group of thick setae. IV, tibia v p1-p1-p1; metatarsus v p1–2-comb (preening comb not in a definite line). Abdomen length 1.61, width 0.55, spiracle close to spinnerets. Color: types badly faded. Abdomen described by Tullgren (1902: 57): “The color seems to have been greenish-yellow on the back with two longitudinal reticulated brown-violet bands confluent at the hind-end. On the sides there are longitudinally arranged scattered short stripes of the same color. The venter is one-colored light yellow-brown.” Epigyne (fig. 22C, D): lateral lobes separate, median field and copulatory openings in epigastric fold, all weakly sclerotized. Spermathecae irregular, contiguous, copulatory ducts contorted. Accessory bulbs voluminous.

    Male:

    Unknown.

    Morphological Remarks:

    The course of the copulatory ducts, although not clearly discernible, suggests some coiling on oblique axes.

    Other Material Examined:

    None.

    Aysenia segestrioides, new species Figure 23

    Type:

    Male holotype from Chile, Región X, Valdivia province, Las Lajas, W La Unión, ca. 40°46′S, 73°42′W, 19–20.XI.1990, L. Peña, deposited in AMNH.

    Etymology:

    The specific name refers to the striking similarity with segestriid spiders, because of the elongate body and the third leg directed forward.

    Diagnosis:

    Easily distinguished by having a huge primary conductor occupying most of the male copulatory bulb, and greatly coiled female copulatory ducts.

    Female (Valdivia, AMNH):

    Total length 8.10. Carapace elevated, cephalic area flat; length 2.93, width 1.57, wider behind chelicerae. Length of tibia/metatarsus: I, 0.90/0.79; II, 0.67/0.77; III, 0.62/0.66; IV, 1.02/0.99. Palpal tarsus length 0.69. Chelicerae very strong, with two teeth on retromargin, fang short, thick (fig. 23G). Sternum length 1.27, width 0.73. Spines: femora I–IV d 1–1–1. Leg I, tibia v r1-r1–2, p v1–1–0 (these are the ventrals displaced); metatarsus v 2–0. II, tibia v r1-r1-r1, p 0-d1; metatarsus v 2bas, p 1. III, tibia v r1ap, p and r d1; metatarsus v 2–0–2, p 0-d1–1, r 1ap, d 2ap. IV, tibia v p1-p1–2; metatarsus v 2-p1–2, p and r 1ap, d r1ap. Scopulae on tarsi I and II, and metatarsus II, denser on prolateral faces. Leg III oriented forward (fig. 23F). Femora I–III narrow, IV very strong. Abdomen length 5.30, width 2.33, spiracle–epigastrium 3.13, spiracle–spinnerets 0.30. Color: carapace dark brown, ocular area almost black. Legs brown, I and II darker. Sternum uniform brown, endites and labium dark brown. Abdomen grayish brown with cream pattern, venter grayish uniform. Epigyne (fig. 23D, E): lateral lobes well sclerotized, separate, elevated above median field. Copulatory ducts very long, coiled.

    Male (holotype):

    Total length 5.00. Carapace length 2.40, width 1.37 (fig. 23A). Length of tibia/metatarsus: I, 1.05/1.00; II, 1.05/0.97; III, 0.73/0.82; IV, 1.05/1.00. Chelicerae smaller than those of female. Sternum length 1.08, width 0.68. Spines as in female, except: leg I, tibia v r1-r1–2 or r1-r1-p1, p d1-0-d1-0; metatarsus v 2–2–0, p 0–1-d1, d p1. II, tibia v r1-r1–2; metatarsus v 2bas, p 0–1-d1. III, tibia p d1 or d1-d1–0. IV, metatarsus p 0–1–1. Abdomen length 2.60, width 1.30, spiracle–epigastrium 1.20, spiracle–spinnerets 0.23. Color as in female. Palp (fig. 23B, C): tibia short, width/length 0.93, RTA sharp, long. Cymbial conductor wide. Copulatory bulb extremely modified, distal region occupying ventral face, primary conductor covering tegulum. Sperm duct with two conspicuous loops at anterior dorsal margin, one at base of secondary conductor. Embolus very long, thin, basal process rounded. Median apophysis apical, hooked, tip pointing basally and retrolaterally. Primary conductor huge, crescent-shaped, projecting at both ends; canal area covered by thin projections. Secondary conductor flattened, partly fused to anterior margin of tegulum, without canal. Paramedian apophysis with three cusps, two heavily sclerotized, close to median apophysis, third one less sclerotized, flattened, close to primary conductor.

    Natural History:

    Unknown. The prolateral displacement of both ventral spines on first tibiae, and of scopulae of first and second legs, suggest further adaptations for living in tubes.

    Distribution:

    Known only from Valdivia province.

    Other Material Examined:

    CHILE: Región X (Los Lagos): Valdivia: Valdivia, no date, collection E. Simon, 1♂ (MHNP 18235); Valdivia, 12.X.1976, E. Krahmer, 1♀ (AMNH).

    Aysenia cylindrica, new species Figure 24

    Type:

    Male holotype from Chile, Región X, Valdivia province, Rincón de Piedra, south of Valdivia, 23–26.II.1979, L. Peña, deposited in AMNH, and female paratype from Valdivia, 1983, deposited in MHNS 837.

    Etymology:

    The specific name refers to the thin, elongate body.

    Diagnosis:

    Resembles A. araucana in body shape and genitalia, but can be distinguished by having relatively short, not coiled, female copulatory ducts and a shorter embolus.

    Female (paratype):

    Total length 4.75. Carapace length 1.43, width 0.75, wider on leg II. Length of tibia/metatarsus: I, 0.63/0.52; II, 0.59/0.51; III, 0.33/0.38; IV, 0.66/0.47. Palpal tarsus length 0.28. Chelicerae (fig. 24D) very strong, with three teeth on retromargin, basal one larger; fang short, thick. Sternum length 0.73, width 0.47. Spines: femora I–IV d 1–1–1. Leg I, tibia v 2–2–2 (the x-p1-x displaced to prolateral, fig. 24E), p d1-0-1-0; metatarsus v 2-r1 (the basal pair advanced), p 1. II, tibia v r1-r1–2, p 0-d1; metatarsus = I. III, tibia v r1ap; metatarsus v 0-r1-0-2 or 0-r1-r1-2, p and r 1ap, d p1ap. IV, tibia v p1-p1–2; metatarsus v 2–2–2, p and r 1ap, d p1ap. Femora I–III narrow, IV strong. Abdomen length 2.80, width 0.47, spiracle–epigastrium 2.00, spiracle–spinnerets 0.24. Color: carapace pale brown, with bright cuticle, cephalic area darker. Legs, femora brownish violet with grayish longitudinal stripes; patella and tarsus I grayish, II pale gray, III cream; leg IV, patella and tarsus cream, tibia brownish violet, distally cream. Abdomen brownish violet, dark, with three whitish dorsal spots, covered by white hairs, two patches of white hairs on posterior end. Leg coxae pale, sternum, labium, endites, and venter dark. Epigyne (fig. 24F, G): lateral lobes widely separated, slightly elevated above median field. Copulatory ducts not coiled.

    Male (holotype, fig. 24A):

    Total length 4.00. Carapace length 1.47, width 0.87, wider at leg II. Length of tibia/metatarsus: I, 0.96/0.74; II, 0.72/0.68; III, 0.44/0.51; IV, 0.77/0.61. Chelicerae slightly smaller than those of female. Sternum length 0.77, width 0.49. Spines as in female, except: leg II, metatarsus v 2–2. III, tibia v 2ap, p d1bas or 0, metatarsus p 0-d1–1. IV, tibia v 0-p1-p1 or 0-(p1-r1)-p1, r d1-d1-d1; metatarsus p 0-d1–1. Abdomen length 2.17, width 0.73, spiracle–epigastrium 1.30, spiracle–spinnerets 0.10. Color (fig. 24A) as in female. Palp (fig. 24B, C): tibia short, width/length 0.93, RTA sharp, long. Cymbium relatively small, globose, cymbial conductor wide. Tegulum basal. Embolus with basal process flat, short, wide. Median apophysis apical, hook-shaped. Primary conductor heavily sclerotized, with sharp apex. Secondary conductor with short apical tip. Paramedian apophysis heavily sclerotized, forming longitudinal shallow ridge.

    Variability:

    Female spines: III, tibia v 0-r1-r1; metatarsus v 2-r1–2 or 2–0–2, p 0-d1–1. IV, tibia v p1-(p1-r1)-2, r 0-d1; metatarsus r d1–0–1, d r1ap. Some specimens with anterior abdominal spot tenuous, divided longitudinally, or absent.

    Natural History:

    Unknown.

    Distribution:

    Known only from a few localities in Valdivia and Río Negro provinces.

    Other Material Examined:

    ARGENTINA: Río Negro: San Carlos de Bariloche, Colonia Suiza, 800 m, 19.IX.1981, Nielsen and Karsholt, 1 immature (ZMK). CHILE: Región X (Los Lagos): Valdivia: Valdivia, 1983, E. Krahmer, 2♀ 1 immature (MHNS 837), 1♀ (MHNS 799), XI–XII.1982.

    Aysenia araucana, new species Figure 25

  • Aysenia sp.: Ramírez, 1995a: 366.

  • Types:

    Male holotype and female paratype from Chile, Región VIII, Biobío province, El Manzano, nr. Contulmo, ca. 38°01′S, 73°20′W, 3–5.III.1986, L. Peña, deposited in AMNH.

    Etymology:

    The specific name refers to the region where this species lives.

    Diagnosis:

    Resembles A. segestrioides and A. cylindrical in body shape and genitalia, but can be distinguished by having the embolus intermediate in length, and by the moderately coiled female copulatory ducts.

    Female (paratype):

    Carapace length 2.27, width 1.17, wider at chelicerae bases and at leg III. Length of tibia/metatarsus: I, 1.00/0.93; II, 0.93/0.83; III, 0.53/0.70; IV, 1.00/0.80. Palpal tarsus length 0.53. Chelicerae very strong, with 3–4 teeth on retromargin, increasing in size to basal. Sternum length 1.03, width 0.72. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–2 (the x-p1-x slightly displaced to prolateral), p 1-d1-1-0; metatarsus v 2-r1–0 (the basal pair advanced), p 1–0. II, femur = I; tibia v r1-r1–2, p d1–1; metatarsus = I. III, femur d 1–1–1, p and r d1ap; tibia v 0-p1–2, p d1–0, d r1bas; metatarsus v 2–0–2, p 0-d1–1, r 1ap, d 0-p1–2. IV, femur d 1–1–1; tibia v p1-p1–2, r d1–1; metatarsus v 2–2–2, p 0-d1–1 or 1ap, r d1–0–1, d 0-p1–2. Leg III oriented forward. Femora I–III narrow, IV strong. Color: dark brown, cephalic area paler, ocular area almost black. Legs dark brown, patellae and dorsum of tibiae, metatarsi, and tarsi III–IV pale gray. Abdomen digested behind epigastrium, tracheae exposed. Epigyne (fig. 25D, E): lateral lobes slightly elevated, arched, copulatory ducts moderately coiled.

    Male (holotype):

    Total length ca. 4.25. Carapace length 1.73, width 1.07. Length of tibia/metatarsus: I, 1.27/1.20; II, 1.10/1.07; III, 0.60/0.83; IV, 1.10/0.90. Chelicerae slightly smaller than those of female, with three teeth on retromargin. Sternum length 0.88, width 0.64. Spines as in female, except: leg I, tibia r 1-0-1-0; metatarsus v 2–2–0, r 1. II, metatarsus v 2–2–0 or 2–2–1, p 0-1-0-1, r 1. III, tibia v 2ap, p d1–1–0. Abdomen (deformed) length ca. 2.50, spiracle–epigastrium 1.40, spiracle–spinnerets 0.27. Color as in female, except legs I and II pale gray from tibiae. Abdomen brownish violet, with whitish dorsal spots, two anteriors, two medians larger, two posteriors covered by whitish hairs. Sternum, labium, and endites dark. Palp (fig. 25A, B): tibia short, width/length 1.08, RTA sharp, long. Cymbial conductor large, triangular. Copulatory bulb greatly modified, distal region occupying ventral face. Tegulum basal. Sperm duct with two conspicuous loops at dorsal anterior margin. Embolus very long, basal process flat, rounded. Median apophysis apical, hook-shaped. Primary conductor with narrow apex. Secondary conductor with short apical tip. Paramedian apophysis with two heavily sclerotized cusps, close to median apophysis.

    Variability:

    The female from Valdivia has the lateral lobes of epigyne more arched.

    Natural History:

    Unknown.

    Distribution:

    Only known from a few localities, in Cordillera Nahuelbuta and X Region.

    Other Material Examined:

    CHILE: Región IX (Araucanía): Cautín: Coicoicura, 5.XII.1992, T. Cekalovic, 1♀ (AMNH). Región X (Los Lagos): Valdivia: Valdivia, E. Krahmer, XI–XII.1982, E. Krahmer, 1♀ 1 immature (MHNS 685). Osorno: Pucatrihue, 4.III.2001, T. Cekalovic, 1♂ (AMNH).

    Aysenoides, new genus Table 10

    Type Species:

    Aysenoides terricola, new species.

    Etymology:

    The generic name is a derivation of the close relative Aysenia, proposed to me by John Kochalka (IBNP). Gender is masculine.

    Diagnosis:

    Resembles Aysenia in having an elongate body, recurved posterior eye row, and the third legs directed forward, but distinguished by having spherical spermathecae (fig. 28F) and a spinelike embolar process (fig. 26B).

    Description:

    Carapace very narrow, posterior eye row recurved, ocular area projecting (fig. 27A). Chelicerae strong, with three teeth on promargin and 2–3 regular teeth on retromargin. Leg III directed forward. Tibia I with ventral x-p1-x spine displaced prolaterally. Male palp with RTA long, pointed, cymbium relatively small. Median apophysis slender, paramedian with two cusps (fig. 26C, D). Primary conductor well developed, with canal (fig. 26A). Secondary conductor fused to tegulum, small, forming wide inconspicuous canal, or absent. Embolus short, with basal process thin, pointed (fig. 26B). Epigyne with median field not elevated, copulatory ducts short, spermathecae spherical.

    Natural History:

    Little is known of the behavior and habitat of these spiders, and most species are rare in collections. As in Aysenia, they may live in some kind of tubes.

    Distribution:

    Southern forests of Chile and Argentina.

    Composition:

    Three species here newly described, and at least three undescribed.

    Aysenoides terricola, new species Figure 27

    Types:

    Male holotype and female paratype from Chile, Región V, Valparaíso province, Cuesta Cavilolén, ca. 31°46′S, 71°19′W, in soil crevices, 6.XI.1988, P. Goloboff, E. Maury, C. Szumik, deposited in MACN-Ar 9808.

    Etymology:

    The specific name refers to the habitat where the types were collected.

    Diagnosis:

    Resembles A. colecole in body shape and genitalia, but can be distinguished by having curved elevations on the epigyne anterior of the spermathecae, and by a longer cusp of the paramedian apophysis on the primary conductor.

    Female (paratype, fig. 27A):

    Total length 4.55. Carapace length 2.03, width 1.08, wider on leg II. Length of tibia/metatarsus: I, 1.02/0.91; II, 0.87/0.81; III, 0.58/0.66; IV, 1.00/0.77. Palpal tarsus length 0.41. Chelicerae with three teeth on retromargin, basal one largest. Sternum length 1.06, width 0.64. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2-2-0-2 (the x-p1-x displaced prolaterally); metatarsus v 2–2–0 (the x-p1-x displaced prolaterally). II, femur = I; tibia v 2-2-0-2, p 0–1; metatarsus v 2-r1–0. III, femur = I; tibia v r1ap or 0-p1-r1, p and r d1–1; metatarsus v 2–0–2 or r1–0–2, p and r 0-d1–1, d 2ap. IV, femur d 1–1–1; tibia v p1-p1–2; metatarsus v 2-p1–2, p and r 1ap, d r1ap. Leg III directed forward. Femora I–III narrow, IV strong. Abdomen (slightly bowed ventrally) length 2.50, width 1.07. Color: carapace grayish, legs pale gray with brownish violet spots, I and II darker. Leg coxae pale, endites, labium, and sternum dark. Abdomen brownish violet with yellow dorsal pattern, venter brownish violet uniform. Epigyne (fig. 27D, E): lateral lobes anteriorly curved, widely separate, closer posteriorly, limiting T-shaped median field. Limit between lateral lobes and median field unclear close to copulatory openings. Spermathecae spherical, copulatory ducts heavily sclerotized.

    Male (holotype):

    Total length 4.00. Carapace length 1.87, width 1.07. Length of tibia/metatarsus: I, 1.63/1.47; II, 1.13/1.17; III, 0.73/0.87; IV, 1.23/1.03. Chelicerae smaller than those of female. Sternum length 0.98, width 0.64. Spines as in female, except: leg I, tibia v 2–2–2, p 1-0-1-0. II, tibia p 1-d1-1-0; metatarsus v 2–2–0, p d1. III, tibia v 0-p1-r1; metatarsus v 2–2–2 or 2-r1–2. IV, femur r d1ap; tibia v p1–2–2 or p1-p1–2, r 1-d1-1-0; metatarsus p 0-d1–1, r d1-d1–1. Abdomen length 2.13, width 0.97, spiracle–epigastrium 1.10, spiracle–spinnerets 0.11. Color as in female. Palp (fig. 27B, C): tibia width/length 0.58, RTA long, sharp. Cymbium relatively small, globose, cymbial conductor wide. Tegulum basal. Sperm duct with two conspicuous loops at dorsal anterior margin, one at base of secondary conductor. Embolus with basal process long, narrow. Median apophysis retrolateral, long, sinuous. Primary conductor with basal portion hyaline, without canal, not fitted to embolus; apical portion long, heavily sclerotized, with long canal, connected to basal portion by thin translucent vertical lamina. Secondary conductor triangular, with acute apex and membranous ventral area. Paramedian apophysis apparently with two cusps, one retrolateral, triangular, flat, another ventral, on primary conductor, long, narrow.

    Variability:

    Male spines: III, tibia v 0–2–2.

    Natural History:

    The types were collected in crevices on dry soil, in a steep ravine (Pablo Goloboff, personal commun.).

    Distribution:

    Central Chile, in Elqui and Valparaíso provinces.

    Other Material Examined:

    CHILE: Región IV (Coquimbo): Elqui: 20 km N La Serena (Rt. 5 km 491), 120 m, 7.X.1992, N. Platnick, P. Goloboff, K. Catley, 1♂ (AMNH). Región V (Valparaíso): Valparaíso: Same data as types, 3 immatures (MACN-Ar); Viña del Mar, I.1979, A. Tobar, 1♂ 3♀ 4 immatures (AMNH).

    Aysenoides colecole, new species Figures 26, 28, 35B

    Types:

    Male holotype (in MHNS) and one male paratype (in MACN-Ar 9809) from Chile, Región X, Chiloé province, Isla de Chiloé, Arroyo Cole Cole, 25 km N Cucao, ca. 200 m, ca. 42°30′S, 54°11′W, 8–11.II.1991, M. Ramírez.

    Etymology:

    The specific name is a noun in apposition taken from the type locality.

    Diagnosis:

    Resembles A. terricola in body shape and genitalia, but can be distinguished by having two epigynal pouches with their openings directed forward, and by a shorter cusp of the paramedian apophysis on the primary conductor.

    Female (Antillanca, fig. 28D):

    Total length 5.30. Carapace length 1.87, width 0.93, wider at leg II. Length of tibia/metatarsus: I, 0.88/0.81; II, 0.74/0.69; III, 0.43/0.51; IV, 0.90/0.69. Palpal tarsus length 0.39. Chelicerae with 3 strong teeth on retromargin, basal one larger. Sternum length 0.97, width 0.58. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–2 (the x-p1-x slightly displaced prolaterally); metatarsus v 2bas. II, femur = I; tibia v r1-r1–2, p 0–1; metatarsus = I. III, femur d d1-d1-d1, p and r d1ap; tibia v r1ap, p d1–1 or 0–1, r 0–1; metatarsus v r1–0–2, p 0-d1–1, r 1ap, d 0-p1–2. IV, femur d 1–1–1; tibia v 0-p1–2, r 0–1; metatarsus v 2-p1–2, p 1ap, r d1–0–1, d 2ap. Leg III directed forward. Femora II and III narrow, IV very strong. Abdomen length 3.40, width 1.27, spiracle–epigastrium 2.10, spiracle–spinnerets 0.16. Color: carapace and legs grayish, posterior legs paler. Abdomen yellow with grayish stripes; dorsum with median stripe on anterior half, two lateral ones; epigastrium grayish, venter with median stripe from epigyne to spinnerets. Dark arch-shaped mark surrounding spinnerets ventrally and laterally. Epigyne (fig. 28E, F): median field sclerotized, longitudinal median ridge close to lateral pouches. Limit between lateral lobes and median field unclear anterior to copulatory openings. Two lateral pouches with openings directed forward, apparently formed by elevated margins of lateral lobes. Spermathecae spherical, accessory bulbs voluminous, contiguous.

    Male (holotype):

    Total length 5.45. Carapace length 2.13, width 1.15. Length of tibia/metatarsus: I, 2.00/1.80; II, 1.30/1.23; III, 0.64/0.82; IV, 1.27/1.13. Chelicerae slightly narrower than those of female, with more evenly spaced teeth. Sternum length 1.09, width 0.61. Spines as in female, except: leg I, p 1-d1-1-0; metatarsus p d1–0 or 0-d1-0-1. II, tibia p 1-d1-1-0, v r1–2–2; metatarsus p d1-d1-0-d1. III, femur r 0-d1-d1; tibia v 0-r1–2 or 0-p1–2, r 1-d1-1-0; metatarsus v 2–0–2, p d1-d1–1, r 0-d1–1. IV, femur r d1ap or 0; tibia v p1–2–2, r 1-d1-1-0. Abdomen length 3.27, width 0.93, spiracle–epigastrium 1.97, spiracle–spinnerets 0.13. Color as in female, with abdominal stripes darker, except median stripe, diffuse. Palp (figs. 26, 28A–C): tibia width/length 0.77, RTA very long, thin. Cymbium relatively small, globose, cymbial conductor wide. Tegulum basal. Sperm duct with two conspicuous loops at dorsal anterior margin, one at base of secondary conductor. Embolus with basal process long, narrow (fig. 26B). Median apophysis retrolateral, long, sinuous. Primary conductor with basal portion hyaline, without canal, not fitted to embolus (fig. 26A); apical portion long, heavily sclerotized, with long canal, connected to basal portion by thin translucent vertical lamina. Secondary conductor triangular, pointed, with ventral membranous area and dorsal patch of denticles (fig. 26C, D). Paramedian apophysis apparently with two cusps, one retrolateral, conical, curved, heavily sclerotized, another ventral, on primary conductor, narrow, small.

    Variability:

    Male spines: III, tibia v r1ap, p and r d1–1, or 1-d1-1-0. IV, tibia v p1-(p1-r1)-2, r d1–1; metatarsus p 0–1–1.

    Natural History:

    The types were collected by beating the endemic “colihue” bamboos (Chusquea spp.).

    Distribution:

    Forests in southern Chile, from Cautín to Chiloé. At the type locality I failed to find them close to sea level.

    Other Material Examined:

    CHILE: Región IX (Araucanía): Cautín: Chacamo, NW Nueva Imperial, W Temuco, 16–24.II.1981, L.E. Peña, 1♂ (AMNH). Región X (Los Lagos): Valdivia: Valdivia, 1984, E. Krahmer, 1♀ (MHNS 849). Osorno: Puyehue Natl. Park: Antillanca rd, 470–720 m, valdivian rainforest, screen-sweeping at dusk, 18–24.XII.1982, A. Newton and M. Thayer, 1♀ (AMNH). Chiloé: Isla de Chiloé: Same data as holotype, 2♂ 2 immatures (MACN-Ar 9811), 1♂ (MACN-Ar 9810 photos MJR 512–513).

    Aysenoides parvus, new species Figures 29, 30

    Types:

    Male holotype and female paratype from Chile, Región IX, Malleco province, Monumento Natural Contulmo, ca. 38°01′S, 73°11′W, 11.XII.1984–13.II.1985, FIT, 350 m, S. and J. Peck, deposited in AMNH. One male paratype from Argentina, Chubut province, Los Alerces Natl. Park, Río Arrayanes, II.1986, M. Ramírez, deposited in MACN-Ar 9812.

    Etymology:

    The specific name refers to the small size.

    Diagnosis:

    Easily distinguished from other Aysenoides by having a relatively small cymbium and copulatory bulb, with the distal sclerites crowded behind the anterior margin of the cymbium, and a flat epigyne without ridges.

    Female (paratype):

    Total length 4.80. Carapace length 1.90, width 1.20, wider on legs II–III. Length of tibia/metatarsus: I, 1.10/0.88; II, 0.94/0.81; III, 0.86/0.63; IV, 1.07/0.92. Palpal tarsus length 0.48. Chelicerae with two teeth on retromargin. Sternum length 1.00, width 0.68. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–2 (the x-p1-x slightly displaced prolaterally); metatarsus v 2bas. II, femur = I; tibia v r1-r1–2, p 0–1; metatarsus = I. III, femur = I; tibia v 0-p1-r1, p d1–1, r 0–1; metatarsus v 2–0–2 and some distal thick setae, p and r 0-d1–1, d 0-p1–2. IV, femur d 1–1–1; tibia v p1-p1–2, r 0–1; metatarsus v r1-r1–2, p 1ap, r d1–0–1, d r1ap. Spines on tibiae and metatarsi I and II long, thick. Leg III directed forward. Abdomen length 2.87, width 1.50, spiracle–epigastrium 1.70, spiracle–spinnerets 0.10. Color: carapace and legs grayish, posterior legs paler. Abdomen pale gray with dorsal pattern grayish violet, venter grayish violet uniform. Epigyne (figs. 29A, 30C–E) flat, lateral lobes separate, median field narrow, slightly rugose, weakly sclerotized. Copulatory ducts short, spermathecae spherical.

    Male (holotype):

    Total length 3.99. Carapace length 1.83, width 1.17. Length of tibia/metatarsus: I, 1.50/1.27; II, 1.27/1.10; III, 0.78/0.88; IV, 1.23/1.10. Chelicerae slightly narower than those of female. Sternum length 0.96, width 0.68. Spines as in female (but weaker), except: leg I, tibia p d1–1. II, tibia v r1–2–2, p d1–1; metatarsus p d1–0. III, tibia v p1–2–2 or p1-p1–2, p and r d1–1. IV, tibia r d1–1; metatarsus p 0-d1–1, d 2ap. Abdomen (slightly bowed) length 2.07, spiracle–epigastrium 1.11, spiracle–spinnerets 0.13. Color as in female, but abdominal pattern more heavily contrasting. Palp (figs. 29B–D, 30A, B): tibia width/length 0.70, RTA long, narrow. Cymbium relatively small, globose, cymbial conductor wide. Sperm duct with pronounced loop at dorsal anterior margin, secondary conductor absent. Embolus with basal process acute but greately reduced (fig. 29D). Other apical sclerites crowded at bulb apex, small, difficult to observe. Median apophysis apical, triangular, hyaline. Primary conductor with basal portion short, with canal; apical portion curved, conical, without canal. Paramedian apophysis apparently with two flattened triangular cusps.

    Variability:

    Female, spines: III, tibia v 0-p1–2, p1–2–2, r1–2–2, or 0-(p1-r1)-2; metatarsus r 1ap. IV tibia v p1-p1–2. Males, III, tibia v 2–2–2; metatarsus v 2-p1–2, p d1–1–1. IV, tibia v 2-p1–2 or 2–2–2; metatarsus v 2–2–2.

    Natural History:

    Mostly unknown, but some specimens were collected by beating foliage. The paratype from Los Alerces was collected on the bark of a Nothofagus sp. tree, at 1.7 m high. They may otherwise live on the ground, because most specimens were collected in pitfall traps.

    Distribution:

    Forests in southern Chile, from Ñuble to Aisén, and adjacent humid mountain passes in Argentina.

    Other Material Examined:

    CHILE: Región VIII (Biobío): Ñuble: 60 km SE Chillán, Termas Road, beech forest, FIT, 1300 m, 7.XII.1984–19.II.1985, S. and J. Peck, 1♂ 1 immature (AMNH); 72 km SE Chillán, Trancas, nr. Termas, FIT, 1700 m, Nothofagus forest, 6.XII.1984–19.II.1985, S. and J. Peck, 2♂ (AMNH). Región IX (Araucanía): Malleco: 17 km W Angol, 800 m, FIT, mixed Nothofagus, 8.XII.1984–16.II.1985, S. and J. Peck, 2♂ (AMNH); 40 km W Angol, Nahuelbuta Natl. Park, FITS, 1200–1500 m, Nothofagus/Araucaria forest, 9.XII.1984–17.II.1985, S. and J. Peck, 6♂ 1♀ (AMNH); Monumento Natural Contulmo, same data as types, 1♂ 1♀ (AMNH), 19–21.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1♀ (MACN-Ar). Cautín: Monte Verde, Cavahue, 31.I.1993, L. Peña, 1♀ (AMNH); 15 km NE Villarrica, Flor del Lago, 500 m log spraying, 10.II.1985, S. and J. Peck, 2♂ (AMNH); 300 m, 2 FITS, Nothofagus forest, 14.XII.1984–10.II.1985, S. and J. Peck, 5♂ 4♀ (AMNH). Región X (Los Lagos): Osorno: 36 km W La Unión, 600 m, 25–28.III.1987, L. Peña, 1♀ (AMNH); Puyehue Natl. Park, 7.7 km NE Termas de Puyehue, 200 m, site 664, window trap, 19–25.XII.1982, A. Newton and M. Thayer, 1♀ (AMNH). Llanquihue: Lago Chapo, 13.5 km E Correntoso, site 656, window trap, 310 m, valdivian rainforest, 16–27.XII.1982, A. Newton and M Thayer, 1♀ (AMNH); 34 km E Puerto Montt, 300 m, FIT, 2nd growth Nothofagus, 24.XII.1984–2.II.1985, S. and J. Peck, 4♂ 1♀ (AMNH). Chiloé: Isla de Chiloé: Lago Tepuhueco, 33 km SW Chonchi, 25 m, 42°49′S, 73°55′W, 26.XI.1994, no. 163, beating vegetation, 1♀, no. 167, fogging fungusy logs, 1 immature, R. Leschen and C. Carlton, 1♀ (AMNH). Palena: 4 km NW Chaitén, 30.I.1985, S. and J. Peck, 1♂ (AMNH); 37 km SE Chaitén, 28.XII.1984–30.I.1985, S. and J. Peck, 6♂ (AMNH). Región XI (Ibáñez del Campo): Aisén: 15 km S La Junta, FIT, 100 m, Nothofagus forest, 30.XII.1984–29.I.1985, S. and J. Peck, 1♂ 1♀ (AMNH).

    Acanthoceto Mello-Leitão Table 11

  • Acanthoceto Mello-Leitão, 1944: 352 (type species by original designation Acanthoceto adelae Mello-Leitão, 1944). Ramírez, 1995a: 381, 1997: 186. Revised by Ramírez, 1997.

  • Diagnosis:

    Distinguished from other Amaurobioidinae by the male abdomen with a terminal projection over the anal tubercle (Ramírez, 1997: fig. 16).

    Description:

    Redescribed by Ramírez (1997). See below for additional data and reinterpretation of male palpal sclerites, principally the reinterpretation of the conductors.

    Distribution:

    South America.

    Composition:

    Seven species, all included here and in Ramírez (1997), and perhaps an additional, undescribed one, very close to A. pichi.

    Acanthoceto pichi Ramírez Figure 33B

  • A. pichi Ramírez, 1997: 181.

  • Description and Diagnosis:

    See Ramírez (1997). Additional data are provided below.

    Female:

    Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2-p1 or 2–2–2, p 0 or 0–1; metatarsus v 2–0. II, femur d 1–1–1, p d1ap; tibia v r1–2-p1, r1–2–2 or 2–2–2, p d1–1; metatarsus v 2–0, p 1. III, femur d 1–1–1, p and r d1ap; patella r d1; tibia v p1-p1–2, p and r d1–1, d r1bas; metatarsus v 2–0–2, p d1–1–1 or 0–1–1, r d1–1–1, d p1–2. IV, femur d 1–1–1, r d1ap; patella r d1; tibia = III; metatarsus v 2–0–2 or 2-p1–2, p and r d1–1–1, d p1–2.

    Male:

    Spines as in female but: III, tibia v p1–2–2; metatarsus v 2–0–2 or 2-p1–2. IV, tibia v p1-p1–2 or 2–2–2. Palp: tibia short, width/length 0.89, RTA sharp, short, sinuous. Cymbial conductor wide. Tegulum small, restricted to bulb base. Embolus long, basal process well developed into translucent blade. Median apophysis apical, hyaline, not articulated, with flattened, blunt tip. Paramedian apophysis heavily sclerotized, with conical cusp (but see note below). Median and paramedian apophyses arising from wide, heavily sclerotized sclerite. Primary conductor with basal portion translucent, elongate, with canal where embolus fits; apical portion conspicuous, sclerotized, with long canal, arising from median part of basal portion. Membranous areas that should separate primary conductor, median and paramedian apophysis, all quite sclerotized. Short, pointed cusp arises from basal part of sperm duct, close to apex of embolar process, may be relict of secondary conductor (fig. 33B).

    Note:

    Several males, including some examined in Ramírez (1997), have two cusps on PMA, instead of one. They probably belong to a separate species, but I have not yet associated this with corresponding variability in females. As both forms share all the autapomorphies of A. pichi, I leave this problem unresolved (and scored the species as being polymorphic for character 68).

    New Records:

    ARGENTINA: Neuquén: Lanín Natl. Park: Lago Lolog, 4 km N San Martín de los Andes, FIT, Nothofagus forest, ca. 950 m, Gentili property, 25.XI–1.XII.1989, S.A. Marshall, 1♀ (AMNH); 8 km N San Martín de los Andes, 1000 m, Malaise trap, 16–22.XI.1997, C. and M. Vardy, 1♂ (BMNH/MACN-Ar). Río Negro: El Bolsón, 28.X.1961, A. Kóvacs, 1♂ (AMNH). CHILE: Región VIII (Biobío): Ñuble: Las Trancas, E Chillán, 29–30.XI.1990, L. Peña, 1♂ 1♀ (AMNH), 1200 m, 24–27.XI.1994, L. Peña, 1♀ (AMNH). Región IX (Araucanía): Malleco: Cordillera Nahuelbuta, 18–20.XII.1993, L. Peña, 1♀ (AMNH). Región X (Los Lagos): Osorno: Puyehue Natl. Park: Aguas Calientes, 480 m, 40°44′S, 72°18′W, 21.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH); Los Derrumbes road, Aguas Calientes, 480 m, 40°44′S, 72°18′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH). Chiloé: Isla de Chiloé: Butalcura, 21.II.1997, T. Cekalovic, 2♀ (AMNH), Coico, 8.II.1994, T. Cekalovic, 2♀ (AMNH), Piroquina, 16.II.1995, T. Cekalovic, 1♀ (AMNH), intersection road to Piroquina, 22.II.1991, T. Cekalovic, 1♀ (AMNH), Puente Trainel, 9.II.1993, T. Cekalovic, 1♀ (AMNH), no date, Skottsberg, 1♀ (NRS).

    Acanthoceto cinerea Group

    Description and Diagnosis:

    See Ramírez (1997). Additional data are provided below.

    Male:

    Palp: tibia long, width/length about 0.45, RTA sharp, short, very thin, cymbium relatively small. Tegulum small, restricted to base of bulb. Embolus with basal process flattened, translucent (fig. 33C). Paramedian apophysis conical, heavily sclerotized. Median apophysis small, apical, hook-shaped, with rounded, weakly sclerotized flat extension close to its base (fig. 32A). Primary conductor with weakly sclerotized basal portion; apical portion long, thin, with long, relatively wide canal, arising from median part of basal portion (canal, restricted to apical portion, was overlooked in Ramírez, 1997). Secondary conductor wide, fused to anterior dorsal margin of tegulum.

    Distribution:

    Southern forests of Chile and Argentina.

    Composition:

    Three very similar species listed below. They have almost identical genitalia, but can be distinguished by the spines on the anterior legs and by the cheliceral teeth.

    Acanthoceto cinerea (Tullgren) Figures 32A, 33C

  • Gayenna cinerea Tullgren, 1901: 244.

  • A. cinereus: Ramírez, 1997: 153.

  • A. cinerea: Platnick, 1997: 684 (emendation of A. cinereus).

  • Description and Diagnosis:

    See Ramírez (1997). Additional data are provided below.

    Spines, male and female (those on femora weak in some specimens): leg I, femur d 1–1–1, p (1-d1)ap; tibia v 2–2–0; metatarsus v 2bas. II, femur d 1–1–1, p 0–1-(1-d1), 0-d1-d1 or d1ap; tibia v 2–2–0, p 0–1; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p and r 0-d1-d1; tibia v 2–2–2, p 1-d1-1-0, r d1–1; metatarsus v 2–2–1(slightly p) and an apical group of thick setae, p and r d1–1–1, d 0-(p1-r1)-2 or 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; tibia v 2–2–2, p and r 1-d1-1-0; metatarsus = III.

    New Records:

    CHILE: ARGENTINA: Tierra del Fuego: road to Glaciar Le Martial, XII.1989, A. González, 1♂ 2♀ 2 immatures (MLP). Región X (Los Lagos): Osorno: 500 m, 26.I.1969, L. Peña, 1♀ (MCZ). Chiloé: Isla de Chiloé: Lago Huillinco, 9.II.1981, T. Cekalovic, 6 immatures (AMNH), Vilupulli, TC-99, 7.II.1981, or Piroquina, TC-101, 10.II.1981, T. Cekalovic, 1♀ (AMNH). Specimen misidentified as A. marina by Ramírez (1997): CHILE: Región VIII: Concepción: Hualqui, 18.XII.1988, R. Vergara, 1♀ (AMNH).

    Acanthoceto ladormida Ramírez

  • A. ladormida Ramírez, 1997: 185.

  • New Records:

    CHILE: Región IV (Coquimbo): Elqui: 9 km S Cruz Grande, beach, 5 m, 11.XI.1993, 29°29′S, 71°19′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1 immature (AMNH).

    Note:

    The apparently disjunct distribution in the central littoral and Cuesta La Dormida is similar to that of Gayennoides molles.

    Acanthoceto marina Ramírez Figure 31B

  • A. marinus Ramírez, 1997: 184 (incorrect gender, here emended).

  • Description and Diagnosis:

    See Ramírez (1997). Additional data are provided below.

    Spines, male and female (those on femora weak in some specimens): leg I, femur d 1–1–1, p (1-d1)ap; tibia v 2–2–0; metatarsus v 2–2–0. II, femur d 1–1–1, p (1-d1)ap or d1ap; tibia v 2–2–0, p 0–1; metatarsus v 2–2–0, p 1–0. III, femur d 1–1–1, p and r d1ap; patella r d1 or 0; tibia v p1–2–2 or p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–2–1(slightly p) and an apical group of thick setae, p and r d1–1–1, d 0-(p1-r1)-2 or 0-p1–2. IV = III; patella r d1; tibia v 2–2–2 or p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus = III.

    New Records:

    CHILE: Región VIII (Biobío): Concepción: Lirquen, 5.VII.1992, T. Cekalovic, 4♂ 1♀ 4 immatures (AMNH).

    Acanthoceto acupicta Group

    Description and Diagnosis:

    See Ramírez (1997). Additional data are provided below.

    Male:

    Palp: tibia long, width/length 0.35–0.60, RTA extremely thin, oblique. Cymbium relatively small. Tegulum small, restricted to basal part of bulb. Embolus with basal process weakly sclerotized, thick, concave, inflated on artificial expansion (fig. 33E). Paramedian apophysis forming concave sclerotized plate, with arch of several conical cusps in variable number, up to seven, most ventral on primary conductor. Median apophysis small. Primary conductor with basal portion massive, with conspicuous canal; apical portion (“secondary conductor” in Ramírez [1997]) small, without canal, contiguous to median apophysis (figs. 32B, 33D). Sperm duct lacking loop on anterior dorsal margin (wrongly interpreted as present in Ramírez, 1997; apical part of bulb extremely modified).

    Distribution:

    South America.

    Composition:

    Two species listed below, and Acanthoceto septentrionalis (Berland). They have mostly identical genitalia, but can be distinguished by the spines on the anterior legs and by the cheliceral teeth.

    Acanthoceto acupicta (Nicolet) Figures 31A, 32B, 33D, E

  • Clubiona acupicta Nicolet, 1849: 420 (female holotype from Chile, San Carlos, in MHNP 4223, not reexamined; probably from Ñuble province, San Carlos, 25 km NE Chillán).

  • A. acupictus: Ramírez, 1997: 186 (incorrect gender, here emended).

  • Description and Diagnosis:

    See Ramírez (1997). Additional data are provided below.

    Spines: female: leg I, femur d 1–1–1, p 2ap; tibia v 0-p1-p1, 0–2-p1 or 0-p1–2 (rarely 0–2–2), short; metatarsus v 2bas. II, femur d 1–1–1, p and r d1ap or r 0; tibia v p1ap, 2ap or 0-r1–2; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p and r d1ap; tibia v 0-p1–2, r 0–1 or d1–1; metatarsus v 2–0–2, p 0–1–1 or 1ap, r 0–1–1, d 2ap. IV, femur d 1–1–1, r d1ap; tibia v p1-p1–2, r d1–1; metatarsus v 2-p1–2 or 2–2–2, p 0–1–1, r d1–1–1, d 2ap. Male: spines as in female, except: I, femur p d1ap.

    New Records:

    ARGENTINA: Entre Ríos: Arroyo Brazo Largo, V.1939, Castillo, 1 immature (MACN-Ar); Rosario (?), illegible, 1♂ (ZMK). Buenos Aires: San Pedro, 2.XI.1991, M. Ramírez, 1♂ (MACN-Ar). CHILE: Región VIII (Biobío): Concepción: Fundo El Manzano, 23.XI.1990, T. Cekalovic, 1♀ (AMNH); 7.XII.1996, T. Cekalovic, 1♂ (AMNH).

    Acanthoceto riogrande Ramírez

  • A. riogrande Ramírez, 1997: 189.

  • Description and Diagnosis:

    See Ramírez (1997). Additional data are provided below.

    Spines: Female: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–2 long; metatarsus v 2bas. II, femur = I; tibia v r1-r1–2, p 0–1; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p and r d1ap; tibia v 0-p1–2, r d1–1; metatarsus v 2–0–2, p and r 0–1–1, d 0-p1–2. IV, femur d 1–1–1, r d1ap; tibia v p1–2–2, p 0–1 or 0, r d1–1; metatarsus v 2-p1–2, p 0–1–1, r d1–1–1, d 0-p1–2.

    Ferrieria Tullgren Table 12

  • Ferrieria Tullgren, 1901: 247 (type species by monotypy Ferrieria echinata Tullgren, 1901). Ramírez, 1995a: 381. Ramírez, 1997: 190 (revision of the genus).

  • Terupis Simon, 1904: 103 (type species by monotypy Terupis bicolor Simon, 1904). First synonymized by Ramírez, 1997: 190.

  • Diagnosis:

    The single known species resembles some Aysenia and Acanthoceto in its small size and recurved posterior eye row, but it can be distinguished by the combination of large spines on the anterior legs, a short, acute RTA, and copulatory ducts coiled along a longitudinal axis. Immatures are very similar to those of Acanthoceto pichi Ramírez, but are distinguished by having larger spines on the anterior legs.

    Description:

    Redescribed by Ramírez (1997). Palp (figs. 33A, 34): tibia short, as long as wide, RTA very short, acute. Cymbium with wide conductor, conspicuous translucent lamina opposed to RTA. Tegulum large, with rectangular basal notch. Sperm duct loops at base of secondary conductor, at anterior ventral margin of tegulum, just before entering embolus, and at embolar base. Embolus with basal process well developed, laminar. Paramedian apophysis well sclerotized, formed by two irregular protuberances, one approximately conic. Median apophysis small, slender, connected to primary conductor by sclerotized stripe. Primary conductor with wide canal where embolus fits. Secondary conductor small, with canal, apex acute. Epigyne with separate lateral lobes, bearing depressions at posterior margins. Copulatory openings close to epigastric furrow; copulatory ducts colied along longitudinal axes, spermathecae separate from each other.

    Composition:

    Only the type species.

    Ferrieria echinata Tullgren Figures 33A, 34, 35C

  • Ferrieria echinata Tullgren, 1901: 247. Redescribed by Ramírez, 1997: 191.

  • Terupis bicolor Simon, 1904: 103. Synonymized by Ramírez, 1997: 191.

  • Diagnosis and Description:

    See Ramírez (1997). Additional data are provided below.

    Variability:

    Female spines: III, tibia v 0-p1-r1. IV, tibia r 1ap; metatarsus v p1–0-comb, p 1ap. Male spines: III, tibia v 0-p1-r1.

    Distribution:

    Southern forests of Chile and Argentina.

    New Records:

    Argentina: Neuquén: Puerto Blest, 7–20.I.2000, L. Lopardo and A. Quaglino, 6 immatures (MACN-Ar). Chile: Región VII (Maule): Talca: Alto de Vilches, elev. 1180 m, 35°36′S, 71°04′W, 14–15.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH). Región VIII (Biobío): Concepción: Hualpén, 7.VI.1996, T. Cekalovic, 1♀ (AMNH). Malleco: Monumento Natural Contulmo, 19–21.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1 immature (MACN-Ar). Arauco: 10 km N Curanilahue, 21.XI.1992, T. Cekalovic, 2♀ (AMNH); Los Morongos, E Los Niches, 600 m, 17–20.XI.1994, L. Peña, 1♂ (AMNH). Biobío: W Ralco, Santa Bárbara, 400 m, 22–23.XI.1994, L. Peña, 1♂ (AMNH). Región X (Los Lagos): Valdivia: Las Lajas, W La Unión, 13–15.I.1991, L. Peña, 1♂ (AMNH), 19–20.XI.1990, L. Peña, 1♀ (AMNH); 37 km SE Panguipulli, 39°45′S, 72°20′W, 300 m, beating vegetation, 14.XI.1994, R. Leschen and C. Carlton no. 097, 1♀ (AMNH); Parque Oncol: intersection Tepual trail with Rio Cruces, 8.I.2001, T. Cekalovic, 1♂ 1♀ (AMNH), Quitaqui trail, 19.I.2001, T. Cekalovic, 1♂ 1♀ (AMNH), Punucapa trail, TC 642, 15.I.2001, T. Cekalovic (AMNH). Osorno: Puyehue Natl. Park: Aguas Calientes, 700 m, 21.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ 1♀ (AMNH), 13–17.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1♂ 1♀ 12 immatures (MACN-Ar), 1♂ 1♀ (MHNS). Llanquihue: Alerce Andino Natl. Park, elev. 100 m, 41°35′S, 72°41′S, 23.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 2♂ 1♀ 1 immature (AMNH).

    Coptoprepes Simon Table 13

  • Coptoprepes Simon, 1884: 130, 136 (type species by monotypy Coptoprepes flavopilosus Simon, 1884), 1887: E24, 1897a: 93, 96, 102. Ramírez, 1995a: 369, 1997: 178.

  • Diagnosis:

    Distinguished from other Amaurobioidinae by the combination of a greatly developed, often bifid median apophysis, and by the apical cymbial notch, retrolateral to the cymbial conductor, where the median apophysis fits (fig. 42A, B).

    Description:

    Color generally dark, with pattern diffuse or absent. Carapace narrowed in front, posterior eye row procurved, ocular area not projecting. Chelicerae relatively small, unmodified, slightly smaller in males, three to five teeth on promargin, a series of small teeth (four to seven) on retromargin. Anterior legs with few spines, lacking prolateral spines on tibia II. Male palpal tibia short, RTA variable, even absent. Cymbium large, with retrolateral apical notch, contiguous to cymbial conductor, where median apophysis fits. Tegulum displaced basally, median apophysis large, apical, often bifid. Primary conductor with long canal. Secondary conductor of variable shape, separate or fused to anterior dorsal margin of tegulum. Paramedian apophysis small or absent. Embolus long, basal process flattened. Epigyne flattened, displaced posteriorly, copulatory openings in or very close to epigastric fold. Copulatory ducts long in species with long embolus, with tortuous course, not colied along any axis; fertilization duct generally separated from posterior epigynal margin.

    Distribution:

    Southern forests of Chile and Argentina.

    Composition:

    C. flavopilosus Simon, three species newly described below, and at least five undescribed species.

    Coptoprepes flavopilosus Simon Figures 35A, 36, 37

  • Coptoprepes flavopilosus Simon, 1884: 137 (male holotype from Chile, Cabo de Hornos, in MHNP 6672, examined), 1887: E25, 1897a: 97, 102 (fulvopilosus, lapsus), 1902: 29. Tullgren, 1901: 245, 260. Merian, 1913: 12. Ramírez, 1995a: 366, 369.

  • Note:

    In the same vial as the holotype there is a female, but Simon only described the male. He reported that the male palpal tibia lacks any apophysis. The RTA is slightly translucent and may have been overlooked.

    Diagnosis:

    Females are asily distinguished from those of other Coptoprepes by the lateral curved ridges on the epigyne; males resemble those of C. nahuelbuta by having a curved tibial apophysis, concave dorsally, but can be distinguished by the much smaller secondary conductor.

    Female (Ushuaia, Castellanos and Gómez, MACN-Ar 9822):

    Total length 5.35. Carapace length 2.23, width 1.50, wider on legs II–III. Length of tibia/metatarsus: I, 1.03/0.63; II, 0.94/1.00; III, 0.97/1.08; IV, 1.30/1.63. Palpal tarsus length 0.64. Chelicerae with 5 teeth on promargin and 5 or 6 denticles on retromargin. Sternum length 1.17, width 0.93. Spines: leg I, femur d 1–1–1, p 2ap; tibia v r1–2–0 or 0–2–0; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v 0-r1-p1; metatarsus v 2bas. III, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–2-comb, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus = III, but d 0–2–2. Dorsal, long, thin, erect bristles on patellae (d 1–0–1) and tibiae (d r1–0–1). Abdomen length 3.17, width 1.70, spiracle–epigastrium 1.50, spiracle–spinnerets 0.22. Color: grayish uniform, darker dorsally (fig. 35A). Epigyne (fig. 36D, E): lateral lobes separate, slightly projecting above epigastric fold. Copulatory ducts irregular. Spermathecae fused to each other.

    Male (Punta Remolino, MACN-Ar 9821):

    Total length 4.40. Carapace length 2.20, width 1.50 (fig. 36A, B). Length of tibia/metatarsus: I, 1.33/1.27; II, 1.17/1.17; III, 0.96/1.23; IV, 1.40/1.70. Chelicerae slightly smaller than those of female, with 6 denticles on retromargin (fig. 36C). Sternum length 1.17, width 0.83. Spines as in female, except: leg I, tibia v 0–2–2. II, tibia v r1-r1–2, p 0–1. III, tibia p and r 1-d1-1-0. IV, tibia v 2–2–2. Abdomen length 2.33, width 1.30, spiracle–epigastrium 1.20, spiracle–spinnerets 0.20. Color as in female. Palp (figs. 36F, G, 37): femur short, laterally compressed. Tibia very short, width/length 1.33, RTA flattened, tip concave dorsally. Cymbium relatively large, with apical retrolateral notch where median apophysis fits, cymbial conductor wide. Tegulum basal. Sperm duct with loop at dorsal anterior margin, close to base of secondary conductor (fig. 37B). Embolus with basal process ample, flattened, rounded. Median apophysis apical, wide, with apical projection long, curved. Primary conductor hyaline, tip simple. Secondary conductor well developed, apex acute, partially separate from tegulum by ventral membranous area. Paramedian apophysis with one triangular cusp, flattened, close to base of median apophysis; rounded, ventral protuberance, may also be part of paramedian apophysis.

    Variability:

    Female spines: I, tibia v p1–2-p1. II, tibia v 0-r1–2. III, tibia v p1–2–2.

    Natural History:

    This species constructs retreats under logs in very humid localities.

    Distribution:

    Forests in southern Argentina and Chile, from Osorno to Tierra del Fuego.

    Other Material Examined:

    ARGENTINA: Chubut: Los Alerces Natl. Park: Río Arrayanes, II.1985, M. Ramírez, 2♀ (MACN-Ar). Santa Cruz: Ventisquero Moreno, 18–24.I.1971, J. Vellard, 1♀ (MACN-Ar). Tierra del Fuego: Bahía Buen Suceso, 16–31.I.1986, E. Maury, 1 immature (MACN-Ar); Cabo de Hornos, same vial as holotype, 1♀ (MHNP); Isla de los Estados, Arroyo Goffre, 20.X.1971, Menéndez, 1♀ (MACN-Ar); Laguna Negra, XII.1989, A. González, 1 immature (MLP); Punta Remolino 24, 24.II.1959, J. Vellard, 1♂ (MACN-Ar 9821); Ushuaia, 1–14.XII.1932, Castellanos and Gómez, 1♀ (MACN-Ar 9822); Río Pipo, XII.1989, A. González, 1♀ (MLP); Ushuaia, no date, J. Vellard, 1♀ (MACN-Ar); Valle Carbajal, 17.II.1961, B. Malkin, 1♀ (AMNH). CHILE: Región X (Los Lagos): Osorno: Puyehue Natl. Park, Los Derrumbes, 18.I.1989, M. Ramírez, 1♀ (MACN-Ar, photo MJR 53), Los Mallines, 40°46′0″S, 72°17′00″W, 700 m, bog, pitfall 514T1, 2.XII.2000–2.I.2001, 1♀, 12.XII.2000–2.I.2001, pitfall 514T1, 1♂ 2♀, J. Miller, I. Agnarsson, Alvarez, J. Coddington, G. Hormiga, (USNM), Antillanca, 40°46′30″S, 72°11′30″W, 1050–1350 m, alpine meadow, 2.XII.2000, J. Miller, I. Agnarsson, Alvarez, J. Coddington, G. Hormiga, pitfall 57T1, 1♀, pitfall 511T1, 1♂, pitfall 512T2, 1♀ (USNM); Antillanca road, 40°46′30″S, 72°12′00″W, 1150 m, Nothofagus pumilio forest, 24.XII.2000, J. Miller, I. Agnarsson, Alvarez, J. Coddington, G. Hormiga, pitfall 511T1, 1♂ (USNM). Llanquihue: Lago Chapo, 13.5 km E Correntoso, site 656, window trap, 310 m, valdivian rainforest, 16–27.XII.1982, A. Newton and M Thayer, 1♂ (AMNH); 35 km W Río Negro, 240 m, disturbed forest, 24.I.1986, N. Platnick and T. Schuh, 1♀ 1 immature (AMNH). Chiloé: Arroyo Cole Cole, 25 km N Cucao, 8–11.II.1991, M. Ramírez, 1♀ (MACN-Ar). Región XI (Ibáñez del Campo): Aisén: Laguna San Rafael, II.1957, N. Codoceo, 2♀ (MACN-Ar); Río Simpson Natl. Park, N margin, 17.II.1991, M. Ramírez, 4♀ 2 immatures (MACN-Ar); 85–89 km S Puerto Puyuguapi, 220–270 m, burned forest, 19.I.1986, N. Platnick, P. Goloboff, T. Schuh, 1♀ (AMNH); 102 km S Puerto Puyuguapi, 220 m, wet forest, 19.I.1986, N. Platnick, P. Goloboff, T. Schuh, 4♀ (MACN-Ar). Región XII (Magallanes y Antártica): Ultima Esperanza: Torres del Paine Natl. Park: near Refugio Chileno, 50°56′45″S, 72°55′0″W, 400–600 m, 8–9.XII.2000, J. Miller, I. Agnarsson, 1♂, 1♀, 2♀ (USNM); near Refugio Pudeto, 51°3′45″S, 72°58′45″W, 100 m, 7.XII.2000, pitfall in scrub, J. Miller, I. Agnarsson, 1♂, pitfall 51T1, 1♂ (USNM); Laguna Parrillar Natl. Res., 53°24′15″S, 71°15′45″W, 1–10.XII.2000, 350 m, J. Miller, I. Agnarsson, forest Berlese, 1♀, dry grass near Chorio Hermoso, pitfall 55T2, 1♀, scrub in bog, pitfall 54T1, 1♀, scrub, grass in bog, pitfal 54T2, 4♀ 2 immatures, Chorio Hermoso, flood plain, 55T1, 2♀ (USNM). Magallanes: Cabo de Hornos, 1844–84, P. Hahn, 1♀ (MHNP); Camerón, 14–17.XI.1960, L. Peña, 2♀ (MCZ); Estancia La Vicuña, SE Camerón, 1–6.XII.1960, L. Peña, 1♀ (MCZ); Isla Nueva, 4.II.1896, O. Nordenskjöld, 2♀ (NRS); Isla Picton (1896, O. Nordenskjöld), 1♀ (NRS); Puerto Bridges, 9.I.1893, Michelsen, 3♀ (ZMH); Puerto Toro, Isla Navarino, 19.XII.1992, Michelsen, 1 immature (ZMH); Río Rubens, 1956, J. Vellard, 1♂ 2♀ (MACN-Ar); Rubens, 13.XII.1960, L. Peña, 1♀ (MCZ); Rusffin, SE Cameron, 17–20.XI.1960, L. Peña, 1♀ (MCZ); Tres Vientos, Puerto Arturo, 53°34′S, 73°23′W, 25–28.XI.1960, L. Peña, 1♀ (MCZ). Mistaken Locality: Argentina, La Pampa, Santa Rosa, V.1962, Aravena, 1♀ (MACN-Ar), a tentative transcription made by M.E. Galiano of an illegible label, is here considered inaccurate.

    Coptoprepes nahuelbuta, new species Figure 38

    Types:

    Female holotype and male paratype from Chile, Región IX, Malleco province, Nahuelbuta Natl. Park, FITS, 1200–1500 m, Nothofagus/Araucaria forest, ca. 38°S, 73°W, 9.XII.1984–17.II.1985, S. and J. Peck, deposited in AMNH.

    Etymology:

    The specific name is a noun in apposition, referring to the area where this species lives.

    Diagnosis:

    Distinguished from other Coptoprepes by having a projecting secondary conductor, articulated and heavily sclerotized, and an epigynal median field hidden in the epigastric furrow.

    Female (holotype):

    Total length 5.30. Carapace length 2.03, width 1.43, wider on leg II. Length of tibia/metatarsus: I, 0.92/0.84; II, 0.83/0.80; III, 0.72/0.89; IV, 1.10/1.07. Palpal tarsus length 0.57. Chelicerae with five teeth on promargin, seven denticles on retromargin. Sternum length 1.08, width 0.87. Metatarsi III and IV with preening comb. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2–0; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v r1-r1–2; metatarsus = I. III, femur d 1–1–1, p 0-d1-d1 or d1ap, r d1ap or 0; patella r 1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–0-comb, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus v 2–2-comb, p and r d1–1–1, d 0–2–2. Dorsal bristles as in C. flavopilosus. Abdomen (with cuticle partially detached) length 3.55. Color: holotype quite faded; other specimens with carapace, legs, sternum, mouthparts brown, abdomen dark grayish, dorsum paler, cardiac area dark. Epigyne (fig. 38F–H) displaced posteriorly, in ventral view only visible as elevation of epigastrium. Lateral lobes separate in posterior view. Copulatory ducts contorted, asymmetrical, spermathecae contiguous, lumen small.

    Male (paratype):

    Total length 4.12. Carapace length 1.90, width 1.37. Length of tibia/metatarsus: I, 1.23/1.09; II, 1.04/0.99; III, 0.81/1.00; IV, 1.17/1.47. Chelicerae smaller than those of female. Sternum length 1.06, width 0.78. Spines as in female, except: leg I, tibia v 2–2–2. II, tibia p 0–1. Abdomen (badly preserved) length ca. 2.00. Color as in female. Palp (fig. 38A–D): femur short; tibia very short, width/length 1.39, RTA long, flattened, distally bent dorsally. Cymbium relatively large, with deep apical retrolateral notch where median apophysis and secondary conductor fit; cymbial conductor wide. Tegulum basal. Sperm duct with pronounced loop at dorsal anterior margin, between secondary conductor and median apophysis. Embolus with basal process ample, flattened, rounded. Median apophysis apical, sinuous, forming right angle, apex very thin. Primary conductor with deep canal, apical portion projecting as straight, heavily sclerotized prong, where canal ends. Secondary conductor conspicuous, heavily sclerotized, almost totally surrounded by membranous area (fig. 38B). Paramedian apophysis absent, or represented only by a longitudinal ridge close to base of median apophysis; rounded, ventral protuberance, may also be part of paramedian apophysis.

    Variability:

    Females spines: I, tibia v 2–2-p1 or 2–2–2. II, tibia v r1-r1-p1. IV, metatarsus d 0-p1–2.

    Natural History:

    Unknown. All specimens were collected in pitfall traps or in leaf litter.

    Distribution:

    Forests in southern Chile, in Malleco and Aisén provinces, probably also in intermediate localities.

    Other Material Examined:

    CHILE: Región IX (Araucanía): Malleco: Same data as types, 2♂ 3♀ (AMNH); 17 km W Angol, 800 m, FIT, mixed Nothofagus, 8.XII.1984–16.II.1985, S. and J. Peck, 1♂ (AMNH); Monumento Natural Contulmo, 11.XII.1984–13.II.1985, S. and J. Peck, 1♂ 1♀ (AMNH); Nahuelbuta Natl. Park, 1250 m, 19.XI.1981, N. Platnick and T. Schuh, mossy forest litter, Nothofagus, Araucaria, 1♀ (AMNH). Región XI (Ibáñez del Campo): Aisén: 102 km S Puerto Puyuguapi, 220–270 m, burned forest, 19.I.1986, N. Platnick, P. Goloboff, T. Schuh, 2♀ (AMNH).

    Coptoprepes campanensis, new species Figures 3941

    Types:

    Male holotype from Chile, Región V, Quillota province, Palmas de Ocoa, La Campana Natl. Park, unburned site, 23.VIII.1985, pitfall 1, R. Calderón; female paratype from the same locality, trap 5, 22.VI.1984, R. Calderón, deposited in AMNH.

    Etymology:

    The specific name refers to the type locality, where this species seems to be very common.

    Diagnosis:

    Distinguished from other Coptoprepes by having a huge embolus and primary conductor, and long, contorted female copulatory ducts.

    Female (paratype):

    Total length 3.17. Carapace length 1.40, width 1.02, wider between legs II and III. Length of tibia/metatarsus: I, 0.72/0.61; II, 0.66/0.59; III, 0.57/0.63; IV, 0.98/1.07. Palpal tarsus length 0.41. Chelicerae with three teeth on promargin, four on retromargin. Sternum length 0.81, width 0.61. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2–2–0; metatarsus v 2bas. II = I. III, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–0–1 (and apical preening comb, fig. 40), p and r d1–1–1, d 0-p1–2. IV, femur = III; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus v 2–2–1, p and r d1–1–1, d 0-p1–2. Abdomen length 1.87, width 1.07, spiracle–epigastrium 1.00, spiracle–spinnerets 0.16. Color: grayish, spotted in dark brown. Sternum and mouthparts dark brown, leg coxae paler. Abdomen with dark dorsum, darker on cardiac area, with three pairs of paler spots covered by whitish hairs, spots in two posterior pairs closer to each other. Venter grayish, slightly paler than dorsum. Epigyne (fig. 41C, D) partially displaced posteriorly, lateral lobes widely separate, copulatory openings near epigastric fold. Copulatory ducts long, convoluted, fused with those of opposite side though in part of their length. Spermathecae contiguous, lumen small, fertilization ducts separate from posterior border.

    Male (holotype):

    Total length 3.00. Carapace length 1.43, width 0.97. Length of tibia/metatarsus: I, 1.11/1.01; II, 0.96/0.93; III, 0.73/0.90; IV, 1.11/1.37. Chelicerae slightly narrower than those of female. Sternum length 0.83, width 0.67. Spines as in female, except: leg I, tibia v 2–2–2. II, tibia v r1-r1–2. IV, patella r 1. Abdomen length 1.60, width 0.91, spiracle–epigastrium 0.80, spiracle–spinnerets 0.18. Color as in female, but darker carapace, with paler central strip. Palp (figs. 39, 41A, B): femur short; tibia short, as wide as long, RTA absent. Cymbium relatively large, flattened, with deep apical retrolateral notch where median apophysis and secondary conductor fit; cymbial conductor wide. Tegulum basal. Sperm duct with pronounced loop at base of secondary conductor. Embolus with basal process ample, flattened, rounded. Median apophysis apical, large, heavily sclerotized, bifid, apical projection forming right angle (fig. 39B–D). Primary conductor huge, canal deep (fig. 39A–C), tip simple. Secondary conductor triangular (fig. 39B). Additional projection of tegulum near tip of primary conductor. Paramedian apophysis absent.

    Natural History:

    Unknown. Most specimens were collected in pitfall traps.

    Distribution:

    Relict forests in central Chile.

    Other Material Examined:

    CHILE: Región IV (Coquimbo): Elqui: 34 km SE La Serena, 29°58′S, 70°57′W, 300 m, riparian litter, 23.X.1994, R. Leschen, C. Carlton no. 002, 1♂ (AMNH). Limarí: Many vials from type locality and Fray Jorge Natl. Park, pitfall traps, R. Calderón (AMNH). Región V (Valparaíso): Petorca: Cuesta El Melón, 520 m, chaparral, 10.I.1985, N. Platnick and O. Francke, 3♀ (AMNH); Los Molles, Rt. 5, km 188, elev. 10 m, 9.XI.1993, 32°14′S, 71°30′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 2♀ (AMNH). Valparaíso: Quintero, 9.III.19??, 1♀ (MACN-Ar). Región Metropolitana (Santiago): Santiago: Batuco, 1964, 3♀ 1 immature (MHNS); Quebrada La Plata, fundo La Rinconada de Maipú, 8.X.1958–10.V.1960, W. Noodt, many specimens (MHNS); Valle del Río Mapocho between El Arrayán and Farellones (Barber traps), 15.X.1958–8.VI.1960, W. Noodt, many specimens (MHNS). Cordillera: Río Clarillo Natl. Res., 940 m, 26.XI.1993, 33°44′S, 70°28′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH); Río Clarillo, flight intersection trap, 3.XI–7.XII.1989, S.A. Marshall, 1♀ (AMNH).

    Coptoprepes valdiviensis, new species Figure 42

    Types:

    Male holotype from Chile, Región X, Llanquihue province, Lago Chapo, 13.5 km E Correntoso, carrion trap (squid), site 656, 310 m, valdivian rain forest, ca. 41°33′S, 71°57′W, 16.27.XII.1982, A. Newton and M Thayer; female paratype with same data, window trap, deposited in AMNH.

    Etymology:

    The specific name refers to the valdivian forest where this species lives.

    Diagnosis:

    Distinguished from other Coptoprepes by having a sharp and slender tibial apophysis and a rectangular median epigynal field.

    Female (paratype):

    Total length 4.66. Carapace length 1.77, width 1.17, wider at leg III. Length of tibia/metatarsus: I, 1.07/0.82; II, 0.89/0.81; III, 0.74/0.90; IV, 1.17/1.37. Palpal tarsus length 0.59. Chelicerae with three teeth on promargin, four on retromargin. Sternum length 1.17, width 0.75. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2-p1; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v r1-r1-p1; metatarsus = I. III, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r 1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–0-comb, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus = III, but v 2–2-comb. Dorsal bristles as in C. flavopilosus, but tibiae d 1ap. Abdomen length 2.83, width 1.60, spiracle–epigastrium 2.25, spiracle–spinnerets 0.17. Color: grayish, abdomen slightly darker on cardiac area and several chevrons extending to spinnerets. Epigyne (fig. 42D, E) partially displaced posteriorly, lateral lobes separate, copulatory openings close to epigastric fold. Spermathecae contiguous, lumen small, copulatory ducts fused with those of opposite side though in part of their length.

    Male (holotype):

    Total length 4.12. Carapace length 1.90, width 1.27. Length of tibia/metatarsus: I, 1.33/1.10; II, 1.02/0.93; III, 0.86/1.00; IV, 1.23/1.47. Chelicerae slightly narrower than those of female. Sternum length 1.00, width 0.81. Spines as in female, except: leg I, tibia v 2–2–2. II, tibia p 0–1. Abdomen length 2.13, width 1.17, spiracle–epigastrium 0.37, spiracle–spinnerets 0.16. Color as in female. Palp (fig. 42A–C): femur short, flattened. Tibia short, width/length 1.06, RTA sharp, slender. Cymbium relatively large, with apical retrolateral notch where median apophysis fits; retrolateral margin basally extended into thin lamina; cymbial conductor wide. Tegulum basal/retrolateral. Sperm duct with pronounced loop at dorsal anterior margin, close to secondary conductor (fig. 42C). Embolus with basal process ample, flattened, rounded. Median apophysis apical, well developed, bifid, long, narrow projection forming straight angle, another projection long, thick, directed apically. Primary conductor with canal. Secondary conductor triangular with acute apex, with ventral membranous area, fused to anterior dorsal margin of tegulum. Paramedian apophysis reduced to sclerotized piece at base of median apophysis, slightly elevated in rounded mound.

    Variability:

    Male spines: III, tibia v p1–2–2, p and r 1-d1-1-0.

    Natural History:

    Unknown. Several specimens were collected in leaf litter.

    Distribution:

    Forests of southern Chile and Argentina, in Valdivia, Cautín, Osorno and Neuquén provinces, plus one isolated record in Ultima Esperanza province, Magallanes.

    Other Material Examined:

    ARGENTINA: Neuquén: Lanín Natl. Park: Lago Lolog, nr. San Martín de los Andes, pans nr. stream, ca. 900 m, 23–30.XI.1989, S. Marshall 1♂ (AMNH); Lago Lolog, 4 km N San Martín de los Andes, FIT, Nothofagus forest, ca. 950 m, Gentili property, 23.XI–1.XII.1989, S.A. Marshall, 1♂ (AMNH); San Martín de los Andes, Gentili Cabin, pans and FIT along streambed, 18–21.XI.1989, S. Marshall, 1♂ (AMNH), forest and meadow, 18–21.XI.1989, S.A. Marshall, 1♂ 1♀ (AMNH); 4 km N San Martín de los Andes, FIT, Nothofagus forest, ca. 950 m, Gentili property, 25.XI–1.XII.1989, S.A. Marshall, 1♂ (AMNH); Villa La Angostura, pans nr. Laguna Verge, 26–28.XI.1989, S.A. Marshal, 1♀ (AMNH). CHILE: Región IX (Araucanía): Cautín: Volcán Villarrica, 1250 m, site 653, window trap, Nothofagus domb.-pumilio forest with Chusquea, 15–29.XII.1982, A. Newton and M. Thayer, 1♀ (AMNH); site 654, Nothofagus domb.-pumilio forest with Drimys, A. Newton and M. Thayer, 1♀ (AMNH). Región X (Los Lagos): Osorno: Puyehue Natl. Park: Antillanca rd, 965 m, trap site 658, window trap, Nothofagus pumilio forest, 18–25.XII.1982, A. Newton and M. Thayer, 1♂ (AMNH); 40°46′30″S, 72°12′00″W, 1000 m, 12.XII.2000–2.I.2001, Nothofagus pumilio forest, J. Miller, I. Agnarsson, Alvarez, J. Coddington, G. Hormiga, 2♂ 2♀ (USNM). Llanquihue: 10–14 km E Correntoso, 305 m, disturbed forest, 3.II.1985, N. Platnick and O. Francke, 1♂ 2 immatures (AMNH); NE Puerto Montt, 22–28.XII.1985, L. Peña, 1♂ (AMNH). Región XII (Magallanes y Antártica): Ultima Esperanza: Torres del Paine Natl. Park: near Refugio Pudeto, 51°3′45″S, 72°58′45″W, 100 m, 7.XII.2000, pitfall in scrub, J. Miller, I. Agnarsson, 1♀ (USNM).

    Gamakia, new genus Table 14

    Type Species:

    Gamakia hirsuta, new species.

    Etymology:

    Gamakia is the supreme being of the Septentrional Tehuelches. Gender is feminine.

    Diagnosis:

    The single known species can be distinguished from other Amaurobioidini by having a relatively long male palpal tibia with relictual apophysis, the chelicerae and palpal femora covered with thick, long setae, and the epigyne with a double depression anterior of copulatory openings, which are filled by a copulatory plug in mated females.

    Description:

    Carapace narrowed in front, posterior eye row slightly procurved, ocular area not projecting. Chelicerae unmodified, slightly smaller in males, with three teeth on promargin, a series of six or seven small denticles on retromargin; males have long, thick setae on anterior face of chelicerae. Anterior legs with unmodified spines, more spinose in males. Male palp (figs. 44B–D, 45C–E, 46) with relatively long tibia, wider distally, RTA reduced to small dorsal/retrolateral peak (fig. 46), femur with long, thick setae on ventral/retrolateral face (fig. 45D). Cymbium relatively small. Copulatory bulb: median apophysis long, slender, sinuous. Primary conductor short, with canal. Secondary conductor long, with marked canal, fused to apical dorsal margin of tegulum, additional projection prolateral to secondary conductor. Paramedian apophysis well developed, with multiple cusps (fig. 44C). Embolus with simple, flattened basal process. Membranous area between paramedian apophysis and tegulum lined with thin projections (fig. 44C, D). Epigyne (figs. 44A, 45A, B) with two hemispheric depressions anterior to copulatory openings, filled by copulatory plug in mated females. Spermathecae irregular, copulatory ducts not coiled.

    Composition:

    Only the type species.

    Gamakia hirsuta, new species Figures 4346

    Types:

    Male holotype and female paratype from Chile, Región V (Valparaíso), Petorca province, Los Molles, Rt. 5, km 188, elev. 10 m, 9.XI.1993, 32°14′S, 71°30′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, deposited in AMNH.

    Etymology:

    The specific name refers to the thick hairs on chelicerae and male palp.

    Diagnosis:

    See generic diagnosis.

    Female (paratype):

    Total length 4.92. Carapace length 2.10, width 1.47, wider on legs II–III. Length of tibia/metatarsus: I, 1.20/1.00; II, 1.13/0.83; III, 0.90/1.05; IV, 1.28/1.28. Palpal tarsus length 0.52. Chelicerae with 6 teeth on retromargin, slightly decreasing in size to basal. Sternum length 1.05, width 0.80. Spines: leg I, femur d 1–1–1, p d2 ap, r d1ap; tibia v 2–2–2 or 2–2-p1, p d1–1; metatarsus v 2bas, p d1 or 0. II, femur d 1–1–1, p 0-d1-d2 or d2ap, r 0-d1-d1; tibia v 2–2–2, p d1–1, r 0–1; metatarsus p v 2bas, 0-d1-0-1, r d1–0. III, femur d 1–1–1, p and r 0-d1-d1; patella r d1; tibia v p1-p1–2, p and r 1-d1-1-0, d r1bas; metatarsus v 2-p1–2, p and r d1-d1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia = III or v p1–2–2; metatarsus = III or v 2–2–2. Abdomen length 3.00, width 1.90, spiracle–epigastrium 1.67, spiracle–spinnerets 0.22. Color: carapace gray, ocular area darker, eyes bordered black. Legs, femora pale gray with darker spots, darker from patella to tarsus. Sternum gray, darker on margins. Endites brown, labium dark brown. Abdomen cream with gray pattern, venter with irregular gray spots. Epigyne: see generic description.

    Male (holotype):

    Total length 4.79. Carapace length 2.27, width 1.50. Length of tibia/metatarsus: I, 1.87/1.69; II, 1.70/1.60; III, 1.57/1.48; IV, 1.67/2.03. Chelicerae long, narrow, and vertical, anterior face with short, thick hairs; promargin with 7 teeth grouped at base, forming short, sinuous line, basals slightly smaller; fang long, sinuous. Endites with external angle prominent. Sternum length 1.10, width 0.87. Spines as in female, except: leg I, femur r 0-d1-d1; tibia v 2–2–2, p and r 1-d1-1-0; metatarsus v 2-0-2-0, p and r d1–0, d 2ap. II, femur p and r 0-d1–2; tibia = I; metatarsus =I, but p d1-d1-0-0. III, tibia v p1–2–2; metatarsus v 2–0–2. IV, femur r 0-d1-d1; tibia v 2–2–2; metatarsus v 2–2–2. Abdomen length 2.50, width 1.43, spiracle–epigastrium 1.37, spiracle–spinnerets 0.25. Color as in female but darker in general, abdomen with denser pattern. Palp: see generic description.

    Variability:

    The abdominal pattern is extremely variable (fig. 43), as is the length of the male chelicerae, which vary from similar to much larger than those of female. Spines: tibia III, IV, v p1-p1–2. Metatarsus III, v 2–0–2.

    Natural History:

    This species builds retreats on foliage of forest and chaparrals.

    Distribution:

    Southern and central Chile, from Elqui to Chiloé provinces.

    Other Material Examined:

    CHILE: Región IV (Coquimbo): Elqui: 9 km S Cruz Grande, beach, 5 m, 11.XI.1993, 29°29′S, 71°19′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ 3 immatures (AMNH); 34 km SE La Serena, 29°58′S, 70°57′W, 300 m, riparian litter, 23.X.1994, R. Leschen, C. Carlton no. 002, 1♀ (AMNH). Choapa: El Bato (chacra en montaña), E Illapel, 10.X.1985, L. Peña, 4♀ (AMNH); 22 mi N Los Vilos, 13.XII.1950, E.I. Schlinger, 2♀ (CAS); Ñagué, 10 km N Los Vilos, Rt. 5, km 236, elev. 40 m, 31°50′S, 71°31′W, 13.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 2♀ (AMNH), 1♀ (MACN-Ar); Hacienda Illapel, 900–1800 m, 5.XI.1954, L.E. Peña, 1♀ (IG 20275 IRSN); Pichidangui, 32°08′S, 71°32′W, 12.VIII.1966, E. Schlinger, 3♂ (CAS). Región V (Valparaíso): Petorca: Same data as types, 1♀ (AMNH), 1♀ (MACN-Ar); E La Ligua, relict forest, 27.IX.1980, L. Peña, 1♀ (AMNH); Zapallar, 27.XI.1950, Ross and Michelbacher, 4♀ (CAS). Quillota: Cuesta El Melón, nr. La Calera, 15.XI.1985, L. Peña, 1♂ 1♀ (AMNH); Cuesta La Dormida, N Tiltil, 800–1300 m, 13–18.XI.1982, L. Peña, 1♀ (AMNH); La Campana Natl. Park, 29.XII.1973, J. Solervicens, 1♂ (UC). Valparaíso: 10 mi N Concón, 16.XII.1950, Ross and Michelbacher, 1♂ 2♀ (CAS); Central coast (no specific locality), 31.X.1982, no collector, 11♂ 9♀ (AMNH); Tunquén, S Quintay, X.1982, M. Pino, 2♀ (MHNS 619); Valparaíso, 15.VIII.1961, J. Kothmann, 1♀ (AMNH). San Felipe de Aconcagua: Cachagua, 14.II.1980, L. Peña, 2♀ (AMNH); SW Catapilco, 30.IX.1964, L. Peña, 1♂ (MCZ); Los Hornos, 20 km E Huaquén, 2–4.XII.1986, L. Peña, 1♀ (AMNH); Pullalli, coastal town, 16.XII.1980, L. Peña, 1♂ 4♀ (AMNH). San Antonio: Quebrada de Córdoba, 1–4.XI.1985, L. Peña, 1♂ 4♀ 1 immature (AMNH), 15–20.II.1979, L. Peña, 2♀ (AMNH); 5 km E El Tabo, 6.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ 1 immature (AMNH). Región Metropolitana (Santiago): Santiago: Bucalemi, San Antonio, 23–24.X.1994, L. Peña, 2♂ 2♀ (AMNH); Pirque, 20.XI.1982, L. Peña, 1♀ (AMNH); Quilicura, VIII.1979, L. Peña, 2♂ 11♀ (AMNH); Pilay, 800 m, 23–25.XI.1981, L.E. Peña, 1♀ (AMNH). Región VII (Maule): Curicó: Las Tablas, E Curicó, II.1985, L. Peña, 11♀ 28 immatures (AMNH); Los Queñes (Ladera Sur), 27.IV.1980, J. Soler, R. Calderón, 2♀ (UC). Talca: Alto de Vilches, 18.25.X.1964, L. Peña, 1♂ (MCZ), 17–24.X.1964, L. Peña, 1♀ (MCZ). Gil de Vilches, 7–8.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH). Cauquenes: Los Ruiles Natl. Park, 25.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♂ 3 immatures (AMNH); Tregualemu, 300–500 m, 10.XII.1953, L. Peña, 2♀ (IRSN IG 19.736); 500 m, 7.XI.1993, L. Peña, 2♂ 1♀ (AMNH); 520 m, 6–7.XI.1993, L. Peña and A. Ugarte, 2♂ (AMNH); 4–5.XI.1995, L. Peña, 1♀ (AMNH). Linares: Bullileo, Parral, 5–8.XII.1990, L. Peña, 2♂ 5♀ (AMNH); Fundo Malcho, Andes in Parral, 11–20.XI.1964, L. Peña, 5♂ 10♀ 3 immatures (MCZ). Región VIII (Biobío): Ñuble: Cobquecura, 8–9.XI.1993, L. Peña, 1♀ (AMNH), 12–14.II.1959, L. Peña, 1♀ 1 immature (IRSN IG 19.736). Concepción: road Chome-Ramuntcho, 8.XI.1996, T. Cekalovic, 4♂ (AMNH); Cerro Caracol, Concepción, elev. 200 m, 36°51′S, 73°02′W, 17.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH); Escuadrón, 10.IV.1988, 1♂ 1♀, elev. 5 m, 36°57′S, 73°09′W, 18.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 2♂ 1 immature (AMNH); Escuadrón, 10.IV.1988, 2♀ 1 immature, 3.IX.1988, 1♀, T. Cekalovic (AMNH); Estero Nonguén, 10.II.1996, T. Cekalovic, 1♀ (AMNH); 7.XI.1992, 2♀, Fundo El Manzano, 8.XI.1992, 1♂ 1♀, 12.X.1996, 1♀, 23.IX.1996, 1♂ 2♀, T. Cekalovic (AMNH); Fundo El Venado, 6.I.1996, T. Cekalovic, 1♀ (AMNH); Hualpén, 3.IV.1988, 1♀, 7.VI.1996, 4♀ 3 immatures, T. Cekalovic (AMNH), 11.I.1989, M. Ramírez, 4♀ (MACN-Ar, photos MJR 68–70); Hualqui, 4.XI.1989, T. Cekalovic, 1♀ (AMNH); Las Escaleras, 18.XI.1989, 2♂, 24.XI.1989, 1♀, 6.I.1991, 1♀, T. Cekalovic (AMNH); Patagual, 29.XI.1993, T. Cekalovic, TC-369, 1♂ 1♀ (AMNH); Periquillo, 22.XI.1992, 2♂ 2♀, 7.X.1994, 1♂, 6.XI.1994, 1♀, 8.XII.1994, 3♀, 4.I.1997, 1♀, 22.III.1997, 1♀, T. Cekalovic (AMNH); Laguna Chica de San Pedro, 5.XII.1994, T. Cekalovic, 1♀ (AMNH); Tomé, 8.X.1983, 1♂, 1.I.1992, 1♀, 10.I.1992, 1♀, T. Cekalovic (AMNH). Biobío: Caledonia, E Mulchen, 700–900 m, 6.15.II.1981, L. E, Peña, 1♀ (AMNH); Los Morongos, E Los Niches, 600 m, 17–20.XI.1994, L. Peña, 3♂ 2♀ 1 immature (AMNH); W Ralco, Santa Bárbara, 400 m, 22–23.XI.1994, L. Peña, 2♀ (AMNH). Región IX (Araucanía): Malleco: 18 km W Angol, 13.II.1991, M. Ramírez, N. Platnick, P. Goloboff, 2♀ (AMNH); Fundo María Ester, 15 km W de Victoria, 14.I.1989, M. Ramírez, 1♀ 2 immatures (MACN-Ar, photo MJR 121, 122); Monumento Natural Contulmo, 12.I.1989, M. Ramírez, 4♀ (MACN-Ar, photo MJR 92); 340 m, 38°01′S, 73°11′W, 18.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 2♂ (AMNH). Cautín: Cerro Ñielol, Temuco, I.1989, M. Ramírez, 2♀ (MACN-Ar); Chacamo, NW Nueva Imperial, W Temuco, 16–24.II.1981, L.E. Peña, 1♀ (AMNH); 15–30 km S Cherquenco, 26.II.1989, L. Peña, 1♀ (AMNH); Pucón at Lago Villarrica, 39°16′S, 71°58′W, 14.XII.1988, V. and B. Roth, 1♂ (CAS); Fundo La Selva, W Temuco, NW Nueva Imperial, 700 m, 9–12.XII.1981, L.E. Peña, 1♂ (AMNH); Villarrica, 250 m, crest of river gorge, 3–4.III.1965, H. Levi, 1 immature (MCZ); 14 km N Villarrica, elev. 250 m, 39°10′S, 72°12′W, 20.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH), 30 km NE Villarrica, 1–30.I.1965, L. Peña, 3♂ 4♀ 1 immature (MCZ); NE Villarrica, 16–31.XII.1964, L. Peña, 2♀ 1 immature (MCZ). Región X (Los Lagos): Valdivia: Nancul, Fundo “El Lingue”, 8.II.1993, T. Cekalovic, 2♀ (AMNH); Valdivia, 1984, E. Krahmer, 1♀ (MHNS 848). Osorno: Fundo Campolindo, Casablanca, 9.XII.1971, R. Calderón, 2♀ (UC); Puyehue Natl. Park: Aguas Calientes, 480 m, 40°44′S, 72°18′W, 21.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH, photo MJR 1416), Pucatrihue, I–III.1968, L. Peña, 1♂ (MCZ); Río Bueno, no. 13, L. Peña, 5♀ (IRSN IG 19.736). Chiloé: Arroyo Cole Cole, 25 km N Cucao, 8–11.II.1991, M. Ramírez, 1♀ (MACN-Ar); Chepu, 21.II.1997, T. Cekalovic, 1♀ (AMNH). Mistaken Locality: Prov. Santiago, Malleco, XI.1979, L.E. Peña, 8♂ 8♀ (AMNH) (see Ramírez, 1995b: 83).

    Negayan, new genus Table 15

  • Gayenna strigosa group”: Ramírez, 1997: 179.

  • Type Species:

    Gayenna tridentata Simon, 1886.

    Etymology:

    The generic name is an anagram of Gayenna; gender is feminine.

    Diagnosis:

    Easily distinguished from all other Amaurobioidinae by the characteristic shape of the retrolateral tibial apophysis, which is long, thick, and sinuous at the tip (fig. 48F, G). The female genitalia resemble those of Ferrieria echinata and Acanthoceto pichi in having coiled copulatory ducts (fig. 49C–F).

    Description:

    Carapace narrowed in front, posterior eye row straight, ocular area not projecting. Chelicerae unmodified, slightly smaller in males, three teeth on promargin, two or three teeth, or series of small denticles, on retromargin. Anterior legs with unmodified spines. Male palp with RTA long, thick, distally sinuous. Cymbium short, wide, cymbial conductor small, with elevated borders. Shallow notch prolateral to cymbial conductor fits tip of primary conductor. Tegulum with extended anterior ventral border, in the same plane with primary conductor. Median apophysis reduced, triangular or absent. Primary conductor well developed, basal portion massive, with long canal where embolus fits (fig. 47D); apical portion heavily sclerotized, thick, curved, canal ending in acute tip (fig. 47A, B). Secondary conductor absent. Paramedian apophysis heavily sclerotized (fig. 47C). Embolus long, basal process simple, flattened (fig. 47D). Epigyne: median field elevated in some species, copulatory openings close to epigastric furrow. Copulatory ducts coiled 360°. Spermathecae contiguous.

    Natural History:

    Most species live on the ground, under stones, or in leaf litter. Some species are common under stones at beaches in lakes, or at the sides of mountain streams.

    Distribution:

    Most species from Argentina and Chile, in a wide variety of habitats and climates. Northern limit of distribution seems to be in Peru.

    Composition:

    In addition to the species detailed below: Gayenna excepta Tullgren, 1901 (female holotype in NRS, examined, new combination), Gayenna exigua Mello-Leitão, 1940 (male holotype in MLP 14404, examined, new combination), Tomopisthes lebruni Simon, 1886 (two females syntypes in MHNP 7733, examined, new combination). Also several undescribed species. The genus is being revised by L. Lopardo (in prep.).

    Negayan tridentata (Simon), new combination Figure 48

  • Gayenna tridentata Simon, 1886: 570 (female lectotype and one immature paralectotype here designated, from Argentina, Santa Cruz, in MHNP 2189, examined), 1897a: 91 (tridens, lapsus).

  • Diagnosis:

    Distinguished from other Negayan by having the posterior borders of epigynal lateral lobes relatively close to each other, converging over the median field; males resemble those of N. coccinea in having a bifid conductor, but the tibial apophysis is more sinuous at the tip. Both sexes commonly have three teeth on the cheliceral retromargin.

    Female (Lago Roca, MACN-Ar 9820):

    Total length 4.12. Carapace length 1.65, width 1.15, wider on leg III. AME smaller than ALE (fig. 48C). Length of tibia/metatarsus: I, 0.83/0.70; II, 0.77/0.67; III, 0.70/0.80; IV, 1.08/1.27. Palpal tarsus length 0.43. Chelicerae with two teeth on retromargin (variable, fig. 48B). Sternum length 0.92, width 0.72. Spines (quite strong): leg I, femur d 1–1–1, p 2ap; tibia v 2–2–0; metatarsus v 2bas. II, femur d 1–1–1, p d1 ap, r 0 or d1ap; tibia v 2–2–0 or 2–2-p1, p 0–1; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p and r d1ap; patella r d1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–0–2, p and r d1–1–1, d 0-p1–2. IV, femur = III; patella r d1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus = III, but v 2–2–2. Dorsal, long, thin, erect bristles on patellae (d 1–0–1) and tibiae (d r1–0–1). Abdomen length 2.30, width 1.33, spiracle–epigastrium 1.12, spiracle–spinnerets 0.17. Color: carapace pale brown with two longitudinal brown bands (fig. 48A). Sternum pale with lateral brown bands. Abdomen pale with dorsal brown pattern, three ventral lines of specks. Epigyne (fig. 48D, E): lateral lobes slightly projecting over epigastric furrow, projections concave, close to each other. Copulatory ducts coiled about 360°, spermathecae with contorted lumen.

    Male (Lago Roca, MACN-Ar 9820):

    Total length 3.59. Carapace length 1.67, width 1.30, relatively wider than in female, but narrowed anteriorly. Length of tibia/metatarsus: I, 1.18/1.02; II, 1.03/0.95; III, 0.90/0.97; IV, 1.30/1.48. Chelicerae slightly smaller than those of female, with three teeth on retromargin, apical one smaller. Sternum length 0.97, width 0.77. Spines as in female, except: leg I, tibia v 2–2–2, p 0–1. II, tibia = I. III, tibia v p1–2–2, p and r 1-d1-1-0. Abdomen length 2.03, width 1.17, spiracle–epigastrium 1.12, spiracle–spinnerets 0.50. Color as in female. Palp (fig. 48F–H): RTA with narrow, sinuous tip. Median apophysis triangular, hyaline. Apical portion of primary conductor bifid, with thick curved tip where canal ends, and ventral prolateral elongate projection (fig. 48H). Paramedian apophysis with two triangular, flattened cusps, retrolateral cusp longer, curved, heavily sclerotized, ventral cusp close to base of primary conductor, weakly sclerotized.

    Variability:

    Two or most commonly three teeth on cheliceral retromargin.

    Natural History:

    Unknown.

    Distribution:

    Patagonian forests in Argentina, from Río Negro to Tierra del Fuego provinces. Probably also in Chile.

    Other Material Examined:

    ARGENTINA: Río Negro: San Carlos de Bariloche, IV.62, Havrylenko, 1♀ (MACN-Ar); Caviahue, 12–15.II.1968, E. Maury and N. Müller, 1♀ (MACN-Ar); Lago Pellegrini, 853, 16.II.1972, L. Herman, 2♀ (AMNH). Chubut: Cholila, 25.VIII.1962, A. Kovács, 3♂ 8♀ (AMNH); Epuyén, 5.VIII.1966, A. Kovács, 2♂ 1♀ (AMNH); Esquel, road to La Hoya, 42°54′S, 71°19′W, 16.XI.1988, V.D. Roth, 3♀ (CAS). Santa Cruz: Los Glaciares Natl. Park, II.1975, E. Fernández, 1♂ 1 immature (MACN-Ar). Tierra del Fuego: Lago Roca, Nothofagus antarctica forest, 27.I.1971, J. Vellard, 2♂ 3♀ 4 immatures (MACN-Ar 9820).

    Negayan paduana (Karsch), new combination Figure 49, 50

  • Clubiona paduana Karsch, 1880: 379 (female holotype from Chile, Punta Arenas, Magallanes, Exp. Gazelle, in ZMB 2622, examined).

  • Tomopisthes magellanicus Simon, 1887: E32 (female holotype from Chile, Punta Arenas, in MHNP 6685, examined), 1895: 168, 1896a: 142, 1897a: 91, 1902: 32. new synonymy.

  • Gayenna strigosa Tullgren, 1901: 237, 259 (male lectotype Chile, Magallanes, Gente Grande, and several females paralectotypes, plus one female Sanogasta maculosa, and one immature cf. Amaurobiidae, paralectotypes designated by Platnick, 1977: 196, in NRS, examined). Schiapelli and Gerschman, 1974: 91. Ramírez, 1997: 178. new synonymy.

  • Tomopisthes strigosus: Simon, 1902: 34.

  • Philisca colulata Hogg, 1911: 42 (female holotype from Islas Malvinas, in BMNH, not examined). Synonymized with G. strigosa by Platnick, 1977: 196.

  • Gayenna magellanica: Merian, 1913: 13.

  • Synonymy:

    The holotypes or lectotypes of the species here synonymized were compared, together with extensive collections from the same areas; no relevant differences were found.

    Diagnosis:

    Distinguished from other species of Negayan by having the posterior elevations of epigynal lateral lobes just below the median field (fig. 49C), and by lacking a bifurcate primary conductor, or a ventral cusp on tegulum, at the end of the tegular notch. Both sexes commonly have two teeth on the cheliceral retromargin.

    Female (holotype):

    Total length 6.92. Carapace length 3.17, width 2.17, wider on legs II–III. Length of tibia/metatarsus: I, 1.60/1.45; II, 1.55/1.37; III, 1.37/1.53; IV, 2.04/2.47. Chelicerae with two teeth on retromargin. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v p1–2–2; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p 0-d1-d1, r d1ap; patella 0; tibia v p1–2–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–2-(2+thick setae), p and r d1–1–1, d 0–2–2. IV, femur d 1–1–1, p and r d1ap; patella 0; tibia v 2–2–2, p and r 1-d1-1-0, d r1bas; metatarsus = III. Abdomen length 4.10, width 2.25. Spiracle–epigastrium ca. 1.90, spiracle–spinnerets ca. 0.60. Color: holotype faded. Female from Magallanes, Estancia Gazy Harbour (AMNH): carapace brown, darker toward ocular area, margins dark. Legs brown, with some dark spots, more distinct on legs III and IV. Endites, labium, and sternum dark brown. Abdomen cream with dark dorsal pattern brownish violet, venter cream, with large violet patch anterior of tracheal spiracle, prolonged to epigastric furrow in three lines of dots; epigastrium dark between pulmonary plates and epigyne. Epigyne (fig. 49C–F): median field wide, elevated, lateral lobes elevated posteriorly.

    Male (Magallanes, Estancia Gazy Harbour):

    Total length 5.05. Carapace length 2.67, width 1.70. Length of tibia/metatarsus: I, 1.87/1.77; II, 1.80/1.70; III, 1.07/1.63; IV, 1.97/2.33. Chelicerae slightly smaller than those of female. Sternum length 1.47, width 1.00. Spines as in female, except: leg I, femur r d1; tibia p 1-d1-1-0, r 1-0-1-0; metatarsus p 1. II, femur r 0-d1-d1; tibia and metatarsus = I. III, femur r 0-d1-d1; tibia v 2–2–2, p and r 1-d1-1-0; metatarsus d 0-p1–2. IV, femur p 0-d1-d1. Abdomen length 2.67, width 1.40, spiracle–epigastrium 1.28, spiracle–spinnerets 0.25. Color as in female. Palp (fig. 50): RTA with apical constriction. Median apophysis hyaline, hook-shaped. Sperm duct wide, suddenly narrowed before reaching embolar base (fig. 50D). Apical portion of primary conductor with only one apical cusp, longitudinally striated, where canal ends. Paramedian apophysis with two cusps, retrolateral cusp heavily sclerotized, short, curved, ventral cusp close to base of primary conductor, rounded, flat, weakly sclerotized.

    Distribution:

    Argentina and Chile, from Neuquén and Aisén provinces, respectively, to Tierra del Fuego and Islas Malvinas.

    Natural History:

    This species builds retreats under stones or barks of fallen logs, in areas of rigorous climate.

    Variability:

    The ocular area is relatively narrower in larger specimens (as in fig. 49B). The abdominal pattern is quite variable, from almost uniform dark to slightly contrasting (fig. 49A, B). Some males have only one tooth on cheliceral retromargin. Female spines: I, tibia I v 2–2–0; metatarsus p 0. III, tibia v 2–2–2, r 1-d1-1-0; metatarsus d 0-p1–2.

    Other Material Examined:

    ARGENTINA: Neuquén: San Martín de los Andes, Cerro Chapelco, 1700 m, II.1961, M.E. Galiano, 1♂ (MACN-Ar). Río Negro: El Bolsón, Cerro Piltriquitrón, 3–4.II.1985, II.1986, M. Ramírez, 1♀ (MACN-Ar). Chubut: La Hoya, 800–1350 m, 24.II.1979, Misión Científica Danesa, 2♂ 1 immature (ZMK). Santa Cruz: Calafate, II.1963, E. Maury, 1♂ 1♀ (MACN-Ar); 30 km S Caleta Olivia, 6.V.1974, M. Rumboll, 1♀ (MACN-Ar); 35 km S Caleta Olivia, 6.V.1974, M. Rumboll, 1♀ (MACN-Ar); Lago Belgrano, 14.II.1973, M. Rumboll, 1♀ (MACN-Ar); Lago Argentino, “estepa”, nr. Calafate, 15–30.I.1971, Vellard, 1♀ 2 immatures (MACN-Ar); Lago Argentino, III.1900, Excursión Silvestri, 2♀ (MACN-Ar); Lago Cóndor, Río Turbio, 28.I.1976, M. Rumboll, 1♀ (MACN-Ar); Lago Frías, no date, E. Maury, 1♀ (MACN-Ar); Puerto Bandera, II.1963, Margheritis and Rizzo, 1♂ (MACN-Ar); Puerto Coyle, 10 m, 26.XI.1966, M. Irwin and E. Schlinger, 1♂ (CAS); Río Gallegos, Cerro Aymond, 26.III.1949, Núñez and Partridge, 1♀ (MACN-Ar); San Julián, XI.1973, M. Rumboll, 1♀ (MACN-Ar); Ventisquero Moreno, 18–24.I.1971, J. Vellard, 1♂ 2♀ (MACN-Ar). Tierra del Fuego: No specific locality, “lote 39”, M.P. Gómez, 1933, 1♀ (MACN-Ar 32052); Estancia Viamonte, Auricosta, 2 m, 23.I.1979, Misión Científica Danesa, 1♀ (ZMK); Lapataia, II.1963, E. Maury, 1♂ 2♀ (MACN-Ar); Río Grande, 26–31.X.1973, M. Rumboll, 1♀ (MACN-Ar); Ushuaia, 8.II.1961, B. Malkin, 8♂ 18♀ 1 immature (AMNH); 8–26.II.1961, B. Malkin, 7♂ 12♀ (AMNH); Río Pipo, XII.1989, A. González, 1♀ (MLP); Valle Carbajal, 17.II.1961, B. Malkin, 1♀ (AMNH). Islas Malvinas: All specimens reported by Schiapelli and Gerschman (1974: 91), except 2♀ from Puerto Stanley, belong to other Negayan species not included here. CHILE: Región XI (Ibáñez del Campo): Aisén: Balmaceda, 17–22.I.1961, L. Peña, 1♀ (IRSN IG 23.077). Región XII (Magallanes y Antártica): Ultima Esperanza: Torres del Paine Natl. Park, 150 m, 10.II.1985, N. Platnick and O. Francke, 1♀ (AMNH), near Refugio Chileno, 50°56′45″S, 72°55′0″W, 400–600 m, 8–9.XII.2000, J. Miller, I. Agnarsson, 1♀ (USNM); Puerto Natales, Seno Ultima Esperanza, 51°41′45″S, 72°31′30″W, 50 m, 7.XII.2000, pasture, shoreline, J. Miller, I. Agnarsson, 2♀ (USNM). Magallanes: Cabo Negro, 29.I.1976, T. Cekalovic, 1♀ (AMNH); Estancia La Vicuña, 1956, J. Vellard, 1♀ (MACN-Ar); Cañadón Bombalot, 29.I.1976, T. Cekalovic, 3♀ (AMNH); El Sombrero, 1956, J. Vellard, 1♂ 1♀ 1 immature (MACN-Ar); Espora, 52°29′S, 69°28′W, 29.XI.1966, M. Irwin and E. Schlinger, 1♀ (CAS); Estancia Gazy Harbour, 10.II.1990, T. Cekalovic, 8♂ 5♀ 1 immature (AMNH); Estancia Virgen de Lourdes (Sector Dinamarquero), 6.II.1990, 2♂ 6♀, 8.II.1990, 2♂ 5♀, T. Cekalovic (AMNH); Estancia Virgen de Lourdes, Gazy Harbour, 23.III.1991, T. Cekalovic, 1♀ 1 immature (AMNH); Gallegos Chico, 8.II.1990, T. Cekalovic, 2♂ 2♀ (AMNH), 10.II.1990, S. Cekalovic, 6♂ 2♀ 4 immatures (AMNH); Gobernador Philippi, 29.I.1976, T. Cekalovic, 2♀ (AMNH); Isla Riesco, Posomby, 31.I.1976, T. Cekalovic, 1♂ 4♀ (AMNH); Isla Riesco, Vaquería, 31.I.1976, T. Cekalovic, 2♂ 2♀ 2 immatures (AMNH); Laguna Amarga, 21.IV.1962, T. Cekalovic, 1♀ (AMNH); Laguna Figueroa, 18, 28.I.1976, T. Cekalovic, 1♀ (AMNH); Manantiales, 1956, J. Vellard, 3♂ 1♀ (MACN-Ar); Península Brunswick, Barranco Amarillo, 27.I.1976, T. Cekalovic, 1♀ (AMNH); Península Brunswick, Tres Brazos, 9.III.1961, T. Cekalovic, 1♀ (UC), 28.II.1981, T. Cekalovic, 1♀ (AMNH); Río Chico, Estancia Brazo Norte (Cueva Fall), 12.II.1972, V. Pérez, 1♀ (UC); Río Verde, 31.I.1976, T. Cekalovic, 1♀ (AMNH), 29.VIII.1976, T. Cekalovic, 3♀ (MCZ), 3♀ (AMNH); 15.2 km NE San Gregorio (dunes), 5 m, 36°17′S, 71°49′W, 27.XI.1966, E. Schlinger and M. Irwin, 3♂ (CAS); Silla del Diablo, 28.I.1976, T. Cekalovic, Tres Vientos, Puerto Arturo, 53°34′S, 73°23′W, 25–28.XI.1960, L. Peña, 1♀ (MCZ), 1♀ (AMNH); Aserradero Yendegaia, no. 5, 14.II.1957, J. Vellard, 1♂ 7♀ 6 immatures (MACN-Ar).

    Negayan coccinea (Mello-Leitão), new combination Figures 47, 51

  • Axyracrus coccineus Mello-Leitão, 1943b: 115 (female holotype from Argentina, Córdoba province, Bajo Grande, I.1940, M. Birabén, in MLP 15800, examined).

  • Diagnosis:

    Distinguished from other Negayan by the small size, the absence of median apophysis, and by having the copulatory openings in the epigastric furrow. Males resemble those of N. tridentata in the bifid conductor, but the tibial apophysis is less sinuous at the tip.

    Female (Cabana):

    Total length 3.13. Carapace length 1.27, width 0.92, wider on legs II–III. Length of tibia/metatarsus: I, 0.73/0.60; II, 0.65/0.57; III, 0.57/0.60; IV, 0.90/1.02. Palpal tarsus length 0.37. Chelicerae with four teeth on promargin, four on retromargin, basal one slightly larger. Sternum length 0.70, width 0.58. Spines (those of tibiae and metatarsi I, II relatively long, thick): leg I, femur d 1–1–1, p d1ap; tibia v 2–2–0; metatarsus v 2bas. II, femur = I; tibia v 2–2–0, p 0–1; metatarsus v 2bas, p 0–1. III, femur d 1–1–1, p and r d1ap; patella r 0; tibia v p1-p1–2, p and r d1–1, d r1bas; metatarsus v 2–0-comb, p and r d1-d1–1 or 0-d1–1, d 0-p1–2. IV, femur = III; patella r d1; tibia = III, but v p1–2–2; metatarsus = III, but v 2-p1-comb. Abdomen length 1.93, width 1.20, spiracle–epigastrium 0.93, spiracle–spinnerets 0.13. Color: carapace pale brown with dark brown band at each side, margins almost black. Legs pale brown with dark brown spots. Chelicerae with basal two-thirds dark, basal pale patch, endites and labium dark, sternum with central pale patch. Abdomen dorsally brownish violet, with small pale dots, two pale anterior patches, some chevrons at posterior end, fusing on pale spot above spinnerets. Venter with dark rectangle anterior of spiracle, prolonged to epigastric furrow in median band plus some asymmetrical spots; epigastrium dark between pulmonary plates and epigyne. Epigyne (fig. 51C, D): median field triangular, slightly elevated, copulatory ducts complex, difficult to observe, colied along oblique axes; copulatory openings not seen.

    Male (Cabana):

    Total length 2.67. Carapace length 1.20, width 0.88. Length of tibia/metatarsus: I, 0.80/0.68; II, 0.72/0.67; III, 0.58/0.67; IV, 0.95/1.07. Chelicerae smaller than those of female, with five teeth on retromargin. Sternum length 0.68, width 0.58. Spines as in female. Abdomen length 1.50, width 0.80, spiracle–epigastrium 0.63, spiracle–spinnerets 0.13. Color as in female, but wider dark bands on carapace, median pale band narrowing between eyes and thoracic groove. Anterior pale patches on abdomen extending in lateral oblique lines of pale dots. Palp (figs. 47, 51A, B): RTA with flattened tip, only slightly widened. Tegulum with ventral conical protuberance at end of tegular notch (fig. 47A). Median apophysis absent. Apical portion of primary conductor bifid, with thick tip where canal ends, and ventral/prolateral elongate projection (fig. 47C). Paramedian apophysis with two cusps connected by ridge, retrolateral cusp longer, curved, heavily sclerotized, ventral cusp close to base of primary conductor, weakly sclerotized. Base of primary conductor projecting anteriorly in globose lobe, weakly sclerotized, close to secondary conductor (fig. 47C).

    Natural History:

    The specimens from La Cumbre were collected in leaf litter in a mesophytic forest.

    Distribution:

    Central Argentina.

    Other Material Examined:

    ARGENTINA: Córdoba: Cabana, VII.1950, M. Birabén, 6♂ 13♀ 21 immatures (MLP); Calamuchita, II.1959, J.M. Viana, 1♀ (MACN-Ar); La Cumbre, 8.XI.1990, M. Ramírez, 3♀ 1 immature (MACN-Ar). Entre Ríos: Arroyo Gualeyán and Ruta Nac. 14, nr. Gualeguaychú, 13.XI.1982, P. Goloboff, 1♂ (MACN-Ar). Buenos Aires: Atucha, 27.VI.1991, M. Ramírez, 1♂ (MACN-Ar); Ciudad de Buenos Aires, 17.V.1981, M. Ramírez, 1♂ 1♀ 1 immature (MACN-Ar), 1.VIII.1981, F. Miranda, M. Ramírez, 1♂ (MACN-Ar); Lima, 19.IX.1981, P. Goloboff, M. Ramírez, 1♂ (MACN-Ar); Tigre, 19.VIII.1951, J.M. Viana, 1♂ (MACN-Ar). La Pampa: Lihuel Calel Natl. Park, XI.1975, A. Toth and E. Maury, 1♀ (MACN-Ar).

    Selknamia, new genus Table 16

    Type Species:

    Selknamia minima, new species.

    Etymology:

    The generic name refers to the Selk'nam, the now extinct inhabitants of Tierra del Fuego and Magallanes. Gender is feminine.

    Diagnosis:

    The single known species of the genus is distinguished from other Amaurobioidini by having an elongate, relatively thick tibial apophysis, and by the combination of spherical spermathecae and copulatory openings on the epigastric furrow.

    Description:

    Carapace narrowed in front, posterior eye row straight, ocular area not projecting. Chelicerae unmodified, slightly smaller in males, with three teeth on retromargin, growing larger to basal. Anterior leg spines unmodified. Male palp (figs. 52, 53A, B) with elongate tibia, RTA long, straight, thick. Cymbium very small, cymbial conductor subterminal. Copulatory bulb small. Tegulum with anterior margin at same level with primary conductor. Median apophysis simple, close to apical portion of primary conductor. Primary conductor with basal portion well developed, massive, with canal where embolus fits; apical portion with two tips, heavily sclerotized, retrolateral tip bearing canal, prolateral tip flattened, triangular. Secondary conductor absent. Paramedian apophysis not well developed, with two shallow cusps. Embolus without basal process. Epigyne (fig. 53C–F) slightly projecting posteriorly, copulatory openings in the epigastric furrow. Spermathecae spherical, copulatory ducts short.

    Composition:

    Only the type species.

    Selknamia minima, new species Figures 52, 53

    Types:

    Female holotype from Argentina, Tierra del Fuego province, Bahía Lapataia, ca. 54°52′W 68°32′S, II.1963, E. Maury, deposited in MACN-Ar 9850; male paratype from Tierra del Fuego, Ushuaia, dung traps in Sphagnum, lakeside bog, 20 pans, ca. 54°48′S, 68°18′W, 12–14.II.1982, S. Marshall, deposited in AMNH.

    Etymology:

    The specific name refers to the small body size.

    Diagnosis:

    See generic diagnosis.

    Female (holotype):

    Total length 4.83. Carapace length 2.23, width 1.60, wider on legs II–III. Length of tibia/metatarsus: I, 1.12/0.92; II, 1.00/0.87; III, 0.83/1.00; IV, 1.30/1.53. Palpal tarsus length 0.40. Chelicerae with three teeth on retromargin, median one slightly smaller. Sternum length 1.13, width 0.98. Spines: leg I, femur d 1–1–1 thin bristles, p d1ap; tibia v 2–2–0; metatarsus v 2bas. II, femur d 1–1–1 or 1–1–0 thin bristles, p d1ap; tibia v 2–2–0, p 0–1 or d1–1; metatarsus = I. III, femur d 1–1–1, p d1ap or 0; patella 0; tibia v p1–2–2, p 1-d1-1-0, r d1–1; metatarsus v 2–2-comb, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, r d1ap or 0; patella 0; tibia v p1–2–2, p and r 1-d1-1-0; metatarsus = III. Abdomen length 2.80, width 1.45, spiracle–epigastrium 1.08, spiracle–spinnerets 0.12. Color: grayish brown almost uniform. Abdomen with two paler bands at sides of cardiac area, diffusing posteriorly. Epigyne: see generic description.

    Male (paratype):

    Total length 3.17. Carapace slightly narrower in front than that of female, length 1.37, width 0.95. Length of tibia/metatarsus: I, 0.82/0.67; II, 0.68/0.62; III, 0.55/0.67; IV, apparently regenerated. Chelicerae slightly narrower than those of female. Sternum length 0.75, width 0.63. Spines as in female, except: leg II, femur d 1–1–1 bristles; tibia p 0–1; metatarsus p 0–1. III, femur p 0; tibia p d1–1, v p1-p1–2; metatarsus v 2–2-comb or 2-p1-comb. IV, badly developed, apparently regenerated, with reduced spination. Abdomen length 1.60, width 0.90, spiracle–epigastrium 0.82, spiracle–spinnerets 0.27. Color: carapace grayish brown, legs paler. Sternum and mouthparts brown, darker than coxae. Abdomen grayish brown, dorsum yellow, dark grayish on cardiac area and several dark diffuse chevrons up to posterior end. Palp: see generic description.

    Variability:

    Some males with RTA slightly narrower and sharper. Female, spines: III, tibia p d1–1. IV, tibia r d1–1.

    Natural History:

    Ground dwellers. Several specimens were collected under stones or logs near the seashore.

    Distribution:

    Chile, from Osorno to Magallanes, and Argentina, in Tierra del Fuego.

    Other Material Examined:

    ARGENTINA: Tierra del Fuego: Bahía Buen Suceso, 13.X.1971, Menéndez, 1♀ 3 immatures (MACN-Ar); shore at Canal de Beagle, I.1933, Castellanos and Gómez, 1 immature (MACN-Ar); road to Glaciar Le Martial, on moss Polytrichum strictum, XII.1989, A. González, 2♀ 1 immature (MLP); Isla de los Estados, Puerto Flinders, 7.X.1971, Menéndez, 1♀ (MACN-Ar); Ushuaia, dung traps in Sphagnum, 2♀, dry bog, pans, 2♂ 1♀ 1 immature, lakeside bog, 20 pans, 1♂, 1♀, 1 immature, 12–14.II.1982, S. Marshall (AMNH). CHILE: Región X (Los Lagos): Osorno: Puyehue Natl. Park: Antillanca rd., 965 m, trap site 658, Berlese, leaf and log litter, 18–25.XII.1982, A. Newton and M. Thayer, 1♀ (AMNH). Llanquihue: Huelmo, superior level of beach, 30.XII.1986, J. Kochalka, 1♂ 1♀ (IBNP). Región XII (Magallanes y Antártica): Ultima Esperanza: Laguna Parrillar Natl. Res., 53°24′15″S, 71°15′45″W, 1–10.XII.2000, 350 m, J. Miller, I. Agnarsson, 1♂, pitfall, 3♀, pitfall 55T3, bog, Berlese in moss, 1♀, 1♀, 1♀, 2♀, pitfall 54T3, Sphagnum, 1♂ 1♀, pitfall 55T3, Sphagnum, 2♀, pitfall 55T4, scrub, grass in bog, 3♀, pitfall 55T6, forest, 2♀, bog, moss, 2♀, grass near Churio Hermoso, pitfall 55T2, 3♀, moss near Churio Hermoso, 2♀, pitfall 55T3, 2♀ (USNM). Magallanes: Isla Lennox, Lennox Cave, 5.II.1896, O. Nordenskjöld, 1♀ (NRS).

    Josa Keyserling Table 17

  • Tetromma Keyserling, 1878: 608 (type species by monotypy Tetromma lutea Keyserling, 1878; preoccupied by Déjean, 1834). new synonymy.

  • Josa Keyserling, 1891: 83 (type species Anyphaena pilosa Keyserling, 1880; earliest available name for Tetromma Keyserling, preoccupied). Simon, 1897a: 104. Brescovit, 1993: 129. Ramírez, 1995a: 381, 1997: 178.

  • Pelayo O.P.-Cambridge, 1896: 194 (type species by monotypy Pelayo laetus O.P.-Cambridge, 1896); Simon, 1903a: 1032. F.O.P.-Cambridge, 1900: 94, 107. Synonymized by Brescovit, 1993: 129.

  • Haptisus Simon, 1897a: 100 (type species by original designation Anyphaena nicoleti Simon, 1897). new synonymy.

  • Olbophthalmus Simon, 1904: 98 (type species Olbus personatus Simon, 1897, designated by Petrunkevitch, 1928: 173). Ramírez, 1995a: 381, 1997: 178. new synonymy.

  • Gayennella Berland, 1913: 102 (type species by monotypy Gayennella riveti Berland, 1913). new synonymy.

  • Synonymy:

    The type species of Tetromma, Olbophthalmus, and Gayennella are here considered typical members of Josa. Anyphaena pilosa and A. nicoleti are here considered junior synonyms of Josa lutea.

    Note:

    Simon (1880) described the genus Olbus for the poorly preserved type specimen of Olios sparassoides Nicolet, a corinnid (Ramírez et al., 2001). However, his subsequent description of Olbus (Simon, 1897a), and the specimens identified by him all correspond to Josa species with a distinctly recurved posterior eye row. Simon only later (1904) examined fresh specimens of O. sparassoides, when he clarified the point, and proposed the genus Olbophthalmus in Anyphaenidae.

    Diagnosis:

    Easily recognized from other Amaurobioidinae by having a ventral apical palpal femoral apophysis (fig. 60E). Females have copulatory ducts coiled along longitudinal axes, and the epigynal median field at the same level or lower than the posteriorly projecting lateral lobes.

    Description:

    Carapace narrowed in front, posterior eye row variable, ocular area not projecting. Chelicerae unmodified, slightly smaller in males, with three teeth on promargin, two on retromargin, occasionally four promarginal, three retromarginal. Anterior legs with unmodified spines, more spinose in males. Male palp with ventral apical femoral apophysis, hook-shaped; tibia short, RTA absent. Cymbium large. Tegulum placed basally and prolaterally in cymbium, median apophysis wide, apical, bifid. Anterior dorsal margin of tegulum with dorsal lobe, sperm duct without anterior dorsal loop. Primary conductor fused to tegulum, without canal. Secondary conductor extremely modified, semicircular, not associated with embolus when unexpanded (fig. 54A, B). Paramedian apophysis with shallow cusps, fused to tegulum. Embolus long, hidden between cymbium and bulb, with complex basal process (fig. 54B). Epigyne variable, median field mostly visible in posterior view (fig. 54C, D), copulatory openings in epigastric furrow. Lateral lobes with posterior depressions, slightly projecting posteriorly, limiting commonly narrow notch. Copulatory ducts long, coiled along longitudinal axes.

    Distribution:

    South and Central America, with most species occurring in Andean cloud forests and paramos.

    Composition:

    The genus is extremely diverse, with most species being still undescribed and undersampled in collections. The morphology of the male copulatory bulb is remarkably constant within several clusters of species, making identification problematic. In addition to the species detailed below: Anyphaena keyserlingi L. Koch, 1866 (several males, females, and immatures syntypes, from Colombia: Santa Fe de Bogotá, in BMNH, examined, new combination), Josa bryantae (Caporiacco, 1955), Josa laeta (O.P.-Cambridge, 1896), Olbophthalmus lojensis Berland, 1913 (new combination, see Note under Josa personata), Gayenna andesiana Berland, 1913 (male and female syntypes in MHNP, examined, new combination), Gayenna simoni Berland, 1913 (male and female syntypes in MHNP, examined, new combination), Haptisus analis Simon, 1897 (male and female syntypes in MHNP 11265, examined, new combination), Haptisus maurus Simon, 1897 (penultimate female holotype in MHNP 17554, examined, new combination), Olbus gounellei Simon, 1897 (male and female syntypes should be in MHNP 8166, not found, examined by Kochalka [1980], new combination), Tomopisthes chazaliae Simon, 1897, new combination (three females syntypes in MHNP 18296, B.1811, examined by Kochalka [1980], belonging to two different species of Josa; the types were not found in subsequent years; the very superficial description [Simon, 1897c] and the type locality [Colombia, Sierra Nevada de Santa Marta] are compatible with the genus).

    Nomen Dubium:

    Clubiona nigricans Nicolet, 1849 (male and female syntypes from Chile, Valdivia, presumably in MHNP, not found; transferred to Haptisus by Simon, 1897a: 100). The reference to the elongate abdomen (Nicolet, 1849: 447) suggests that this species might actually belong to Acanthoceto, Aysenia, or Aysenoides.

    Josa lutea (Keyserling), new combination Figure 55

  • Anyphaena citrina: L. Koch, 1866: 194, 199 (many specimens from Colombia, Santa Fe de Bogotá, in BMNH, examined, misidentification).

  • Tetromma luteum Keyserling, 1878: 608 (female holotype from Nueva Granada [Colombia], in BMNH, examined by John Murphy, in litt.).

  • Anyphaena pilosa Keyserling, 1880: 327 (male and female syntypes, from Nueva Granada [Colombia], in BMNH, examined). new synonymy.

  • Josa pilosa: Keyserling, 1891: 83. Simon, 1897a: 104.

  • Anyphaena nicoleti Simon, 1897a: 92 (name for the specimens misidentified by L. Koch, 1866 as “Anyphaena citrina Nicol.?”).

  • Haptisus nicoleti: Simon, 1897a: 92, 95, 100. Berland, 1913: 104.

  • Gayenna riveti Berland, 1913: 100 (female holotype from Ecuador, Borma, 1905, in MHNP, examined). new synonymy.

  • Tetromma lutea: Bonnet, 1959: 4364 (emendation of T. luteum Keyserling).

  • Synonymy:

    The description by L. Koch (1866) is headed “Anyphaena citrina Nicol.?”, with a reference to Clubiona citrina Nicolet (1849: 433). This is not the description of a new species (Bonnet, 1957: 2098; contra Roewer, 1954: 540). Clubiona citrina Nicolet (see Nomen Dubium under Monapia) was transferred by Simon (1897a: 92) to Gayenna. Simon gave the name Anyphaena nicoleti (1897a: 92) to the species that L. Koch identified as “Anyphaena citrina Nicol.?”, and seven lines below named the same species as Haptisus nicoleti. Hence, the types of Anyphaena nicoleti Simon, 1897 are the specimens identified by L. Koch (1866) as “Anyphaena citrina Nicol.?” (Bogotá, BMNH). I have seen drawings of the holotype of Tetromma luteum (thanks to John Murphy and Norman Platnick, in litt.). It is a teratological female, with only three eyes (both AME and left ALE). The epigyne is normal, allowing acceptable identification. The holotype of Gayenna riveti Berland also does not show differences in the epigyne.

    Diagnosis:

    Very similar to J. riveti in the epigyne, distinguished by the sinuous posterior borders of epigynal lateral lobes.

    Female (Bogotá, MHNP 3510):

    Total length 10.08. Carapace length 3.50, width 2.83, wider on legs II–III. Length of tibia/metatarsus: I, 2.25/2.75; II, 2.20/2.05; III, 1.80/1.90; IV, 2.15/2.32. Chelicerae unmodified, with two teeth on retromargin (fig. 55D). Spines: leg I, femur d 1–1–1, p 0-d1-(1-d1) or 0-d1-d1; tibia v 2–2–2; metatarsus v 2bas. II, femur = I; tibia v 2–2–2, p 0–1; metatarsus v 2bas, d p1bas. III, femur d 1–1–1, p and r 0-d1-d1; patella rd1; tibia v 2–2–2 or p1–2–2, p and r 1-d1-1-0; metatarsus v 2–0–2, p and r 1-d1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia = III; metatarsus v 2–2–2, p and r 1-d1–1, d 0-p1–2. Abdomen length 6.58, width 4.67, spiracle–epigastrium 2.86, spiracle–spinnerets 1.35. Color: carapace grayish with dark median longitudinal band, slightly darker on cephalic area, ocular area dark gray, clypeus with white hairs. Legs gray with dorsal dark longitudinal stripes as: I and II, tibiae d 3–1–1 short; metatarsi d 1–1, basal long. III, IV tibiae d p2-r2-r1 short; metatarsi = I. Abdomen yellowish with grayish dorsal pattern, venter with grayish median longitudinal band. Epigyne (fig. 55A–C): median field wide, oval in posterior view. Lateral lobes approaching each other, limiting narrow notch between two rounded projections, each with anterior depression. Copulatory ducts long, coiled. Ducts of accessory bulbs long, converging.

    Male:

    Unknown.

    Natural History:

    Unknown.

    Distribution:

    Known only from Bogotá and probably from Tungurahua, Ecuador.

    Other Material Examined:

    COLOMBIA: Bogotá, no date, no collector, 1♀ (MHNP 3510). ECUADOR: Tungurahua: (two labels) Mt. Tungurahua or Baños (?), 1850 or 3800 m (?), 6.I.1938 or 1.XI.1937 (?), W.M. Clarke-Macintyre, 1♀ (AMNH).

    Josa riveti (Berland), new combination Figure 56

  • Gayennella riveti Berland, 1913: 100 (female immature holotype from Ecuador, Borma, El Pelado, G. Rivet, 1905, in MHNP, examined).

  • Note:

    Berland (1913: 102) described an immature female, and he seemed clear in that the specimen from El Pelado was the holotype (“Je crée un genre noveau pour une araignée de El Pelado”). He mentioned also one female and an immature male from Yana-Urcu (in MHNP, examined). Both vials in MHNP are labeled “Type du genre et de l'especie”. The present knowledge of the genus does not permit the identification of immatures, so I provisionally distinguished the species based on the female from Yana-Urcu.

    Diagnosis:

    Very similar to J. lutea in the epigyne, distinguished by the slightly curved, almost straight posterior borders of the epigynal lateral lobes. Males can be distinguished from similar Josa species by the small apical bifurcation of the conductor.

    Female (Zumbahua, MACN-Ar 9813):

    Total length 12.40. Carapace length 5.05, width 3.46, wider between legs II and III. Length of tibia/metatarsus: I, 2.73/2.27; II, 2.73/2.27; III, 2.27/2.53; IV, 3.30/3.78. Palpal tarsus length 1.63. Chelicerae with two teeth on retromargin (fig. 56F). Sternum (fig. 56G) length 2.50, width 1.83. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 0–2–2 or p1–2–2 or 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v r1–2–2 or 2–2–2, p 0 or 0–1; metatarsus v 2bas. III, femur d 1–1–1, p and r 0-d1-d1 or p d1-d1-d1; patella d d1; tibia v 2–2–2 (the r1bas smaller), p and r 1-d1-1-0, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0–2–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia and metatarsus = III. Dorsal long, thin, erect bristles on patellae (d 1–0–1) and tibiae (d r1–0–1). Abdomen length 7.95, width 4.50, spiracle–epigastrium 3.60, spiracle–spinnerets 1.83. Color (from several specimens): carapace grayish with darker sides, legs grayish, coxae pale, sternum grayish, paler at center, mouthparts dark. Abdomen with pattern of grayish spots on paler background, dorsal pattern variable, often with anterior dark bands on cardiac area, one anterior band at each side, several chevrons extending to spinnerets; venter variable from pale to dark, may have diffuse median longitudinal dark band. Epigyne (fig. 56H–J): median field wide, sclerotized, trapezoidal in posterior view, ventral angles rounded. Lateral lobes close to each other, forming narrow notch between posterior projections, shallow depression anterior to each projection of lateral lobes. Copulatory ducts very long, coiled. Ducts of accessory bulbs long, converging.

    Male (Zumbahua, MACN-Ar 9813):

    Total length 8.40. Carapace length 3.99, width 2.87. Length of tibia/metatarsus: I, 2.57/2.13; II, 2.50/2.07; III, 2.07/2.17; IV, 2.83/3.17. Chelicerae narrower than those of female. Sternum length 2.00, width 1.47. Spines as in female, except: leg I, femur r 0 or d1 or 0-d1-d1; tibia v 2–2–2, p 1-d1-1-0, r 1-0-1-0 or 1–0; metatarsus p 0 or d1. II, femur p 0-d1-d1 or 0-d1–2, r d1 or 0-d1-d1; tibia v 2–2–2, p 1-d1-1-0, r 1-0-1-0; metatarsus p = I. Abdomen length 4.66, width 2.28, spiracle–epigastrium 1.63, spiracle–spinnerets 0.94. Color as in female, often darker, more heavily contrasting. Copulatory bulb (fig. 56A–E): median apophysis wide, with two tips, apical tip with ventral canal. Secondary conductor bifid just at apex. Paramedian apophysis heavily sclerotized, with one rounded cusp with sharp border, fused to tegulum and primary conductor. Embolus very long, apex describing complete loop between bulb and cymbium.

    Variability:

    Female spines: III, IV, metatarsus d 0-p1–2. Males from Volcán Chiles have slightly different MA (compare fig. 56D and E).

    Natural History:

    Unknown. Some specimens were collected under rocks in high-altitude grasslands.

    Distribution:

    Highlands of Ecuador and Bolivia.

    Other Material Examined:

    ECUADOR: Carchi: Volcán Chiles, 0°47′40″N, 77°57′00″W, above Naranjal, páramo grassland, 4050 m, sample 286 (pitfall), 10.VIII.1997, N. Atkins, 1♀ (UPBS); sample 285, 1♂ 1♀; sample 284, 0°47′15″N, 77°56′45″W, 2♂. Cotopaxi: 5 km E Zumbahua, W Latacunga, ca. 3500 m, 18.IV.1982, A. Roig, 3♀ 2♂ 2 immatures (MACN-Ar 9813); Lacatunga to Quevedo, 3600 m, 15.VIII.1965, L. Peña, 1♂ (MCZ). Pichincha: 10 km E Pifo, 30.VII.1978, A. Roig, 1♂ 4♀ 1 immature (MACN-Ar); 15 km E Pifo, grassland with cattle, under stones, 3.V.1982, A. Roig, 2♂ 7♀ 4 immatures (MACN-Ar); 22 km E Pifo, 3.V.1982, under stones, A. Roig, 5♀ 1 immature (MACN-Ar). BOLIVIA: La Paz: Botijlaca, 3600 m, 2.XI.1984, L.E. Peña, 1♀ (AMNH).

    Josa personata (Simon), new combination Figure 57

  • Olbus personatus Simon, 1896b: 506 (female holotype from Ecuador Meridional, Amazula, Gaujon coll., in MHNP 9776, examined), 1897a: 93, 102.

  • Olbophthalmus personatus: Petrunkevitch, 1928: 173. Simon, 1904: 98.

  • Note:

    Olbophthalmus lojensis Berland (1913: 104; one female and one male without palps syntypes, from Ecuador, Loja, 2200 m, in MHNP, examined) may be a junior synonym. Because I have seen other very similar species represented only by small samples, a synonymy is postponed until more conclusive evidence is available.

    Diagnosis:

    Resembles J. calilegua in having a pale body and recurved posterior eye row, but distinguished by having two teeth on the cheliceral retromargin and simple ridges on the epigynal lateral lobes.

    Female (holotype):

    Total length 4.73. Carapace length 2.38, width 2.02, wider between legs II and III. Posterior eye row recurved, AME larger than ALE (fig. 57A, B). Length of tibia/metatarsus: I, 2.08/1.63; II, 2.06/1.83; III, 1.45/1.45; IV, 1.78/1.70. Chelicerae with two teeth on retromargin. Sternum length 1.48, width 1.20. Spines: leg I, femur d 1–1–1, p 0–2-d1; tibia v 2–2–0; metatarsus v 2bas. II, femur d 1–1–1, p 0-d1-d1; tibia and metatarsus = I. III, femur d 1–1–1, p 0-d1-d1, r d1ap; patella 0; tibia v 2–2–2, r d1–1 or d1–0, d r1bas; metatarsus v 2–0–2, p and r d1–1, d 0–2. IV, femur d 1–1–1, p d1ap; patella 0; tibia v 2–2–2, p d1(small)-1, r 1-d1-1-0, d r1bas; metatarsus v 2–0–2, p and r d1-1-0-1, d 1–2. Abdomen length 2.35, width 1.18, spiracle–epigastrium 1.35, spiracle–spinnerets 0.60. Color in alcohol: carapace and abdomen yellow, eyes bordered black. Chelicerae with anterior face dark gray, sternum and mouthparts yellow. Legs yellow with dorsal longitudinal lines dark gray: patellae, d 2bas, short; tibiae, d p2bas extending up to one-third or one-fourth of tibial length, d r2bas extending up to one-half of tibial length, d 2bas short, d 1, and d r1. Metatarsi III, IV with apical bundle of white hairs on each side. Epigyne (fig. 57C–F) weakly sclerotized, spermathecae visible through cuticle; two posterior depressions, oriented forward. Lateral lobes separate, limiting shallow notch at posterior border of epigyne. Median field wide, oval in posterior view. Copulatory ducts very long, coiled. Ducts of accessory bulbs long, converging. Lumen of spermatheca elongate, coiled along with coils of copulatory duct. Fertilization ducts long, also coiled.

    Male:

    Unknown.

    Variability:

    Spines: I, femur p 0-1-d1-1 or 0-1-d1-0. III, tibia p 0–1, r 0–1; metatarsus p and r d1-1-0-1, d p1–2.

    Natural History:

    Unknown.

    Other Material Examined:

    ECUADOR: Loja: Malacatos, 1900 m, 21–22.VIII.1977, L. Peña, 1♀ (AMNH).

    Josa calilegua, new species Figures 54, 58

    Types:

    Female holotype and male paratype from Argentina, Jujuy province, Calilegua Natl. Park, 11.3 km from Park entrance, ca. 23°30′S, 64°50′W, 23–24.IX.1995, M. Ramírez, P. Goloboff, C. Szumik, deposited in MACN-Ar 9814.

    Etymology:

    The specific name is a noun in apposition taken from the type locality.

    Diagnosis:

    Resembles J. personata in having a pale body and recurved posterior eye row, but distinguished by having three teeth on the cheliceral retromargin and four on the promargin, lacking lateral depressions anterior of the epigynal lateral lobes, and the characteristic shape of the apex of conductor.

    Female (holotype):

    Total length 6.25. Posterior eye row recurved. Carapace length 2.80, width 2.17, wider between legs II and III. Length of tibia/metatarsus: I, 1.70/1.57; II, 1.67/1.50; III, 1.50/1.47; IV, 1.90/2.03. Palpal tarsus length 0.90. Chelicerae unmodified, with four teeth on promargin, three on retromargin. Sternum length 1.87, width 1.10. Spines (all tibiae d r1–0–1 bristles): leg I, femur d 1–1–1, p 0-d1-1-1, r 0-d1-d1; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p and r 0-d1-d1; tibia = I (the v p1bas thinner; one additional spine close to the x-r1-x on left leg); metatarsus v 2bas. III, femur = II; tibia v 2–2–2, p 1-d1-1-0 or d1–1, r 1-d1-1-0; metatarsus v 2–0–2, p and r d1-1-0-1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; tibia = III; metatarsus v 2-p1–2, p and r d1-1-0-1-0, d p1–2. Color: yellowish white, legs darkening distally, abdomen with white guanine reticulum, less dense along median lines of dorsum and venter. Ocular area black, many white hairs on clypeus, margin of carapace, dorsum of coxae, and sides of abdomen. Epigyne (figs. 54C, D, 58C, D): median field wide, oval in posterior view, lateral lobes with oblique posterior borders, projecting over median field, limiting rectangular notch. Copulatory ducts long, coiled, ducts of accessory bulb long, converging. Lumen of spermatheca irregular, partially coiled along with coils of copulatory ducts. Fertilization ducts long.

    Male (paratype):

    Total length 4.90. Carapace length 2.37, width 1.93. Length of tibia/metatarsus: I, 1.93/1.77; II, 1.90/1.83; III, 1.53/1.53; IV, 1.77/2.00. Chelicerae small, vertical. Sternum length 1.30, width 0.93. Spines as in female, except: leg I, tibia v 2–2–2, p and r 1-d1-1-0, d r1–0–1; metatarsus v 2–2–0, p and r d1. II, tibia and metatarsus = I. III, tibia = I. IV, tibia = I. Abdomen reduced by starving, length 2.50, width 1.10, spiracle–epigastrium 0.83, spiracle–spinnerets 0.57. Color as in female, but with small brownish violet spot above anal tubercle. Copulatory bulb (figs. 54A, B, 58A, B): median apophysis wide, apical tip narrow, basal tip represented only by ridge. Secondary conductor bifid apically, one tip blunt, another long, acute. Paramedian apophysis sclerotized, with short, bifid cusp, fused to tegulum, partially separated from conductor base by small membranous area. Embolus very long, apex describing complete loop between bulb and cymbium.

    Natural History:

    Most specimens were collected by beating foliage.

    Distribution:

    Moderate altitude rainforests in Argentina, Tucumán and Jujuy provinces.

    Other Material Examined:

    ARGENTINA: Jujuy: Same data as types, 2♂ 4♀ 2 immatures (MACN-Ar). Tucumán: Horco Molle, II.1965, A. Bachmann, 1♀ (MACN-Ar); Cerro San Javier, 11.II.1951, Ross, 1♀ (CAS).

    Josa nigrifrons (Simon), new combination Figures 59, 60

  • Haptisus nigrifrons Simon, 1896b: 505 (male lectotype and 4 females paralectotypes here designated, from Venezuela, Aragua, Colonia Tovar, in MHNP, examined), 1897a: 92, 100.

  • Pelayo insignis Banks, 1909a: 199 (female lectotype and male immature paralectotype here designated, from Costa Rica, Volcán Poás, Alajuela, J.F. Tristán coll., in MCZ, examined). new synonymy.

  • Josa insignis: Brescovit, 1993: 129.

  • Synonymy:

    The paralectotypes of Haptisus nigrifrons were compared with the lectotype of Pelayo insignis and with numerous specimens from the same area; no relevant differences were found.

    Diagnosis:

    Easily distinguished from other Josa by the extreme development of the secondary conductor and by the female epigyne projecting posteriorly, with anterior lateral sclerotizations.

    Female (paralectotype):

    Total length 5.27. Carapace length 2.24, width 1.62, wider between legs II and III. Anterior eye row procurved, AME slightly smaller than ALE, posterior eye row slightly recurved. Length of tibia/metatarsus: I, 1.10/0.98; II, 1.08/0.98; III, 0.88/0.92; IV, 1.22/1.28. Chelicerae with two teeth on retromargin. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p 2ap, r d1ap; tibia v r1-r1–2; metatarsus v 2bas. III, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1–1 (bristles); metatarsus v 2–0–2, p d1–1–1, r 0–1–1, d 0-p1–2. IV, femur d 1–1–1, p and r d1ap; patella r d1; tibia v p1–2–2, p and r 1-d1-1-0, d r1–1 (bristles); metatarsus v 2–0–2, p and r d1–1–1, d 0-p1-r1. Abdomen length 3.06, width 1.78, spiracle–epigastrium 1.04, spiracle–spinnerets 0.69. Color: pale yelow, carapace with pale brown diffuse lateral bands, ocular area black, clypeus with white hairs. Chelicerae dark gray and black, sternum pale, labium dark brown. Abomen with dark brownish violet dorsal pattern (fig. 59A), and three posterior black marks, epigastrium with reddish spots at sides of epigyne, venter with brownish violet band from epigastrium to spinnerets. Epigyne (fig. 59C–G) projecting posteriorly, epigastrium sclerotized in two patches lateral and anterior of epigyne. Epigynal posterior border formed by projecting lateral lobes, each with depressed area and notch in between. Median field and copulatory openings hidden in epigastric fold, visible only after dissection. Median field flattened, sclerotized, with two lobes directed backward (fig. 59E–G). Copulatory ducts coiled, most anterior loop with ample lumen (probably functioning as reservoir), lumen of spermathecae tortuous, ducts of accessory bulbs long, slender, converging.

    Male (lectotype):

    Total length 4.73. Carapace length 2.28, width 1.84. Length of tibia/metatarsus: I, 1.55/1.35; II, 1.48/1.32; III, 1.12/1.10; IV, 1.50/1.55. Chelicerae slightly narrower than those of female. Spines as in female, except: leg I, femur p d1ap or 2ap, r 0-d1-d1; tibia p and r 1-d1-1-0; metatarsus v 2–2–0. II, femur p and r 0-d1-d1; tibia p and r 1-d1-1-0, v r1–2–2; metatarsus v 2–2–0, p d1, r d1 or 0. III, femur = II; tibia v p1–2–2, r 1-d1-1-0, d r1–0–1; metatarsus d 0–2–2. IV, femur = II; tibia = III; metatarsus v 2-p1–2, d 0–2–2. Abdomen length 2.45, width 1.43, spiracle–epigastrium 0.86, spiracle–spinnerets 0.61. Color: similar to female, lateral band on carapace sharper, band on venter narrower, extending anterior of spiracle to spinnerets. Palp (fig. 60): median apophysis large, concave. Secondary conductor very complex, one acute tip bearing canal, other tip bifid, much more developed, plus additional cusp on base. Paramedian apophysis absent, apparently reduced to shallow ridge parallel to tegular margin. Embolus very long, apex describing complete loop between bulb and cymbium.

    Variability:

    Genitalia quite variable. Some females with lobes of epigynal median field hardly visible in ventral view (fig. 59E, G). Males with secondary conductor also variable in details. Some specimens with longitudinal black lines on legs, from patella to metatarsus, similar to those of J. personata. Abdomen may have short dorsal band, anterior of chevrons. Abdominal ventral band varying in length and width. Female spines: III, tibia v p1–2–2; metatarsus p and r d1–1–1, d 0–2–2. IV, metatarsus v 2–2–2, p and r d1–1–1, d 0–2–2. Male spines: III, IV, metatarsus = female.

    Natural History:

    Unknown.

    Distribution:

    Andean rainforests from Costa Rica to Bolivia.

    Other Material Examined:

    PANAMA: El Volcán, 12.VIII.1950, A.M. Chickering, 3♂ 5♀ 3 immatures (MCZ); 9–14.VIII.1950, A.M. Chickering, 14 immatures, 9♀ 8♂ 1 immature (MCZ), 10.VIII.1950, A.M. Chickering, 1♀ (MCZ). ECUADOR: Loja: Malacatos, 1900 m, 21–22.VIII.1977, L. Peña, 1♂ 3♀ 9 immatures (AMNH). Prov. Tungurahua: Baños, 2200 m, 29.IV.1939, W. Clarke-Macyntre, 1♂ (MCZ); Tungurahua, 2600 m, 6.V.1939, W.M. Clarke-Macintyre, 1♂ (AMNH). BOLIVIA: La Paz: La Paz, Unduavi to Coroico, yungas, 2500–3200 m, 18–22.XI.1984, L. Peña, 1♀ (AMNH).

    TRIBE GAYENNINI, NEW RANK Table 18

    Type Genus:

    Gayenna Nicolet, 1849.

    Note:

    Gayennini is equivalent to the “Gayenna-Oxysoma group” of Kochalka (1980: 36) and Ramírez (1995b: 72).

    Diagnosis:

    Distinguished from Amaurobioidini and Josa by having an anterior epigynal pouch, spherical spermathecae well differentiated from the relatively slender copulatory ducts, and a primary conductor being vestigial or absent.

    Description:

    Posterior eye row procurved or straight. Chelicerae usually with two teeth on retromargin. Male palp without RTA. Cymbial conductor narrow (fig. 61). Primary conductor reduced to small sclerite, hidden between distal sclerites or absent. Secondary conductor well developed, canal of variable shape (absent in some groups). Median apophysis slender, tip hooked. Paramedian apophysis with distinct, elongate cusp. Embolus with short, simple basal process, formed by extension of narrow sclerotized stripe, partially bordered by membranous areas. Epigyne with anterior pouch on median field, lateral lobes usually separate but contiguous or fused to each other in some groups. Spermathecae spherical, outline and lumen well differentiated from those of copulatory ducts.

    Distribution:

    Mainly South America, but Arachosia extending also to Central and North America. Sanogasta maculatipes probably introduced to Eastern Island.

    Composition:

    Eleven genera, two of them newly proposed here: Arachosia O.P.-Cambridge, Araiya, n. gen., Gayenna Nicolet, Gayennoides, n. gen., Monapia Simon, Oxysoma Nicolet, Phidyle Simon (here removed from the synonymy of Oxysoma), Philisca Simon, Sanogasta Mello-Leitão, Tasata Simon, and Tomopisthes Simon.

    Nomina Dubia:

    The following species belong to Gayennini, but cannot be assigned with certainty to a genus: Clubiona albiventris Nicolet, 1849 (immature presumably holotype in MHNP 4225, examined); Clubiona gibbosa Nicolet, 1849 (two immature syntypes in MHNP 4233, examined); Clubiona lepida Nicolet, 1849 (small immature presumably holotype in MHNP 4228, examined); Clubiona puella Nicolet, 1849 (immature presumably holotype in MHNP 4229, examined); Clubiona pulchella Nicolet, 1849 (two immatures syntypes in MHNP 4236, examined).

    Gayenna Nicolet Table 19

  • Gayenna Nicolet, 1849: 450 (type species by monotypy Gayenna americana Nicolet, 1849). Simon, 1884 and 1887: E24, E26, 1897a: 91, 94, 96, 98–99. Keyserling, 1891: 83, 137. F.O.P.-Cambridge, 1900: 94, 107. Tullgren, 1901: 230, 1902: 58. Banks, 1905: 307, 1907: 723 (Gavenna lapsus). Merian, 1913: 76. Mello-Leitão, 1925: 456. Kochalka, 1980: 98–100. Ramírez, 1995a: 1, 1997: 177. Ramírez and Kochalka, 1993: 164.

  • Mezenia Simon, 1897a: 92, 96, 98, 101. Tullgren, 1902: 67. Bonnet, 1957: 2828.

  • Mezenina Strand, 1932: 141 (new name for Mezenia Simon, 1897, preoccupied by Stuckenberg, 1895). Synonymized by Ramírez and Kochalka, 1993: 164.

  • Diagnosis:

    The single known species is easily distinguished from all other Amaurobioidinae by having a characteristic abdominal pattern (fig. 62A) and by the anterior median eyes being much larger than the lateral ones.

    Description:

    See Ramírez and Kochalka (1993: 164). Additional data: posterior eye row procurved. Chelicerae with three teeth on promargin, two on retromargin. Male palp with short tibia. Copulatory bulb (fig. 63): embolus with basal process flattened, associated with distal membranous area. Paramedian apophysis thick, apex bifid, with retrolateral, short, curved cusp, small pointed prolateral cusp. Secondary conductor striated transversely-obliquely, well separated from anterior margin of tegulum by membranous area; retrolateral portion with basal prong; tip of prolateral portion elongate, acute; canal deep. Primary conductor thick, blunt, heavily sclerotized. Epigyne with lateral lobes close to each other, median field very narrow, separated from lateral lobes by shallow sutures. Lateral lobes elevated at sides of median pouch. Copulatory ducts long.

    Composition:

    Only the type species.

    Types Not Examined:

    Gayenna brasiliensis Roewer, 1951 (replacement name for Gayenna alticola Mello-Leitão, 1926, perhaps in MNRJ, not found). Gayenna chrysophyla Mello-Leitão, 1926 (male and female syntypes not examined, perhaps in MNRJ, not found): a schematic illustration of the epigyne (Mello-Leitão, 1926: fig. 4) suggests that if may belong in Sanogasta or Arachosia. Gayenna ignava Banks, 1898 (type originally in CAS, lost, C. Griswold, personal commun.; most probably Anyphaeninae) and Gayenna orizaba Banks, 1898 (same as G. ignava).

    Gayenna americana Nicolet Figures 12, 13E–G, 16, 17, 62, 63, 68A–C

  • Gayenna americana Nicolet, 1849: 450. Ramírez and Kochalka, 1993: 164. Ramírez, 1995a: 366, 1997: 177.

  • Mezenia dorsalis Simon, 1897a: 101, 1904: 103; Tullgren, 1902: 103; Merian, 1913: 13.

  • Mezenina dorsalis: Strand, 1932:141.

  • Diagnosis:

    See generic diagnosis.

    Description:

    See Ramírez and Kochalka (1993: 164) and generic description.

    Variability:

    Female spines: IV, tibia v p1-p1–2 or p1–2–2; metatarsus v 2-p1–2 or 2–2–2, p d1–1–1 or 0-d1–1, r d1–1–1. Male spines: III, tibia v p1–2–2.

    Distribution:

    Forests in southern Argentina and Chile.

    New Records:

    ARGENTINA: Neuquén: Puerto Blest, 7–20.I.2000, L. Lopardo and A. Quaglino, 15 immatures (MACN-Ar). CHILE: Región IV (Coquimbo): Choapa: Los Vilos, Cariloleu, 11.X.1994, L. Peña, 1♂ (AMNH). Región VII (Maule): Talca: Alto de Vilches, 17–24.X.1964, L. Peña, 3♂ 3 immatures (MCZ); elev. 1180 m, 35°36′S, 71°04′W, 14–15.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 5♂ 2♀ (AMNH), 3♂ (MACN-Ar), 2♂ (MHNS); Parque Gil de Vilches, 7–8.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH). Cauquenes: Tregualemu, 520 m, 6–7.XI.1993, L. Peña and A. Ugarte, 1♀ (AMNH). Linares: Bullileo, Parral, 5–8.XII.1990, L. Peña, 1♂ (AMNH); Fundo Malcho, Andes en Parral, 11–20.XI.1964, L. Peña, 7♂ 5♀ 3 immatures (MCZ), 10–11.XI.1993, L. Peña, 1♂ (AMNH). Región VIII (Biobío): Ñuble: Las Trancas, 1–10.XII.65, L. Peña, 4♂ 1 immature (MCZ); Punta El Zorro, 20.XII.1992, T. Cekalovic, 1♂ (AMNH). Concepción: Cerro Caracol, Concepción, elev. 200 m, 36°51′S, 73°02′W, 17.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH); Estero Nonguén, 10.II.1996, 1♂, 5.X.1996, 1♂ 1♀, T. Cekalovic (AMNH); Fundo El Manzano, 7.XI.1992, 2♂ 2♀, 8.XI.1992, 1♂, T. Cekalovic (AMNH); Periquillo, 22.XI.1992, 1♂ 2♀ 2 immatures, 7.X.1994, 1♀, 6.XI.1994, 1♀, T. Cekalovic (AMNH). Biobío: Alto Caledonia, 42 km SE Mulchen, 14.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (MACN-Ar). Arauco: 2 km S Cruce Camino Colicó Norte, 20.X.1996, T. Cekalovic, 1♂ (AMNH); 10 km N Curanilahue, 21.XI.1992, T. Cekalovic, 1♀ (AMNH); Isla Mocha, Quebrada La Hacienda, Sendero Travesía, 3.5 km faro Torrecillas, 16.I.1995, T. and S. Cekalovic, 1♀ (AMNH). Región IX: Malleco: Monumento Natural Contulmo, 12.I.1989, M. Ramírez, 5#h, 1♂ (MACN-Ar, photo MJR 42, 43 R2), 19–21.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1♀ (MHNS); Pata de Gallina, S of Contulmo, 5.I.2001, T. Cekalovic, 1♀ (AMNH). Cautín: Monte Verde, Cavahue, 31.I.1993, L. Peña, 1♀ (AMNH); Conguillio Natl. Park, 23.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH); 14 km N Villarrica, elev. 250 m, 39°10′S, 72°12′W, 20.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH). Región X (Los Lagos): Valdivia: Las Lajas, W La Unión, 19–20.XI.1990, L. Peña, 2♂ 1♀ (AMNH); Parque Oncol, road intersection from Renoval Canelos to Los Tepuales, 12.II.2001, T. Cekalovic, 1♀ (AMNH). Osorno: Puyehue Natl. Park, 10.III.1965, H. Levi, 3 immatures (MCZ); Termas de Puyehue, 14.III.1965, H. Levi, 4 immatures (MCZ); 12 km SE Aguas Calientes, P. N. Puyehue, elev. 700 m, 21.XI.1993, N. Platnick, K. Catley, M, Ramírez, T. Allen, 1♂ 1♀ (AMNH), 3♂ (MACN-Ar); Derrumbes road, Aguas Calientes, P. N. Puyehue, elev. 480 m, 40°44′S, 72°18′W, 22.XI.1993, N, Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (MHNS, photo MJR 1415); Aguas Calientes, 13–17.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 4♂ 4♀ 9 immatures (MACN-Ar), 2♂ 3♀ (MHNS); Bosque del Volcán Osorno, 6 km N Ensenada, 250 m, 17.III.1965, H. Levi, 2 immatures (MCZ, photographed). Llanquihue: Chamisa, 13.XII.1968, L. Peña, 1♀ (MCZ); Correntoso, XII.1968, L. Peña, 1♀ (MCZ); El Pangal, Correntoso, 15.XII.1958, L. Peña, 1♀ (MCZ); Alerce Andino Natl. Park, elev. 100 m, 41°35′S, 72°41′W, 23.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH), 1♂ (MACN-Ar). Chiloé: Isla de Chiloé, Chepu, 21.II.1992, M. Ramírez, P. Goloboff, N. Platnick, 3♀ (MACN-Ar), Puente La Caldera, 6.II.1994, T. Cekalovic, 1♀ (AMNH); Isla Quinchao, Quetro, 19.II.1997, T. Cekalovic, 1♀ (AMNH).

    Gayennoides, new genus Table 20

    Type Species:

    Gayennoides molles, new species.

    Etymology:

    The generic name is a derivative of the close relative Gayenna; gender is masculine.

    Diagnosis:

    Males of Gayennoides resemble those of Gayenna, Sanogasta, and Arachosia in having a long, deep canal on the secondary conductor, arising under the paramedian apophysis, but the genus can be distinguished by having a basal notch on the retrolateral margin of cymbium (figs. 65E, 66D).

    Description:

    Carapace narrowed in front, posterior eye row slightly procurved, ocular area not projecting. Chelicerae unmodified, slightly longer in males, with three teeth on promargin, two or three on retromargin. Anterior leg spines unmodified. Male palp with elongate tibia. Cymbium with basal notch on retrolateral margin. Anterior margin of tegulum compressed over secondary conductor. Embolus with basal process flattened, separated by ample ventral membranous area. Paramedian apophysis complex, heavily sclerotized, with one retrolateral cusp, additional prolateral cusp or ridges. Secondary conductor large, striated transversely-obliquely, not fused to anterior margin of tegulum; retrolateral portion with basal prong, apex of prolateral portion prominent, acute; canal very deep, arising under paramedian apophysis. Primary conductor superficial. Epigyne with lateral lobes separate, median field elevated, with ample anterior pouch. Copulatory ducts heavily sclerotized, relatively thick.

    Distribution:

    Central Chile.

    Composition:

    Two species newly described below.

    Gayennoides molles, new species Figures 64B, 65

    Types:

    Female holotype and male paratype from Chile, Región V, Petorca province, Los Molles, 2 m, under succulent rock cover along coast, ca. 32°14′S, 71°30′W, 9.I.1985, N. Platnick, O. Francke, deposited in AMNH.

    Etymology:

    The specific name is a noun in apposition taken from the type locality.

    Diagnosis:

    Similar to G. losvilos, but distinguished by having only two teeth on the cheliceral retromargin, and a larger, rectangular epigynal median field.

    Female (holotype):

    Total length 10.65. Carapace length 4.05, width 2.93, wider on legs II–III. Length of tibia/metatarsus: I, 2.80/2.40; II, 2.63/2.33; III, 2.10/2.27; IV, 2.97/3.46. Palpal tarsus length 1.25. Chelicerae unmodified, with two teeth on retromargin. Sternum length 2.10, width 1.60. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p 0-d1-d1; tibia v r1–2–2, p 0–1; metatarsus v 2bas. III, femur d 1–1–1, p and r 0-d1-d1; patella r 1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r 1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus = III. Abdomen length 5.72, width 3.05, spiracle–epigastrium 2.57, spiracle–spinnerets 0.77. Color (fig. 65A): carapace reddish brown, with lateral dark stripes. Chelicerae reddish brown. Legs pale grayish, with very dark dots at dorsal and lateral spine bases, some spots on patellae and tibiae. Sternum grayish, with tenuous dark spots in front of coxae I–III, median dark spot. Abdomen yellow with dark spots, anterior half of cardiac area gray, two large spots at insertions of posterior dorsoventral muscles, several median spots extending extending to anal tubercle. Venter pale with a few dark dots. Epigyne (fig. 65F, G): median field rectangular. Copulatory ducts joining at posterior margin.

    Male (paratype):

    Total length 7.60. Carapace length 3.75, width 2.80. Length of tibia/metatarsus: I, 3.85/3.35; II, 3.13/2.90; III, 2.30/2.50; IV, 3.17/3.53. Sternum length 2.00, width 1.57. Chelicerae longer than those of female, median promarginal tooth quite larger, closer to apical tooth. Spines as in female, except: leg II, metatarsus p 1–0. III, tibia v p1–2–2 or p1-p1–2. Abdomen length 4.40, width 2.00, spiracle–epigastrium 1.57, spiracle–spinnerets 0.60. Color as in female (fig. 64B). Palp (fig. 65B–E): tibia long, width/length 0.55, cymbium with conspicuous notch at retrolateral basal margin. Copulatory bulb: paramedian apophysis with blunt cusp, two ridges at sides.

    Natural History:

    Many specimens collected under the Crassulaceae lining stones near seashore.

    Distribution:

    Chilean coast, from Antofagasta to Quillota provinces. Also collected in two noncoastal localities, Cuesta La Dormida and Cuesta El Melón, in Region V. This apparently disjunct distribution is similar to that of Acanthoceto ladormida.

    Other Material Examined:

    CHILE: Región II (Antofagasta): Antofagasta: Caleta Hueso, W of Taltal, 28.I.1941, Junius Bird, 1♀ 1 immature (AMNH); 4 km N Paposo, 20–50 m, 11.X.1992, N. Platnick, P. Goloboff, K. Catley, 9♀ 2 immatures (AMNH); Cerro Moreno, 900 m, 18.IV.1992, H. Larrain, 1♀ (AMNH); 6 km E Paposo, 480 m, 12.X.1992, N. Platnick, P. Goloboff, K. Catley, 2♂ 13♀ (AMNH), 520 m, 25°01′S, 70°27′W, N. Platnick, K. Catley, R. Calderón, R.T. Allen, 1♀ (AMNH); Paposo, 54 km N Taltal, II.1957, W. Toodt, 1♀ (MHNS 532); Cerro Moreno, 900 m, 8.IV.1992, H. Larrain, 1♀ (AMNH); Taltal, La Quinta (bajo piedras), 5–7.X.1983, M. Elgueta, 1♀ (MHNS 766). Región III (Atacama): Chañaral: N Chañaral, 10.X.1992, L. Peña, 2♂ 1 immature (AMNH); Isla de Chañaral, ca. 4 km from coast, 30.X.1980, L. Peña, 1♂ (AMNH); Pan de Azúcar, 10–12.X.1992, L. Peña, 1♀ (AMNH), 12.X.1992, L. Peña, 2♂ 1♀ 1 immature (AMNH). Copiapó: Copiapó, 6.V.1964, G. Mann F., 1♀ (MHNS); Puerto Viejo, 15–16.X.1992, L. Peña, 1♂ 2♀ 4 immatures (AMNH), 17.IV.1980, L. Peña, 1♀ (AMNH); Puerto Viejo, S La Caldera, 15–16.X.1992, L. Peña, 3♂ 1♀ 2 immatures (AMNH); E of Puerto Viejo, 9–10.X.1980, L. Peña, 1♀ 2 immatures (AMNH). Región IV (Coquimbo): Elqui: Choros Bajos, 21.X.1992, L. Peña, 1♀ (AMNH); 6 km S Cruz Grande, 20 m, 7.X.1992, N. Platnick, P. Goloboff, K. Catley, 7♀ (AMNH); 6.X.1992, N. Platnick, P. Goloboff, K. Catley, 5♀ (AMNH); 9 km S Cruz Grande, beach, 5 m, 11.XI.1993, 29°29′S, 71°19′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ 1 immature (AMNH), 1♂ 1 immature (MACN-Ar); 16 km S Cruz Grande, 140 m, 7.X.1992, N. Platnick, P. Goloboff, K. Catley, 3♀ (AMNH); Puerto Pichidangui, Isla de los Locos, 19.VII.1961, R. Donoso, A.F. Archer, 1♀ (AMNH); La Serena, 150 m, coastal scrub matorral, 4.XI.1981, N. Platnick, T. Schuh, 1♀ (AMNH); 19 km N La Serena, 150 m, coastal scrub matorral, 1.XI.1980, N. Platnick, T. Schuh, 1♀ (AMNH). Limarí: Fray Jorge Natl. Park, 400 m, 3.X.1992, N. Platnick, P. Goloboff, K. Catley, 1♀ (AMNH); Termas de Socos, 300 m, 2.X.1992, N. Platnick, P. Goloboff, K. Catley, 1♀ (AMNH). Choapa: Huentelauquén, coastal town, 27.XII.1980, L. Peña, 1♂ 1♀ (AMNH), 3.X.1990, L. Peña, 1♀ (AMNH); 20 km N La Serena (Rt. 5 km 491), 120 m, 7.X.1992, N. Platnick, P. Goloboff, K. Catley, 2♂ 4♀ (AMNH); Los Vilos, 25.IX.1966, E.I. Schlinger, 1♂ (CAS); 5 km N Los Vilos, 3 m, under succulent rock cover along cove, 5.I.1985, N. Platnick, O. Francke, 3♀ 5 immatures (AMNH); 19 km N Los Vilos, Rt. 5, km 244, 5 m, 9.XI.1993, 31°45′S, 71°31′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ 1♀ (AMNH); Isla de Los Locos, Pichidangui, 29.IV.1961, R. Donoso, 1♀ 1 immature (AMNH). Coquimbo: Caleta Oscuro, 31°26′S, 71°37′W, 23.IX.1966, E.I. Schlinger, 1♂ 8♀ 1 immature (CAS), 2.X.1983, E. Maury, 3♀ (MACN-Ar); 5 km S Coquimbo, 30 m, coastal scrub matorral, 31.X.1981, N. Platnick, T. Schuh, 1♂ 1 immature (AMNH); Corral de Julio, Quebrada La Madera, 3.X.1972, N. Figueroa, 1♀ (UC); La Herradura, 80 m, 3.X.1992, N. Platnick, P. Goloboff, K. Catley, 1♀ (AMNH). Región V (Valparaíso): Petorca: Los Molles, 3.VII.1975, R. Pérez, 1♂ (UC); Los Molles, Rt. 5, km 188, elev. 10 m, 9.XI.1993, 32°14′S, 71°30′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ 2♀ (AMNH), 2♂ 1♀ 1 immature (MACN-Ar), 1♂ 1♀ (MHNS, photos MJR 1331–1332), 2 m, under succulent rock cover along coast, 9.I.1985, N. Platnick, O. Francke, 5♀ 10 immatures (AMNH); 13 km S Los Molles, 32°12′S, 71°27′W, beach dunes, 22.IX.1966, E.I. Schlinger, 4♀ 2 immatures, 1♀ (CAS); Quebrada del Chivato, 1 km S Los Molles, 10 m, 2.X.1992, N. Platnick, P. Goloboff, K. Catley, 5♀ 3 immatures (AMNH), 30.X.1988, P. Goloboff, E. Maury, C. Szumik, 1♀ 2 immatures (MACN-Ar); Quebrada Huaquén, Pichicuy, 10 m, 2.X.1992, N. Platnick, P. Goloboff, K, Catley, 1♂ 4♀ (AMNH); Cuesta El Melón, 430 m, 8.XI.1993, 32°37′S, 71°14′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH). Quillota: Cuesta La Dormida (east side), 33°04′S, 71°02′W, 750–1000 m, 20.IX.1966, E.I. Schlinger, 1♂ 2♀ (CAS). No Locality: Loc. 280 (presumably from Chile), L. Peña, 1♀ (IRSN IG 19736).

    Gayennoides losvilos, new species Figures 64A, 66

    Types:

    Female holotype from Chile, Región IV, Choapa province, 5 km N Los Vilos, 3 m, under succulent rock cover along cove, ca. 31°55′S, 71°31′W, 5.I.1985, N. Platnick, O. Francke, deposited in AMNH.

    Etymology:

    The specific name is a noun in apposition taken from the type locality.

    Diagnosis:

    Similar to G. molles, but distinguished by having three teeth on the cheliceral retromargin, the male palpal cymbium with a marked retrolateral notch, and a small, procurved epigynal median field.

    Female (holotype):

    Total length 13.30. Carapace length 5.72, width 4.00, wider on legs II–III. Length of tibia/metatarsus: I, 3.72/3.13; II, 3.46/3.00; III, 2.77/2.90; IV, 3.86/4.40. Palpal tarsus length 1.67. Chelicerae unmodified, with three teeth on retromargin. Sternum length 2.93, width 2.07. Spines: leg I, femur d 1–1–1, p 2ap; tibia v p1–2–2; metatarsus v 2bas. II, femur d 1–1–1, p 0-d1-d1, r d1; tibia v r1–2–2, p 0–1; metatarsus v 2bas. III, femur d 1–1–1, p and r 0-d1-d1; patella r 1; tibia v p1–2–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r 1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus = III. Abdomen length 7.70, width 4.40, spiracle–epigastrium 3.70, spiracle–spinnerets 1.45. Color (fig. 64A): carapace brown, reddish to cephalic area, with lateral diffuse dark bands. Chelicerae reddish brown. Legs pale grayish with dark dots at dorsal and lateral spine bases, some spots on patellae and tibiae. Sternum grayish, slightly darker on margins. Abdomen pale gray with dark spots, anterior third of cardiac area gray, posterior third pale, rest of dorsum with diffuse pattern of dark spots. Venter pale with small spots, more concentrated on median stripe. Epigyne (fig. 66F, G): median field elevated, small, short, with ample anterior pouch of hemispheric lumen; two diverging ridges anterior of and at sides of anterior pouch. Copulatory ducts short.

    Male (Carrizal Bajo, not type):

    Total length 11.15. Carapace length 4.50, width 3.50. Length of tibia/metatarsus: I, 4.52/4.00; II, 3.86/3.30; III, 2.83/3.00; IV, regenerated. Chelicerae longer than those of female, median promarginal tooth quite larger, closer to apical tooth. Sternum length 2.87, width 1.83. Spines as in female, except: leg II, femur p 0-p1–2; tibia v 0–2–2. III, tibia v p1-p1–2 or p1–2–2. IV, right absent, left regenerated, with abnormal spines. Abdomen length 6.12, width 3.33, spiracle–epigastrium 2.90, spiracle–spinnerets 1.07. Color as in female. Palp (fig. 66A–E): tibia long, width/length 0.48, cymbium with conspicuous notch at retrolateral basal margin, slightly advanced compared to that of G. molles. Copulatory bulb: embolus with basal process conspicuous (fig. 66C). Paramedian apophysis with retrolateral cusp squared, prolateral cusp with ventral peak. Primary conductor low, triangular.

    Natural History:

    Collected near seashore, under the Crassulaceae lining stones.

    Distribution:

    Known only from seashore of Chile, in Huasco, Choapa, and Elqui provinces.

    Other Material Examined:

    CHILE: Región III (Atacama): Huasco: N of Carrizal Bajo, 22.X.1980, L. Peña, 1♂ (AMNH). Región IV (Coquimbo): Elqui: 6 km S Cruz Grande, 20 m, beach, 5 m, 11.XI.1993, 29°29′S, 71°19′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH, photos MJR 1329–1330); 5 km N Los Vilos, 3 m, under succulent rock cover along cove, 5.I.1985, N. Platnick, O. Francke, 3♀ 1 immature (AMNH); 20 km N La Serena (Rt. 5 km 491), 120 m, 7.X.1992, N. Platnick, P. Goloboff, K. Catley, 2♀ (AMNH).

    Arachosia O.P.-Cambridge Table 21

  • Arachosia O.P.-Cambridge, 1882: 425 (type species by monotypy Arachosia anyphaenoides O.P.-Cambridge, 1882). Keyserling, 1891: 83, 126. Simon, 1897a: 92, 94, 96, 98, 100. F.O.P.-Cambridge, 1900: 93. Mello-Leitão, 1922: 22. Ramírez, 1995a: 381, 1997: 178.

  • Samuza Keyserling, 1891: 83, 134 (type species Samuza praesignis Keyserling, 1891, designated by Petrunkevitch, 1928: 174). new synonymy.

  • Abuzaida Keyserling, 1891: 83, 132 (type species Abuzaida striata Keyserling, 1891, designated by Simon, 1897a: 104). new synonymy.

  • Eusamuza Mello-Leitão, 1915: 144 (type species by original designation Samuza praesignis Keyserling, 1891).

  • Gayennina Gertsch, 1935: 21 (type species by monotypy Gayennina britcheri Gertsch, 1935). new synonymy.

  • Synonymy:

    The type species of Samuza and Abuzaida are here considered typical members of Arachosia. Gayennina britcheri is a junior synonym of Oxysoma cubana, also transferred to Arachosia here. Arachosia corresponds well with the idea that previous authors had of Oxysoma (Simon, 1897a; Mello-Leitão, 1922; Barnes, 1953; Platnick, 1974).

    Note:

    Mello-Leitão (1915) seemed to have committed a lapsus by creating the genus Eusamuza for Samuza praesignis Keyserling.

    Diagnosis:

    Easily distinguished from other Amaurobioidinae by having thick, erect setae on the anterior lateral spinneret bases (compare fig. 68A–F) and a tracheal spiracle relatively advanced. Only some Philisca have similar setae on the spinnerets, but they have a normal, deep anterior epigynal pouch (fig. 94E) instead of the transverse, more superficial pouch found in Arachosia (fig. 75C, D).

    Description:

    Generally with flattened body, partially covered by whitish hairs (fig. 67). Legs well scopulate, even on posterior tibiae. Carapace narrowed in front, posterior eye row strongly procurved. Males with smaller chelicerae, carapace narrower in front, wider behind, legs more spinose. Chelicerae with three teeth on promargin, generally two, occasionally three, on retromargin. Tracheal spiracle around midpoint between spinnerets and epigastric furrow, may be slightly advanced from midpoint, mostly in males. Base of anterior lateral spinnerets with thick erect setae. Male palp with short tibia, large cymbium. Embolus thin, basal process flattened, weakly sclerotized, associated membranous area folded. Paramedian apophysis simple, with small tip close to median apophysis (fig. 70). Secondary conductor large, fused to anterior dorsal margin of tegulum, with deep canal arising at base of paramedian apophysis. Primary conductor absent or reduced. Epigyne with anterior pouch transverse, M- or inverted U-shaped (fig. 69), with shallow or double lumen. Median field weakly sclerotized behind anterior pouch. Insertions of epigastric muscles depressed. Copulatory openings in deep depressions, close to epigastric furrow. Copulatory ducts long, slender, ducts of accessory bulbs long.

    Note:

    Mello-Leitão (1922: 18–19) noted the thick, erect setae on the anterior lateral spinnerets of Oxysoma polytrichium, but believed these to be a diagnostic character of this species only, hence its name.

    Distribution:

    South and Central America, and Southwestern U.S.

    Composition:

    In addition to the species detailed below: Anyphaena oblonga Keyserling, 1878 (female holotype from Mexico, Veracruz, in BMNH 1890.7.1.617, examined, new combination; see also Note under Arachosia cubana), Arachosia albiventris Mello-Leitão, 1922 (7 females and 8 immatures, probably syntypes, of two different Arachosia species, in MNRJ 674, examined), Arachosia arachosia Mello-Leitão, 1922 (male probably holotype, in MNRJ 675, should be 390, examined), Arachosia freiburgensis Keyserling, 1891 (male holotype in BMNH 1890.7.1.406, examined), Arachosia mezenioides Mello-Leitão, 1922 (two females, probably syntypes, in MNRJ 677, should be 41, examined), Arachosia minensis (Mello-Leitão, 1926), Arachosia puta O.P.-Cambridge, 1892 (female holotype in BMNH 1901.3.3.325, examined), Gayenna bonneti Mello-Leitão, 1947 (female holotype in MNRJ, examined, new combination), Gayenna duplovittata Mello-Leitão, 1942 (male immature holotype in MLP 15580, examined, new combination), Gayenna proseni Mello-Leitão, 1944 (female probably holotype in MLP 15115, and female probably paratype in MNJR, examined, new combination), Oxysoma bifasciatum Mello-Leitão, 1922 (male holotype in MNRJ 662, examined, new combination), Oxysoma dubium Berland, 1913 (three penultimate syntypes [one male, two females], in MHNP, examined, new combination), Oxysoma polytrichium Mello-Leitão, 1922 (penultimate female holotype, pharate, in MZUSP 541 ex 721, examined, new combination). Many probable undescribed species similar to A. bergi or A. cubana, some may be intraspecific variants instead.

    Type Not Examined:

    Arachosia sulfurea Mello-Leitão, 1922 (presumably in MNRJ, not found).

    Arachosia anyphaenoides O.P.-Cambridge

  • Arachosia anyphaenoides O.P.-Cambridge, 1882: 426 (female holotype from Brazil, Amazonas, Prof. Trail coll., probably in BMNH, not found). Simon, 1897a: 101. Mello-Leitão, 1922: 22.

  • Note:

    I could not examine the type, but from the illustrations in the original description it is clear that this species is closely related to (and perhaps a senior synonym of) A. honesta Keyserling.

    Arachosia praesignis (Keyserling), new combination Figures 61A, 67A, 69B, 71, 72

  • Samuza praesignis Keyserling, 1891: 135 (female holotype from Brazil, state of Rio Grande do Sul, no specific locality, V. Ihering coll., in BMNH, examined).

  • Gayenna praesignis: Petrunkevitch, 1911: 485. Mello-Leitão, 1925: 457. Di Caporiacco, 1948: 678.

  • Eusamuza praesignis Mello-Leitão, 1915: 144 (see Note under Arachosia).

  • Tomopisthes tripunctatus Mello-Leitão, 1945: 265 (female holotype from Argentina, Corrientes province, Manantiales, no date, Apóstol, MLP 16595, examined). new synonymy.

  • Synonymy:

    The holotypes of the species synonymyzed were compared; no relevant differences were found.

    Diagnosis:

    Distinguished from other Arachosia by the absence of a prolateral projection on the male secondary conductor (fig. 71A), the narrow anterior epigynal pouch, and the characteristic course of the female copulatory ducts (fig. 71C).

    Female (holotype):

    Total length 6.65. Carapace length 2.67, width 2.00. Chelicerae with two teeth on retromargin. Length of tibia/metatarsus: I, 1.30/1.17; II, 1.28/1.18; III, 1.07/1.08; IV, 1.52/1.67. Spines: leg I, femur d 1–1–1, p 0-d1-(1d1), r 0-d1-d1; tibia v 2–2–2, d r1–1 bristles; metatarsus v 2bas. II, femur = I; tibia v 2–2–2, p 0–1, d r1–1 bristles; metatarsus = I. III, femur = I or p (2-d1)ap; patella r d1; tibia v p1–2–2, p 1-d1–1, r d1–1, d r1–1; metatarsus v 2–0–2, p and r d1-d1–1, d 0-p1–2. IV, femur d 1–1–1, p (1-d1)ap, r d1ap; patella r d1; tibia v p1–2–2, p and r 1-d1–1, d r1–1; metatarsus v 2–2–2 or 2-r1–2, p and r d1-d1–1, d 0-p1–2. Abdomen length 4.12, width 2.43. Spiracle–epigastrium 1.50, spiracle–spinnerets 1.27. Color: type faded, anterior part of dorsal abdominal stripe remains. In fresh specimens (fig. 67A), pattern similar to that of male, but paler: lateral stripes on carapace narrow, on abdomen discontinuous; venter pale. Entire abdomen with white guanine reticulum. Legs yellow with brownish violet dots. Epigyne (figs. 69B, 71C): anterior pouch transverse, inverted U-shaped, with double lumen. Copulatory ducts long, thin, ducts of accessory bulbs long, parallel.

    Male (Pelotas):

    Total length 2.70. Carapace wide, globose, with thoracic groove depressed, length 2.80, width 2.33. Length of tibia/metatarsus: I, 2.17/1.93; II, 2.03/1.83; III, 1.60/1.47; IV, 2.03/2.13. Chelicerae small, narrow. Sternum length 1.38, width 1.08. Spines as in female, except: leg I, tibia p and r 1-d1-1-0, d r1-0-1-0; metatarsus p and r (d1–1)bas, d 0-p1-0-2. II = I. III, tibia = I, but v p1–2–2; metatarsus v 2–2–2. IV, femur = I; tibia and metatarsus = III. Abdomen length 3.00, width 1.73, spiracle–epigastrium 1.27, spiracle–spinnerets 0.77. Color: carapace brown with three dark brown longitudinal stripes, one median, one on each margin, ocular area black, with median longitudinal brown spot. Clipeum with two dark stripes diverging toward chelicerae, and white triangle in between. Legs grayish with dark brown longitudinal lines, spots. Chelicerae with anterior internal dark spot, extending up to two-thirds of their length. Coxae, endites, and sternum yellow, labium brown. Abdomen yellow, with brownish violet pattern on sides, cardiac area, dorsoventral muscle insertions, plus several chevrons on posterior half, ending in spot above anal tubercle. Venter pale, with diffuse dark band from spiracle to spinnerets. Palp (figs. 71A, B, 72): tibia short, as long as wide, cymbium globose. Embolus thin, basal process flattened, weakly sclerotized, associated membranous area extended, folded. Paramedian apophysis simple, straight, parallel to median apophysis. Primary conductor absent. Secondary conductor large, striated obliquely, fused to anterior margin of tegulum; canal conspicuous, arising at base of paramedian apophysis.

    Variability:

    Males can be much darker than females, with entire dorsal and ventral median abdominal bands, and the sternum with dark margins. One specimen from Buenos Aires is completely brownish violet, with two small white spots at sides of the ocular area, white endites, and yellow pulmonary plates. Spines: tibiae III, IV, v 2–2–2.

    Natural History:

    This species builds retreats on foliage, occasionally also under bark.

    Distribution:

    Southeastern Brazil and northeastern Argentina (in Misiones, Chaco, Corrientes, Entre Ríos, and Buenos Aires provinces).

    Other Material Examined:

    BRASIL: Rio de Janeiro: Nova Iguaçu, no date, Blanc, 4♀ (MNRJ). Rio Grande do Sul: Cachoeira do Sul, Alto dos Casemiros, 26.IX.1992, R. Buss, 1♀ 1 immature (MCTP 3369); Cachoeira do Sul, Cordilheira, 27.VIII.1992, 2♂ (MCTP 3366), 27.X.1992, 1♀ penultimate (MCTP 3358), 1♂ 1♀ (MCTP 3364), R.G. Buss; Erval Grande, I.1994, A. Braul, 1♀ (MCTP 4484); Pelotas, XII.1960, C. Biezanko, 3♂ 3♀ 1 immature (AMNH); no specific locality or date, B. Rambo, 1♂ 12 immatures (MNRJ 41661). ARGENTINA: Misiones: Loreto, no date, A.A. Ogloblin, 1♂ (MACN-Ar); Iguazú Natl. Park, Cataratas, XI.1989, M. Ramírez, 1♂ 1♀ (MACN-Ar); Puerto Rico, XII.1943, J.M. Viana, 1 immature (MACN-Ar); Refugio Piñalito, XI.1954, R. Schiapelli and M.E. Galiano, 1♀ (MACN-Ar); Río Uruguaí, km 30, W. Partridge, 1♂ (MACN-Ar); Ruta Nac. 11 and Arroyo Garuhapé, VII.1980, P. Goloboff, 1♀ (MACN-Ar); San Ignacio, 1.IX.1963, M.E. Galiano, 1 immature (MACN-Ar); San Javier, XII.1948, M. Birabén, 2♂ 3♀ (MLP); Santa María, IX.1956, J.M. Viana, 2♀ (MACN-Ar); XII.1947, J.M. Viana, 2♂ (MACN-Ar 2554). Chaco: Resistencia, X.1943, Freiberg, 1♀ (MACN-Ar). Catamarca: Estancia El Chorro, 20–31.I.1953, W. Partridge and Núñez, 1♀ (MACN-Ar), 1–15.II.1953, W. Partridge and Núñez, 1♀ (MACN-Ar). Corrientes: Paso de la Patria, 29.VIII.1963, M.E. Galiano, 1♀ (MACN-Ar); Santiago Alcorta, VI.1943, M. Birabén, 1♀ (MACN-Ar). Entre Ríos: Concordia, no date, Hayward, 1♀ (MACN-Ar); El Palmar Natl. Park, 14.X.1984, M. Ramírez, 2♂ 1♀ penultimate (MACN-Ar); Villaguay, XI.1988, M.E. Galiano, 1♀ (MACN-Ar); no specific locality, 1942, Haedo, 1♀ (MACN-Ar). Buenos Aires: Atucha, 27.VII.1985, P. Goloboff, M. Ramírez, 1♂ 1 immature (MACN-Ar); Boulogne, X.1938, A. Prosen, 1♀ (MLP); Capital Federal, Ciudad de Buenos Aires, IV.1940, F. Monrós, 1♀ (MACN-Ar); X.1940, H. Gavio, 1 immature (MACN-Ar); I.1952, G. Casal, 4♀ (MACN-Ar), 1.IV.1983, E. Maury, 1♀ (MACN-Ar), 19.IX.1990, M. Ramírez, 1♀ (MACN-Ar, photo MJR 353–357), 15.VIII.1998, M. Ramírez, 1♂ (MACN-Ar); winter 1979, P. Goloboff, 1♀ (MACN-Ar); Caseros, X.1947, no collector, 1♀ (MACN-Ar 196); Castelli, X.1960, J.M. Viana, 1♀ penultimate (MACN-Ar); Delta, Arroyo Carancho, 30.XII.1951, A. Bachmann, 1♀ (MACN-Ar); Delta del Paraná, 20.XI.1940, J.B. Daguerre, De Carlo, 1♀ (MACN-Ar 32904); VII.1940, F. Monrós, 1♀ (MACN-Ar); Delta, Paraná de Las Palmas, III.1942, H. Hepper, 1 immature (MACN-Ar); Escobar, V.1938, A. Prosen, 1♂ (MLP); Estación San Alfonso, Pieres, II.1973, Bejarano, 1♀ (MACN-Ar); Florencio Varela, F.C.S., XII.1939, F. Monrós, 1♂ 1♀ 1 immature (MACN-Ar); Hudson, 2.X.1984, M. Ramírez, 1♂ (MACN-Ar); Ing. Maschwitz, XI.1941, A. Prosen, 1♀ (MLP); Isla Martín García, I.1938, J.M. Viana, 1♀ (MACN-Ar 326); IV.1938, J.M. Viana, 1 immature (MACN-Ar 404), 1940, J.M. Viana, 1♀ (MACN-Ar); 25.V.1990, M. Ramírez, 8 immatures (MACN-Ar); La Plata, 22.XII.1978, P. Goloboff, 1♂ (MACN-Ar); no date, Birabén, 1♀ (MACN-Ar); Los Talas, XII.1985, Scioscia, 1♂ (MACN-Ar); Mar del Plata, 4.XI.1988, J. Farina, 2♀ 1 immature (MMLS); Necochea, V.1975, Balech, 1♀ (MACN-Ar); Reserva Otamendi, 10.VI.1997, M. Ramírez, L. Compagnucci, C. Grismado, F. Uehara, 1♂ penultimate (MACN-Ar); Paraná de Las Palmas, 17.IX.1963, M.E. Galiano, 1♂ (MACN-Ar), A. Bachmann, 1 immature (MACN-Ar); Punta Lara, Ensenada, III.1943, A. Moreno, 1♀ (MLP), 19.III.1943, 4♀ (MLP), 1.I.1947, W. Partridge, 1♂ (MACN-Ar), 16.XI.1947, no collector, 2♀ (MACN-Ar 198); III.1961, M.E. Galiano, 1♀ (MACN-Ar); 28.VII.1979, P. Goloboff, 1♀ (MACN-Ar); 28.XI.1985, M.E. Galiano, C. Scioscia, 1♂ (MACN-Ar), 18.IX.1986, M. Ramírez, 1♀ penultimate (MACN-Ar); San Fernando, no date, J.B. Daguerre, 1♀ 1 immature (MACN-Ar); San Isidro, Punta Chica, 5.XI.1941, A. Prosen, 2♀ (MLP); Sierra de la Ventana, Cerro Negro, 12.IV.1974, Cesari, 1 immature (MACN-Ar); Tandil, 1939, S. Holmberg, 2♀ (MACN-Ar 2614); Tigre, I.1938, J.M. Viana, 1♀ (MACN-Ar 301).

    Arachosia striata (Keyserling), new combinationFigure 73

  • Abuzaida striata Keyserling, 1891: 133 (female lectotype here designated from Brazil, state of Rio de Janeiro, Nova Friburgo, in BMNH, examined; the immature paralectotype mentioned by Keyserling was not found). Simon, 1897a: 104.

  • Gayenna striata: Mello-Leitão, 1925: 457.

  • Diagnosis:

    Resembles A. bergi in having the epigynal median field relatively small and short, but can be distinguished by having the shorter copulatory ducts.

    Female (lectotype):

    Total length 5.30. Carapace length 2.17, width 1.60. Length of tibia/metatarsus: I, 1.12/0.97; II, 1.17/1.00; III, 1.05/1.05; IV, 1.37/1.63. Chelicerae with three teeth on retromargin. Spines: leg I, femur d 1–1–1, p 0-d1–2, r 0-d1-d1; tibia v 2–2–2; metatarsus v 2bas. II = I. III, femur = I; patella r d1; tibia v p1–2–2, p 1-d1–1, r d1–1, d r1–1; metatarsus v 2–0–2, d 0-p1–2, p and r d1–1–1. IV, femur = I; patella r d1; tibia = III; metatarsus v 2-p1–2, d 0-p1–2, p and r d1–1–1. Abdomen length 2.83, width 1.67. Spiracle–epigastrium 1.10, spiracle–spinnerets 0.80. Color (lectotype faded; from MCTP 0481): yellowish brown, with median dorsal brown band on carapace and abdomen, ocular area darker. Chelicerae brown, with darker anterior distal area. Sternum with three dark patches in front of coxae I–III. Legs yellow with brown dots, mostly on femora and tibiae. Abdomen with brown ventral longitudinal band, brown dots on sides. Epigyne (fig. 73): anterior pouch transverse, shallow. Copulatory ducts relatively short, ducts of accessory bulbs long, arched.

    Male:

    Unknown.

    Natural History:

    Unknown.

    Distribution:

    Only known from the type locality.

    Other Material Examined:

    BRASIL: Rio Grande do Sul: Barracão, 20.I.1989, F. Franco, 1♀ (MCTP 0481).

    Relationships:

    This species was not included in the analysis, because males are unknown, and for the remaining characters it is practically identical to the other representatives of the genus. The relatively short copulatory ducts (not scored in this analysis) suggest a basal position relative to other Arachosia.

    Arachosia honesta Keyserling Figure 74

  • Arachosia honesta Keyserling, 1891: 127 (female holotype from Brazil, state of Rio Grande do Sul, no specific locality, in BMNH, examined). Mello-Leitão, 1922: 22.

  • Oxysoma ramboi Mello-Leitão, 1943c: 238 (female presumably type, from Brazil, Rio Grande do Sul, B. Rambo coll., in MNRJ 42237, examined). new synonymy.

  • Synonymy:

    According to the original description, the holotype of Oxysoma ramboi should be 41379. In any case, the illustration of the epigyne (Mello-Leitão, 1943c: fig. 64) allows reliable identification. No relevant differences were found between the presumed type of Oxysoma ramboi and the holotype of Arachosia honesta.

    Diagnosis:

    Distinguished from other Arachosia by having a large, triangular epigynal median field and a characteristic course of the copulatory ducts. Males have genitalia very similar to that of A. bergi, but differ by having the AME larger than the ALE, and a larger paramedian apophysis. Also distinguished from A. praesignis by having a prolateral projection on the secondary conductor.

    Female (MNRJ 42237):

    Total length 8.25. Carapace length 3.23, width 2.57. Chelicerae with three teeth on retromargin (fig. 74E). Length of tibia/metatarsus: I, 1.83/1.57; II, 1.86/1.63; III, 1.63/1.60; IV, 2.30/2.43. Palpal tarsus length 1.03. Sternum length 1.67, width 1.27. Spines: leg I, femur d 1–1–1, p 0-p1–2, r 0-p1-p1; tibia v 2–2–2; metatarsus v 2bas. II, femur = I; tibia v 2–2–2, p 1-0-1-0; metatarsus v 2 bas, p 1 or 0. III, femur = I; patella r d1; tibia v 2–2–2, p and r 1-d1–1, d r1–1–0; metatarsus v 2–0–2 (right 2–0–1), p and r d1-1-0-1, d p1–2. IV, femur = I; patella r d1; tibia = III; metatarsus v 2–2–2, p and r d1-1-0-1, d p1–2. Abdomen length 4.92, width 2.79. Spiracle–epigastrium 1.33, spiracle–spinnerets 1.73. Color (slightly faded): pale brown, carapace and abdomen with dorsal median dark band, margins of carapace with dark line. Sternum with margins dark. Femora with dark dots. Some specimens darker, with dark brown areas at sides of abdomen, median dark band from spiracle (or epigastrium) to spinnerets. Epigyne (fig. 74F): median field hourglass-shaped, anterior half larger, wider; anterior pouch wide, transverse, forming cavities at lateral ends. Epigastric muscle insertions deeply depressed. Copulatory ducts long, thin, ducts of accessory bulbs long, parallel.

    Male (Iguazú, MACN-Ar 9823):

    Total length 6.90. Carapace globose, thoracic groove on depressed area, length 3.27, width 2.70. Length of tibia/metatarsus: I, 2.30/2.07; II, 2.30/2.03; III, missing; IV, 2.43/2.60. Chelicerae slightly narrower than those of female. Sternum length 1.63, width 1.27. Spines as in female, except: leg I, femur d 1–1–1, p 0-d1–2, r 0-d1-d1; patella r d1, d 1–0–1; tibia v 2–2–2, p and r 1-d1-1-0, d r1-0-1-0; metatarsus v 2bas, p and r d1–1–0, d 0-p1–2. II, femur = I or r 0-d1–2; patella, tibia and metatarsus = I. III absent. IV, femur, patella, and tibia = I; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. Abdomen length 3.59, width 2.13, spiracle–epigastrium 0.87, spiracle–spinnerets 1.33. Color: carapace brown with dark brown stripes (fig. 74A). Legs grayish, almost totally covered by brownish violet pattern. Abdomen brownish violet, dorsum yellow with median band brownish violet. Palps pale gray from femur to tibia, endites, labium, and sternum brownish violet, sternum with median cream patch. Chelicerae dark, with oblique dark band and pale apical internal area. Palp (fig. 74B–D): tibia short, width/length 0.88, cymbium globose. Embolus thin, basal process flattened, weakly sclerotized, associated membranous area ample, folded. Median apophysis short. Paramedian apophysis very short, apex close to tip of median apophysis. Secondary conductor large, striated obliquely, fused to anterior margin of tegulum, with conspicuous canal, arising at base of paramedian apophysis; prolateral portion with conspicuous finger-shaped apophysis (fig. 74D). Primary conductor absent.

    Variability:

    May have three or two teeth on cheliceral retromargin (only one female from São Paulo MZUSP 10167 with four). Some males with longer median and paramedian apophysis, prolateral projection on conductor.

    Natural History:

    This species builds retreats on foliage.

    Distribution:

    South and southeastern Brazil, states of Rio de Janeiro, São Paulo, and Rio Grande do Sul (probably also in Paraná and Santa Catarina), Mato Grosso, and northeastern Argentina, in Misiones province.

    Other Material Examined:

    BRASIL: Mato Grosso: Utiariti, VII.1961, Lenko, 1♀ (MZUSP DZ3551). São Paulo: Cocaia, 25.XII.1949, H. Urban, 1♀ (MZUSP 10167); Pirassununga, Baguaçú, 28.V.1949, O. Schubart, 1♀ 2 immatures (MZUSP 7094); Pirassununga, Emas, 2.X.1948, O. Schubart, 2♀ (MZUSP DZ7594), 15.II.1949, O. Schubart, 1♀ 2♂ (MZUSP DZ7595); Onda Verde, Fazenda São João, I.1946, E. Leme, 1♀ (MZUSP 14053). Rio Grande do Sul: Pinhal, 40 X Ar, I.1949, A. Maller, 1♂ (AMNH). ARGENTINA: Misiones: Iguazú Natl. Park, XI.1948, M. Birabén, 1♀ (MACN-Ar), X.1985, P. Goloboff, 1♀ (MACN-Ar), Cataratas, 30.VIII.1986, M. Ramírez, 1♂ (MACN-Ar 9823). Mistaken Locality: Río Negro, El Bolsón, 1965–1966, A. Kovács, 1♂ (AMNH), probably from Argentina, Misiones province (see Platnick and Ewing, 1995: 7).

    Arachosia bergi (Simon), new combination Figures 67B, 68D–F, 69A, 70, 75

  • Phidyle bergi Simon, 1880: 345 (four males and three females probably syntypes, labeled “mission B. Aires, Paraguay”, in MHNP 4013, examined).

  • Oxysoma bergi: Simon, 1897a: 100. Berland, 1913: 103.

  • Note:

    According to the original publication (Simon, 1880: 346), the type series comes from “territorio des Missions (coll. E. Simon, reçu du Dr C. Berg)” and should include at least one adult male and one penultimate female (“epigyne non developée”), which disagrees with the three vials found in MHNP (4013 referred before; 22736 from La Plata, Silvestri coll., one male; 21201 from Buenos Aires, one female, and one female of A. cf. cubana). The identity of the types is hence problematic. I decided to identify provisionally as syntypes the sample in MHNP 4013, which better agree with the type locality as was published. It is not clear, however, that the forms identified here as A. bergi are different from A. cubana, because there are many specimens with variants of epigynal shapes, including intermediates. Wide variability is also found in specimens from a single locality. Variability in the male copulatory bulb seems to be even more problematic. Only those specimens most similar to figure 75 are listed below.

    Diagnosis:

    Provisionally distinguished from the very similar A. cubana by having a narrower epigynal median field, and thinner, shorter paramedian apophysis. A. striata is similar in having a similarly shaped median epigynal field but differs by the course of copulatory ducts.

    Female (MHNP 4013):

    Carapace length 3.60, width 2.83. Ocular diameters: AME 0.13, ALE 0.19, PME 0.15, PLE 0.15. Chelicerae with two teeth on retromargin. Length of tibia/metatarsus: I, 2.37/2.07; II, 2.17/1.97; III, 1.80/1.77; IV, 3.00/3.00. Spines: leg I, femur d 1–1–1, p 2ap, r 0-d1-d1 or d1; tibia v 2–2–2, p 1-d1–1; metatarsus v 2bas. II, femur d 1–1–1, p d2ap, r 0-d1-d1; tibia = I; metatarsus = I. III, femur = II; patella r d1; tibia v p1–2–2 or 2–2–2, p and r 1-d1–1, d r1-(1 bristle); metatarsus v 2–0–2, p and r d1-1-0-1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d2 or d2ap, r d1ap; patella r d1; tibia = III; metatarsus v 2–2–2 p and r p1-1-0-1, d 0-p1–2. Metatarsus III with scopula. Abdomen length 6.40, width 2.43. Anterior spinnerets with thick setae. Spiracle–epigastrium 2.07, spiracle–spinnerets 2.40. Color as in male. Epigyne (figs. 69A, 75C, D): anterior pouch transverse, wide, M-shaped. Copulatory ducts long, ducts of accessory bulbs long, sinuous.

    Male (MHNP 4013):

    Carapace length 3.80, width 2.97. Ocular diameters: AME 0.14, ALE 0.19, PME 0.15, PLE 0.15. Chelicerae slightly smaller than those of female. Length of tibia/metatarsus: I. 3.17/3.10; II, 2.83/2.77; III, 2.27/2.10; IV, 3.33/3.40. Spines as in female, except: leg I, femur r 0-d1-d1; tibia d r1–1; metatarsus p and r d1-1-0-0, d p1–0. II, tibia and metatarsus = I. III, patella d 1–0–1; tibia = I. IV, femur p d2ap; patella d 1–0–1; tibia = I. Metatarsus III with scopula. Abdomen length 5.00, width 1.50. Anterior spinnerets with thick setae. Spiracle–epigastrium 1.50, spiracle–spinnerets 1.87. Color (fig. 67B): pale brown, legs darkening distally, with small darker spots. Dorsal darker band diffuse on carapace, fading on abdomen. Three small brown spots between median band and margins of carapace. Abdomen with dark posterior dots, venter and sternum pale. Palp (figs. 75A, B, 70): tibia short, width/length 0.81, cymbium globose. Embolus thin, basal process flattened, weakly sclerotized, associated membranous area ample, folded. Paramedian apophysis very short, base flattened, close to median apophysis. Primary conductor triangular, flattened, arising from same sclerotized piece as paramedian apophysis. Secondary conductor large, fused to anterior margin of tegulum, with conspicuous canal arising close to base of paramedian apophysis; prolateral portion with conspicuous finger-shaped apophysis.

    Variability:

    Some females have only faint sutures between median field and lateral lobes, and shallow anterior pouch. The dorsal band may be solid or double. The posterior half of abdomen may become gradually brown uniform. Some specimens from São Paulo, Botucatu, have median band weakly marked. The venter may have a diffuse band.

    Natural History:

    Very common in grasslands and periodically flooded areas. Females make retreats on “serrucheta” (Eryngium spp.), the large pampa grass (“cortadera”, Cortaderia selloana), or between the leaf bases of regular grasses. Occasionally they try to escape collection by diving into the water accumulated between Eringyum leaves. They are exasperatingly fast.

    Distribution:

    Southeastern Brazil (states of São Paulo and Rio Grande do Sul, probably also in Paraná and Santa Catarina), Uruguay, and Northeastern Argentina, extending up to Buenos Aires province. Sympatric with Arachosia cubana (or similar forms).

    Other Material Examined:

    BRASIL: São Paulo: Botucatu, Distrito Vitoriana, Fazenda Goldfarm C., 5.XI.1987b, I. Rinaldi and L. Forti, 1♀ (UNESP); Botucatu, Usina São Manoel, 10.XII.1986, 1♂, 4.VIII.1987b, 1♂, 5.XI.1987b, 1♂, 6.XI.1986b, 2♂ 1♀, I. Rinaldi and L. Forti (UNESP); Botucatu, Fazenda Nossa Senhora da Conceição, “parte aérea de Saccharum officinarum”, 23.IX.1998, L.I. Rinaldi, B. Mendez do P, 1♂ 1♀ (UNESP); Botucatu, Sítio Novelli, 9.IX.1987b, 1♂, 8.VI.1988b, 2♀, I. Rinaldi and L. Forti (UNESP); Botucatu, 23.IX.1998, I. Rinaldi, 1♂ 1♀ (UNESP). Rio Grande do Sul: Pelotas, 27.III.1956, C. Biezanko, 1♀ (AMNH). URUGUAY: Departamento Rocha: Laguna de Castillos, 19.V.1993, M. Ramírez and F. Pérez-Miles, 1♀ (MACN-Ar). ARGENTINA: Formosa: Pilcomayo Natl. Park, Laguna Blanca, XI.1990, M. Ramírez, 1♀ (MACN-Ar); Puerto Pilcomayo, V.1943, Fourcade (?), 1♀ (MLP). Corrientes: Arroyo Mandisoví Grande and Ruta Nac. 14, 15.VII.1985, M. Ramírez, 2♂ (MACN-Ar). Tucumán: ruta 307, 10 km NW El Indio, 24.XI.1994, M. Ramírez and P. Goloboff, 1♀ (MACN-Ar). Entre Ríos: Arroyo Brazo Largo, 16.XI.1979, P. Goloboff, 1♂ (MACN-Ar). Buenos Aires: Allen, VIII.1945, Cuccioli, 1♀ (MACN-Ar); Atucha, IV.1982, P. Goloboff, M. Ramírez, 1♂ (MACN-Ar); 23.VI.1985, P. Goloboff and M. Ramírez, 2♂ (MACN-Ar), 15.IX.1990, M. Ramírez, 1♀ (MACN-Ar); Delta, no date, no collector, 1♀ (MACN-Ar 36228); Delta, Abra Vieja, V.1944, F. Monrós, 3♀ (MACN-Ar); Delta, Arroyo Carancho, on Eryngium pandanifolium, 30.XII.1951, A. Bachmann, 1♂ (MACN-Ar); Delta, Arroyo Carreras, VIII.1941, F. Monrós, 6♂ 5♀ (MACN-Ar 908); Delta, Canal Arias, VI.1941, F. Monrós, 1♂ (MACN-Ar); Paraná de Las Palmas, 19.XII.1963, A. Bachmann, 1♀ (MACN-Ar); Delta, Río Esperita, X.19?? (illegible), F. Monrós, 1♀ (MACN-Ar); Delta, Río Luján, 9.VI.1941, F. Monrós, 1♂ (MACN-Ar); Delta, Tigre, Río Luján and Arroyo Guayracá, VI.1982, M. Ramírez, 1♂ 1♀ (MACN-Ar); Dique Luján, VI.1938, F. Monrós, 1♂ (MACN-Ar); Escobar, 23.VII.1984, M. Ramírez, 1♀ (MACN-Ar); Estancia El Tonelero, Pdo. Gral. Lavalle, cerca canal 2, 15–21.XII.1951, J. Cranwell, 1♀ (MACN-Ar); Hudson, VIII.1982, P. Goloboff, M. Ramírez, 1♂ 1♀ (MACN-Ar); IV.1984, M. Ramírez, 1♂ (MACN-Ar); 2.IX.1984, M. Ramírez, 1♀ (MACN-Ar), 13.XI.1988, M. Ramírez, 1♂ (MACN-Ar, photos MJR 257, 258), 1♀ (MACN-Ar, photos MJR 259, 260), 1♀ (MACN-Ar); Isla Talavera, Las Palmas, FCGM, 2.XI.1980, P. Goloboff, A. Zanetic, 1♀ (MACN-Ar); Magdalena, no date, P. Goloboff, M. Ramírez, 1♀ (MACN-Ar), 20.V.1989, M. Ramírez, 1♀ (MACN-Ar, photo MJR 287); Otamendi, 5.IX.1980, A. Zanetic, P. Goloboff, 1♀ (MACN-Ar); Reserva Otamendi, 10.VI.1997, M. Ramírez, L. Compagnucci, C. Grismado, F. Uehara, 1♂ (MACN-Ar); Paraná de Las Palmas, 7.IV.1963, M.E. Galiano, 1♀ (MACN-Ar), Punta Lara, Ensenada, 2.VIII.1931, J.B. Daguerre, 1♂ (MACN-Ar 27594); 4.XII.1981, F. Miranda, M. Ramírez, 5♀ (MACN-Ar); 6.III.1982, F. Miranda, M. Ramírez, 1♂ 1♀ (MACN-Ar); Río Luján, estación F.C.G.M., marsh with “espadaña”, 5.X.1993, M. Ramírez and A. Pérez, 1♀ (MACN-Ar); San Isidro, VI.1962, A. Martínez, 1♀ (MACN-Ar); San Pedro, 2.XI.1991, M. Ramírez, 2♀ (MACN-Ar); Tigre, IX.1945, J.M. Viana, 1♂ (MACN-Ar); VI.1955, J.M. Viana, 1♀ (MACN-Ar).

    Arachosia cubana (Banks), new combination

  • Oxysoma cubana Banks, 1909b: 157 (male holotype from Cuba, La Habana, in MCZ, examined). Bryant, 1940: 435. Kaston, 1948: 405. Buchkovich, 1995: 13. Platnick, 1974: 260. Oxysoma cubanum: Platnick, 1993: 596 (emendation of O. cubana Banks).

  • Gayennina britcheri Gertsch, 1935: 21 (female holotype from Woods Hole, Massachusetts, in AMNH, examined). Kaston, 1948: 405. Synonymized by Barnes, 1953: 18.

  • Note:

    The North American specimens in AMNH are very similar to Arachosia bergi as provisionally identified here, differing by the wide, V-shaped epigynal median field, and by the larger paramedian apophysis and prolateral projection on the secondary conductor (Platnick, 1974: figs. 105109). However, there are many intermediate or slightly different forms, some of them sympatric. In South America, specimens similar to the Cuban and North American forms were found in Venezuela, Ecuador, and Peru, and in Argentina, from Salta and Tucumán provinces, to Neuquén and the coast of Chubut. Anyphaena oblonga Keyserling, 1878 has an epigyne very similar to that of North American Oxysoma cubana, and hence that name may turn out to be a senior synonym.

    Description:

    See Platnick (1974).

    Sanogasta Mello-Leitão Table 22

  • Sanogasta Mello-Leitão, 1941a: 177 (type species by original designation Sanogasta intermedia Mello-Leitão, 1941). Transferred from the Corinnidae by Ramírez, 1995a: 381, 1997: 178.

  • Note:

    Sanogasta corresponds broadly with the concept that Simon and Tullgren had of Gayenna and Tomopisthes (Ramírez and Kochalka, 1993). In this analysis Sanogasta is paraphyletic in terms of Arachosia, because of the placement of S. pehuenche. Because there are many additional species of Sanogasta and Arachosia not included here, it seems premature to create a new genus only for S. pehuenche, which is instead provisionally placed in Sanogasta.

    Diagnosis:

    Sanogasta resembles Arachosia in having a slender paramedian apophysis associated with the median apophysis (figs. 78C, 82A), but it can be distinguished by lacking thick setae on the anterior lateral spinnerets.

    Description:

    Carapace narrowed in front, posterior eye row slightly procurved or straight. Chelicerae with three teeth on promargin (except S. x-signata, with four), two on retromargin. Males usually with smaller chelicerae, carapace narrower in front and wider behind. Tracheal spiracle closer to spinnerets than to epigastrium. Male copulatory organ with thin embolus bearing small, acute basal process. Median apophysis small, slender, associated with paramedian, very short in S. maculatipes and closest relatives (clade 167). Paramedian apophysis with membranous area dividing part or all of its base, tip slender. Primary conductor absent. Secondary conductor not fused to anterior dorsal margin of tegulum, with deep, long canal arising at base of paramedian apophysis (except S. pehuenche and S. approximata, with canal reduced and secondary conductor fused to tegulum); retrolateral portion with basal prong of variable shape. Epigyne with insertions of epigastric muscles depressed (except S. pehuenche, S. x-signata, and S. tenuis). Copulatory openings in deep depressions on epigastric fold, ducts of accessory bulbs very short.

    Distribution:

    South America.

    Composition:

    In addition to the species detailed below: Gayenna bonariensis Mello-Leitão, 1940 (female holotype in MLP 14400, examined, new combination); Gayenna paucilineata Mello-Leitão, 1945 (male immature holotype in MLP 16.590, examined, new combination); Gayenna rufithorax Tullgren, 1902 (male holotype in NRS, examined, new combination). Several probable undescribed species very similar to S. maculosa, S. maculatipes, S. minuta, and S. backhauseni, some of which may be intraspecific variants instead.

    Nomina Dubia:

    Anyphaena pampa, Holmberg 1881: 145 (female holotype from Argentina, Buenos Aires province, near Sierra de La Ventana, lost). The body pattern and the simple sketch of the epigyne illustrated by Holmberg (1881: fig. 7a, b) are reminiscent of Monapia fierro or M. carolina, but the three pairs of ventral spines on tibia I differ from the four to six pairs found in those species. Clubiona gemella Nicolet, 1849 (several immatures syntypes from Chile, no specific locality, in MHNP 4238, examined, similar to Sanogasta maculosa).

    Sanogasta maculatipes (Keyserling), new combination Figures 61B, 76A, 77E, 78A, B, D, E, 79A, 80A–C, 81D, E

  • Anyphaena maculatipes Keyserling, 1878: 603 (female holotype from Uruguay, in BMNH, examined; also an immature Josa in the same vial, see Note below).

  • Anyphaena argentina Holmberg, 1881: 141 (no type designated, described from two females and one immature female from Buenos Aires, Sierra de la Ventana and Rio Colorado, all lost). Synonymized by Mello-Leitão, 1933: 55.

  • Gayenna maculatipes: Keyserling, 1891: 141. F.O.P.-Cambridge, 1899: 18. Tullgren, 1905: 44. Mello-Leitão, 1925: 456. Berland, 1934: 168.

  • Gayenna argentina: Simon, 1897a: 91.

  • Sanogasta intermedia Mello-Leitão, 1941a: 177 (female holotype from Argentina, La Rioja province, Sañogasta, II.1939, M. Birabén, in MLP 14948, examined; a female without locality, in collection Birabén, labeled “Sanogasta intermedia/Tipi” in MLP, examined, may be a paratype). Ramírez, 1995a: 366. new synonymy.

  • Synonymy:

    The holotypes of the species here synonymized were compared; no relevant differences were found.

    Notes:

    The immature specimen of Josa sp. along with the type of A. maculatipes most probably come from another locality, because there are no records of similar Josa species from Uruguay or nearby localities. The limits between S. maculatipes and S. alticola are problematic and are only provisionally accounted here. I have seen several intermediate forms of male and female genitalia, as well as diverging forms, differing slightly from both species as limited here. The problem of intermediates is less acute for S. mandibularis, but should be also considered in a future revision.

    Diagnosis:

    Provisionally distinguished from the very similar S. alticola by having a less advanced epigynal anterior pouch. Typical males also have a shorter paramedian apophysis.

    Female (Montevideo, Arroyo Carrasco):

    Total length 6.00. Carapace length 2.30, width 1.73, wider on legs II–III. Length of tibia/metatarsus: I, 1.50/1.33; II, 1.50/1.33; III, 1.23/1.33; IV, 1.70/1.97. Palpal tarsus length 0.73. Chelicerae with two teeth on retromargin. Sternum length 1.20, width 0.97. Spines: leg I, femur d 1–1–1, 2ap; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v r1–2–2 or 2–2–2, p 0–1; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p and r d1ap; patella r d1; tibia v p1–2–2, p 1-0-1-0 or 0-d1-1-0, r d1–1 or 0–1, d r1–0-(1 bristle)-0; metatarsus v 2-p1–2 or 2–0–2, p and r d1–1–1, d 0-p1–2. IV, femur, patella = III; tibia v p1–2–2, p and r 1-d1-1-0, d r1–0-(1 bristle)-0; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. Abdomen length 3.15, width 1.60, spiracle–epigastrium 1.03, spiracle–spinnerets 0.77. Color: pale grayish, with grayish brown dorsal pattern. Sternum with gray spot in front of coxae I–III, small posterior spot. Epigyne (figs. 78D, E, 80A–C): opening of anterior pouch facing backward.

    Male (Montevideo, Arroyo Carrasco):

    Total length 5.30. Carapace length 2.40, width 1.83. Length of tibia/metatarsus: I, 2.67/2.60; II, 2.50/2.67; III, 1.83/1.87; IV, 2.27/2.63. Chelicerae slightly smaller than those of female. Sternum length 1.30, width 1.10. Spines as in female, except: leg I, tibia p 1-0-1-0 or 0. II, femur p 0-d1-d1; tibia v 2–2–2, p 1-0-1-0. III, femur p 0-d1-d1; tibia v 2–2–2, p and r 1-d1-1-0; metatarsus v 2–0–2. IV, tibia = III. Abdomen length 3.00, width 1.90, spiracle–epigastrium 1.03, spiracle–spinnerets 0.83. Color as in female (figs. 76A, 77E). Palp (figs. 78A, B, 81D, E): tibia width/length 0.56, cymbium relatively large. Embolus thin, basal process small, acute (fig. 78A). Median apophysis vestigial, short. Paramedian apophysis long, thin, base membranous. Secondary conductor large, not fused to anterior margin of tegulum (fig. 78B), with conspicuous canal arising at base of paramedian apophysis; retrolateral portion with basal triangular prong. Anterior margin of tegulum compressed over base of secondary conductor.

    Development:

    The development of the epigyne is similar to that of other Amaurobioidinae (see Tomopisthes horrendus). The primordium in penultimate females (fig. 79A) has traces of anterior pouch, of depressions of copulatory openings, and (presumably) of spermathecae.

    Natural History:

    This species builds retreats on foliage, grasses, and occasionally on “serruchetas” (Eryngium spp.) and under bark.

    Distribution:

    Peru, northern Chile, Bolivia, southern Brazil, Uruguay, and Argentina. Also found in Eastern Island (Baert et al., 1997), where it was most probably introduced by human activity.

    Variability:

    Specimens with four teeth on cheliceral promargin, or three on retromargin, are extremely rare. Most females lack the prolateral spine on metatarsus II.

    Other Material Examined:

    PERU: Apurimac: 37 km S Andahuaylas, 6.III.1951, Ross and Michelbacher, 1♂ 4♀ (CAS); 47 km N Andahuaylas, 7.III.1951, Ross and Michelbacher, 2♀ (CAS). Arequipa: 100 km NE Arequipa, 4500 m, 14.X.1983, E. Maury, 4♀ (MACN-Ar), A. Roig, 1♀ (MACN-Ar); 150 km W Arequipa, 14.X.1983, A. Roig, 7♀ (MACN-Ar); 170 km NE Arequipa, 4300 m, 15.X.1983, E. Maury, 4♀ (MACN-Ar); San Ignacio, Cailloma, 1.IX.1939, K. Schmidt, 1♂ 1♀ (AMNH). Puno: 15 km W Mañazo, 4100 m, 15.X.1983, A. Roig, 1♀ (MACN-Ar). Localities Not Found: Masocruz, 3800 m, 17.XII.1955, L. Peña, 6♀ (IRSN IG 20275); Perú, no specific locality, L. Peña, 2♀ (IG 20651). BOLIVIA: Cochabamba: Colomi, 21.X.1983, E. Maury, 1♀ (MACN-Ar); E of Cumbre Pass, Coroico Road, 17.VII.1960, B. Malkin, 1♀ (AMNH); road to Illimani, 3700–4000 m, 25.XII.1975, L. Peña (AMNH); Tunari, nr. 1 and 2, lake, 4100–4200 m, 10.X.1953, Forster and Schlinger, 3♀ (AMNH). La Paz: carretera La Paz-Sorata, 1400 m, 25.IV.1972, Bordon, 1♀ (MACN-Ar); La Cumbre, 4800 m, 20.IX.1988, V. and B. Roth, 1♀ (CAS); La Paz, suelo, 4550 m, VIII.1993, A. Brescovit, 3♀ (MACN-Ar, thanks to A. Brescovit); 65 km NE La Paz, Altiplano, nr. Juana de Potosí Mt., 14,500 ft, under flat rock, 10.II.1959, R. Walsh, 2♀ (AMNH); Tiahuanaco, 10–13.VI.1960, B. Malkin, 2♀ (AMNH). Locality Not Found: Chacaltaya, 4700 m, from small field, 24–25.IV.1954, Forster and Schlinger, 1♀ (AMNH). BRASIL: Santa Catarina: Curitibanos, Est. Campos Novos, 12.V.1967, P. Biasi, 1♀ (MZUSP 7032). Rio Grande do Sul: Santa Vitória do Palmar, Estação Ecológica do Taim, 12.IX.1991, A. Lise, 1♂ 1♀ (MCTP 0993). URUGUAY: Departamento Minas: Lavalleja, Cerro Penitente, under stones, 10.XII.1967, P.R. San Martín, 1♀ (MCZ). Departamento Rocha: Arroyo Sarandí del Consejo, ruta 9 km 251, 18.V.1993, M. Ramírez and F. Pérez Miles, 2♂ (MACN-Ar); Laguna de Castillos, 19.V.1993, M. Ramírez and F. Pérez-Miles, 1♀ (MACN-Ar). Departamento Maldonado: Punta Ballenas, 29–30.VIII.1980, P. Goloboff, 1♀ (MACN-Ar). Departamento Montevideo: Montevideo, calle Durazno y Acevedo, VI.1964, R. Capocasale and L. Bruno, 1♂ 1♀ (CAS); Montevideo, Arroyo Carrasco, 20.VIII.1961, R. Capocasale and L. Bruno, 1♂ 6♀ (CAS); Montevideo, no date, no collector, 1♀ (MNRJ 14129). ARGENTINA: Jujuy: Cachinoca, I.1966, E. Maury, 2♀ (MACN-Ar 1034); Humahuaca, 20–21.I.1985, E. Maury, 1♂ (MACN-Ar); Fraile Pintado, X.1967, E. Maury, 2♀ (MACN-Ar 6033); Laguna de Yala, I.1966, E. Maury, 1♀ (MACN-Ar); Mina El Aguilar, Tres Cruces, 16.I.1942, M. Birabén and Scott, 1♀ (MLP); Tilcara, II.1981, P. Goloboff, 1♂ (MACN-Ar). Salta: Iruya, 29.XI.1981, E. Maury, 1♀ (MACN-Ar); Laguna Brealito, 15 km W Seclantás, 29.I.1981, E. Maury, 1♀ (MACN-Ar); Rosario de la Frontera, X.1986, O. Donado, 1♂ (MACN-Ar); San Bernardo, San Lorenzo, 25–31.V.1933, J.B. Daguerre, 3♀ (MACN-Ar). Misiones: Fracrán, 23.XI.1948, M. Birabén, 1♀ (MLP); Santa María, XII.1947, J.M. Viana, 1♂ (MACN-Ar). Tucumán: ruta 40 km 999, Quilmes, 9.I.1995, P.A. Goloboff, 1♂ (IML). Catamarca: Capital, VII.1949, J.M. Viana, 4♀ (MACN-Ar 2943); I.1946, Schaeffer, 2♀ (MLP); El Rodeo, I.1957, M.E. Galiano, 1♀ (MACN-Ar), 11.XII.1951, 1♀ (MLP); Est. Grande, II.1946, M. Vignalli, 1♀ (MLP); Quebrada La Cébila, 21.X.1963, M.E. Galiano, 1♂ (MACN-Ar); Santa María, 18.XII.1994, C. Grismado, 1♂ 1♀ (MACN-Ar). La Rioja: Ascha, Aminogasta, 1947, Cáceres Freyre, 1♀ (MACN-Ar); B. Famatina, Chilecito, II.1953, M.E. Galiano, 1♀ (MACN-Ar). Santiago del Estero: Capital, 4.VI.1963, Havrylenko, 1♀ (MACN-Ar); Colonia Dora, VIII.1940, 1♀ (MACN-Ar 1769). Córdoba: Argüello, XII.1945, J.A. de Carlo, 1♂ (MACN-Ar 1947), 1♀ (MACN-Ar 1948); Calamuchita, Sierras Grandes, X.1970, J.M. Viana, 1♀ (MACN-Ar); Cosquín, 31.X.1898, ″No. frasco 4513”, no collector, 1♂ (MACN-Ar); Departamento San Javier, II.1943, H. Gario and R. Maniglia, 1♀ (MACN-Ar); La Falda, under stones, 23.VIII.1922, A. Frers, 1♂ (MACN-Ar). San Juan: Paso Agua Negra, ca. 3500 m, 1–2.I.1982, A. Roig, 2♂ 2♀ (MACN-Ar). San Luis: Carolina, IX.1970, J.M. Viana and Williner, 1♂ 2♀ (MACN-Ar); Cacheuta, X.1965, E. Maury, 1♀ (MACN-Ar); Papagallos, 9.XI.1982, E. Maury, 1♂ 1♀ (MACN-Ar). Santa Fe: Colonia Macias, Departamento Garay, XI.1942, J.M. Viana, 1♀ (MACN-Ar 1400); El Toba, X.1967, M.E. Galiano, 1♀ (MACN-Ar). Entre Ríos: El Palmar Natl. Park, XI.1988, M.E. Galiano, 1♀ (MACN-Ar); Ibicuicito, 1938, F. Castillo, 3♀ (MACN-Ar); Rosario del Tala, 20.XI.1988, M. Ramírez, 1♀ (MACN-Ar). Buenos Aires: Atucha, 27.VII.1985, P. Goloboff, M. Ramírez, 3♂ (MACN-Ar), 10.V.1987, M. Ramírez, 1♂ (MACN-Ar); 8.IX.1989, M. Ramírez, 1♀ (MACN-Ar); Bahía Blanca, II.1942, S. Monrós, 1♀ (MACN-Ar 1173); Boulogne, X.1938, A. Prosen, 2♂ 13♀ (MLP); many specimens from around Buenos Aires city, in MACN-Ar; Capilla del Señor, 23.I.1942, A. Prosen, 1♀ (MLP); Castelli, X.1960, J.M. Viana, 1♂ 1♀ (MACN-Ar 5155); Chascomús, 16.XII.1984, M. Ramírez and C. Scioscia, 3♂ 7♀ (MACN-Ar), 19.X.1947, N91a, no collector, 2♂ 2♀, 2♀ penultimates (MACN-Ar); Escobar, 1938, A. Prosen, 2♂ 5♀ (MLP); Estancia El Tonelero, Pdo. Gral. Lavalle, cerca canal 2, 15–21.XII.1951, J. Cranwell, 1♀ (MACN-Ar); Ing. Maschwitz, XI.1941, A. Prosen, 1♀ (MLP); Isla Martín García, 25.V.1990, M. Ramírez, 1♂ 1♀ (MACN-Ar); La Plata, 1942, 1♂ 5♀ (MLP); 15 km W Lobería, 4.IX.1972, 1♂ (MACN-Ar); Los Médanos, energía, 8.IV.65, J.M. Gallardo and E. Maury, 5♂ 1♀ (MACN-Ar); Magdalena, 13–14.VIII.1983, P. Goloboff and M. Ramírez, 1♀ (MACN-Ar); Mar del Plata, 20.II.1985, M. Ramírez, 1♀ (MACN-Ar); Paraná de Las Palmas, 17.IX.1963, M.E. Galiano, 1♂ (MACN-Ar); Punta Lara, Ensenada, 28.XI.1985, M.E. Galiano, C. Scioscia, 3♀ (MACN-Ar); III.1943, A. Moreno, 3♂ 4♀ (MLP), 15.XII.1943, 2♀ (MLP); Quequén, II.1960, M. Birabén, 2♀ (MACN-Ar); San Isidro, Punta Chica, 5.XI.1941, A. Prosen, 4♀ (MLP); Sierra de la Ventana, Prov. Park E. Tornquist, Cerro Negro, 12.IV.1974, Cesari, 7♂ 4♀ (MACN-Ar), 18–20.IX.1982, M. Ramírez, 3♂ 1♀ (MACN-Ar); Tandil, excursion J.M. Viana, 2♀ (MACN-Ar). Neuquén: Laguna Blanca Natl. Park, I.1975, E. Maury, 1♀ (MACN-Ar); Piedra Pintada, II.1941, R. Maldonado, 1♀ (MLP); Río Limay, Arroyito (12), 16.XII.1978, Misión Científica Danesa, 1♀ (ZMK); San Martín de los Andes, 3–6.I.1964, no collector, 1♀ (MACN-Ar); Senillosa, I–II.1973, O. de Ferrariis, 1♀ (MACN-Ar). Río Negro: San Carlos de Bariloche, II.1954, M.E. Galiano, 1♀ (MACN-Ar), 1♂ (MACN-Ar 5413); Coronel Gómez, IV.1948, A. Ibarra Grasso, 1♂ (MACN-Ar); Gral. Roca, I.1962, A. Bachmann, 1♂ (MACN-Ar); X.1963, A. Bachmann, 1♀ (MACN-Ar). Chubut: El Hoyo, 1.I.1962, A. Kovács, 1♀ (AMNH); Epuyén, 42°15′S, 71°23′W, A. Kovács, 1♂ 1♀ (AMNH); Languineo, Estancia Manantiales, 6–10.XI.1985, L. Peña, 1♀ (AMNH); Los Cipreses, XI.1982, M. Ramírez, 1♀ (MACN-Ar); Lago Puelo Natl. Park, 10.XI.1969, A. Kovács, 1♂ (AMNH); Viedma, 16.II.1948, M. Birabén, 2♂ 3♀ (MLP). Santa Cruz: Calafate, II.1963, E. Maury, 1♀ (MACN-Ar); Los Cerros, Tres Lagos, IV.1949, Waring, 1♀, 1♂ 2♀ (MLP); 9.III.1948, M. Birabén, 5♂ 2♀ (MLP). CHILE: Región I (Tarapacá): Tarapacá: Chaquiña, 3700–4000 m, L. Peña, 10♀ (IRSN IG19736). Región II (Antofagasta): El Loa: Calama, Río Loa, La Cascada, 10.I.1984, G. Arriagada, 1♀ (MHNS 910); Calama, Vegas del Río Loa, Fundo Soto, 10.VIII.1972, no collector, 4♂ 12♀ (UC); Cautín, N San Pedro de Atacama, 3300 m, 23–31.VIII.1982, L. Peña, 2♀ (AMNH); San Pedro de Atacama, 23.VIII–6.IX.1982, L. Peña, 3♀ (AMNH). EASTER ISLAND: Specimens reported by Baert et al. (1997) were identified by myself from drawings (Pekka Lehtinen, in litt.).

    Note:

    F.O.P.-Cambridge recorded this species from the Juan Fernández Islands (1♂, 1♀ immature, Dr. Plate coll., not seen), indicating that he could not find any difference with the type of Gayenna maculatipes, except that the specimens from the Juan Fernández are larger. This identification is not very reliable, though. The type of G. maculatipes is a female, and in those islands S. maculosa is commonly found, a species very similar to S. maculatipes, which is larger on the Juan Fernández Islands than on the mainland.

    Sanogasta alticola (Simon), new combination Figures 80D, E, 81A–C

  • Gayenna alticola Simon, 1896c: 400 (female lectotype from Bolivia, La Paz, Garlepp coll., and male paralectotype [Meriola cetiformis, misidentification], designated by Platnick and Ewing, 1995: 15; in MHNP 17942, examined), 1897a: 91, 99.

  • Gayenna monticola Chamberlin, 1916: 267 (female holotype from Peru, Cuzco, 11,500 ft, July 1911, Yale Peruvian Expedition, under stones, in MCZ 256, examined). new synonymy.

  • Note:

    The distinction between this species and S. maculatipes is problematic (see note under S. maculatipes).

    Diagnosis:

    Provisionally distinguished from the very similar S. maculatipes by having the epigynal anterior pouch more advanced (fig. 80D). Typical males also have a longer paramedian apophysis (fig. 81B).

    Female (lectotype):

    Total length 6.30. Carapace length 2.52, width 1.78, wider on legs II–III. Length of tibia/metatarsus: I, 1.49/1.34; II, 1.49/1.32; III, 1.25/1.30; IV, 1.62/2.18. Chelicerae unmodified, with two teeth on retromargin. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2-p1 (most females with v 2–2–2); metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v r1–2–2, p 0–1; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p 0-d1-d1, r d1ap; patella rd1; tibia v p1–2–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2-p1–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p and r d1ap; patella dr1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. Abdomen length 4.00, width 1.10. Color: pale grayish with gray pattern, as in S. maculatipes. Epigyne (fig. 80D, E): very similar to that of S. maculatipes, anterior pouch more advanced.

    Male (La Paz, III–IV.1959):

    Total length 6.12. Carapace length 2.53, width 2.00. Length of tibia/metatarsus: I, 2.60/2.63; II, 2.30/2.30; III, 1.73/1.90; IV, 2.23/2.50. Chelicerae slightly smaller than those of female. Sternum length 1.43, width 1.10. Spines as in female, except: leg I, tibia p 1-0-1-0; metatarsus p 1–0. II, femur r 0; tibia = I or p 1-d1-1-0. III, tibia p and r 1-d1-1-0; metatarsus v 2-p1–2. IV, tibia = III. Abdomen length 3.30, width 2.00, spiracle–epigastrium 1.37, spiracle–spinnerets 0.83. Color as in female. Palp (fig. 81A–C): very similar to that of S. maculatipes, often with longer paramedian apophysis. Tibia long, width/length 0.42, cymbium large. Secondary conductor partially fused to anterior margin of tegulum; retrolateral portion with basal rounded projection.

    Variability:

    Some females lack the prolateral spine on metatarsus II. Spines: metatarsus III, v 2–2–2. Male, spines: III, tibia v 2–2–2.

    Natural History:

    Unknown.

    Distribution:

    Puna highlands in Peru, Bolivia, and Argentina.

    Other Material Examined:

    PERU: Cuzco: Cuzco, VIII, no date, Wunderlich, 3♂ 2♀ (AMNH), 7–8.VI.1964, B. Malkin, 2♀ (AMNH); Urubamba, under stone, I. 1965, F. Carrasco, 1♀ (MCZ). Huancavelica: Huancavelica, 10.VIII.19??, 1♀ (sitio 30, IRSN IG 25518); nr. Niñobamba, 3500 m, 2♀ (sitio 36, IRSN IG 25518). Puno: 20 mi N Desaguadero, 27.II.1951, 1♀, 28.II.1951, 1♂, Ross and Michelbacher (CAS). Perú, no specific locality, 42, 1♀ (IRSN IG 25518). BOLIVIA: Cochabamba: 30 mi N Potosí, 22.II.1951, Ross and Michelbacher, 1♂ 1♀ (CAS). La Paz: Apacheta, Cuyu-Cuyu, E Tiahuanaco, 4100 m, L. Peña, 2♀ (AMNH); Huatajata nr. Lake Titicaca, 6.I.1954, Schlinger and Forster, 1♀ (AMNH); hill beyond Huatajata, Lake Titicaca area, 10–15.I.1954, Forster and Schlinger, 1♀ (AMNH); La Paz, III–IV.1959, R. Walsh, 2♂ 1♀ (AMNH); house and garden, IV.1959, R. Walsh, 1♂ (AMNH); La Paz, Avenida Sport Club, 4.I.1959, A. Nadler, 2♀ (AMNH). Locality Not Found: Songo Valley, Cuticucho, 3800 m, 30.I.1954, Forster and Schlinger, 1♂ (AMNH). ARGENTINA: Jujuy: Abra Pampa, III.1966, E. Maury, 5♀ 1 immature (MACN-Ar); Laguna de Yala, X.1967, E. Maury, 1♀ (MACN-Ar); 30.XI.1981, E. Maury, 1♀ (MACN-Ar); V.1983, P. Goloboff, 1♀ (MACN-Ar). Tucumán: Tafí del Valle, 28–30.XII.1994, P. Goloboff, 1♂ 1♀ (MACN-Ar); 27.X.1947, Gobbach, 2♀ (MLP). La Rioja: Alto Carrizal, II.1956, M.E. Galiano, 11♀ (MACN-Ar).

    Sanogasta mandibularis, new species Figures 78C, F, G, 80F, G, 81F, G

    Types:

    Female holotype 9824 from Argentina, Buenos Aires province, Los Talas, reared in captivity, born from female paratype 9826, ca. 34°51′S, 57°55′W, XII.1985, C. Scioscia; male paratype 9825 from Buenos Aires, Partido de Ensenada, Punta Lara, 26.II.1967, M.E. Galiano, deposited in MACN-Ar.

    Etymology:

    The specific name refers to the relatively large chelicerae.

    Diagnosis:

    Resembles S. maculatipes and S. alticola in having very similar genitalia, but typical specimens can be distinguished by having three teeth on the cheliceral retromargin and four on the promargin. Specimens usually have a larger epigynal anterior pouch, situated close to the epigastric furrow, and strong male chelicerae (see Note under S. maculatipes).

    Female (holotype):

    Total length 6.00. Carapace length 2.23, width 1.67, wider on legs II–III. Length of tibia/metatarsus: I, 1.37/1.30; II, 1.33/1.27; III, 1.07/1.17; IV, 1.50/1.83. Palpal tarsus length 0.72. Chelicerae strong, with four teeth on promargin, three on retromargin, apical one slightly smaller. Sternum length 1.17, width 0.92. Spines: leg I, femur d 1–1–1, p (1-d1)ap; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v r1–2–2, p 0–1; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p and r d1ap; patella r d1; tibia v p1-p1–2, p and r d1–1, d r1bas; metatarsus v 2-p1–2, p and r d1–1–1, d 0-p1–2. IV, femur = III; patella r d1; tibia v p1–2–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. All tibiae with d r1-0-1-0 bristles, except III and IV, where basal bristles replaced by spines. Abdomen length 3.70, width 2.33, spiracle–epigastrium 1.40, spiracle–spinnerets 1.00. Color: yellowish cream with grayish spots. Sternum and venter pale. Epigyne (figs. 78F, G, 80F, G): similar to that of S. maculatipes, anterior pouch larger, closer to epigastric furrow.

    Male (paratype):

    Total length 5.54. Carapace length 2.67, width 1.97. Length of tibia/metatarsus: I, 3.37/3.37; II, 2.67/2.70; III, 1.70/1.90; IV, 2.23/2.73. Chelicerae (fig. 81G) strong, longer than those of female, with four teeth on promargin, three on retromargin; middle part of fang thickened. Spines as in female, except: leg II, femur p and r d1ap; tibia v r1–2–2 or 2–2–2 (the p1bas very small), p 0–1 or 0. III, tibia v p1–2–2; metatarsus v 2-p1–2 or 2–2–2. IV, tibia = III. Sternum length 2.67, width 2.33. Abdomen length 2.80, width 1.90, spiracle–epigastrium 0.93, spiracle–spinnerets 0.87. Color as in female, with some diffuse dots in median line on venter. Palp (fig. 81F): very similar to that of S. maculatipes. Tibia width/length 0.76. Median apophysis vestigial, conical, paramedian wide, sinuous. Secondary conductor not fused to anterior margin of tegulum, retrolateral portion with basal rounded projection.

    Variability:

    A female from Punta Lara has three teeth on promargin, two on retromargin.

    Natural History:

    Unknown. All specimens were collected on or close to flooded areas.

    Distribution:

    Northeastern Argentina and Paraguay.

    Other Material Examined:

    PARAGUAY: Central: Asunción, 11.VI.1988, V. and B. Roth, 1 vial (number of specimens not recorded) (CAS). ARGENTINA: Formosa: Capital, IV.1958, Ogueta, 1♀ (MACN-Ar). Corrientes: Apipé, XII.1945, W. Hanke, 1♀ 1 immature (MACN-Ar 1765); Santiago Alcorta, VI.1943, M. Birabén, 1♀ (MACN-Ar). Santa Fe: Alejandra, II.1964, M.E. Galiano, 1♂ 1♀ (MACN-Ar). Entre Ríos: Gualeguaychú, IV.1943, F. Monrós, 1♂ (MACN-Ar); El Palmar Natl. Park, IV.1981, P. Goloboff, 1♂ (MACN-Ar); Salto Grande, III.1964, M.E. Galiano, 1♀ (MACN-Ar). Buenos Aires: Capital Federal, Bañados de Palermo, 31.V.1916, A.G. Frers, 1♂ (MACN-Ar); Saavedra, XII.1935, Castillo, 1♀ (MACN-Ar); Glew, no date, Carpintero, 1♀ (MACN-Ar); Isla Maciel, I.1948, Partridge, 3♀ (MACN-Ar): Isla Martín García, II.1933, J.B. Daguerre, Pérez-Moreau, 1♂ 2♀ 1 immature (MACN-Ar 34338), IV.1938, J.M. Viana, 1♀ (MACN-Ar 421); Punta Lara, Ensenada, III.1943, A. Moreno, 1♂ (MLP), 18.IX.1985, M. Ramírez, 1♀ (MACN-Ar); San Pedro, 2.XI.1991, M. Ramírez, 1♂ 1♀ (MACN-Ar); San Isidro, Reserva Ribera Norte, 16.II.1999, M. Pandolfi, 1♂ (MACN-Ar), 12.VIII.1999, M. Pandolfi, 1♀ (MACN-Ar).

    Sanogasta maculosa (Nicolet), new combination Figures 14B–D, 61C, 76B–D, 77A–D, 79B, 82, 83

  • Clubiona maculosa Nicolet, 1849: 423 (female lectotype and three immature paralectotypes here designated, from Chile, in MHNP 4234, examined). Simon, 1864: 132.

  • Clubiona sternalis Nicolet, 1849: 424 (female lectotype and three immature paralectotypes here designated, from Chile, in MHNP 4237, examined). Simon, 1864: 132, 1887: E4. new synonymy.

  • Gayenna stellata: Simon, 1884: 131 (only male paralectotype, from Chile, Cabo de Hornos, in MHNP, examined).

  • Tomopisthes taeniatus Simon, 1886: 571 (female holotype from Argentina, Santa Cruz province, Patagonia, in MHNP 7729, examined), 1897a: 91, 1902: 31, 1903b: 312, 1903d: 6, 1905b: 12. new synonymy.

  • Gayenna maculosa: Simon, 1887: E4. Mello-Leitão, 1936: 119.

  • Anyphaena ignota Keyserling, 1891 (male holotype from “Possessions Bay, Straits of Magellan”, in MCZ, examined). new synonymy.

  • Gayenna affinis Tullgren, 1901: 241, 259 (male lectotype, two male and one female paralectotypes from Chile, Punta Arenas, 27.XI.1895, O. Nordenskjöld coll., female paralectotype from Puerto Herberton, 14.II.1896, here designated, in NRS, examined), 1902: 59. new synonymy.

  • Gayenna dubia Tullgren, 1901: 243 (female lectotype from Chile, Punta Arenas, 29.XI.1895, and two females paralectotypes, from Ultima Esperanza, 2.IV.1896, O. Nordenskjöld coll., here designated, in NRS, examined). new synonymy.

  • Tomopisthes conspersus Simon, 1902: 33 (female holotype from Chile, Punta Arenas, in MHNP 21816, examined). new synonymy.

  • Gayenna conspersa: Merian, 1913: 13.

  • Tomopisthes injucundus Simon, 1902: 33 (female lectotype, three female and one male paralectotypes here designated, from Tierra del Fuego, MHNP 21782, examined; the male paralectotype belongs to a different, presumably undescribed Sanogasta species), 1903b: 312. new synonymy.

  • Tomopisthes modestus Simon, 1902: 35 (female holotype from Chile, Punta Arenas, IX.1892, Michaelson coll., in MHNP, examined). new synonymy.

  • Gayenna modesta: Merian, 1913: 13.

  • Gayenna skottsbergi Berland, 1924: 434 (male and female syntypes and 19 female paratypes from the Juan Fernández Islands, Mas a Tierra, in NRS, examined). new synonymy.

  • Monapia andina: Gerschman and Schiapelli, 1970: 131 (misidentification, male allotype subsequently designated, in MACN-Ar 6269, examined).

  • Synonymy:

    The primary types of all species synonymized were compared, together with extensive samples from the same areas. The slight differences in epigyne (see Variability below) were not found to be correlated with any differences in the male palp, which is remarkably uniform. The specimens from the Juan Fernández Islands (and some from Central Chile as well) are larger, but their genitalia are otherwise indistinguishable from those of other specimens of typical size. There are many intermediate forms in epigyne conformation and body size; these differences are here regarded as intraspecific variability.

    Note:

    There are many vials in NRS identified by Tullgren as Gayenna affinis. I considered syntypes only those whose locality is listed in the original description.

    Diagnosis:

    Typical specimens can be distinguished by having the epigynal anterior pouch close to the epigastric furrow, with the opening facing backward; some specimens have a small pit in place of the pouch. Males are recognized by the shape of the paramedian apophysis, with a thin, curved apex.

    Female (spines from paralectotype of Gayenna affinis, other data from Chubut, Lago Menéndez):

    Total length 7.45. Carapace length 2.73, width 1.93, wider on legs II–III. Length of tibia/metatarsus: I, 1.52/1.20; II, 1.32/1.20; III, 1.13/1.25; IV, 1.55/0.95. Palpal tarsus length 0.82. Chelicerae unmodified, with two teeth on retromargin. Sternum length 1.52, width 1.07. Spines: leg I, femur d 1–1–1, p (1-d1)ap, r d1ap; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p 0-d1-(1-d1), r 0-d1-d1; tibia v r1–2–2; metatarsus v 2bas. III, femur d 1–1–1, p and r 0-d1-d1; patella r d1; tibia v p1-p1–2, p and r 1-d1-1-0 or r 1-d1-1-0, d r1bas; metatarsus v 2–0–2, p and r d1-1-0-1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia v p1–2–2, p and r 1-d1-1-0, d r1bas; metatarsus v 2–2–2, p and r d1-1-0-1, d 0-p1–2. Abdomen length 4.52, width 2.83, spiracle–epigastrium 2.30, spiracle–spinnerets 0.83. Color (cf. figs. 76B, 77B): carapace grayish, margins dark. Legs pale gray with grayish spots, coxae pale. Endites and labium grayish, sternum pale with darker sides. Abdomen grayish, darker to posterior end, dorsal pattern on cream background, cardiac area dark, venter pale with small dark dots, forming median band. Epigyne (figs. 82D, 83D–K): anterior pouch small, close to epigastric furrow, opening facing backward. Copulatory openings in deep depressions, in epigastric furrow, hidden by membrane (fig. 82D). Ducts of accessory bulbs short, ventral.

    Male (spines from lectotype of Gayenna affinis, other data from Chubut, Río Arrayanes):

    Total length 6.12. Carapace length 2.77, width 2.20, relatively wider and more rounded than that of female. Length of tibia/metatarsus: I, 2.30/2.17; II, 2.20/2.10; III, 1.77/1.83; IV, 2.17/2.60. Chelicerae quite smaller than those of female. Sternum length 1.40, width 1.08. Spines as in female, except: leg I, femur r 0-d1-d1; tibia p and r 1-d1-1-0; metatarsus p d1-1-0-0, r 1. II, femur p and r 0-d1-d1; tibia and metatarsus = I. III, tibia v 2–2–2. IV, femur r d d1ap or 0-d1-d1; tibia = III. Abdomen length 3.45, width 1.97, spiracle–epigastrium 1.53, spiracle–spinnerets 0.77. Color (cf. fig. 76D): as in female, but carapace with long dark patches on median band, plus four radial lines from thoracic groove to palps and hindlegs. Sternum with small dark spots in front of coxae I–III. Palp (figs. 82A–C, 83A–C): tibia short, width/length 0.92, cymbium relatively large. Embolus thin, basal process small, acute. Median apophysis very thin, closely associated with paramedian (fig. 82A). Apex of paramedian apophysis short, thin, sinuous. Primary conductor apparently absent, only low mound in that place. Secondary conductor not fused to anterior margin of tegulum, canal conspicuous, arising at base of paramedian apophysis (fig. 82B); retrolateral portion with internal conical projection, and external ridge. Anterior margin of tegulum compressed over base of secondary conductor.

    Variability:

    Size is also very variable: the specimens from the Juan Fernández Islands are especially large. Body color and pattern is very variable (figs. 76B–D, 77A–D). Female spines: femur I = II. Male spines: metatarsi I, p 1–0, r 0; II, r 0; III; v 2-p1–2. Epigyne quite variable in details, some extreme forms illustrated in figure 83D–I.

    Natural History:

    This species builds retreats on foliage, under bark, and occasionally under stones.

    Distribution:

    All austral forests of Chile and Argentina, including semiarid and littoral areas. In Chile, from Parinacota Province, in Argentina from Neuquén, to Tierra del Fuego.

    Other Material Examined:

    ARGENTINA: Neuquén: Collón Curá, 750 m, 19.IX.1981, Nielsen and Karsholt, 3♀ (ZMK); Estancia San Ramón, Rincón Chico, Río Limay, X–XII.1962, 3♂ 7♀ 1♀ penultimate, I.1962, 2♀, 1♀, Junín de los Andes, II.1968, E. Maury and N. Müller, 1♀ 1♀ penultimate (MACN-Ar); Lanín Natl. Park: Lago Hermoso, 15.I.1985, M. Ramírez, 1♀, 1♀ (MACN-Ar); Lago Moquehue, 10.I.1985, E. Maury, 3♀ (MACN-Ar); Lago Lácar, X.1955, A. Giai, 3♀, 1♂, 1♀ (MACN-Ar); 5 km E Hua Hum, 640 m, 5.XI.1981, 1♀, 16.XI.1981, 1♂, Nielsen and Karsholt (ZMK); Pucará, 5–16.II.1956, A. Ogloblin and Sra., 2♀ (MLP), 1.II.1971, Schajovskoy, 1♀ (MACN-Ar); XI.1971, L. Yinoff, 1♂ (MACN-Ar); XII.1973, S. Schajovskoy, 1♀ (MACN-Ar); 750 m, 25.XI.1978, 1♂, 1♂, 1♂, 750 m, 1.XII.1978, 1♂, 750 m, 3.XII.1978, 1♂, Misión Científica Danesa (ZMK); Lago Lolog, Gentili Cabin, above town, pan and FIT, forest and meadow, 18–21.XI.1989, S. Marshal, 1♂ 1♀ (AMNH); Lago Lolog, nr. San Martín de los Andes, pans nr. stream, ca. 900 m, 23–30.XI.1989, S. Marshall, 3♂ 3♀ (AMNH); dung trap, 1♀ (AMNH); fit nr. pond, 23.XI–1.XII.1989, S. Marshall, 3♂ 4♀ (AMNH); Lago Lolog, 4 km N San Martín de los Andes, FIT, Nothofagus forest, ca. 950 m, Gentili property, 23.XI–1.XII.1989, S.A. Marshall, 6♂ 3♀ (AMNH); 8 km N San Martín de los Andes, 1000 m, Malaise trap, 16–22.XI.1997, C. and M. Vardy, 6♂ 10♀ (BMNH/MACN-Ar); Lago Tromen, Rodeo Grande, 900 m, 30.XI.1978, Misión Científica Danesa, 2♂ (ZMK); mouth of Río Blanco in Lago Huechulaufquen, 6.I.1985, under bark, M.J. Ramírez, 3♀ (MACN-Ar); San Martín de los Andes, 40°10′S, 71°21′W, 20–21.XI.1988, V. and B. Roth, 1♂ (CAS), 1000 m, XI–XII.1985, Gentili, 32♂ 19♀ (MACN-Ar); 640 m, 2.XI.1981, 1♀, 5♂, Nielsen and Karsholt (ZMK); San Martín de los Andes, Cerro Chapelco, 1400–1600 m, 2–19.XII.1981, 3♂ 3♀, 12–23.XII.1981, 16♂, 1♀, 17♂, Nielsen and Karsholt (ZMK); I.1961, M. Galiano, 1♀ (MACN-Ar 5294); Quillén, I.1986, Duprés, 1♀ (MACN-Ar); Termas de Epulaufquen, 9.I.1986, M. Ramírez, 1♂ (MACN-Ar); Nahuel Huapi Natl. Park: Isla Victoria, IV.1945, Havrylenko, 1♂ 2♀ (MLP); XII.1959, I. de Orfila, 1♀ penultimate (MACN-Ar); 41°S, 71°W, 1.V.1965, A. Kovács, 2♀ (AMNH); Isla Victoria, 28.III.1961, A. Kovács, 1♂ 1♀ (AMNH). Río Negro: San Carlos de Bariloche, II.1954, M.E. Galiano, 2♂ 8♀ (MACN-Ar), 18.VIII.1961, A. Kovács, 2♀ (AMNH), 1964, Monrós, 1♀ 3 immatures (MACN-Ar); 11 km W San Carlos de Bariloche, Cerro Otto, 14.I.1972, L. Herman, 1♀ (AMNH); Colonia Suiza, 800 m, 19.IX.1981, 5♀, 10.X.1981, 1♀, 8♂ 1♀, 1–7.XI.1981, 4♂, 1♂, 12–20.XI.1981, 2♂, 7.XII.1981, 1♂, 21–22.XII.1981, 1♂, Nielsen and Karsholt (ZMK); 810 m, 22.XI.1978, Misión Científica Danesa, 1♀ (ZMK); Pampa del Toro, 900 m, 9–10.XI.1981, 1♂, 1♂, 900 m, 22–23.XI.1981, 5♂, Nielsen and Karsholt (ZMK); El Bolsón, 24.XI.1962, M. Birabén, 6♀, 1♂ (MACN-Ar); II.1963, Birabén, 2♀ (MACN-Ar); X.1963, Birabén, 1♀ (MACN-Ar), 15.I.1961, 1♀, 13.III.1961, 3♀, 4.IX.1961, 1♀, 27.IX.1961, 1♀, 10.X.1961, 1♀, 28.X.1961, 1♀, 28.X.1961, 1♀, 7.X.1962, 1♀, 31.X.1966, 1♂, no date, 2♀, A. Kovács (AMNH); 24.XI.1962, 1♂ 2♀, 25.II.1963, 1♀, M. Birabén (MLP); Paso Flores, 3.X.1965, A. Kovács, 1♀ (AMNH); Río Azul, 15.X.1961, 1♀, A. Kovács (AMNH); Ñorquinco, 11.X.1961, 1♂, 27.VIII.1962, 1♀, 5♀ 3♂, 20.VI.1966, 1♀, 3.VII.1966, 1♂ 2♀, A. Kovács (AMNH); Nahuel Huapi Natl. Park: 880 m, 11.I.1986, N. Platnick, P. Goloboff and R. Schuh, 1♀ (AMNH). Chubut: Cushamen, 14.IX.1966, A Kovács, 2♀, 1♂ (AMNH); Cholila, 25.VIII.1962, A. Kovács, 1♂ 1♀ (AMNH); El Hoyo, 30.IX.1961, 1♀, 2.X.1962, 3♂ 4♀, 26.V.62, 7♀, VII.1962, 1♀, VIII.1962, 3♀, A. Kovács (AMNH); El Maitén, IX.1961, A. Kovács, 1♀ (MLP); 20.VI.1962, 1♂, 13.IX.1962, 1♀, A. Kovács (AMNH); Epuyén, 2.VII.1962, 1♀, 12.VI.1962, 17♀, 2.VIII.1962, 4♂, 18.XI.1962, 42°15′S, 71°23′W, 2♂, 1♂ 1♀, 5.VIII.1966, 1♀, A. Kovács (AMNH), 18.XI.1962, M. Birabén, 1♂ 1♀ (MLP); Esquel, road to La Hoya, 42°54′S, 71°19′W, 16.XI.1988, V.D. Roth, 6♀ (CAS); 17 km E Esquel, on Rt. 40, muddy shore of pond with scattered vegetation, 22.I.1986, R. Schuh and N. Platnick, 1♀ (AMNH); 35 km E Esquel, 720 m, 18.IX.1966, E. Schlinger and M. Irwin, 1♀ (CAS); Poto Cahuel, 8.X.1966, A. Kovács, 1♀ (AMNH); Languineo, Estancia Manantiales, 6–10.XI.1985, L. Peña, 1♀ (AMNH); Los Cipreses, XI.1982, M. Ramírez, 2♂, 1♀, 1♂ (MACN-Ar); Lago Puelo Natl. Park, 220 m, 18.XI.1978, Misión Científica Danesa, 1♂ (ZMK); 250 m, 22–23.X.1981, Nielsen and Karsholt, 1♂ (ZMK); Los Alerces Natl. Park: Lago Escondido, 19.XI.1981, A. Kóvacs, 3♀ (AMNH); Lago Futalaufquen, II.1985, 1♂ 6♀ 4♀ penultimates, II.1986, 2♂ 3♀, 9.II.1986, 3♀ M. Ramírez (MACN-Ar); I.1990, M. Ramírez, 1♀ (MACN-Ar); Lago Menéndez, I.1990, M. Ramírez, 1♀ (MACN-Ar); Lago Verde, II.1986, 1♂, I.1990, 1♀, Río Arrayanes, I.1990, 1♂, W margin, II.1986, 3♀, M. Ramírez (MACN-Ar); Río Turbio, 11.VI.1961, 4♀, no date, 1♀, A. Kovács (AMNH); I.1976, M. Rumboll, 1♀ (MACN-Ar); 19.5 km W Shaman, 650 m, 19.XI.1966, E. Schlinger and M. Irwin, 1♂ (CAS); Tecka, Corcovado, 750 m, 17.II.1979, Misión Científica Danesa, 1♀ (ZMK). Santa Cruz: Calafate, II.1963, E. Maury, 1♀, 1♂ (MACN-Ar); Estancia Cóndor, Río Gallegos, 28.IV.1974, M. Rumboll, 1♀ (MACN-Ar); Lago Argentino, III.1900, Excursión Silvestri, 1♂ (MACN-Ar); Lago Belgrano, 15.II.1973, 1♀, 21.II.1973, M. Rumboll, 2♀ (MACN-Ar); Lago Frías, no date, E. Maury, 1♀ (MACN-Ar); Lago Musters, SW margin, 20.I.1977, E. Maury and Patrick, 1♀ (MACN-Ar); Lago San Martín, X.1939, S. Radone, 1♀ (MACN-Ar 599); Los Cerros, Tres Lagos, IV.1949, Waring, 2♂ 3♀, 4♂ 4♀, 9♂ 9♀, 2♂ 2♀ (MLP); Morro Chico, Río Turbio, 28.I.1976, M. Rumboll, 2♀ (MACN-Ar); Los Glaciares Natl. Park, 11.II.1973, M. Rumboll, 1♂ (MACN-Ar), 18.I.1980, P. Goloboff, 1♂ 5♀, 1♀ (MACN-Ar); Los Glaciares Natl. Park, Estancia La Oriental, 14.II.1973, M. Rumboll, 1♀ (MACN-Ar); Los Glaciares Natl. Park, Península Magallanes, in front of Glaciar Moreno, II.1977, D. Pepe and M. Rumboll, 1♀, 1♂ (MACN-Ar); Puerto Coyle, 10 m, 26.XI.1966, M. Irwin and E. Schlinger, 1♀ (CAS); Puerto Deseado, on house wall, IV.1963, Pujals, 1♂ (MACN-Ar); II.1961, Pallares and Zapata, 2♂ (MACN-Ar); X.1964, Pallares, 1♀ (MACN-Ar); II.1966, Pallares, 1♀ (MACN-Ar); XII.1971, A. Gosztonyi, 1♀ (MACN-Ar); Río Seco and Ruta 3, X.1973, M. Rumboll, 1♂ (MACN-Ar); San Julián, XI.1973, M. Rumboll, 1♀ (MACN-Ar); Ventisquero Moreno, 18–24.I.1971, J. Vellard, 2♀ (MACN-Ar). Tierra del Fuego: Bahía Thetis, hanging from roof, no date, Gosztonyi, 1♀ (MACN-Ar); Estancia Herberton, 25.I.1979, Misión Científica Danesa, 1♀ (ZMK); Lago Fagnano, Kaiken, 100 m, 18–19.I.1979, 1♀, 1♂, 21.I.1979, 1♀, Laguna Negra, XII.1989, A. González, 1♀ (MLP); Nueva Herberton, 16.II.1965, M. Birabén, 4♀ (MLP); Río Ewan, I.1975, M. Rumboll, 1♀ (MACN-Ar); Río Grande, XI.1973, M. Rumboll, 1♀ (MACN-Ar); Ushuaia, 1–14.XII.1932, Castellanos and Gomez, 1♂ 2♀ (MACN-Ar); I.1960, A. Bachmann, 1♀ (MACN-Ar); 8–26.II.1961, B. Malkin, 1♀, 1♀ (AMNH); XII.1989, A. González, 1♀ (MLP); Monte Olivia, XII.1989, A. González, 1♀, 1♀, 2♀ (MLP); Nueva Herberton, 16.II.1965, M. Birabén, 4♀ (MLP); Río Ewan and Ruta 3, I.1975, M. Rumboll, 1♀ (MACN-Ar 6802); Ushuaia, II.1951, J. Boero, 1♂ (MLP); Ushuaia, no date, A. del Pino, 2♀ (MACN-Ar 29952). CHILE: Región I (Tarapacá): Parinacota: Parinacota, 2 km S Zapahuira, 3400 m, 18°20′S, 69°34′W, 3400 m, 3.II.1994, N. Platnick, K. Catley, R. Calderón, R. Allen, 1♂ 1♀ penultimate (AMNH). Región III (Atacama): Huasco: El Tránsito to Pinte, 1100–1600 m, 25–27.X.1980, L. Peña, 1♀ (AMNH); Huasco beach, elev. 5 m, 8.X.1992, N. Platnick, P. Goloboff, K. Catley, 1♀ (AMNH). Región IV (Coquimbo): 8.X.1992, N. Platnick, P. Goloboff, K. Catley, 1♂ (AMNH). Elqui: Choros Bajos, 21.X.1992, L. Peña (AMNH); 200 m, 11.XI.1993, 29°33′S, 71°19′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH), 16 km S Cruz Grande, 140 m, 7.X.1992, N. Platnick, P. Goloboff, K. Catley, 3♀ (AMNH); Diaguitas, 18.XI.1963, Gleisner, 1♂ (UC); El Pangue, 20 km S Vicuña, elev. 1500 m, 4.X.1992, N. Platnick, P. Goloboff, K. Catley, 3♀ (AMNH); La Serena, III.1947, L.E. Peña, 1♀ (IG 19736 IRSN); 79 km N La Serena, Rt. 5, km 553, elev. 300 m, 15.X.1992, N. Platnick, P. Goloboff, K. Catley, 3♀, 1♀ (AMNH); Quilacan, 16 km E La Serena, 2.X.1961, R. Wagenknecht, 1♂ (AMNH); 10 mi W Vicuña, “larva ex Puya Castamidae moth”, 3.XII.1950, no collector, 1♀ (CAS). Limarí: 100–500 m, 21.X.1966, E. Schlinger, M. Irwin, 1♂ (CAS); Fray Jorge, Pachingo, 30°27′S, 71°32′W, 29.XII.1966, E. Schlinger and M. Irwin, 1♂ (CAS). Choapa: El Bato (farm in mountains), E Illapel, 10.X.1985, L. Peña, 2♀ (AMNH); Céspedes, Illapel, 1100 m, 13–14.X.1990, L. Peña, 3♂ 1♀ (AMNH); Ñagué, 10 km N Los Vilos, Rt. 5, km 236, elev. 40 m, 31°50′S, 71°31′W, 13.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH), Hacienda Illapel, 600–900 m, 31°36′S, 71°07′W, 19.X.1966, M. Irwin, E. Schlinger and L. Peña, 2♀, 1♂ (CAS); 2 km S Pichidangui, Rt. 5, km 193, elev. 40 m, 32°10′S, 71°31′W, 9.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH), 1♀ (MACN-Ar, photos MJR 1274–1277); Quebrada Los Vilos, 6.XI.1988, P. Goloboff and E. Maury, 1♀ (MACN-Ar). Región V (Valparaíso): Petorca: Quebrada Huaquén, Pichicuy, 6–7.I.1984, P. Goloboff, 1♀ (MACN-Ar); Cuesta El Melón (Metropolitana), 10–12.X.1986, L. Peña, 1♂ (AMNH); 430 m, 8.XI.1993, 32°37′S, 71°14′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ 1♀ (AMNH). Quillota: Cuesta El Melón, nr. La Calera, 15.XI.1985, L. Peña, 1♀ (AMNH); Cuesta La Dormida, N Tiltil, 800–1300 m, 13–18.XI.1982, L. Peña, 1♂ 2♀ (AMNH); 610 m, under rocks, 11.I.1985, N. Platnick and O. Francke, 1♀ (AMNH); Palmas de Ocoa, La Campana Natl. Park, unburned site, 23.VIII.1985, pitfall 1, R. Calderón G., 1♀ (AMNH); Quillota, I.1979, A. Tobar, 1♀ (AMNH). Valparaíso: Archipiélago Juan Fernández, Mas a Fuera (Isla Alejandro Selkirk): 30.I.1935, no collector, 1♀ (AMNH); XII.1965, O. Solbrig, 1♀ (MCZ), 19.I–21.II.1955, G. Kuschel, 2♀ (MLP); Plano de Chosa, 800–1000 m, 29.III.1962, 1♀, 1♂ 1♀, Quebrada Casa, 13–31.III.1962, 1♂ 4♀, 1♀, Quebrada Pangal, Monte Oscuro, 100 m, 9.III.1962, 1♀, Quebrada Vaca, 22.III.1962, 1♂ 3♀, B. Malkin (AMNH). Mas a Tierra (Isla Robinson Crusoe): El Camote, 600–650 m, 25.IV.1962, 1♀, 1♀, 600 m, 19.IV.1962, 1♀, Galpón, Valle Villagra, 23–24.IV.1962, 4♀, Portezuelo, 500 m, 7.IV.1962, 1♂, Portezuelo trail, 7.IV.1962, 1♀, Quebrada Damajuana, 5.IV.1962, 1♀, Valle Anson, Plazoleta del Yunque, 200–250 m, camote side, 1–28.IV.1962, 1♂ 6♀, 1♂ 9♀, Valle Villagra, Portezuelo trail, 400–450 m, 19.IV.1962, 1♀, B. Malkin (AMNH), wet areas near Plazoleta, pans, 24–28.I.1992, S. Marshall, 1♀ (AMNH); Plazoleta, Malaise trap, 24–29.I.1992, S. Marshall, 1♀ (AMNH); Plazoleta El Yunque, 26.I.1992, 2♀, 28.I.1992, 1♀, S. and C. Marshall (AMNH); pans in wet area, 23–29.I.1992, S. Marshall, 4♀ (AMNH); pans nr. Plazoleta campsite, 23–29.I.1992, S. Marshall, 4♀ (AMNH); slopes of El Yunque, fern covered, pan traps, 28.I.1992, S. and C. Marshall, 1♂ (AMNH). Cerro Alegre, 12.IV.1971, Lineros, (UC); Cerro Las Vizcachas, 1800–2200 m, 1–12.XII.1982, L. Peña, 2♂ 5♀ (AMNH); 35 km SE Lago Peñuelas, 350 m, mauco Quintero, II.1979, A. Tobar, 1♀ (AMNH); thorn forest, mesquite flower sweep, 4.XII.1984, S. and J. Peck, 1♀ (AMNH); Río Marga Marga, Los Perales, 33°09′S, 71°19W, 330 m, 13.X.1966, E. Schlinger, M. Irwin, 1♂ (CAS); Valparaíso, II.1954, E. Reed, 1♀, 1♂ (AMNH); XII.1968, H. Sielfeld, 1♀ (UC); Viña del Mar, VIII.1978, A. Tobar, 1♀ (AMNH). San Felipe de Aconcagua: Guardia Vieja, 4.II.1951, Ross and Michelbacher, 2♀ (AMNH); Jahuel (120), no date, L. Peña, 1♀ (IRSN IG 19736); Llay-Llay, 4.II.1951, Ross and Michelbacher, 1♀ (CAS); Los Andes, 30.IX.1983, E. Maury, 1♀ (MACN-Ar). Región Metropolitana (Santiago): Santiago: Aculeo, Cerro San Cristóbal, nr. Santiga City, 500–800 m, 30.XI.1982, L. Peña, 4♀; Aculeo, Las Canchas, 8–11.XII.1983, L. Irrazaval, 1♀, 1♀ (AMNH); Baños de Morales, 3.XI.1995, no collector, 10♀ (AMNH); El Canelo, XII.1958, M. Toro, 1♀ (MACN-Ar); 800–1000 m, 1980, L. Peña, 1♀ (AMNH); El Convento, 18.IX.1966, 33°48′S, 71°43′W, L. Peña, 1♀ (CAS); El Portezuelo, nr. Colina, 500 m, IX–X.1983, L. Peña, 2♂ 1♀ (AMNH); 16 km N La Colina, 30.IX.1992, P. Goloboff, 1♂ (AMNH); Lo Cañas, 29.XI.1982, L.E. Peña, 1♂ (AMNH); 2 km E Embalse El Yeso, 2800 m, 1.III.1974, no collector, 3♀ (UC); Melipilla, La Viluma, 13–14.V.1980, L. Peña, 2♂ (AMNH); flat road before Cuesta Barriga, lado E, 25.I.1971, R. Calderón, 1♀ (UC); Melipilla, San Manuel, 13–14.V.1980, L. Peña, 1♂ (AMNH); Pirque, 30.XI.1982, 1♂, 20.XI.1982, 1♀, L. Peña (AMNH); Quebrada La Plata, near Maipú, 510 m, 33°30′S, 70°55′W, Malaise, 26.I.1966, 1♂, 3–4.X.1966, 1♀, M. Irwin (CAS); Rapel, Estación Hidrobiológica, IV.1977, 1♂ (MHNS 670); Valle del Río Mapocho between El Arrayán and Farellones (Barber traps), 15.X.1958–8.VI.1960, W. Noodt, 1♀, 1♀, 1♂ (MHNS). Cordillera: Río Clarillo Natl. Res., 940 m, 26.XI.1993, 33°44′S, 70°28′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH). Región VII (Maule): Curicó: Las Tablas, E Curicó, II.1985, L. Peña, 3♀ (AMNH); Río Teno, 25–28.XI.1981, L.E. Peña, 5♂ 1♀ (AMNH). Talca: Alto de Vilches, 17–24.X.1964, L. Peña, 1♀ (MCZ), 1.XI.1971, R. Calderón, 1♀, 2♀ (UC); Gil de Vilches, 7.I.1989, M. Ramírez, 1♂, 1♀ (MACN-Ar, photos MJR 5–7), 1200 m, 7–8.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (MACN-Ar, photos MJR 768–771); 5 km W Laguna del Maule, 27.XI.1970, R. Calderón, 1♀ (UC); 10 km Laguna del Maule, 22.XII.1970, R. Calderón, 2♀, 1♀ (UC). Cauquenes: 25 km ESE Cauquenes, 25.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 3♀ (AMNH). Linares: Linares, I.1947, L.E. Peña, 2♀ (IRSN IG 19736); hotel room, 18.I.1985, N. Platnick and O. Francke, 1♂ (AMNH); 16.5 km E Linares, 8.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH). Región VIII (Biobío): Ñuble: Atacalco, just SW Recinto, Chillán area, 17.III.1983, L. Peña, 1♀ (AMNH); Chillán, 31.XII.1975, 2♀, 2.I.1976, 17♂ 18♀, G. Moreno, 2♀ (AMNH), 6♂ 3♀ (MCZ); Chillán, Cuesta de Quilmo, 13.XI.1976, G. Moreno, 3♀ (AMNH); Cobquecura, 8–9.XI.1993, L. Peña, 2♀ (AMNH); Las Cabras, Cordillera de Chillán, 8–15.II.1958, L. Peña, 1♀ (IRSN IG 19736); Las Trancas, 1–10.XII.1965, L. Peña, 1♀, 3♂ (MCZ); E Chillán, 29–30.XI.1990, L. Peña, 1♀ (AMNH); Las Trancas, E Recinto, 1100 m, II.1987, L. Peña, 16♀ (AMNH); Los Lleuques, 5–20.XII.1985, L. Umaña (AMNH); 4 km E road to Pinto, 4.I.1976, B. Moreno, 1♂ 1♀ (AMNH); 1000 m, 1–3.X.1983, L. Peña, 1♀ (AMNH); 40 km E San Carlos, 24.XII.1950, Ross and Michelbacher, 1♂ (CAS). Concepción: Bajada Chivilingo, 15.XI.1992, T. Cekalovic, 1♀ (AMNH); Caleta Chome, 10.I.1997, T. Cekalovic, 2♀ (AMNH); Estero Nonguén, 11.XI.1996, 2♀, 2♀, 13.I.1997, 1♀, T. Cekalovic (AMNH); Fundo El Manzano, 7.XII.1996, T. Cekalovic, 1♀ (AMNH); Fundo El Venado, 6.I.1996, T. Cekalovic, 1♀ (AMNH); Hualqui, 11.VIII.1996, T. Cekalovic, 1♀ (AMNH); Laguna San Pedro, 23.XI.1994, T. Cekalovic, 1♀ (AMNH); Laraquete, 8.XII.1988, T. Cekalovic, 1♀ (AMNH); Lomas Colorada, 24.XI.1996, T. Cekalovic, 2♀, 1♂ (AMNH); Penco, 23.III.1980, T. Cekalovic, 1♀ (AMNH); Periquillo, 3.XI.1996, 4♀, 21.XII.1996, 1♀, 29.XII.2000, 2♀, 10♀, T. Cekalovic (AMNH). Arauco: 2 km S 5 km N Curanilahue, 20.X.1996, T. Cekalovic, 1♀ (AMNH); Cruce Camino Colicó Norte, 20.X.1996, T. Cekalovic, 1♀ (AMNH). Biobío: 2.5 km E El Abanico, 760–975 m, scrubby mountainside, under rocks, 20–21.XI.1981, N. Platnick and R. Schuh, 2♀ (AMNH); Caledonia, E Mulchen, 700–900 m, 6–15.II.1981, L. Peña, 1♀ (AMNH); Guallali, Lag. El Barco, 1200 m, 21–28.II.1981, L. Peña, 3♀ (AMNH); W Ralco, Santa Bárbara, 400 m, 22–23.XI.1994, L. Peña, 2♀ (AMNH); 5 km W Tucapel, 28.XII.1950, Ross and Michelbacher, 4♀ (CAS). Región IX (Araucanía): Malleco: 40 km W Angol, Nahuelbuta Natl. Park, FITS, 1200–1500 m, Nothofagus/Araucaria forest, 9.XII.1984–17.II.1985, S. and J. Peck, 1♂ (AMNH); Cordillera de Las Raíces, 1600–1800 m, 13–18.II.1980, L. Peña, 1♂ 1♀ 3♀ penultimates (AMNH); El Manzano (Arauco/Malleco), Cordillera Nahuelbuta, 3–5.III.1986, L. Peña, 1♀ (AMNH); Estero Huemul, tributary of Lago Gualletué, nr. Marimeneuco, 11.XII.1963, G. Edwards, 1♂ 1♀ (AMNH); Lago Gualletué, nr. Marimenuco, 10.XII.1963, G.F. Edmunds, 1♂ (AMNH); Malalcahuello, 9–15.XII.1985, L. Peña, 7♂, 2♀, 2♀, 3♂ (AMNH); Nahuelbuta Natl. Park, 1200 m, 23.I.1951, Ross and Michelbacher, 2♀, 1♀ (CAS), 1200–1400 m, mixed forest with Araucaria, 26.I.1985, N. Platnick and O. Francke, 1♀ (AMNH), 1300 m, 1–6.II.1979, L. Peña, 1♀, 1♀ (AMNH); 4350 m, 38°01′S, 73°13′W, 24.I.1967, M. Irwin, 1♀ (CAS); Cordillera Nahuelbuta, 15–20.XII.1993, L. Peña, 1♀ (AMNH), 18–20.XII.1993, L. Peña, 1♀, 1♀ (AMNH); Tolhuaca, 15–23.III.1986, L. Peña, 2♀ (AMNH); Tolhuaca, Laguna Malleco, 4.III.1978, T. Cekalovic, 5♀ (AMNH). Cautín: Estero Molco, 24.II.1988, T. Cekalovic 1♀ (AMNH); Volcán Villarrica, FIT in “tundra”, 8.XI–3.XII.1989, S. Marshall, 1♀ (AMNH); Volcán Villarrica, FIT nr. edge of old lava flow, 10.XI–3.XII.1989, S. Marshall, 2♂ (AMNH); Pucón, 15.XI–2.XII.1980, Malaise trap in peninsula, S.A. Marshall, 1♂, 1♂ (AMNH); Pucón, lakeshore FIT, 15.XI–2.XII.1989, 1♂, FIT nr. lake, 8–13.XI.1989, 1♂, S.A. Marshall (AMNH); Fundo La Selva, W Temuco, NW Nueva Imperial, 700 m, 9–12.XII.1981, L.E. Peña, 2♀ (AMNH); Villarrica, 1–30.I.1965, L. Peña, 3♀ (MCZ); NE Villarrica, 16–31.XII.1964, L. Peña, 1♂ 1♀ (MCZ); Volcán Llaima, 16.I.1972, R. Ladrón de Guevara, 1♀ (UC). Región X (Los Lagos): Valdivia: Corral, dead inside retreat, 17.I.1989, M. Ramírez, 1♀ (MACN-Ar); Las Trancas, 1200 m, 24–27.XI.1994, L. Peña, 1♀, 2♂ 3♀ (AMNH); Peulla, Río Nahuilán, 24 km SE Corral, 16.I.1989, M. Ramírez and E. Maury, 1♀ (MACN-Ar); Santo Domingo, en Blechnum magellanicum, 26.X.1984, D. Jackson, 1♀ (MHNS 886); Valdivia, 20 m, 23.IX.1981, Nielsen and Karsholt, 1♂ (ZMK); XI–XII.1982, E. Krahmer, 1♀ (MHNS 700); 30 km S Valdivia, Ross and Michelbacher, 1♀ (CAS). Osorno: 2 km E Puente Río Golgol, 14.II.1992, T. Cekalovic, 2♀ (AMNH); Puyehue Natl. Park: Aguas Calientes, 500 m, 2–5.V.1988, L. Peña, 1♀ (AMNH); 600 m, 12–20.II.1979, L.E. Peña, 1♀ (AMNH); Antillanca rd., 470–720 m, valdivian rainforest, screen-sweeping at dusk, 965 m, trap site 658, window trap, Nothofagus pumilio forest, 18–24.XII.1982, A. Newton and M. Thayer, 1♂ (AMNH), 18–25.XII.1982, A. Newton and M. Thayer, 1♂ (AMNH); 40°46′30″S, 72°11′30″W, 1050–1350 m, alpine meadow, pitfall 59T1, 30.XII.2000, J. Miller, I. Agnarsson, Alvarez, J. Coddington, G. Hormiga, (USNM); Los Derrumbes road, Aguas Calientes, 480 m, 40°44′S, 72°18′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH); 10 km E Puyehue, 24.I.1951, Ross and Michelbacher, 1♀ (CAS); Volcán Casablanca, 1130–1180 m, site 665, pan traps above tree line, valdivian rainforest, 20–25.XII.1982, A. Newton and M. Thayer, 1♂ (AMNH); Termas de Puyehue, 260 m, logs, stones, 12.III.1965, H. Levi, 1♀ (MCZ); Osorno, X.1977, 1♀, 1♀, XII.1978, 1♀, A. Tobar (AMNH); II.1967, L. Peña, 1♀ (MCZ); Pucatrihue, II.1967, L. Peña, 1♀ (MCZ). Llanquihue: Los Muermos, S. Chile, forest, 19.I.1951, Ross and Michelbacher, 1♀ (CAS); nr. Petrohué, 200 m, vegetation, 21.III.1965, H. Levi, 1♀ (MCZ); 5 km S Puerto Montt, 17.III.1991, T. Cekalovic, 1♀ (AMNH); Puerto Montt, Río Blanco, 24.XI.1995, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH); 24–29.I.1983, G. Arriagada, 1♀ (MHNS 718). Chiloé: Isla de Chiloé: no date, Skottsberg, 1♂ (NRS); 15 km S Chepu, beach of round stones, over upper tide line, 3.II.1991, M. Ramírez, 1♀ (MACN-Ar); Pid-Pid, 10–12.III.1987, L. Peña, 3♀ (AMNH); Isla Lemuy, Ichuac, 20.II.1996, T. Cekalovic, 1♀ (AMNH). Palena: Chaitén, XII.1985, L. Peña, 1♀ (AMNH). Región XI (Ibáñez del Campo): Aisén: Balmaceda, 17–22.I.1961, L. Peña, 9♀ (IRSN IG 23077); Murta, Lago General Carrera, 29–30.I.1990, L. Peña, 1♀ (AMNH); Río Simpson Natl. Park, N margin, 17.II.1991, M. Ramírez, 2♀ (MACN-Ar); 20 km E Puerto Aisen, 26.I.1961, L. Peña, 1♀ (IRSN IG 23077); Río Cisnes, 1–28.II.1961, L. Peña, 1♀ (IRSN IG 23077); Río Ibáñez, 27–28.I.1990, L. Peña, 2♀ (AMNH). Región XII (Magallanes y Antártica): Ultima Esperanza: Torres del Paine Natl. Park: Lago Sarmiento de Gamboa, 51°2′0″S, 72°46′15″W, 100 m, steppe, 6–9.XII.2000, J. Miller, I. Agnarsson, 1♀ (USNM), same, ground, 3♀ (USNM), near Refugio Chileno, 50°56′45″S, 72°55′0″W, 400–600 m, 8–9.XII.2000, J. Miller, I. Agnarsson, 4♀, 1♀, 1♀, 2♀ (USNM), Laguna Larga, 51°1′30″S, 72°52′45″W, 300 m, under rocks in steppe, 7.XII.2000, J. Miller, I. Agnarsson, 6♀, 2♀ (USNM). Magallanes: Cañadón Bombalot, 29.I.1976, T. Cekalovic, 1♀ (AMNH); Estancia Gazy Harbour, 10.II.1990, T. Cekalovic, 2♂ 2♀ (AMNH); Estancia La Vicuña, 1956, 1♀, 1♀, 2♀, 2♀, no. 14, 4.III.1957, 1♀, no. 17, 15.II.1959, 1♀, no. 17, monte seco, 400 m, 15.II.1959, 4♀, J. Vellard (MACN-Ar); Estancia La Vicuña, SE Camerón, 1–6.XII.1960, L. Peña, 4♀ (MCZ); 35 km SW Camerón, Nothofagus assoc., 2.XII.1966, E. Schlinger and M. Irwin, 2♀ (CAS); Estancia Virgen de Lourdes (Sector Dinamarquero), 6.II.1990, 1♀, 8.II.1990, 3♀, T. Cekalovic (AMNH); Isla Dawson, Puerto Harry, 8.VIII.1976, T. Cekalovic, 1♀ (AMNH); Isla Grande, NW Tierra del Fuego, 12–15.XI.1961, L. Peña, 1♀ (MCZ); Isla Navarino, 16.III.1961, B. Malkin, 1♀ (AMNH); Laguna Amarga, 14–21.XII.1960, L. Peña, 1♀, 2♀, 3♀ (MCZ); 21.IV.1962, T. Cekalovic, 1♂ (AMNH); 4 km W Laguna Amarga, 8.XII.1966, E. Schlinger and M. Irwin, 10♀, 2♀, 1♀, 3♀ 3♀ (CAS); Magallanes, XI.1960, T. Cekalovic, 1♀ (MACN-Ar); Torres del Paine Natl. Park, 150 m, scrub, 10.II.1985, N. Platnick and O. Francke, 2♀ (AMNH); Península Brunswick, Barranco Amarillo, 27.I.1976, T. Cekalovic, 2♀ (AMNH); Puerto Hambre, 25.III.1991, T. Cekalovic, 2♀ (AMNH); Punta Arenas, 17.IX.1963, 1♀, (La Turba), 27.VIII.1976, 1♀, T. Cekalovic (AMNH), Punta Arenas, Quinta Pittet, 29.II.1969, T. Cekalovic, 1♀ (AMNH); 102.2 km NNW Punta Arenas, 430 m, Nothofagus assoc., 6.XII.1966, E. Schlinger and M. Irwin, 2♀, 3♀, 1♀ (CAS); Punta El Monte, 27.I.1976, T. Cekalovic, 1♀ (MCZ); Cerro Castillo Natl. Res., 500–600 m, dry forest, 7.II.1985, N.I. Platnick and O.F. Francke, 1♀ (AMNH); Cerro Castillo, Natales, 13.XII.1960, L. Peña, 2♀, 1♂ 6♀ (MCZ); Río Chico, 1956, 3♀, 1♀, 17.II.1956, 2♀, J. Vellard (MACN-Ar); Laguna Parrillar Natl. Res., 53°24′15″S, 71°15′45″W, beating, 1–10.XII.2000, 350 m, J. Miller, I. Agnarsson, 1♂ 3♀ (USNM); Río San Juan, 25.I.1976, T. Cekalovic, 1♀ (AMNH); Rubens, 22.III.1948, M. Birabén, 4♀ (MLP), 13.XII.1960, L. Peña, 2♀, 1♂ 8♀ (MCZ); Rusffin, no. 11, III.1957, J. Vellard, 1♀, 1♀, 3♀ (MACN-Ar); Rusffin, SE Cameron, 17–20.XI.1960, L. Peña, 9♂ 39♀ (MCZ); Tres Vientos, Puerto Arturo, 53°34′S, 73°23′W, 25–28.XI.1960, L. Peña, 1♂ (MCZ); Aserradero Yendegaia, no. 2, 12.II.1957, 10♀, 1♂ 5♀, no. 3, 13.II.1957, 1♀, J. Vellard (MACN-Ar). Mistaken Locality: Santiago Prov., Malleco, XI.1979, L. Peña, 4♂, 1♀, 1♀, 1♀ (AMNH) (see Ramírez, 1995b: 83).

    Sanogasta backhauseni (Simon), new combination Figures 61D, 77F, G, 84C, D, 85A–E

  • Tomopisthes backhauseni Simon, 1895: 168, 172 (female holotype from Tierra del Fuego, in MHNP 16093, examined; male allotype from Tierra del Fuego, not paratype, designated subsequently by Simon, 1896a, but in the same vial 16093, examined), 1896a: 142, 144, 1897a: 91, 1902: 32.

  • Gayenna trilineata Tullgren, 1901: 240, 259 (four syntypes from Tierra del Fuego: one female from Porvenir, 23.XII.1895, one male from Cordillera 54°S, I.1896, two females from Páramo, 10.I.1897, in NRS, examined). Synonymized by Simon, 1902: 32.

  • Tomopisthes backhauseni patagonicus Simon, 1905b: 12 (female holotype from Argentina, Santa Cruz, Silvestri coll., in MHNP 12689, examined). I did not attempt to review the status of subspecies in this contribution.

  • Diagnosis:

    Resembles S. minuta in genitalia, but distinguished by having three dorsal longitudinal stripes on the abdomen, and by lacking a narrow epigynal posterior notch between the lateral lobes.

    Female (holotype, measurements of specimen from Los Glaciares Natl. Park):

    Total length 6.90. Carapace length 3.07, width 2.17, wider on legs II–III. Length of tibia/metatarsus: I, 1.70/1.53; II, 1.70/1.57; III, 1.63/1.73; IV, 2.33/2.73. Palpal tarsus length 0.92. Chelicerae with two teeth on retromargin. Sternum length 1.60, width 1.20. Spines: leg I, femur d 1–1–1, p 0-d1–2, r 0-d1-d1; tibia v 2–2–2, p 0–1; metatarsus v 2bas. II, femur = I; tibia v r1-r1–2, p 0–1 or 1-0-1-0; metatarsus = I. III, femur = I; patella r d1; tibia v p1–2–2, p and r 1-d1-1-d1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia = III; metatarsus = III or d 2ap. Abdomen length 4.25, width 2.25, spiracle–epigastrium 1.57, spiracle–spinnerets 1.00. Color: pale gray with darker pattern. Legs with small dots at bases of lateral and dorsal spines. Femora with ventral longitudinal dark spot. Patellae with dark spots p and r 1-0-1-0. Sternum, labium, and endites pale. Abdomen with dark cardiac area, continued in two dark stripes (fig. 77F), venter with three longitudinal bands from epigastrium to tracheal spiracle. Epigyne and spermathecae (figs. 84C, 85C–E) very similar to those of S. minuta, anterior pouch very close to epigastric furrow, hidden between lateral lobes.

    Male (allotype, measurements of specimen from Calafate):

    Total length 6.65. Carapace length 2.97, width 2.33. Length of tibia/metatarsus: I, 2.33/2.13; II, 2.20/2.10; III, 2.00/1.10; IV, 2.67/3.13. Chelicerae smaller than those of female. Sternum length 1.53, width 1.20. Spines as in female, except: leg I, femur p 2ap; tibia p and r 1-d1-1-0; metatarsus p d1–1–0. r 1. II, femur p 0-d1–2 or 0-d1-d1; tibia v 2–2–2 (the p1bas smaller), p 1-d1-1-d1 or 1-d1-0-1, r 1-d1-1-0; metatarsus p and r d1–1–0. IV, femur = II; tibia p and r 1-d1-1-d1. Abdomen length 3.85, width 2.25, spiracle–epigastrium 1.17, spiracle–spinnerets 1.13. Color similar to female (fig. 77G). Palp (figs. 61D, 84D, 85A, B): tibia short, width/length 0.87, cymbium relatively large. Copulatory bulb very similar to that of S. minuta.

    Variability:

    Several specimens have supernumerary spines (e.g., a pair where only one is expected). Females spines: II, tibia v r1–2–2. III, IV, tibia v 2–2–2, p and r 1-d1-1-0. Specimens from the seashore in Chubut and Buenos Aires provinces are pale yellow, almost lacking any pattern. A male from Ñorquinco, Rio Negro province, has a dark abdomen with pale venter. Some females have epigynal lateral lobes not fused, limiting deep notch with parallel borders, similar to S. minuta.

    Natural History:

    Unknown.

    Distribution:

    In Argentina, Neuquén, Río Negro, and Chubut provinces, seashore of Buenos Aires. In Uruguay only known from Montevideo, in Chile only from Magallanes.

    Other Material Examined:

    URUGUAY: Departamento Montevideo: Canelones, Marindia, 8.IV.1976, F. Costa, R. Capocasale, 1♂ (CAS). ARGENTINA: Buenos Aires: Carmen de Patagones, no date, excursion J.B. Daguerre and Carcelles, 1♂ (MACN-Ar 36829), 1♀ (MACN-Ar); Mar del Tuyú, costa atlántica (walking over sand beach), 2.V.1981, M. Ramírez, 1♀ (MACN-Ar); Quequén, no date, J.B. Daguerre, 1♀ (MACN-Ar); San Blas, Patagones, Carcelles, 1♂ 1 immature (MACN-Ar 36833); Sierra de la Ventana, 22.XI.1972, E. Maury, 1♀ (MACN-Ar). Neuquén: Laguna Blanca Natl. Park, N26, 30.IV.1964, no collector, 1♀ (MACN-Ar); Piedra del Aguila, V.1972, Gentili, 2♂ (MACN-Ar). Río Negro: San Carlos de Bariloche, II.1954, M.E. Galiano, 1♀ (MACN-Ar 5412), Ñorquinco, 11.X.1961, 1♀, 11.V.1962, 1♂ 8♀, 27.VIII.1962, 1♀ 2 immatures, 3.VII.1966, 3♂ 6♀, VIII.1962, 1♀, A. Kovács (AMNH). Chubut: El Maitén, 20.VI.1962, A. Kovács, 1♂ 4♀ (AMNH); Leleque, 71°06′W, 42°28′W, 12.II.1965, A. Kovács, 1♀ (AMNH); Puerto Madryn, V.1975, M. Rumboll, 1♂ 1♀ (MACN-Ar 6852); Península de Valdez, Puerto Pirámides, 11.V.1970, M. Rumboll, 1♀ (MACN-Ar); II.1980, P. Goloboff, 1♂ (MACN-Ar); 42°34′S, 64°17′W, 12.XI.1988, V. and B. Roth, 1♂ (CAS); Río Turbio, 28.V.1976, M. Rumboll, 1♀ (MACN-Ar). Santa Cruz: Calafate, 23.I.1980, P.A. Goloboff, 1♂ 2♀ (MACN-Ar); Lago Argentino, Estancia La Cristina, 200 m, 17.III.1953, A. Willink, 1♂ (MACN-Ar); Las Cruces, Tres lagos, 9.III.1948, M. Birabén, 2♀ (MLP); Los Glaciares Natl. Park: 2.II.1973, M. Rumboll, 1♀ (MACN-Ar); Los Glaciares Natl. Park, Estancia La Oriental, 840 m, Y. p. tr., 14–17.II.1998, C. and M. Vardy, 1♀ (BMNH/MACN-Ar). Tierra del Fuego: Río Grande, 21.I.1960, A. Bachmann, 1♂ (MACN-Ar); 3.II.1965, 1♀ (MLP). CHILE: Región XII (Magallanes y Antártica): Magallanes: Estancia Virgen de Lourdes (Sector Dinamarquero), 6.II.1990, S. Cekalovic, 2♀ (AMNH); Isla Riesco, Posomby, 31.I.1976, T. Cekalovic, 1♂ (AMNH); Magallanes, no specific locality, II.1957, 1♂ (MHNS 50).

    Sanogasta x-signata (Keyserling), new combination Figures 61E, 84E, F, 85F–H

  • Gayenna x-signata Keyserling, 1891: 138 (two females syntypes from Brazil, state of Rio Grande do Sul, no specific locality, should be in BMNH, not found). Mello-Leitão, 1925: 457.

  • Sanogasta sp.: Ramírez, 1995a: 361.

  • Samuza sp.: Ramírez, 1995a: 382 (lapsus).

  • Identification:

    The illustration of the epigyne by Keyserling (1891) is sufficient to identify this distinctive species.

    Diagnosis:

    Easily distinguished from other Sanogasta by having the epigynal posterior margin projecting over the epigastric furrow, and by the complex-shaped paramedian apophysis and the secondary conductor.

    Female (Atucha, 27.VII.1985):

    Total length 4.10. Carapace length 1.70, width 1.18, wider on legs II–III. Length of tibia/metatarsus: I, 0.75/0.65; II, 0.75/0.65; III, 0.50/0.72; IV, 0.92/1.13. Palpal tarsus length 0.47. Chelicerae unmodified, with two teeth on retromargin. Sternum length 0.92, width 0.70. Spines: leg I, femur d 1–1–1, p d1ap; tibia v 2–2-p1; metatarsus v 2bas. II, femur = I; tibia v r1–2-p1; metatarsus v 2bas. III, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–0–2 and an apical group of thick setae, p and r d1–1–1, d p1–2. IV, femur = III; patella r 1; tibia v p1–2–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2-p1–2 and an apical group of thick setae, p and r d1–1–1, d p1–2. Abdomen length 2.43, width 1.50, spiracle–epigastrium 0.92, spiracle–spinnerets 0.42. Color: carapace and legs pale gray with darker pattern. Abdomen cream with dorsal gray pattern, venter with a few dark dots, recurved transverse mark anterior of tracheal spiracle. Epigyne (fig. 85G, H): posterior margin projecting over epigastric furrow, with two contiguous pouches where copulatory openings lie. Insertions of epigastric muscles superficial. Copulatory ducts arising contiguously. Ducts of accessory bulbs very short, ventral.

    Male (Atucha, 10.V.1987):

    Total length 4.15. Carapace globose, length 2.00, width 1.60. Length of tibia/metatarsus: I, 1.52/1.35; II, 1.47/1.37; III, 1.18/1.28; IV, 1.50/1.80. Chelicerae small, vertical. Sternum length 1.02, width 0.80. Spines as in female, except: leg I, femur p 2ap; tibia v 2–2–2. II, tibia v r1–2–2. III, tibia v 0-p1–2 or p1-p1–2. IV, metatarsus v 2–2–2. Abdomen length 2.17, width 1.33, spiracle–epigastrium 0.80, spiracle–spinnerets 0.47. Color: similar to female, mark anterior of spiracle diffuse. Palp (figs. 61E, 84E, F, 85F): tibia short, width/length 0.80, cymbium relatively large. Apical hematodocha rugose, prolonged between margin of tegulum and paramedian apophysis. Embolus thin, basal process triangular. Median apophysis thin, wide at base, narrow at apex. Tip of paramedian apophysis long, sinuous. Secondary conductor wide, thin, not fused to anterior margin of tegulum (fig. 84F); canal conspicuous, arising at base of paramedian apophysis; anterior border projecting, thin; retrolateral portion with rugose external projection, internal, conical projection. Anterior margin of tegulum compressed over base of secondary conductor.

    Variability:

    Female spines: III, metatarsus v 2–2–2.

    Natural History:

    This species builds retreats under bark or on dry leaves, and has also been collected on epiphytic bromeliads of the genus Tillandsia.

    Distribution:

    Southwestern Brazil, Uruguay, and Argentina, from Buenos Aires to the north; one isolated southern record from Argentina, Río Negro, suggest a wider distribution.

    Other Material Examined:

    BRASIL: São Paulo: São José do Barreiro, Serra da Bocaína, 1960 m, XI.1968, M. Alvarenga 1♀ (AMNH). Rio Grande do Sul: Cachoeira do Sul, Capanézinho, 12.V.1993, R.G. Buss, 1♀ (MCTP 3613); Cachoeira do Sul, Cordilheira, 14.X.1992, 1♂ (MCTP 3266), 12.X.1993, 1♀ (MCTP 4276), R.G. Buss; Cachoeira do Sul, Porteira Sete, 13.XI.1993, R.G. Buss, 1♂ (MCTP 4217); Pelotas (após a ponte), 9.V.1967, Biasi, 1♀ (MZUSP 14087); Pelotas, V.1964, C. Biezanko, 3♀ (AMNH); Quaraí, 24–28.V.1991, A. Lise, 2♀ (MCTP 0456). URUGUAY: Departamento Río Negro: Arroyo Negro, 15 km S Paysandú, 3.I.1963, R.G. Van Gelder, 1♀ (AMNH). ARGENTINA: Salta: El Alisal, 45 km W Salta, 1950 m, cañadón húmedo, Malaise-FIT trap, 1–29.XII.1987, S. and J. Peck, 1♀ (AMNH); El Rey Natl. Park, 880 m, Río Los Puertos, Prosopis forest, Malaise-FIT traps 87–145, 6–16.XII.1987, S. and J. Peck, 1♀ (AMNH). Jujuy: Las Capillas, XI.1956, Fritz, 1♂ (MACN-Ar). Corrientes: Paso de los Libres, II.1971, Bejarano, 1♀ (MACN-Ar). Catamarca: Estancia El Chorro, 1–15.II.1953, Partridge, Rodríguez, 1♀ (MACN-Ar). Córdoba: Alta Gracia, 3er paredón, XI.1985, Galiano, 1♀ (MACN-Ar); Calamuchita, III.1954, J.M. Viana, 1♀ (MACN-Ar); III–IV.1958, J.M. Viana, 1♀ (MACN-Ar); II.1959, J.M. Viana, 1♂ (MACN-Ar); Horco Molle, 6–13.XI.1966, M.E. Galiano 1♀ (MACN-Ar); La Cumbre, 8.XI.1991, M. Ramírez, 1♀ 4 immatures (MACN-Ar); Lago San Rafael, 20.XI.1983, M. Galiano, 1♀ (MACN-Ar). Santa Fe: Las Gamas, 20 km W Vera, 27–30.X.1994, M. Ramírez and J. Faivovich, 2♂ 1♀ (MACN-Ar). Entre Ríos: Río Gualeguaychú and Ruta Nac. 14, 10.XII.1982, M. Ramírez and P. Goloboff, 1♂ (MACN-Ar); Rosario del Tala, Ruta Prov. 38 and Río Gualeguay, 11.I.1988, P. Goloboff, C. Szumik, 1♀ (MACN-Ar); Salto Grande, IV.1977, J.M. Viana, 1♀ (MACN-Ar); no specific locality, 1942, Haedo, 1♀ (MACN-Ar). Buenos Aires: Atucha, on Tillandsia sp., 27.VII.1985, P. Goloboff, M. Ramírez, 4♀ (MACN-Ar), 10.V.1987, M. Ramírez, 1♂ 1♀ (MACN-Ar); Isla Martín García, 25.V.1990, M. Ramírez, 2♂ 2♀ (MACN-Ar); Ciudad de Buenos Aires, 27.XII.1983, M. Ramírez, 1♀ (MACN-Ar); La Plata, 22.XII.1978, P. Goloboff, 1♂ (MACN-Ar); San Antonio de Areco, IV.1982, M. Ramírez, 1♂ (MACN-Ar); Punta Lara, Ensenada, V.1954, J.M. Viana, 1♀ (MACN-Ar); Santa Teresita, II.1984, M. Ramírez, 1♀ (MACN-Ar); Tandil, IV.1963, Ogueta, 1♂ (MACN-Ar); Tigre, I.1938, J.M. Viana, 2♀ (MACN-Ar); Tigre, VI.1955, J.M. Viana, 13♂ 6♀ (MACN-Ar); Zelaya, 22.IX.1968, H. Hepper, 1♂ (MACN-Ar). Río Negro: El Bolsón, Cerro Piltriquitrón, 3–4.II.1985, M. Ramírez, 1♀ (MACN-Ar).

    Sanogasta minuta (Keyserling), new combination Figures 84A, B, 86A–D

  • Samuza minuta Keyserling, 1891: 139 (syntypes in two vials: two females, one male, and one immature male, and five females, two males, and four immatures, from Brazil, state of Rio Grande do Sul, no specific locality, in BMNH, examined).

  • Gayenna minuta: Petrunkevitch, 1911: 485. Mello-Leitão, 1925: 456.

  • Diagnosis:

    Resembles S. backhauseni in genitalia, but distinguished by having almost contiguous epigynal lateral lobes, limiting narrow notch. Males are smaller, their abdomen lacks the three definite dorsal longitudinal stripes.

    Female (Huerta Grande):

    Total length 6.65. Carapace length 2.63, width 1.90, wider on legs II–III. Length of tibia/metatarsus: I, 1.33/1.15; II, 1.27/1.17; III, 1.10/1.20; IV, 1.53/2.17. Palpal tarsus length 0.77. Chelicerae with two teeth on retromargin. Sternum length 2.67, width 1.93. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2–2; metatarsus v 2bas. II, femur = I; tibia v r1–2–2; metatarsus v 2 bas. III, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1-p1–2, p 1-d1-1-0, r d1–1, d r1bas; metatarsus v 2–0–2, p and r d1-1-0-1, d 0-p1–2. IV, femur = III; patella r 1; tibia v p1–2–2, p 1–1, r 1-d1-1-0, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. Abdomen length 4.00, width 2.60, spiracle–epigastrium 1.77, spiracle–spinnerets 0.83. Color: pale grayish with dark pattern. Sternum pale. Venter with line of small dots anterior of tracheal spiracle. Epigyne (figs. 84A, B, 86C, D): anterior pouch small, very close to epigastric furrow, reduced between lateral lobes. Copulatory ducts arising contiguously. Ducts of accessory bulbs very short, ventral.

    Male (Huerta Grande):

    Total length 5.70. Carapace length 2.80, width 2.07. Length of tibia/metatarsus: I, 1.70/1.43; II, 1.60/1.37; III, 1.35/1.52; IV, 1.83/2.07. Chelicerae slightly smaller than those of female. Sternum length 1.32, width 0.98. Spines as in female, except: leg I, tibia p 1-0-1-0. II, tibia = I. III, femur p 0-d1–2, r 0-d1-d1; tibia v 2–2–2, p and r 1-d1-1-0; metatarsus v 2-p1–2. IV, femur p 0-d1–2; tibia p and r 1-d1-1-0. Abdomen length 2.90, width 1.40, spiracle–epigastrium 1.00, spiracle–spinnerets 0.50. Color as in female, but darker cardiac area. Palp (fig. 86A, B): tibia short, width/length 0.93, cymbium relatively large. Embolus thin, basal process small, triangular. Median apophysis very thin, closely associated with paramedian. Apex of paramedian short, sinuous. Secondary conductor not fused to anterior margin of tegulum, canal conspicuous, arising at base of paramedian apophysis; retrolateral portion with external rectangular rugose projection. Anterior margin of tegulum compressed over base of secondary conductor.

    Variability:

    Female spines: III, tibia v p1–2–2; metatarsus v 2-p1–2.

    Natural History:

    Unknown.

    Distribution:

    Southwestern Brazil and central Argentina. Probably also in northeastern Argentina and Uruguay.

    Other Material Examined:

    BRASIL: Rio de Janeiro: Petrópolis, no date, Mello-Leitão, 1♂ 3♀ (MNRJ 671). São Paulo: Botucatu, Rubião Júnior, 15.VII.1964, V.C. Jesus, A. Montover, 1♀ (MZUSP 14064), 24.V.1978, M. Carneiro, 1♀ (MZUSP 14.051); São Paulo, Jabaquara, 20.VI.1943, A. Zoppei, 1♀ (MZUSP 9845); Monte Alegre do Sul, Amparo, Fazenda de Santa María, 18–19.XII.1942, B.A.M. Soares, 1♀ (MZUSP 14077); São Bernardo, Repressa Nova, 12.X.1041, F. Leme, 3♂ 6♀ 2 immatures (MZUSP 14075); Santo André, 19.XI.1941, J. Domingo, 1♀ (MZUSP 14055). Paraná: Guarapuava, ponte do Rio Coutinho, 28.IV.1967, P. Biasi, 1♀ (MZUSP 7035). Rio Grande do Sul: Capaõ Novo, Capaó da Canoa, 17–18.IV.1993, A. Lise, 2♀ (MCTP 3135). ARGENTINA: Catamarca: Ruta 4 km 9–24, road to El Rodeo, IX.1981, A. González, 1♀ (MACN-Ar). Córdoba: Calamuchita, XI.1941, J. Viana, 1♀ (MACN-Ar); Calamuchita, “El Sauce”, XII.1941, 6♀ (MACN-Ar 1890); Huerta Grande, II.1943, 1♂ 6♀ (MACN-Ar). San Luis: Merlo, XI.1985, M.E. Galiano, Miranda, 3♀ (MACN-Ar); Papagallos, 9.XI.1982, E. Maury, 1♀ (MACN-Ar); Villa Elena, X.71, J. Viana, 1♀ with embolus inserted (MACN-Ar). La Pampa: Lihuel Calel Natl. Park, XI.1969, E. Maury, 1♀ (MACN-Ar). Buenos Aires: La Plata, 1942, 1♂ (MLP); Rosas, F.C.G.R., 1♀ (MACN-Ar); Sierra de la Ventana, Pque. Prov. E. Tornquist, 18–20.IX.1982, M. Ramírez, 1♀ (MACN-Ar); Tandil, III.1963, Ogueta, 1♀ (MACN-Ar); IV.1963, Ogueta, 2♀ (MACN-Ar); V.1967, E. Maury, 1♀ (MACN-Ar); Tandil, La Cascada, 16.V.1973, C. Cesari, 1♀ (MACN-Ar).

    Sanogasta puma, new species Figures 76E, 86E–H, 87

    Types:

    Female holotype and male paratype from Argentina, Buenos Aires province, Isla Talavera, 2 km E Zárate, ca. 34°06′S, 59°02′W, 3.XI.1996, M. Ramírez, deposited in MACN-Ar 9815.

    Etymology:

    The specific name is a noun in apposition taken from the felid Puma concolor, referring to the uniformly grayish brown color of the body.

    Diagnosis:

    Resembles S. tenuis in having an elongate body without pattern, but can be distinguished by having ovate, voluminous spermathecae, with pointed anterior ends, and a relatively large secondary conductor.

    Female (holotype):

    Total length 6.12. Carapace flattened, without thoracic groove, length 2.47, width 1.50, wider at leg II. Length of tibia/metatarsus: I, 1.35/1.13; II, 1.18/0.98; III, 0.83/0.82; IV, 1.55/1.22. Palpal tarsus length 0.60. Chelicerae unmodified, with two teeth on retromargin. Sternum length 1.57, width 0.82. Spines: leg I, femur d 1–1–1, p 0-d1–2, r d1ap; tibia v 2–2–2; metatarsus v 2bas. II = I. III, femur d 1–1–1, p and r 0-d1-d1; tibia v p1–2–2, p v1-v1 + 0–1–0 bristle, r d1–1, d r1–0–1 bristles; metatarsus v 2–0–2, p 1–1, r 1–1, d 0-(p1 bristle)-2. IV, femur d 1–1–1, p and r d1ap; tibia v p1–2–2, p v1-v1–0 + 0–1–0 bristle, r 1-d1–1, d r1–0–1 bristles; metatarsus v 2–2–2 or 2-p1–2, p 0-v1-0-v1, r d1–1–1, d 0-(p1 bristle)-0–2 (the p1ap is a bristle). Prolateral spines on tibiae III and IV ventrally displaced. Abdomen length 3.65, width 1.47, spiracle–epigastrium 1.90, spiracle–spinnerets 0.73. Color: grayish brown uniform (fig. 76E), paler on venter, abdomen darkening posteriorly, membranous base of anterior spinnerets dark. Epigyne (figs. 86G, H, 87): anterior pouch advanced, longitudinal, without definite cavity. Copulatory ducts arising contiguously. Ducts of accessory bulbs very short, ventral. Spermathecae ovate, anteriorly acute.

    Male (paratype):

    Total length 5.45. Carapace with thoracic groove scarcely marked, length 2.43, width 1.60. Length of tibia/metatarsus: I, 1.73/1.40; II, 1.35/1.13; III, 0.98/0.95; IV, 1.70/1.37. Chelicerae small, vertical. Sternum length 1.47, width 0.78. Spines as in female, except: leg I, tibia p 1-d1-1-0, r 1-0-d1-0, d r1–0–1 bristles; metatarsus p d1. II, tibia p 1-d1-1-0, r 1-0-d1-0, d r1–0–1 bristles; metatarsus p d1. III, tibia, r 1-d1-0-1-0; metatarsus v 2-p1–2. Abdomen length 2.93, width 1.33, spiracle–epigastrium 1.50, spiracle–spinnerets 0.58. Color as in female. Palp (fig. 86E, F): tibia short, width/length 1.39, cymbium relatively large. Embolus thin, basal process small, triangular. Median apophysis very thin, short, closely associated with paramedian. Apex of paramedian apophysis short, sinuous, translucent. Secondary conductor not fused to anterior margin of tegulum; canal conspicuous, arising at base of paramedian apophysis; retrolateral portion with basal, conical projection. Anterior margin of tegulum compressed over base of secondary conductor.

    Natural History:

    This species lives in the bases of large grasses, mainly pampa grasses (“cortadera”, Cortaderia selloana), in periodically flooded areas. The elongate, flattened body with uniform color and the way the spider aligns the legs with the grass leaves make a strikingly cryptic effect.

    Distribution:

    Southeastern Brazil, Uruguay, and northeastern Argentina.

    Other Material Examined:

    BRASIL: São Paulo: Botucatu, Fazenda Nossa Senhora da Conceição, “parte aérea de Saccharum officinarum”, 4.II.1999, L.I. Rinaldi, B. Mendez, 1♂ (UNESP), 20.IV.1999, 1♀ (UNESP); Botucatu, Usina São Manoel, 2.III.1988, I. Rinaldi and L. Forti, 1♀ (UNESP), 29.IX.1986, 1♂ (UNESP). URUGUAY: Departamento Rocha: Arroyo Sarandí del Consejo, ruta 9 km 251, 18.V.1993, M. Ramírez and F. Pérez Miles, 1♂ (MACN-Ar). ARGENTINA: Santa Fe: El Toba, 4.IV.1968, M. Galiano, 1♀ (MACN-Ar). Entre Ríos: Rosario del Tala, XI.1988, M. Ramírez, 1♀ (MACN-Ar, photos MJR 62, 63, 269). Buenos Aires: Same data as types, 1♂ 4♀ 1 immature; Delta, estación experimental INTA, 3.II.1983, M. Carbajal and M. Ramírez, 1♂ (MACN-Ar); Delta, Paraná de Las Palmas, 17.VII.1964, M. Galiano, 2♀ (MACN-Ar); Dique Luján, 26.IX.1982, P. Goloboff and M. Ramírez, 1♂ (MACN-Ar); Mar del Tuyú, costa atlántica, 2.V.1981, M. Ramírez, 1♀ 1 immature (MACN-Ar); Reserva Otamendi, 10.VI.1997, M. Ramírez, L. Compagnucci, C. Grismado, F. Uehara, 1 immature (MACN-Ar); Punta Lara, Ensenada, 10.X.1984, M. Galiano, C. Scioscia and P. Goloboff, 1♂ (MACN-Ar), 11.X.1985, F. Miranda and M. Ramírez, 1♂ (MACN-Ar); Río Luján, estación FCGM, marsh with “espadaña”, 5.X.1993, M. Ramírez and A. Pérez, 2♂ (MACN-Ar); Zelaya, VII.1938, J.B. Daguerre, 2♀ (MACN-Ar).

    Sanogasta tenuis, new species Figures 76F, 88

    Types:

    Female holotype and male paratype from Argentina, Buenos Aires province, Río Luján, estación Ferrocarril General Mitre, grassland, ca. 34°17′S, 58°54′W, photos MJR 1242–1254, 5.X.1993, M. Ramírez and A. Pérez, deposited in MACN-Ar 9816.

    Etymology:

    The specific name refers to the slender body.

    Diagnosis:

    Resembles S. puma in having an elongate body without pattern (fig. 88A, B), but can be distinguished by having small, spherical spermathecae and a relatively small secondary conductor.

    Female (holotype):

    Total length 4.12. Carapace without thoracic groove, flattened, elongate, length 1.37, width 0.80, wider on legs II–III. AME larger than ALE, with ocular cones visible through cuticle, black. Length of tibia/metatarsus: I, 0.78/0.55; II, 0.53/0.38; III, 0.33/0.32; IV, 0.95/0.60. Palpal tarsus length 0.32. Chelicerae unmodified, with two teeth on retromargin. Sternum length 0.95, width 0.52. Spines (all tibiae with d r1–0–1 bristles): leg I, femur d 1–1–1, p 2ap, r d1ap; tibia v 2–2–2; metatarsus v 2bas (0-2-0-0). II, femur d 1–1–1, p d1ap, r 0-d1-d1; tibia v r1-r1–2; metatarsus = I. III, femur d 1–1–1, p and r 0-d1-d1; tibia v p1-p1–2, p v1-v1 + 0–1–0 bristle, r d1, d r1–0–1 bristles; metatarsus v 2–0–1, p 0–1, r 0-d1–1, d r1ap. IV, femur d 1–1–1, p and r d1ap; tibia v p1–2–2, p v1-v1–0 bristles + 0–1–0 bristle, r 0-d1–1 bristles, d r1–0–1 bristles; metatarsus v 2–0–1, p d1–1, r 0-d1-0-d1 bristles, d 0-p1-0-2 bristles. Prolateral spines on tibiae III and IV ventrally displaced. Abdomen length 2.70, width 1.27, spiracle–epigastrium 1.08, spiracle–spinnerets 0.83. Color: uniforma pale grayish brown. Epigyne (fig. 88C–E): anterior pouch small, close to epigastric furrow, with transverse elevation limiting depressed area anterior of pouch. Insertions of epigastric muscles superficial. Copulatory ducts arising contiguously. Ducts of accessory bulbs very short, ventral.

    Male (paratype):

    Total length 2.90. Carapace without thoracic groove, length 1.22, width 0.72. AME projecting above clypeus. Length of tibia/metatarsus: I, 0.85/0.63; II, 0.53/0.42; III, 0.33/0.32; IV, 0.82/0.57. Chelicerae narrow, vertical. Sternum length 0.82, width 0.45. Spines as in female, except: leg I, tibia p 1-d1-1-0, r 1-0-v1-0; metatarsus p d1. II, tibia p 1-d1-1-0; metatarsus p d1. III, tibia, r 0-d1-1-0; metatarsus p and r 0-d1-0-1. IV, metatarsus p and r d1–1, d r1–0-r1. Abdomen length 1.62, width 0.60, spiracle–epigastrium 0.70, spiracle–spinnerets 0.93. Color as in female. Palp (fig. 88F, G): tibia short, width/length 1.20, cymbium relatively large. Embolus thin, basal process small, triangular. Median apophysis very thin, closely associated with paramedian. Apex of paramedian apophysis short, sinuous. Secondary conductor not fused to anterior margin of tegulum; canal well defined, arising slightly distal to base of paramedian apophysis; retrolateral portion with rounded basal projection, external margin forming elevated, thin ridge. Anterior margin of tegulum pronounced, compressed over base of secondary conductor.

    Natural History:

    This species lives in the bases of grasses. The spider places its legs in a line with the body on grass leaves, and is extremely cryptic.

    Distribution:

    Collected in São Paulo, Buenos Aires, and Córdoba, probably widely distributed through grasslands in central and northeastern Argentina, southeastern Brazil, and Uruguay.

    Other Material Examined:

    BRASIL: São Paulo: Botucatu, Usina São Manoel, 11.II.1987, I. Rinaldi and L. Forti, 1♀ (UNESP). ARGENTINA: Buenos Aires: Same data as types, 5♀ 3 immatures, 30.X.1990, M. Ramírez, 1♀ 4 immatures (MACN-Ar), 14.IX.1991, M. Ramírez, 1 immature (MACN-Ar); no specific locality, J.B. Daguerre, 1♀ (MACN-Ar 31344). Córdoba: Pampa de Achala, 15 km W El Cóndor, 31.VII.1999, M. Ramírez and L. Lopardo, 1♀ (MACN-Ar).

    Sanogasta approximata (Tullgren), new combination Figures 8991

  • Gayenna approximata Tullgren, 1901: 233, 259 (two females syntypes from Tierra del Fuego, Tweedie, Sierra del Toro, 19.III.1899, E. Nordenskjöld, and Río Azopardo, 1.III.1896, O. Nordenskjöld, in NRS, examined).

  • Tomopisthes kraepelini Simon, 1902: 31 (female holotype from Chile, Punta Arenas, in MHNP 20723, examined). new synonymy.

  • Gayenna kraepelini: Merian, 1913: 13.

  • Synonymy:

    The types of the species synonymized were examined, together with specimens from the same area; no relevant differences were found.

    Diagnosis:

    Distinguished from other Sanogasta by having two pairs of ventral spines on metatarsi II, a conspicuous embolar process, and triangular sclerotized areas at the epigastric muscle insertions, flanking the epigyne.

    Female (lectotype, measurements of specimen from Los Glaciares Natl. Park):

    Total length 11.50. Carapace length 5.45, width 4.00, wider on legs II–III. Length of tibia/metatarsus: I, 3.72/3.72; II, 3.59/3.72; III, 3.00/4.00; IV, 3.86/5.72. Palpal tarsus length 1.90. Chelicerae unmodified, with two teeth on retromargin. Sternum length 3.13, width 2.17. Spines: leg I, femur d 1–1–1, p 0-d1–2, r 0-d1-d1; tibia v 2–2–2 or 2–2-p1; metatarsus v 2–2–0. II, femur = I; tibia v 2–2-p1 or 2–2–2 (the r1ap very small), p 0–1; metatarsus v 2–2–0 (the x-r1-x more apical). III, femur d 1–1–1, p and r 0-d1-d1; patella r d1; tibia v 2–2–2, p and r v1-d1-1-0, d r1-0-1-0; metatarsus v 2–2–2, p and r d1-1-0-1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia and metatarsus = III. Abdomen length 6.38, width 3.19, spiracle–epigastrium 2.70, spiracle–spinnerets 1.60. Color: grayish, with darker pattern (fig. 89), sternum grayish, slightly darker at margins. Venter pale. Epigyne (figs. 90, 91D, E): anterior pouch close to epigastric furrow. Insertions of epigastric muscles very deep, associated with triangular sclerotized areas. Median field with pair of shallow depressions on epigastric furrow. Copulatory ducts short, thick. Ducts of accessory bulbs very short, oriented forward.

    Male (Los Glaciares Natl. Park):

    Total length 10.00. Carapace length 4.80, width 3.72. Length of tibia/metatarsus: I, 4.39/4.79; II, 4.12/4.39; III, 3.23/4.26; IV, 4.12/6.25. Chelicerae slightly smaller than those of female. Sternum length 2.67, width 1.93. Spines as in female, except: leg I, tibia v 2–2–2, p and r 1-d1-1-0; metatarsus v 2–2–0 (both r advanced), p and r 1. II, femur = I; tibia = I; metatarsus p d1–1–0, r 1. IV, femur = III. Abdomen length 5.30, width 2.50, spiracle–epigastrium 2.10, spiracle–spinnerets 1.30. Color as in female. Sternum and coxae clothed with short hairs, some of them thicker. Palp (fig. 91A–C): tibia short, width/length 0.70, cymbium relatively large. Tegular notch short (fig. 91A). Embolus short, not associated with canal on conductor, basal process conspicuous, heavily sclerotized. Median apophysis short, hook-shaped. Apex of paramedian apophysis short. Secondary conductor partially fused to anterior margin of tegulum, wide, thin; canal vestigial, ending in low peak; prolateral portion with flattened, rounded projection pointing distally; retrolateral portion concave, with elevated ridge at external margin, small basal internal projection. Anterior margin of tegulum compressed over base of secondary conductor.

    Variability:

    The median ventral spines on metatarsus II (x-2-x) arise gradually in immatures, being absent in younger instars. Most females have metatarsus I, v 2bas, some v 2-p1–0, occasionally v 2–2–0 (as in lectotype). Some penultimate males also have v 2–2–0 on metatarsus I.

    Natural History:

    This species builds retreats under stones at shores of lakes or streams, often sharing a stone with conspecifics, or with Acanthoceto cinerea group species.

    Distribution:

    Forests in southern Argentina (from Neuquén province) and Chile (from Malleco province) to Tierra del Fuego. One isolated record from Santiago (CAS).

    Other Material Examined:

    ARGENTINA: Neuquén: San Martín de los Andes, 3–6.I.1964, no collector, 2♀ (MACN-Ar); Nahuel Huapi Natl. Park: Lago Nahuel Huapi, Península Quetrihué, II.1986, M. Ramírez, 2♀ (MACN-Ar), “arrayan” forest, beach of stones, 24.II.1996, M. Ramírez, 1♂ 2♀ (MACN-Ar); Lago Nahuel Huapi, lado este, I.1989, M. Ramírez and E. Maury, 2♀ 3 immatures (MACN-Ar, photos MJR 50–51). Río Negro: San Carlos de Bariloche, II.1954, M.E. Galiano, 8♀ 5 immatures (MACN-Ar 5412), III.1947, A. Giai, 1♂ 4 immatures (MACN-Ar 2225), 1♂ 3 immatures (MACN-Ar 2219), 1♀ (MACN-Ar 2221); El Bolsón, 71°35′S, 41°59′W, 2.III.1964, A. Kóvacs, 1♀ (AMNH); Los Repollos, 5.III.1962, A. Kovács, 1♀ (AMNH); Río Azul, V.1962, A. Kovács, 4♀ (AMNH). Chubut: Esquel, road to La Hoya, 42°54′S, 71°19′W, 16.XI.1988, V.D. Roth, 8♀ (CAS); Los Alerces Natl. Park: Lago Futalaufquen, I.1990, M.J. Ramírez, 9♀ (MACN-Ar); II.1986, M. Ramírez, 1♂ 4♀ 1 immature (MACN-Ar), 18.I.1977, E. Maury, B. Detricet, 1♀ (MACN-Ar); Lago Futalaufquen, Bahía Rosales, 7.II.1986, M. Ramírez, 7♀ 1♂ 1 immature (MACN-Ar); Lago Verde, II.1985, M. Ramírez, 1♂ 1♀ 6 immatures (MACN-Ar). Santa Cruz: Lago Frías, no date, E. Maury, 1♀ 2 immatures (MACN-Ar); Los Glaciares Natl. Park, II.1977, no collector, 1♂ 1♀ (MACN-Ar), 18.I.1980, P. Goloboff, 1♂ 3♀ (MACN-Ar); Valle Eléctrico, 10–15.II.1949, N.S. Gianollina, 1♂ (MACN-Ar 2691). CHILE: Región Metropolitana (Santiago): Santiago: Cañón del Maipo, 28.XII.1966, L. Campos, 2♂ (CAS). Región IX (Araucanía): Malleco: Tolhuaca, 15.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 2♀ (AMNH). Región X (Los Lagos): Osorno: E Entre Lagos, orilla del Lago Puyehue, 150 m, 24.XI.1981, N. Platnick and T. Schuh, 2♂ (AMNH); Puyehue Natl. Park: Los Derrumbes, 18.I.1989, M. Ramírez, 2♀ (MACN-Ar); Aguas Calientes, 13–17.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1♂ 1♀ (MACN-Ar). Chiloé: Isla de Chiloé: no date, Skottsberg, 1♀ (NRS); Lago Huillinco, 9.II.1981, T. Cekalovic, 1♀ 28 immatures (AMNH). Región XI (Ibáñez del Campo): Aisén: Balmaceda (“Río Negro”, lapsus?), 27.I.1957, M. Cadoceo, 12♀ 2♂ 1 immature (MACN-Ar), 28.III.1957, M. Cadoceo, 4♀ 28 immatures (MACN-Ar); Murta, Lago General Carrera, 29.30.I.1990, L. Peña, 1♀ (AMNH); Puerto Aisén, II.1957, M. Cadoceo, 14♀ 6♂ 1 immature (MACN-Ar). Región XII (Magallanes y Antártica): Magallanes: Magallanes, III.1959, no collector, 2♀ (MACN-Ar), no date, T. Cekalovic, 6♀ (UC); 30 km Natales, road to Magallanes, 21.III.1948, M. Birabén, 2♀ (MLP); Península Brunswick, Tres Brazos, 28.II.1971, T. Cekalovic, 9♀ 2 immatures (AMNH); Punta Arenas, 17.IX.1963, 3♀, 18.IX.1963, 1♀, T. Cekalovic (AMNH), Río Rubens, ca. 52°S, 30.XI–3.XII.1962, P. Darlington, 1♀ (MCZ).

    Sanogasta pehuenche, new species Figure 92

    Types:

    Male holotype and female paratype from Argentina, Neuquén province, Lanín Natl. Park, Lago Curruhé Chico, ca. 39°54′S, 71°21′W, 15–16.I.1983, E. Maury, deposited in MACN-Ar 9817.

    Etymology:

    The specific name refers to the pehuenches, who live in the land of pehuenes (Araucaria araucana), where this species is found.

    Diagnosis:

    Resembles S. approximata in having short, thick copulatory ducts, but can be distinguished by having the epigynal anterior pouch with longer borders, no sclerotized areas flanking the epigyne, and a paramedian apophysis forming a short hook.

    Female (paratype):

    Total length 7.60. Carapace length 3.72, width 2.57, wider on legs II–III. Length of tibia/metatarsus: I, 2.10/1.87 (left smaller, regenerated); II, 2.00/1.90; III, 1.63/1.83; IV, 2.30/2.70. Palpal tarsus length 1.00. Chelicerae unmodified, with two teeth on retromargin. Sternum length 1.87, width 1.40. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2-p1; metatarsus v 2bas. II, femur d 1–1–1, p 0-d1–2, r d1; tibia v r1–2–2 or 0–2–2, p 0–1; metatarsus = I. III, femur d 1–1–1, p and r 0-d1-d1; patella r 1; tibia v p1–2–2, p and r d1–1, d r1bas; metatarsus v 2–0–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r 1; tibia v p1–2–2, p and r 1-d1-0-1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. Abdomen length 4.39, width 2.23, spiracle–epigastrium 1.70, spiracle–spinnerets 0.63. Color: carapace grayish, with dark dorsal pattern. Legs pale grayish with dark spots. Endites brown, labium dark brown. Sternum with dark brown spots at margins, center pale. Abdomen with dense, dark dorsal pattern, venter pale, with median band of small spots, a few dots at sides. Epigyne (fig. 92C, D): anterior pouch close to epigastric furrow. Insertions of epigastric muscles superficial. Copulatory ducts short, thick. Ducts of accessory bulbs very short, oriented forward.

    Male (holotype):

    Total length 6.38. Carapace length 2.90, width 2.23. Length of tibia/metatarsus: I, 2.53/2.30; II, 2.67/2.23; III, 1.73/1.93; IV, 2.10/2.37. Chelicerae slightly narrower than those of female. Sternum length 3.07, width 2.33. Spines as in female, except: leg I, tibia v 2–2-r1 or 2–2–0. II, tibia p 0; metatarsus p d1–0. III, metatarsus v 2–2–2. Abdomen length 3.72, width 2.00, spiracle–epigastrium 1.60, spiracle–spinnerets 0.70. Color as in female, but dorsum of abdomen yellow with dark brownish pattern. Palp (fig. 92A, B): tibia long, width/length 0.55. Tegular notch short. Embolus short, not associated with canal on conductor, with basal process small, superficial. Apex of paramedian apophysis short, hook-shaped. Secondary conductor small, not fused to anterior margin of tegulum, with canal reduced; retrolateral portion concave, posteriorly elevated. Anterior margin of tegulum compressed over base of secondary conductor.

    Natural History:

    Unknown.

    Distribution:

    Known only from Neuquén and Concepción, probably with wider distribution.

    Other Material Examined:

    ARGENTINA: Neuquén: San Martín de los Andes, Cerro Chapelco, 1700 m, II.1961, M. Galiano, 1♀ (MACN-Ar); Lanín Natl. Park, same data as types, 2♀, 1♀ penultimate, 2 immatures (MACN-Ar); Lago Moquehue, 10.I.1985, E. Maury, 2♀ 1 immature (MACN-Ar); Lago Quillén, II.1968, E. Maury, Müller 2♀ (MACN-Ar). CHILE: Región VIII (Biobío): Concepción: Hualqui, 18.XII.1988, R. Vergara, 1♀ (AMNH).

    Philisca Simon Table 23

  • Philisca Simon, 1884: 129 (type species by monotypy Philisca hahni Simon, 1884), 1887: E21, 1897a: 77–78, 82, 84, 86, 1903a: 1031. Transferred from the Miturgidae by Ramírez, 1995a: 381.

  • Liparotoma Simon, 1884: 130, 137 (type species Liparotoma hyadesi Simon, 1884, subsequently designated by Simon, 1897a: 100). Ramírez, 1993: 196. Revised by Ramírez, 1993. new synonymy.

  • Synonymy:

    Some years after the proposal of Cluilius, Simon (1904) considered it a section of Philisca (1904: 98). Unfortunately, Simon designated two type species for the genus Cluilius: “Clubiona elegans Nicolet” and Clubiona chilensis Nicolet. Clubiona elegans is a nomen nudum that Simon made referring to some specimens labeled by Nicolet as “C. elegans” (Simon, 1903a: 1031). The type of Clubiona chilensis Nicolet is an Amaurobioides. I have not found any specimen labeled “C. elegans” or “Cluilius chilensis” in MHNP. Simon had not seen specimens of Amaurobioides (“a Clubiona differt, sec. Cambridge …” [Simon, 1897a: 89]; I have not found any Amaurobioides in Simon's collection, except the type of C. chilensis). However, it seems evident that Simon would have been able to distinguish between Amaurobioides and Philisca, because the type of Clubiona chilensis Nicolet has a label (probably transcribed by Berland) “Axyracrus chilensis Nicolet, (sub Clubiona), Simon det.”. If this label is correct, at some point Simon thought that C. chilensis was not a Philisca, but an Axyracrus, a genus very similar to Amaurobioides.

    In tune with the tradition, Mello-Leitão (1951) described a homonym Cluilius chilensis, which belongs to the genus Philisca, and I will retain the name Philisca chilensis until a revision of the genus is completed.

    Liparotoma hyadesi, the type species of Liparotoma, is here considered a member of Philisca. The names Philisca and Liparotoma were published in the same paper (Simon, 1884). I decided to use the name Philisca, where most species of this group were described. Liparotoma comprises four species of a derivative group, which may constitute a subgenus in the future.

    Note:

    Philisca puconensis is provisionally included in Philisca by the thin branches of the median apophysis. In slightly suboptimal trees it jumps to the base of Tomopisthes.

    Diagnosis:

    Most species are distinguished from other Gayennini by having thin branches on the median apophysis. Similar branches are also found in Tasata centralis, but that species is easily distinguished by having five retrolateral cheliceral denticles, instead of the three ones in Philisca. Philisca ornata lacks those branches, but shares with other Philisca the modified male chelicerae. P. tripunctata also lacks branches, but shares with several Philisca a reduced spination of first and second tibiae and metatarsi.

    Description:

    Carapace wide in front, chelicerae strong; males with chelicerae similar to or larger than those of females; three teeth on promargin, one to three on retromargin. Male palpal tibia relatively long, usually two or more times longer than wide. Cymbium relatively small, narrow. Embolus with tip relatively thick, not associated with secondary conductor. Median apophysis generally with one to several thin basal or medial branches. Paramedian apophysis simple, thick, elongate. Primary conductor present, simple. Secondary conductor with several projections, well separated from anterior margin of tegulum by membranous band wider on prolateral side; canal vestigial or absent. Epigyne with anterior pouch deep or widened, several species with median field heart-shaped.

    Distribution:

    Forests in southern Chile and Argentina.

    Composition:

    In addition to the species detailed below: Philisca accentifera Simon, 1904 (female and immature syntypes in MHNP 22413, examined), Philisca ingens Berland, 1924 (female holotype in NRS 817, and female paratype in MHNP, examined), Cluilius chilensis Mello-Leitão, 1951 (male and female syntypes, in MNRJ, examined), Clubiona gayi Nicolet, 1849 (several males and females syntypes, in MHNP 4226, examined, new combination), and few bizarre undescribed species on the Juan Fernández Islands.

    Philisca hahni Simon Figures 93A, B, 94, 95, 96A

  • Philisca hahni Simon, 1884: 129 (female holotype from Chile, Cabo de Hornos, in MHNP 6679, examined), 1887: E21, 1897a: 78, 82, 86, 1902: 29.

  • Philisca navarinensis Tullgren, 1901: 228, 259 (female holotype from Chile, Magallanes, Puerto Toro, 8.II.1896, in NRS, examined). new synonymy.

  • Synonymy:

    The holotypes of the species synonymized were examined; no relevant differences were found.

    Note:

    The type of Philisca navarinensis is labeled “Philisca toronensis”; the corresponding file in NRS was corrected as “navarinensis (toronensis)”.

    Diagnosis:

    Distinguished from other Philisca by having the epigynal median field rectangular, slightly elevated, with a deep anterior pouch, and the male paramedian apophysis elongate, thick, with rounded tip.

    Female (holotype, measurements of specimen from Pucón):

    Total length 9.05. Carapace (fig. 94B, C) length 4.00, width 2.77, wider on legs II–III. Length of tibia/metatarsus: I, 2.50/2.07; II, 2.30/2.00; III, 1.70/1.87; IV, 2.33/2.63. Palpal tarsus length 1.12. Chelicerae unmodified, with three teeth on retromargin, basal one larger (fig. 96A). Sternum length 1.93, width 1.53. Spines: leg I, femur d 1–1–1, p 2ap; tibia v p1–2–2; metatarsus v 2bas. II, femur = I; tibia v 0–2–2; metatarsus = I. III, femur d 1–1–1, p 0-d1-(1-d1), r d1ap; patella r d1; tibia v 0–2–2, p and r d1–1; metatarsus v 2-p1-p1 and group of apical hairs, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia v p1–2–2, p d1–1, r 1-d1-0-1-0; metatarsus = III, but v 2–2-p1. Abdomen length 5.30, width 3.33, spiracle–epigastrium 3.03, spiracle–spinnerets 0.50. Color: carapace and legs brown. Sternum dark brown with pale center, endites and labium dark. Abdomen yellow, cardiac area violet, dorsum with brown spots, venter with some diffuse dark spots. Epigyne (fig. 94D, E): anterior pouch with deep cavity, median field rectangular, slightly elevated. Insertions of epigastric muscles superficial. Ducts of accessory bulbs short, converging.

    Male (Pucón):

    Total length 7.70. Carapace length 3.07, width 2.27. Length of tibia/metatarsus: I, 2.67/2.37; II, 2.40/2.20; III, 1.67/1.77; IV, 2.20/2.47. Chelicerae very long, fang long, thick; retromarginal teeth small, apical tooth separated from other two. Endites unmodified. Sternum length 1.55, width 1.23. Spines as in female, except: leg II, femur p d1ap. III, femur p 0-d1-(1-d1) or p 0-d1-d1; tibia v 0–2–2, 0-p1–2 or p1–2–2; metatarsus v 2-p1-p1, 2-r1-p1 or 2–0-p1. IV, metatarsus v 2-p1-p1 or 2–2-p1. Abdomen length 2.75, width 1.70, spiracle–epigastrium 1.57, spiracle–spinnerets 0.28. Color (fig. 94A): as in female, but abdomen with more heavily contrasting pattern, dorsal pattern and epigastrium violet (some specimens with a ventral band). Palp (figs. 93A, B, 95): tibia long, width/length 0.37, cymbium relatively small. Embolus very short, base globose, with longitudinal projecting ridge; basal process flattened, rounded. Median apophysis branched at base (fig. 93A, B). Paramedian apophysis wide, tip rounded. Primary conductor triangular, flattened, curved (fig. 93B). Secondary conductor small, canal vestigial, limited to tip, ending in small peak, on rounded projection directed backward; area basal to canal membranous; retrolateral portion wide, thin. Anterior dorsal margin of tegulum projecting as rounded lobe.

    Variability:

    Spines: III, tibia v p1-p1–2 or 0-p1–2. IV, tibia v p1–2–2.

    Natural History:

    Unknown. According to labels of specimens from Pucón collected by S.A. Marshall, this species might live on stony beaches. Many specimens have regenerated legs.

    Distribution:

    Chile, from Cautín province to Cabo de Hornos.

    Other Material Examined:

    CHILE: Región IX (Araucanía): Cautín: Pucón, dung traps nr. lake, 9–16.XI.1989, 3♂, pan traps in drift, nr. lake, 5–8.XI.1989, 4♂, pan traps in lakeside debris, 8–13.XI.1989, 9♂ 1♀ 1♀ penultimate, lakeshore FIT, 15.XI–2.XII.1989, 5♂ 5♀, FIT nr. lake, 8–13.XI.1989, 6♂ 1♀, pan in lake wrack, 15.XI–2.XII.1989, 21♂ 3♀ 1♂, S.A. Marshall (AMNH). Región X (Los Lagos): Chiloé: Isla de Chiloé: Cucao, 12.XII.1985, E. Maury, 1♀ (MACN-Ar). Región XII (Magallanes y Antártica): Magallanes: Cabo Negro, 29.I.1976, T. Cekalovic, 1♀ (AMNH); Isla Navarino, Puerto Toro, 19.XII.1992, Michelsen, 2♀ (ZMH 178).

    Philisca ornata Berland Figures 96B, 97, 98

  • Philisca ornata Berland, 1924: 435 (male and female syntypes, from the Juan Fernández Islands, Mas a Tierra, 28.XII.1916, K. Bäckström coll., in NRS, S.P.E. 188, and male paratype from Mas a Tierra, 1917, Bäckström [labeled “cotype”], in MHNP, examined).

  • Diagnosis:

    Distinguished from other Philisca by having a large yellow body and a very long male palpal tibia; the epigynum is very similar to that of other Chilean species.

    Female (syntype, measurements of specimen from nr. Plazoleta):

    Total length 7.18. Carapace wide in front (fig. 97A, C), length 3.20, width 2.37, wider on legs II–III. Length of tibia/metatarsus: I, 2.33/2.13; II, 2.27/2.10; III, 1.57/1.77; IV, 2.20/2.53. Palpal tarsus length 0.92. Chelicerae strong, with three similar teeth on retromargin. Sternum length 1.55, width 1.18. Scopulae on legs I and II not reaching tibiae. Spines (shorter on legs I and II): leg I, femur d 1–1–1, p (1-d1)ap; tibia v p1–2–2 or p1–2-p1; metatarsus v (p1-r1)bas. II, femur d 1–1–1, p d1ap; tibia v 0–2–2, p 0–1 or 0; metatarsus v (p1-r1)bas, p 1–0. III, femur d 1–1–1, p and r d1ap; patella r d1; tibia v 0-p1–2, p and r 1–1; metatarsus v 2–2–2, p d1–1–1, r d1–0–1, d 0-p1–2. IV, femur, patella and tibia = III; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. Abdomen length 4.12, width 2.40, spiracle–epigastrium 1.90, spiracle–spinnerets 0.40. Color: yellow, cephalic area and chelicerae reddish brown. Sternum pale brown, center paler. Dorsum of abdomen with anterior lateral gray areas and three pairs of small gray spots at posterior half, closing posteriorly; some specimens with white guanine reticulum at sides. Epigyne (fig. 98E, F) small, weakly sclerotized, translucent. Margins of lateral lobes arched, median field heart-shaped, anterior pouch in deep notch. Ducts of accessory bulbs long, with conspicuous gland ducts.

    Male (syntype, measurements of specimen from Valle Anson):

    Total length 8.65. Carapace very wide in front (fig. 97B, D), length 4.00, width 3.13. Legs very thin, especially the metatarsi. Length of tibia/metatarsus: I, 3.72/3.80; II, 3.60/3.60; III, 2.13/2.83; IV, 3.10/3.33. Chelicerae (figs. 96B, 97D) very strong, with internal superior margins projecting, three teeth on promargin, on elevation, and three on retromargin, basals contiguous. Endites with two protuberances at each external angle, most external larger. Sternum length 2.00, width 1.57. Spines as in female, except: leg I, tibia v p1–2–2. II, femur p (1-d1)ap or d1ap; tibia v 0–2–2 or r1–2–2, p 0; metatarsus v (p1-r1)-p1–0. Abdomen length 4.50, width 2.12, spiracle–epigastrium 2.37, spiracle–spinnerets 0.73. Color: yellow, cephalic area, mouthparts, and chelicerae reddish brown. Sternum and coxa I orange. Legs yellow with gray patches, femora with one ventral apical patch, covering part of articulation membrane; patellae with one spot at each side; tibiae with several basal and pair of apical spots. Dorsum of abdomen with brownish violet pattern, venter with diffuse violet area, from epigastrium to tracheal spiracle. Palp (fig. 98A–D) with all segments long, thin, cymbium relatively small. Copulatory bulb small, central into cymbium, well separated from tibia. Tegular notch short. Embolus long, basal process flattened, rounded. Median apophysis long, quite straight, unbranched. Paramedian apophysis elongate, acute. Primary conductor wide, triangular, flattened. Secondary conductor with canal vestigial, limited to hook-shaped peak, on rounded projection directed backward; area basal to canal membranous; retrolateral portion wide, thin, elevated.

    Variability:

    Female spines: I, tibia v 0–2-p1. Male spines: III, tibia v 0–2–2. IV, tibia v p1–2–2.

    Natural History:

    Unknown.

    Distribution:

    Known only from the Juan Fernández Islands, Mas a Tierra (now Isla Robinson Crusoe).

    Other Material Examined:

    CHILE: Valparaíso: Archipiélago Juan Fernández, Mas a Tierra (Isla Robinson Crusoe): Same data as syntypes, 2 immatures (NRS); Portezuelo trail, 7.IV.1962, 1♀ 2 immatures, Quebrada Damajuana, 5.IV.1962, 1♂, Valle Anson, Plazoleta del Yunque, 200–250 m, camote side, 1–28.IV.1962, 6♂ 18♀ 10 immatures, Valle Villagra, Portezuelo trail, 400–450 m, 19.IV.1962, 1♂, B. Malkin (AMNH); Galpón, Valle Villagra, 23–24.IV.1962, 1♂, wet areas near Plazoleta, pans, 24–28.I.1992, S. Marshall, 1♀ (AMNH); Plazoleta, Malaise trap, 24–29.I.1992, S. Marshall, 1♂ (AMNH).

    Philisca huapi, new species Figures 61H, 93C–E, 99

    Types:

    Male holotype from Argentina, Neuquén province, Nahuel Huapi Natl. Park, Puerto Blest, Laguna Los Cántaros, ca. 41°00′S, 71°50′W, 30.I.1985, M. Ramírez, 9818, and female paratype from Puerto Blest, trail to Lago Ortiz Basualdo, I.1990, M. Ramírez, deposited in MACN-Ar.

    Etymology:

    The specific name is a noun in apposition taken from the type locality.

    Diagnosis:

    Resembles P. ornata (and other species not included here) in having anteriorly modified male chelicerae, but can be distinguished by lacking the lateral projections on the male endites and by the long, straight ducts of the female accessory bulbs.

    Female (paratype):

    Total length 5.05. Carapace length 2.10, width 1.48, wider on legs II–III. Length of tibia/metatarsus: I, 1.17/1.00; II, 0.98/0.88; III, 0.80/0.90; IV, 1.18/1.33. Palpal tarsus length 0.58. Chelicerae unmodified, with three teeth on retromargin, basal one largest. Sternum length 1.17, width 0.88. Spines: leg I, femur d 1–1–1, p 2ap; tibia v 2–2–2; metatarsus v 2bas. II, femur d 1–1–1, p d1ap; tibia v r1–2–2, p 0–1; metatarsus v 2bas, p 1. III, femur d 1–1–1, p and r d1ap; patella r d1; tibia v 0-p1–2 or p1-p1–2, p and r d1–1–0, d r1bas; metatarsus v 2–0-p1 and apical group of hairs, p and r d1–1–1, d 0-p1–2. IV, femur = III; patella r d1; tibia v p1-p1–2, p and r d1–1, d r1bas; metatarsus v 2-p1-p1 and apical group of hairs, p and r d1–1–1, d 0-p1–2. Abdomen length 3.07, width 1.80, spiracle–epigastrium 1.70, spiracle–spinnerets 0.25. Color (fig. 99A): yellowish with grayish brown pattern. Legs with gray spots, patellae with spot on each side, tibiae with several basal spots and pair of apical spots. Margins of sternum and labium dark brown, endites brown. Abdomen yellow with white guanine reticulum, dorsum with park pattern, venter with median band dark brown from epigastrium to tracheal spiracle, two lateral lines of small spots. Anterior spinnerets different from those of males, slightly thickened at bases, without modified hairs. Epigyne weakly sclerotized, median field heart-shaped, ducts of accessory bulbs long, stright, slightly converging. Lateral lobes contiguous posteriorly (but slightly separate in some females, fig. 99D).

    Male (holotype):

    Total length 6.38. Carapace very wide in front, ocular area slightly protruding, length 2.30, width 1.57. Length of tibia/metatarsus: I, 2.17/1.97; II, 1.77/1.63; III, 1.15/1.30; IV, 1.63/1.87. Chelicerae very strong (fig. 99C), with internal superior margins slightly protruding in ridges. Three teeth on promargin, median one very thick, on elevation, other two small; three teeth on retromargin, basal one largest. Endites unmodified. Sternum length 1.13, width 0.95. Spines as in female, except: leg II, tibia v 2–2–2. III, femur p 0-d1-d1 or 0-d1–2; tibia v p1–2–2. IV, tibia v p1–2–2. Abdomen length 2.77, width 1.53, spiracle–epigastrium 1.30, spiracle–spinnerets 0.25. Base of anterior spinnerets globose, covered by thick, short hairs. Color (fig. 99B): similar to female. Palp (figs. 61H, 93C–E, 99E, F): tibia long, width/length 0.47. Embolus short, flattened, base globose, with longitudinal flat projecting ridge; basal process flattened, rounded. Median apophysis with series of thin branches (fig. 93C). Triangular sclerotized stripe runs from sperm duct to base of median apophysis. Paramedian apophysis wide, curved at tip. Primary conductor triangular, slightly flattened. Secondary conductor small, canal vestigial, limited to tip, ending in longitudinally ridged peak, on rounded projection directed backward; area basal to canal membranous; retrolateral portion wide, thin, elevated.

    Variability:

    The size of male chelicerae is variable. The lateral epigynal lobes may be separate or contiguous. Spines: metatarsi III, IV, v 2-p1-p1.

    Natural History:

    This species makes retreats on foliage. Most specimens were collected on bamboos (Chusquea spp.), and they resemble Gayenna americana, also common on the same bamboos. Notably, Platnickia elegans (Nicolet) (Zodariidae), which lives exclusively on bamboos (personal obs.), has a similar appearance too.

    Distribution:

    Humid southern forest in Chile, from Malleco to Palena provinces; in Argentina only collected in Puerto Blest and Termas de Epulaufquen, two humid Andean passes at the Chilean border.

    Other Material Examined:

    ARGENTINA: Neuquén: Lanín Natl. Park: Termas de Epulaufquen, 9.I.1986, M. Ramírez, 1♂ (MACN-Ar); Nahuel Huapi Natl. Park: same data as holotype, 2♂ 1♀ 1♀ penultimate (MACN-Ar); same data as paratype, 26♂ 8 immatures (MACN-Ar), 1♂ (MACN-Ar). CHILE: Región IX (Araucanía): Malleco: Monumento Natural Contulmo, 11.XII.1984–13.II.1985, FIT, 350 m, S. and J. Peck, 4♂ 1♀ (AMNH), 19–21.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 3♀ 1♂ (MACN-Ar), 5♀ (MHNS). Región X (Los Lagos): Osorno: Puyehue Natl. Park: Aguas Calientes, 600 m, 12–20.II.1979, L.E. Peña, 1♂ (AMNH), 13–17.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1♂ 1♀ (MACN-Ar), 2♀ 1♂ (MHNS), 500 m, 2–5.V.1988, L. Peña, 1♂ 1♀ (AMNH); 4.1 km W Anticura, 270 m, trap site 663, window trap, valdivian rainforest, 19–25.XII.1982, A. Newton and M. Thayer, 1♂ (AMNH); 7.7 km NE Termas de Puyehue, 200 m, site 664, window trap, 19–25.XII.1982, A. Newton and M. Tayer, 1♂ (AMNH). Chiloé: Guabún, N Ancud, 13–15.I.1980, L. Peña, 1♀ (AMNH). Palena: Río Ventisquero, Lago Yelcho, 5–9.XII.1985, L. Peña, 1♂ (AMNH).

    Philisca hyadesi (Simon), new combination Figures 93F, 101E, 102D, E

  • Liparotoma hyadesi Simon, 1884: 138. Ramírez, 1993: 201.

  • Description and Diagnosis:

    See Ramírez (1993). Additional data are provided below.

    Female:

    Palpal claw modified (fig. 101E). Spines: leg I, femur d 1–1–1, p 2ap; metatarsus v 2bas short. II = I or femur p d1ap. III, femur d 1–1–1, p and r d1ap; tibia v 2ap, p 1, r d1–1; metatarsus v 2–0-r2 and bunch of apical hairs, p and r 0–1–1, d 2ap. IV, femur d 1–1–1, r d1ap; tibia v 2ap or p1–0–2, r d1–1; metatarsus v 2–0-r2 or p1–0-r2 and bunch of apical hairs, p 1ap, r 0–1–1, d 2ap.

    Male:

    Palp (figs. 93F, 102D, E): tibia long, width/length 0.37, cymbium relatively small. Embolus short, base thick, anterior margin longitudinally flattened, slightly projecting as ridge; basal process flattened, rounded. Median apophysis long, thin, with small branches at base. Paramedian apophysis long, sinuous. Sperm duct with apical/retrolateral loop, approaching base of median apophysis. Primary conductor well developed, wide, slightly flattened. Secondary conductor complex, canal completely absent, area corresponding to canal base membranous; prolateral portion with triangular and flattened projection, and thick, rounded projection directed backward; retrolateral portion thin, projecting as acute peak; additional projection flat, triangular, between both portions.

    Distribution:

    Southern forests in Chile, from Cautín, and Argentina, from Neuquén, to Cabo de Hornos; two isolated records from Valparaíso (Quintay) and Petorca (Cuesta El Melón, Ramírez, 1993) might indicate an interesting relict, or a mislabeling (see also Oxysoma punctatum).

    New Records:

    ARGENTINA: Neuquén: Nahuel Huapi Natl. Park, Isla Victoria, IV.1945, Havrylenko, 1♂ (MLP); Península Quetrihué, 24.II.1986, M. Ramírez, 1♀ (MACN-Ar); Puerto Blest, 7–20.I.2000, L. Lopardo and A. Quaglino, 3♀ (MACN-Ar). CHILE: Valparaíso: 8 km SE Quintay, 33°12′S, 71°41′W, 150 m, 17.II.1967, E.I. Schlinger, 1♂ (CAS). Región Metropolitana (Santiago): Santiago: Bucalemi, San Antonio, 23–24.X.1994, L. Peña, 2♀ 1 immature (AMNH). Región VIII (Biobío): Ñuble: Las Trancas, 1–10.XII.1965, L. Peña, 2 immatures (MCZ). Concepción: Cerro Caracol, Concepción, 28.III.1994, T. Cekalovic, 1♀ (AMNH). Cautín: Monte Verde, Cavahue, 800 m, 30.I–2.II.1993, L. Peña, 1♂ (AMNH). Biobío: Los Morongos, E Los Niches, 600 m, 17–20.XI.1994, L. Peña, 1♀ (AMNH). Región IX (Araucanía): Malleco: Nahuelbuta Natl. Park, Pehuenco, 37°49′45″S, 73°0′30″W, 1100 m, 4–9.I.2001, forest with Nothofagus antarctica, Araucaria, and Chusquea, J. Miller, Alvarez, J. Coddington, G. Hormiga, 1♀ (USNM). Región X (Los Lagos): Valdivia: Neltume, II.1987, L. Peña, 2♀ (AMNH); Valdivia, XI–XII.1982, E. Krahmer, 2♀ 1 immature (MHNS 700). Osorno: Puyehue Natl. Park: 19 km E Termas de Puyehue, 40°40′S, 71°14′W, 450 m, fogging rotten branch, 30.XI.1994, R. Reschen and C. Carlton no. 188, 1 immature (AMNH). Llanquihue: Correntoso, XII.1969, L. Peña, 1♂ (MCZ); 8 mi W Puerto Varas, 18.I.1951, Ross and Michelbacher, 1♀ (CAS). Chiloé: Isla de Chiloé: 5 km SW Chonchi, 2.II.2001, T. Cekalovic, 1♀ (AMNH), Coihuin (Púlpito 1), 8.II.2001, T. Cekalovic, 1♀ (AMNH), Dalcahue, II.1967, 1♀ (MCZ); Isla Quinchao, Quetro, 19.II.1997, T. Cekalovic, 1♀ (AMNH). Región XI (Ibáñez del Campo): Aisén: Ventisquero San Rafael, III.1953, I. Bernasconi, 1♂ (MACN-Ar 3682).

    Philisca amoena (Simon), new combination Figures 100B, 101A, B, 102A, B

  • Liparotoma amoenum Simon, 1884: 138. Ramírez, 1993: 202.

  • Description and Diagnosis:

    See Ramírez (1993). Additional data are provided below.

    Female:

    Palpal claw modified (fig. 101A, B). Spines: leg I, femur d 1–1–1, p d1ap; metatarsus v 2bas short. II = I. III, femur d 1–1–1, p and r d1ap; tibia v 0-p1-p1 or 0-p1–2, p 0–1, r d1–1; metatarsus v 2–0-r2 and an apical group of hairs, p and r 0–1–1, d 2ap. IV, femur d 1–1–1, r d1ap; tibia v p1-p1–2 or 0-p1–2, r d1–1; metatarsus v 2–0-r2 and apical group of hairs, p d1–0–1, r 0–1–1, d 2ap.

    Male (fig. 100B):

    Palp (fig. 102A, B): tibia long, width/length 0.48, cymbium relatively small. Embolus short, base thick, basal process shallow. Median apophysis long, thin, with basal branch. Paramedian apophysis long, straight. Sperm duct with conspicuous loop approaching base of median apophysis (fig. 102A). Primary conductor small, thin, pointed. Secondary conductor complex, canal absent, area corresponding to canal base membranous; prolateral portion with triangular, flattened projection, covered at basal side by thick denticles pointing forward; this peak placed on projection as in P. hyadesi; retrolateral portion small, thin, translucent. Anterior margin of secondary conductor with two additional projections: one prolateral, rectangular, curved, with dentate border, other central, triangular.

    New Records:

    ARGENTINA: Neuquén: Puerto Blest, 7–20.I.2000, L. Lopardo and A. Quaglino, 1 immature (MACN-Ar). CHILE: Región IX (Araucanía): Cautín: Monte Verde, Cavahue, 31.I.1993, L. Peña, 2 immatures (AMNH). Región X (Los Lagos): Valdivia: Valdivia, XI–XII.1982, E. Krahmer, 1♂ (MHNS 700). Osorno: Puyehue Natl. Park: 700 m, 9.XII.1994, L. Peña, 1♀ (AMNH); XI.1992, L. Peña, 1♂ (AMNH); Aguas Calientes, 450 m, 11.XII.1981, Nielsen and Karsholt, 1 immature (ZMK); elev. 480 m, 40°44′S, 72°18′W, 21.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (MACN-Ar, photo MJR 1417–1418), 13–17.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 2♀ (MACN-Ar), 1♀ (MHNS). Llanquihue: Vicente Pérez Rosales Natl. Park, Cayutue, road to Calbutue, 19.II.1974, 2 immatures (UC). Chiloé: Chepu, 21.II.1992, M. Ramírez, P. Goloboff, N. Platnick, 2 immatures (MACN-Ar).

    Philisca tripunctata (Nicolet), new combination Figures 100A, 102C

  • Clubiona tripunctata Nicolet, 1849: 138 (two females syntypes from Chile, no specific locality, in MHNP 4219, not reexamined).

  • Liparotoma pardalis Mello-Leitão, 1943a: 405 (holotype immature from Chile, Santiago, J. Carbalho coll., in MNRJ, examined). Synonymized by Ramírez, 1993.

  • Liparotoma tripunctatum: Ramírez, 1993: 198.

  • Synonymy:

    In Ramírez (1993) the synonymy of L. pardalis was based on the description by Mello-Leitão (1943a); it is confirmed here after examination of the holotype.

    Description and Diagnosis:

    See Ramírez (1993). Additional data are provided below.

    Female (fig. 100A):

    Spines: leg I, femur d 1–1–1, p d1ap; metatarsus 0. II = I. III, femur d 1–1–1, p 0-d1-d1, r d1ap or 0-d1-d1; tibia v p1-p1–2 or 2–2–2, p 0–1 or d1–1, r d1–1; metatarsus v 2-p1–2 or 2–0–2, p d1–1–1 or 0–1–1, r d1–1–1, d 0-p1–2. IV, femur, d 1–1–1, r d1ap; tibia v p1-p1–2, r d1–1; metatarsus v 2-p1–2 or 2–2–2, p d1–1–1 or 0–1–1, r d1–1–1, 0–1–1 or d1–0–1, d 0-p1–2.

    Male:

    Palp (fig. 102C): tibia long, length/width 0.60. Embolus short, base short, thick, basal process flattened, rounded. Paramedian apophysis elongate, sinuous. Triangular sclerotized stripe runs from sperm duct to base of median apophysis. Primary conductor vestigial, only sclerotized band remains. Secondary conductor without canal, basal area corresponding to canal membranous; prolateral portion with flattened, slightly pointed peak; retrolateral portion elevated, small, triangular.

    New Records:

    ARGENTINA: Neuquén: San Martín de los Andes, 40°10′S, 71°21′W, 20–21.XI.1988, V. and B. Roth, 1♀ (CAS). Río Negro: San Carlos de Bariloche, 9: Colonia Suiza, 800 m, 27.IX.1981, Nielsen and Karsholt, 1 immature (ZMK), 19.IX.1981, 1♀, 2 immatures (ZMK); 22.XI.1978, Misión Científica Danesa, 1 immature (ZMK); Lago Puelo Natl. Park, nr. intendencia, 205 m, Malaise trap, 31.XII–6.I.1998, C. and M. Vardy, 1♂ (BMNH/MACN-Ar). Chubut: Lago Menéndez, I.1990, M. Ramírez, 1♀ (MACN-Ar). Tierra del Fuego: Isla de los Estados, Bahía Crosby, 18.X.1941, no collector, 2 immatures (MACN-Ar). CHILE: Región IV (Coquimbo): Choapa: Los Vilos, Cariloleu, 11.X.1994, L. Peña, 1♀ (AMNH); Ñagué, 10 km N Los Vilos, Rt. 5, km 236, elev. 40 m, 31°50′S, 71°31′W, 13.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH), Región V (Valparaíso): Petorca: Los Molles, Rt. 5, km 188, elev. 10 m, 9.XI.1993, 32°14′S, 71°30′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH). Quillota: Cuesta La Dormida (east side), 33° 04′S, 71°02′W, 750–1000 m, 20.IX.1966, E.I. Schlinger, 1♀ (CAS). Valparaíso: Quintero, pitfalls in relict forest, 2.X.1968, R. Calderón G., 1♀ (AMNH); 4.IV.19??, R. Calderón, 1♂ (MACN-Ar). Región Metropolitana (Santiago): Santiago: Quebrada El Arbol, Aculeo, X.1969, L. Peña, 1♂ (MCZ). Región VII (Maule): Talca: Alto de Vilches, 17–24.X.1964, L. Peña, 2♂ 2 immatures (MCZ); 3 km E Gil de Vilches, 7.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH, photos MJR 745–747); Las Placetas, San Clemente, 800 m, 19–20.XI.1994, L. Peña, 1♀ (AMNH). Linares: Fundo Malcho, Andes in Parral, 11–20.XI.1964, L. Peña, 2♂ 1 immature (MCZ). Región VIII (Biobío): Ñuble: Las Trancas, 1–10.XII.1965, L. Peña, 1♂ 1♀ penultimate (AMNH); Las Trancas, 1200 m, 24–27.XI.1994, L. Peña, 1♀ (AMNH). Concepción: Curinam, 14.XII.1996, T. Cekalovic, 1♀ (AMNH); Escuadrón, 18.XI.1996, 1♂ 2♀, 18.XI.1996, 1♀ T. Cekalovic (AMNH); Fundo El Manzano, 7.XI.1992, 1♀, 8.XI.1992, 1♀, 22.X.1996, 1♀, 18.XI.1996, 1♀, 23.IX.1996, 4♀, 7.XII.1996, 3♀, T. Cekalovic (AMNH); Mitrihue, 29.XII.1996, T. Cekalovic, 2♀ (AMNH); Periquillo, 22.XI.1992, T. Cekalovic, 1♀ (AMNH). Arauco: 10 km N Curanilahue, 21.XI.1992, T. Cekalovic, 1♀ (AMNH). Biobío: N Ralco/Trapa-Trapa, 21–22.XI.1994, P. Peña, 1♀ (AMNH); W Ralco, Santa Bárbara, 400 m, 22–23.XI.1994, L. Peña, 2♀ (AMNH). Región IX (Araucanía): Malleco: Monumento Natural Contulmo, 19–21.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1♀ (MHNS). Región IX (Araucanía): Cautín: 30 km NE Villarrica, 1–30.I.1965, L. Peña, 1♂ (MCZ); NE Villarrica, 16–31.XII.1964, L. Peña, 1♀ (MCZ); Flor del Lago Ranch, Villarrica, Polo Field, 39°12.300′S, 72°08.367′W, 282 m, canopy fogging GT Nothofagus obliqua roble, 13.XII.2001, Arias et al., 1♀ 1 immature (UCB). Región X (Los Lagos): Valdivia: Huachocopihue, 7.III.1965, H. Levi, 2♂ 1♀ 1♂ penultimate. 1♀ penultimate (MCZ); Isla Teja, 6.III.1965, H. Levi, 1♂ (MCZ). Osorno: Puyehue Natl. Park: 26.I.1969, L. Peña, 1♂ (MCZ). Llanquihue: Alerce Andino Natl. Park, elev. 100 m, 41°35′S, 72°41′S, 23.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1 immature (MACN-Ar). Chiloé: 5 km SW Chonchi, 19.II.1997, T. Cekalovic, 1♀ (AMNH). Región XII (Magallanes y Antártica): Ultima Esperanza: Cerro Castillo, Natales, 13.XII.1960, L. Peña, 1♀ (MCZ); Torres del Paine Natl. Park: near Refugio Chileno, 50°56′45″S, 72°55′0″W, 400–600 m, 8–9.XII.2000, J. Miller, I. Agnarsson, 1♀ (USNM). Magallanes: Cueva del Milodón, 28.I.1976, T. Cekalovic, 1♀ (UC); Gobernador Philippi, 29.I.1976, T. Cekalovic, 1♀ (AMNH); Isla Navarino, Puerto Williams, XII.1962–I.1963, P. Darlington, 1♂ penultimate (MCZ); Manantiales, 1956, J. Vellard, 1♀ (MACN-Ar); Otway, El Canelo, 18.III.1969, L. Peña, 2♀ (MCZ); Aserradero Río Bueno, 8.II.1959, 2 immatures, 10.II.1959, 1♀, 1 immature J. Vellard (MACN-Ar).

    Philisca doilu (Ramírez), new combination Figure 101C, D

  • Liparotoma doilu Ramírez, 1993: 203.

  • Description and Diagnosis:

    See Ramírez (1993). Additional data are provided below.

    Female:

    Palpal claw modified (fig. 101C, D). Spines: leg I, femur d 1–1–1, p 2ap; metatarsus v 2bas short. II = I. III, femur p and r d1ap; tibia v 2ap, p 1, r d1–1; metatarsus v 2–0–2, p and r 0-d1–1, d 2ap. IV, p d1ap; tibia v 2ap, r d1–1; metatarsus v 2–0–2, p 1ap, r 0–1–1, d 0-p1–2.

    New Records:

    ARGENTINA: Neuquén: Nahuel Huapi Natl. Park, Isla Victoria, IV.1945, Havrylenko, 1 immature (MLP).

    Philisca puconensis, new species Figure 103

    Note:

    Provisionally included in the genus, see Note under Philisca.

    Types:

    Male holotype from Chile, Región X, Cautín province, Pucón, ca. 39°16′S, 71°58′W, 15.XI–2.XII.1980, Malaise trap in peninsula, S.A. Marshall, deposited in AMNH.

    Etymology:

    The specific name refers to the type locality.

    Diagnosis:

    Resembles other Philisca species in having the branch of the median apophysis, but easily distinguished by having a bifid paramedian apophysis and a narrow epigynal median field.

    Female (Malalcahuello, not type):

    Total length 6.50. Carapace length 2.80, width 2.03, wider on legs II–III. Diameter AME one-third of ALE. Length of tibia/metatarsus: I, 1.65/1.33; II, 1.62/1.33; III, 1.27/1.25; IV, 1.73/1.83. Palpal tarsus length 0.75. Chelicerae unmodified, with two teeth on retromargin. Sternum length 1.50, width 1.08. Spines: leg I, femur d 1–1–1, p (1-d1)ap; tibia v 2–2–2; metatarsus v 2bas. II, femur = I; tibia v 2–2–2, p d1–1; metatarsus v 2bas, p 1–0. III, femur d 1–1–1, p and r 0-d1-d1; patella r 1; tibia v p1–2–2, p and r d1–1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p and r d1ap; patella r 1; tibia v p1–2–2 or 2–2–2, p and r d1–1 or 1-d1-0-1-0, d r1bas; metatarsus = III. Abdomen length 3.72, width 2.23, spiracle–epigastrium 1.77, spiracle–spinnerets 0.33. Color: carapace grayish, ocular area almost black, two lateral bands and margins dark. Legs grayish with many dark spots. Sternum with two lateral dark bands, made of spots in front of coxae. Abdomen yellow with dark spots, cardiac area dark, pair of dark spots behind median transverse line of dorsum, several diffuse chevrons on posterior third. Venter with a few dark spots, mostly anterior of tracheal spiracle. Epigyne (fig. 103D, E): lateral lobes close to each other, median field elongate. Opening of anterior pouch almost circular, lumen deep.

    Male (holotype, fig. 103A):

    Total length 5.05. Carapace relatively wider than that of female, length 2.67, width 2.03. Length of tibia/metatarsus: I, 2.73/2.33; II, 2.43/2.17; III, 1.73/1.73; IV, 2.13/2.30. Sternum length 1.33, width 1.05. Spines as in female, except: leg I, femur p 0-d1-(1-d1) or 0-d2-(1-d1), r 0-d1-d1 or d1ap; tibia p and r 1-d1–1; metatarsus p 1–0. II, femur p 1–1-(1-d1), r 0-d1-d1; tibia = I. III, tibia v 2–2–2, p and r 1-d1-0-1-0. IV, femur p and r 0-d1-d1 or 0–0-d1; tibia = III. Abdomen length 2.40, width 1.40, spiracle–epigastrium 1.13, spiracle–spinnerets 0.15. Color: similar to female, but abdomen with very dark dorsal stripe, irregular, wider and darker at center of posterior half, sides dark, venter with line of spots along median line. Palp (fig. 103B, C): tibia long, width/length 0.74. Cymbium with retrolateral margin curved at base. Basal process of embolus projecting as flattened ridge, separated by ample ventral membranous area. Rectangular sclerotized stripe runs from sperm duct to base of median apophysis. Relict of primary conductor well developed, concave, with rounded tip. Secondary conductor small, rugose; prolateral portion with rounded projection; canal vestigial, base membranous; retrolateral portion thin, rugose. Median apophysis with large, median branch (fig. 103C). Paramedian apophysis bifid, retrolateral tip bent dorsally, prolateral curved ventrally.

    Variability:

    Female spines: III, metatarsus v 2-p1–2.

    Natural History:

    Unknown.

    Distribution:

    Known only from a few localities in Cautín, Malleco, Valdivia, and Neuquén provinces.

    Other Material Examined:

    ARGENTINA: Neuquén: Pucará, 15.XII.1965, A. Giai, 1♀ 1 immature (MACN-Ar). CHILE: Región IX (Araucanía): Malleco: Malalcahuello, 9–15.XII.1985, L. Peña, 1♀ described above (AMNH). Región X (Los Lagos): Valdivia: Valdivia, XI–XII.1982, E. Krahmer, 1♀ (MHNS 700). Mistaken Locality: Prov. Santiago, Malleco, XI.1979, L.E. Peña, 1♂ (AMNH) (see Ramírez, 1995b: 83).

    Tomopisthes Simon Table 24

  • Heteromma Karsch, 1880: 380 (type species by monotypy Heteromma fueguiana Karsch, 1880). new synonymy.

  • Tomopisthes Simon, 1884: 130, 132 (type species Tomopisthes immanis Simon, 1884, subsequently designated by Simon, 1897a: 99), 1887: E24, 28, 1897a: 90–99, 1903a: 1032. Ramírez, 1995a: 88, 1997: 178.

  • Gayenna: Tullgren, 1901: 240, 259. Merian, 1913: 12.

  • Heterommides Strand, 1912b: 16 (new name for Heteromma Karsch, 1880, preoccupied by Menge, 1856).

  • Synonymy:

    The type species of Heteromma Karsch is here considered a junior synonym of Tomopisthes horrendus.

    Diagnosis:

    Resembles Araiya in having regularly disposed teeth on the apex of the male secondary conductor, but can be distinguished by having a grayish or brown coloration and a wide membranous stripe separating the secondary conductor from the anterior margin of the tegulum, wider on prolateral side.

    Description:

    Color dark, with grayish brown patches, quite cryptic on tree bark or lichens. Chelicerae with three teeth on promargin, two on retromargin. Male palpal tibia longer than wide. Embolus short, thick, not associated with secondary conductor. Paramedian apophysis complex, with two or more apical cusps. Primary conductor present. Secondary conductor well separated from anterior margin of tegulum by membranous stripe, wider on retrolateral side. Apex of prolateral portion of secondary conductor with thick, regularly disposed denticles pointing backward. Triangular sclerotized area extends from sperm duct to base of median apophysis. Epigyne with lateral lobes separate, median field totally occupied by distended anterior pouch, its margin almost reaching epigastric furrow. Copulatory ducts running parallel to sutures between lateral lobes and median field.

    Distribution:

    Forest in southern Chile and Argentina.

    Composition:

    Three species here included.

    Types Not Examined:

    Tomopisthes tullgreni Simon, 1905.

    Nomen Dubium:

    Tomopisthes aethiops Simon, 1903 (female type presumably in MHNP, not found; six immatures in MHNP 21769, probably Sanogasta, are not types, because Simon [1903b: 312] referred to the epigyne).

    Tomopisthes horrendus (Nicolet), new combination Figures 104A, 105A, B, 106, 107

  • Clubiona horrenda Nicolet, 1849: 421 (two females syntypes from Chile, Llanquihue, in MHNP 4235, examined).

  • Amaurobius horrenda: Simon, 1864: 139.

  • Heteromma fuegiana Karsch, 1880: 380 (female immature holotype, from Chile, Punta Arenas, Exp. Gazelle, in ZMB, examined). new synonymy.

  • Tomopisthes horrendus: Simon, 1887: E4, 1897a: 91, 1902: 30, 1904: 102.

  • Tomopisthes immanis Simon, 1884: 133 (two males and three females syntypes, from Chile, Cabo de Hornos, in MHNP 6669, examined), 1887: E28, 1897a: 91, 99, 1901: 21, 1902: 30, 1903d: 5, 1904: 102. Ramírez, 1995a: 368. new synonymy.

  • Philisca sica Strand, 1908: 5 (female holotype from “W. Afrika, Ashanti”, in SMF 4737, examined). new synonymy.

  • Heterommides fuegianus Strand, 1912a: 346.

  • Gayenna horrenda: Merian, 1913: 13.

  • Nonianus argentinus Mello-Leitão, 1940: 42 (female holotype from Argentina, Neuquén province, San Martín de los Andes, III.1938, M. Birabén coll., in MLP 14350, examined). new synonymy.

  • Synonymy:

    The types of the species synonymized were examined, together with numerous specimens from the same areas; no relevant differences were found. The type locality of Philisca sica is most probably a mislabeling.

    Diagnosis:

    This is the largest anyphaenid species, ranging from 12 to 22 mm in total length; can be distinguished from other amaurobioidines by having a distally widened paramedian apophysis, of very characteristic shape, and epigynal median field with parallel margins. Immatures are very similar to those of T. varius, but can be distinguished by having one prolateral and one retrolateral spine on metatarsus I.

    Female

    (Ushuaia): Total length 18.00. Carapace length 8.11, width 6.25, wider on legs II–III. Length of tibia/metatarsus: I, 6.12/3.72; II, 6.00/5.05; III, 4.66/4.60; IV, 5.60/6.00. Palpal tarsus length 2.50. Chelicerae unmodified, with two teeth on retromargin. Sternum length 4.39, width 3.10. Spines: leg I, femur d 1–1–1, p (1-d1)ap, r 0-d1-d1; tibia v 2–2–2, p and r 0–1; metatarsus v 2bas, p and r 1–0. II, femur d 1–1–1, p 0-d1-(1-d1), r 0-d1-d1; tibia and metatarsus = I. III, femur d 1–1–1, p 0-d1-(1-d1), r 0-d1-d1; patella r d1; tibia v 2–2–2, p and r d1–1; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella r d1; tibia and metatarsus = III. Abdomen length 10.00, width 6.00, spiracle–epigastrium 4.90, spiracle–spinnerets 1.33. Color (fig. 104A): carapace and chelicerae reddish brown, legs pale brown with brown spots, metatarsi and tarsi reddish brown. Sternum and mouthparts reddish brown. Abdomen brown with pattern of cream dots. Epigyne (fig. 107F, G): copulatory openings close to anterior end of furrow separating lateral lobes and median field. Ducts of accessory bulbs short, connected very close to copulatory openings.

    Male (Ushuaia):

    Total length 16.90. Carapace length 8.65, width 6.00. Length of tibia/metatarsus: I, 9.58/7.85; II, 9.44/7.58; III, 6.65/6.38; IV, 7.58/8.11. Chelicerae very long, strong (fig. 107A); teeth thick, those of retromargin widely spaced, basal tooth in front of apical promarginal tooth; fang thick, long. Sternum length 4.39, width 3.19. Spines as in female. Abdomen length 8.00, width 4.66, spiracle–epigastrium 4.00, spiracle–spinnerets 0.60. Color as in female. Palp (figs. 105A, B, 107B–E): tibia long, width/length 0.35. Cymbium relatively narrow, retrolateral margin with slight basal notch. Anterior dorsal margin of tegulum projecting as prolateral conical prong. Embolus with basal process lobate, separated by ample membranous ventral area. Median apophysis very long, pointing apically. Paramedian apophysis complex, with retrolateral cusp and fan-shaped tip. Primary conductor slightly wider than long, almost rectangular. Secondary conductor small (fig. 107E), rugose. Anterior dorsal margin of tegulum projecting, partially membranous (fig. 105A, arrow).

    Variability:

    Total size (12–22 mm) and carapace length (6.5–10 mm in females) are very variable. Relative size of male chelicerae is also variable. The abdominal pattern may be contrasting or dark uniform. Male Spines: tibiae III, IV, r 1-d1-1-0. Epigyne (fig. 107F, G) and paramedian apophysis shape (fig. 107B, D) are slightly variable.

    Developmental Remarks:

    Epigyne: The primordium of epigyne in penultimate females is smaller but very similar to the adult structure (fig. 106A). This similarity does not result from because of imprinting by the forming cuticle in the exuviae, because the sizes of adult and penultimate epigyna are very different. The primordium of copulatory opening is evident, leading directly to the primordium of accessory bulb, which bears some primary pores (fig. 106B). Behind the copulatory opening there are two deep furrows (the internal one might be a primordium of copulatory ducts) in an intermediate degree of invagination. Third instar (Neuquén, Quetrihué): This is the dispersing instar, the first with spines. Spines: leg I, femur d 1–1–1, p d1ap; patella d 1–0–1 bristles; tibia v 0–2–0 (plus v 2ap thin bristles), d r1–0–1 bristles; metatarsus v 2bas, p 1. II, femur = I; patella = I; tibia and metatarsus = I. III, femur d 1–1–1, p and r d1ap; patella r d1; tibia v 0-p1–0, p d1–1, r 0, d r1–0–1 bristles; metatarsus v 2–0–1, p 0-d1–1, r 0-d1–1 or 1ap, d 2ap. IV, femur = III; patella r d1; tibia v 0-p1–0, p and r d1–1, d r1–0–1 bristles; metatarsus v 2–0–1 or p1–0–1, p 1ap, r d1–0–1, d 2ap. Carapace, dorsal (from pedicel to eyes), 0-2-2-2-1 (last between AME). Tracheal system well developed, similar pattern as in adults.

    Natural History:

    This species lives under bark or in crevices in rotten logs.

    Distribution:

    Temperate forests in Argentina and Chile, from Huasco to Cabo de Hornos. The locality of the holotyope of Philisca sica (“W. Afrika, Ashanti”) is most probably a mislabeling.

    Other Material Examined:

    ARGENTINA: Neuquén: Estancia San Ramón, Rincón Chico, Río Limay, X–XII.1962, Havrylenko, 1♀ (MACN-Ar); Junín de los Andes, XI.1970, P. Carnotto, 1♀ (MACN-Ar); Lago Nompehuén, NW Aluminé, 12.I.1985, E. Maury and A. Toth, 1♀ penultimate (MACN-Ar); Lago Huechulafquen, Coloradas, 7.I.1985, M. Ramírez, 1♀ (MACN-Ar); Lago Moquehue, 10.I.1985, E. Maury, A. Toth, 1♀ (MACN-Ar); Lago Quillén, 14.I.1985, E. Maury, A. Toth, 1♂ 1♀ (MACN-Ar); Lago Lácar, X.1955, A. Giai, 1♀, 1♀ (MACN-Ar); I.1961, M.E. Galiano, 1♂ (MACN-Ar); Pucará, II.1958, J. Navas, 1♂, 1♀ (MACN-Ar); II.1963, S. Schajovskoy, 3♀ (MACN-Ar); IX.1969, 3♂ 1 immature, 1.II.1971, 1♂ 3♀, Schajovskoy (MACN-Ar); 4.II.1972, L. Herman, 1♀ penultimate (AMNH); X–XII.1972, S. Schak, 2♂ (MACN-Ar); VIII.1973, 1 immature, XII.1973, 2♂, S. Schajovskoy (MACN-Ar), 10.XI.1978, 1♀, 750 m, 1.XII.1978, 1♀, Misión Científica Danesa (ZMK); Lago Lolog, 4 km N San Martín de los Andes, FIT, Nothofagus forest, ca. 950 m, Gentili property, 23.XI–1.XII.1989, S.A. Marshall, 1♀ (AMNH); San Martín de los Andes, XI–XII.1985, Gentili, 3♂ (MACN-Ar); 40°10′S, 71°21′W, 20–21.XI.1988, V. and B. Roth, 2♀ (CAS); 40°10′S, 71°21′W, 13.XI.1990, L. Peña, 1♀ (AMNH); 640 m, 29.IX.1981, 1♀, 17–31.X.1981, 1♂, 26.XI.1981, 2♀, 2–19.XII.1981, 1 immature, Nielsen and Karsholt (ZMK); San Martín de los Andes, Cerro Chapelco, 1700 m, II.1961, M. Galiano, 1♀ (MACN-Ar 5290); 8 km N San Martín de los Andes, 1000 m, Malaise trap, 16–22.XI.1997, C. and M. Vardy, 1♂ (BMNH/MACN-Ar); Nahuel Huapi Natl. Park: Isla Victoria, XII.1959, I. de Orfila, 4♀ (MACN-Ar); Isla Victoria, marmosa, Havrylenko, 1♂ (MLP); Isla Victoria, Piedras Blancas, Lago Frías, 15.I.1940, P. Moreau Guonera, 1♀ 1 immature (MACN-Ar); Lago Nahuel Huapi, Península Quetrihué, 16–25.XII.1984, 1♂ (MACN-Ar); 23.I.1985, M. Ramírez, 1 immature (MACN-Ar); Lago Nahuel Huapi, 1.II.1904, J. Daguerre, 2♀ (MACN-Ar 34332); Brazo Huemul, I.1966, Martínez, 1♂ 1♀ (MACN-Ar); Puerto Blest, 7–20.I.2000, L. Lopardo and A. Quaglino, 1 immature (MACN-Ar). Río Negro: San Carlos de Bariloche, 1944, F. Monrós, 1♀ (MACN-Ar 1696); III.1947, A. Giai, 3♀ 1 immature (MACN-Ar), 1950, 1♂ (MACN-Ar 5507); II.1954, M.E. Galiano, 1♀ (MACN-Ar 5451), 1♀ (MACN-Ar), 16.I.1961, E. Maury, 1♀ (MACN-Ar); 21.IX.1962, A. Kovács, 1♂ (AMNH), 1969, N. Müller, 1♂ (MACN-Ar); Colonia Suiza, 11.XII.1978, Misión Científica Danesa, 1♀ (ZMK); 810 m, 9.XI.1978, Misión Científica Danesa, 1♂ (ZMK); Colonia Suiza, ruta 240 25 km from San Carlos de Bariloche, I.1982, M. Ramírez, 1♂, 2♀ (MACN-Ar); 800 m, 17.IX.1981, 3 immatures, 19.IX.1981, 2♀, 8 immatures, 27.IX.1981, 1♂ 1 immature, 5–7.I.1982, 1 immature, Nielsen and Karsholt (ZMK); El Bolsón, 25.II.1963, M. Birabén, 1♂ (MLP), XI.1957, A. Kóvacs, 1 immature (AMNH), under stones, 25.II.1961, 1♂, 8.VII.1961, 2 immatures, 10.VII.1961, 3♀ penultimates, 3♀ penultimates, 17.VIII.1961, 2♀ penultimates, 2 immatures, 25.IX.1962, 1♂, 15.IX.1966, 1♀, 4.XI.1966, 1♂, A. Kovács (AMNH); El Bolsón, Cerro Piltriquitrón, II.1986, M. Ramírez, 1♀ (MACN-Ar); Lago Fonck, 2.XII.1984, A. Macario, 1♂ (MACN-Ar); Lago Gutiérrez, 41°95′S, 71°24′W, 18.XI.1988, V. and B. Roth, 1♀ (CAS); Lago Mascardi, I.1996, M. Calderón, 1♂ (MACN-Ar); Nahuel Huapi, 1960, Río Azul, 5.XII.1962, 1♂, 19.I.1966, 1♀, A. Kovács (AMNH). Chubut: El Hoyo, 2.X.1962, 3 immatures, VII.1962, 1♀ penultimate, VIII.1962, 1♀ 9 immatures, 15.IX.1962, 6♀ 9 immatures, A. Kovács (AMNH); El Maitén, 2.II.1966, A. Kovács, 1♂ (AMNH); Epuyén, 1.IX.1965, 1♀, 17.X.1966, 2♂ 2♀, A. Kovács (AMNH); Poto Cahuel, 8.X.1966, A. Kovács, 2♂ 5♀ (AMNH); Lago Fontana, I.1944, A. Riggi, 2♂ (MACN-Ar); Los Cipreses, XI.1982, M. Ramírez, 2♀ (MACN-Ar); Lago Futalaufquen, 4.I.1962, Coscarón, 1♀ (MACN-Ar); Lago Verde, II.1985, M. Ramírez, 1♀ (MACN-Ar); área Lago Verde, Lago Menéndez y Río Arrayanes, II.1895, M. Ramírez, 1♀ (MACN-Ar); Río Turbio, 11.VII.1961, 1♀ 1♀ penultimate 1 immature, A. Kovács (AMNH). Santa Cruz: Lago Argentino, II.1963; Lago Argentino, Península Magallanes, 250 m, 11.I.1979, Misión Científica Danesa, 1♂ (ZMK); Margheritis, Rizzo, 2♀ (MACN-Ar); Lago Burmeister, 31.I.1971, J.M. Gallardo, 1♀ (MACN-Ar); Morro Chico, Río Turbio, 28.I.1976, M. Rumboll, 1♀ (MACN-Ar); Los Glaciares Natl. Park, II.1975, E. Fernández, 1♀, 1 immature (MACN-Ar), 18.I.1980, P. Goloboff, 1 immature (MACN-Ar); Los Glaciares Natl. Park, Península Magallanes, in front of Glaciar Moreno, II.1977, D. Pepe and M. Rumboll, 1♂, 1♀ (MACN-Ar); Ventisquero Moreno, 18–24.I.1971, J. Vellard, 6♂ 5♀ (MACN-Ar), 18.II.1971, J. Vellard, 1♂ 6♀ 10 immatures (MACN-Ar). Tierra del Fuego: no specific locality, Castellanos, Gómez, 1 immature (MACN-Ar 32059); no specific locality, R. Dalhene, 1♀ (MACN-Ar 5775); Bahía Aguirre, 4.II.1949, 1♂ (MACN-Ar 2817), 3 immatures, 16.II.1959, 1♂ 1♀ 1 immature (MACN-Ar 2816), S. Núñez; Bahía Buen Suceso, 16–31.I.1986, E. Maury, 1♂ 1♀ 1 immature (MACN-Ar), no date, J.B. Daguerre, 1♂ (MACN-Ar 36735); Bahía Tethys, under tree bark, beach, 19.XI.1969, 1♀, 22.XI.1969, 1♀, Gosztonyi (MACN-Ar); Canal de Beagle, I.1933, Castellanos and Gómez, 3 immatures (MACN-Ar 34340); Isla de los Estados, Bahía San Antonio, 6–13.II.1982, J.C. Chébez, 1♀ 1 immature (MACN-Ar); Isla de los Estados, Punta Roca, no date, J.A. Dagerre and A. Carcelles, 1 immature (MACN-Ar 35069); Isla de los Estados, Puerto Parry, Nothofagus betuloides forest, X.1981, J.C. Chébez, 5♀ (MACN-Ar); Estancia Herberton, 25.I.1979, Misión Científica Danesa, 1♂ (ZMK); Lago Fagnano, no. 25, Nothofagus antartica forest, 26.II.1959, 1♀ (MACN-Ar); no. 26, 27.II.1959, J. Vellard, 1♀ (MACN-Ar); II.1967, Williner, 1♂ (MACN-Ar); Kaiken, 100 m, 18–19.I.1979, Misión Científica Danesa, 1♂, 1 immature, 1♀ penultimate, (ZMK); Lago Roca, II.1967, Williner, 1♀ (MACN-Ar); 27.I.1971, J. Vellard, 1♂ 1♀ (MACN-Ar), 10.I.1972, E. Hernández, 3♂ (MACN-Ar); hanging by thread from edge of roof, 21.I.1998, C. and M. Vardy, 1♀ (BMNH/MACN-Ar); Lapataia, I.1948, J.M. Viana, 3♀ 4 immatures (MACN-Ar 2590); 20–23.II.1961, 1♂ (AMNH); II.1963, E. Maury, 4♀ 1 immature (MACN-Ar); 20 m, 27–28.I.1979, 1♀, 1.II.1979, 1♀, Misión Científica Danesa (ZMK); Puerto Aguirre, no date, no collector, 1♀ (MACN-Ar); Ushuaia, 1–14.XII.1932, Castellanos and Gomez, 2♀ 4 immatures (MACN-Ar); 8–26.II.1961, B. Malkin, 2♂ 1♀ (AMNH); I.1967, Williner, 1♂ 3♀ 1 immature, 1 immature (MACN-Ar); XII.1967, A. Bachmann, 1♂ 1♀ (MACN-Ar); from Ushuaia to Lapataia, 28.I.1960, A. Bachmann, 1♂ (MACN-Ar); Ushuaia, no date, A. del Pino, 6♀ 2 immatures (MACN-Ar 29952). CHILE: Región III (Atacama): Huasco: Freirina, IX.1963, Instituto de Biología, 1♂ 1♀ (UC). Región IV (Coquimbo): Elqui: La Serena, I–II.1961, R. Wagenknecht, 1♀ (AMNH). Limarí: Bosque Talinay, 8.I.1985, N. Platnick and O. Francke, 1♂ 3♀ (AMNH); Bosque Talinay, 35 km S road to Fray Jorge, Panam km 353, 560 m, relict Valdivian fog forest, 6.II.1986, N. Platnick, T. Schuh, 3♀ (AMNH); Fray Jorge Natl. Park, 11.XII.1950, Ross and Michelbacher, 1♀ 5 immatures, (CAS); 21.X.1966, E. Schlinger, M. Irwin, 1♀ penultimate (CAS); 27.IX.1970, R. Calderón G., 2♀ (UC); 27.IX.1970, L. González, 1♀ (UC); 580 m, relict Valdivian forest, 5.I.1985, N. Platnick and O. Francke, 1♀ (AMNH); 560 m, relict Valdivian fog forest, under rocks, 8.II.1986, N. Platnick and R. Schuh, 1 immature (AMNH); 560 m, 3.X.1992, N. Platnick, P. Goloboff, K. Catley, 2 immatures (AMNH); elev. 580 m, 10.XI.1993, 30°40′S, 71°41′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 2♀ (AMNH), 1♀ (MACN-Ar, photo MJR 1296–1298), Fray Jorge, Rancho, 10.XII.1950, Ross and Michelbacher, 3♀, 1♀, 2 immatures (CAS). Choapa: Los Vilos, 25.IX.1971, J. Solervicens, 1♀ (UC); south of Coquimbo province, VII.1960, L. Peña, 2♀ 2 immatures, 4♀ (IRSN IG 23077). Coquimbo: Corral de Julio, 5.VII.1975, H. Hernández, 1♀ (UC). Región V (Valparaíso): Petorca: E La Ligua, relict forest, 27.IX.1980, L. Peña, 1♂ 2♀ (AMNH); Talaquén, 32°33′S, 71°14′W, X.1982, L. Peña, 2♀ (AMNH); Zapallar, 27.XI.1950, Ross and Michelbacher, 1♂ 1♀ 1 immature (CAS). Quillota: Cuesta La Dormida, N Tiltil, 800–1300 m, 13–18.XI.1982, L. Peña, 3♀ (AMNH). Valparaíso: Colliguay, nr. La Retuca, 5.XI.1963, G.F. Edmunds, 1♂ (AMNH); Quintero, 11–12.V.1961, A. Archer, 1♀ (AMNH); pitfalls in relict forest, 31.I.1968, 1♂, R. Calderón G. (AMNH); Quintero, 21 Barber VIII, 2.X.1968, 1♀ (UC). San Felipe de Aconcagua: 10 km E of Zapudo, 28.XI.1950, 1♂ (CAS); El Convento, 18.IX.1966, 33°48′S, 71°43′W, L. Peña, 2♀ 2 immatures, 6♀ 2 immatures (CAS). Región Metropolitana (Santiago): Santiago: Quebrada El Arbol, Aculeo, X.1969, L. Peña, 6♀ (MCZ). Región VII (Maule): Talca: Alto de Vilches, 31.X.1969, Rozen, L. Peña, 2♀, 1♀ (AMNH); Gil de Vilches, 1200 m, I.1984, P. Goloboff and E. Maury, 1♂ (MACN-Ar); 7.I.1989, M. Ramírez, 1♂, 1♂ (MACN-Ar). Cauquenes: Los Ruiles Natl. Park, IX.1985, F. Silva, 1♀ (MHNS). Región VIII (Biobío): Ñuble: Chillán, 31.XII.1975, G. Moreno, 1♀ (AMNH), Fundo El Sauce, San Fabián de Alicó, 8–24.I.1986, L. Peña, 1♀ (AMNH); Las Trancas, 15.VI.1990, C. Carrasco, 1♀ (UC); Las Trancas, 15–21.II.1976, G. Moreno, 1♀ (AMNH); 20–25.II.1980, L. Peña, 1♀ (AMNH), 11–17.I.1983, L. Peña, 1♀ (AMNH); II.1987, L. Peña, 1♀ (AMNH), 1200 m, 24–27.XI.1994, L. Peña, 1♂ (AMNH); Las Trancas, E Recinto, “Shangri la”, 19–30.I.1983, L. Peña, 1♀ (AMNH); 60 km SE Chillán, Termas road, beech forest, FIT, 1300 m, 7.XII.1984–19.II.1985, S. and J. Peck, 1 immature (ANMH); 72 km SE Chillán, Trancas, nr. Termas, FIT, 1700 m, Nothofagus forest, 6.XII.1984–19.II.1985, S. and J. Peck, 1 immature parasitized by a mermithid nematode (AMNH); 77 km SE Chillán, Termas road, 1260 m, 16–25.XI.1993, pitfalls, 36°55′S, 71°27′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH). Concepción: Cerro Caracol, Concepción, elev. 200 m, 36°51′S, 73°02′W, 17.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH); Escuadrón, 16.IV.1988, T. Cekalovic, 1♂ (AMNH); Hualpén, 75 m, moist forest, 22.I.1985, N. Platnick and O. Francke, 4♀ (AMNH); Lomas Colorada, 27.XI.1975, T. Cekalovic, 1♀ (AMNH). Arauco: Río Ibáñez, 27–28.I.1990, L. Peña, 1♀ (AMNH). Biobío: Caledonia, E Mulchen, 600 m, 18–20.II.1990, L. Peña, 1♀ (AMNH); Pemehue (158), L. Peña, 1♂ (IRSN IG 19736); W Ralco, Santa Bárbara, 400 m, 22–23.XI.1994, L. Peña, 1♀ (AMNH). Región IX (Araucanía): Malleco: Angol, 31.XII.1950, Ross and Michelbacher, 1♂ (CAS); Angol, Cordillera Nahuelbuta, 14–24.II.1977, G. Moreno, 1♂ (AMNH); 18 km W Angol, Cordillera Nahuelbuta, 37°48′S, 72°43′W, 10.II.1967, E. Schlinger, 1♂ (CAS); 40 km W Curacautín, Nothofagus/Araucaria forest, 12.XII.1984–16.II.1985, S. and J. Peck, 1♂, 1♀ penultimate (AMNH); Malalcahuello, 22.IX.1968, M. Rancagliolo, 1 immature (UC); 14 km E Malalcahuello, 1570 m, site 649, window trap, Nothofagus pumilio-Araucaria forest, 13–31.XII.1982, A. Newton and M. Thayer, 1♀ (AMNH); Monumento Natural Contulmo, 19–21.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 2 immatures (MACN-Ar), Nahuelbuta Natl. Park, 1200 m, Nothofagus-Araucaria assoc., 9.XI.1966, M. Irwin and E. Schlinger, 1♀ 4 immatures (CAS), 1200–1500 m, 9.XII.1985, S. and J. Peck, FITS forest Nothofagus-Araucaria, 1♂ (AMNH), 1230 m, dry forest, 1.II.1986, N. Platnick and T. Schuh, 1♀ penultimate (AMNH); Pichinahuel, Cordillera Nahuelbuta, 1200 m, 19.I.1988, L. Peña, 1♂ (AMNH); Cordillera Nahuelbuta, 1300–1400 m, 6–12.I.1982, L. Peña, 1♀ (AMNH). Cautín: Chacamó, NW Nueva Imperial, W Temuco, 16–24.II.1981, L. Peña, 1♂ (AMNH); 12.3 km N Loncoche, 280 m, 2.II.1967, I. Schlinger, 1♀ (CAS); Conguillio Natl. Park, 4–5.II.1980, A.E. Quezada, 1♀ (UC); Volcán Villarrica, FIT nr. edge of old lava flow, 10.XI–3.XII.1989, S. Marshall, 1♂ (AMNH); Pucón at Lago Villarrica, 39°16′S, 71°58′W, 14.XII.1988, V. and B. Roth, 1♀ (CAS); 10 mi NE Pucón, 12.I.1951, Ross and Michelbacher, 1♀ (CAS); Temuco, V.1956, Chávez, 1♀ (MACN-Ar); 15 km NE Villarrica, Flor del Lago, 300 m, 2 FITS, Nothofagus forest, 14.XII.1984–10.II.1985, S. and J. Peck, 1♂ (AMNH); Flor del Lago Ranch, Villarrica, Polo Field, 39°12.300′S, 72°08.367′W, 282 m, canopy fogging GT Nothofagus obliqua roble, 13.XII.2001, Arias et al., 2 immatures (UCB); Volcán Llaima, 8.X.1966, J.P. Valdenegro, 1♀ (UC). Región X (Los Lagos): Valdivia: Cudioco, 40°15′S, 73°09′W, 40 m, 10–11.XI.1966, M. Irwin and E. Schlinger, 1♀ (CAS); Isla Teja, farmland, 6.III.1965, H. Levi, 1 immature (MCZ); Las Lajas, W La Unión, 9–13.I.1990, L. Peña, 2♂ 3♀ (AMNH); Peulla, 200 m, 25.III.1965, H. Levi, 1♀ (MCZ), 1 immature (MCZ, photographed); Valdivia (collection E. Simon) 1♂ 1♀ (MHNP), 1983, E. Krahmer, 1♂ (MHNS 807). Osorno: El Mirador, 45 km W La Unión, 900 m, 1–2.III.1987, L. Peña, 1♂ 1♀ (AMNH); 36 km W La Unión, 600 m, 25–28.III.1987, L. Peña, 1♀ (AMNH); Purranque, I–III.1955, E. Reed, 1♂ (AMNH); Puyehue Natl. Park: Aguas Calientes, XII.1981, L. Peña, 1♂ (AMNH); 425 m, Valdivian forest, 31.I.1985, N. Platnick and O. Francke, 1♂ (MACN-Ar); 600 m, Malaise, Nothofagus forest, 18.XII.1984–8.II.1985, S. and J. Peck, 2♂ (AMNH); 40°44′0″S, 72°18′45″W, 450 m, 12.XII.2000–2.I.2001, forest, J. Miller, I. Agnarsson, Alvarez, J. Coddington, G. Hormiga, 1♀ penultimate (USNM); La Picada 450 m, NW Volcán Osorno, 15–20.I.1980, L.E. Peña, 1♂ (AMNH); Osorno, X.1977, A. Tobar, 1♂ 1♀ penultimate (AMNH); Puyehue, 15–16.XI.1990, L. Peña, 1♀ (AMNH); 700 m, 9.XII.1994, L. Peña, 2 immatures (AMNH); Pucatrihue, 25–31.I.1978, L. Peña, 1♀ (AMNH), 1–10.II.1986, L. Peña, 1♂ (AMNH). Llanquihue: Concordia, Fundo Pedernales, 13.II.1994, T. Cekalovic, 1♀ (AMNH); N Correntoso, VI–VII.1989, L. Peña, 1♀ (AMNH); N Correntoso, NE Puerto Montt, VIII.IX.1989, L. Peña, 1 immature (AMNH); Petrohué, 41°08′S, 72°25′W, 100 m, 15.XI.1966, E.I. Schlinger and M.E. Irwin, 1 immature (CAS); Petrohué Norte, III.1974, J. Solervicens, 1♂ (UC). Chiloé: Arroyo Cole Cole, 25 km N Cucao, 8–11.II.1991, M. Ramírez, 1♂, 1♀ (MACN-Ar). Palena: Chaitén, XII.1985, L. Peña, 1♀ (AMNH); 37 km SE Chaitén, FIT, 60 m, riverside secondary forest, 28.XII.1984–30.I.1985, S. and J. Peck, 1♂ (AMNH). Región XI (Ibáñez del Campo): Aisén: Cochrane, Río Backer, 180 m, 30.I.1990, L. Peña, 2♀ (AMNH); 25 km S Cochrane, 1–3.II.1990, L. Peña, 1♂ (AMNH); Lago Polux, 11.II.1983, T. Cekalovic, 1♂ 2♀ (AMNH); Murta, Lago General Carrera, 29–30.I.1990, L. Peña, 1♀ (AMNH); Río Simpson Natl. Park, S margin, 17.II.1991, M. Ramírez, 1 immature (MACN-Ar, photo MJR 551), 1♀ (MACN-Ar); 22 km E Aisén, 300 m, wet forest, 5.II.1985, N. Platnick and O. Francke, 2♀ (AMNH); Puerto Aisén, 2.I.1957, M. Cadoceo, 2♂ (MACN-Ar); 24–26.I.1962, L. Peña, 3♂ (IRSN IG23077); Puerto Ibáñez, 15.I.1882, L. Villavicencio, 1♀ (AMNH); Río Cisnes, 1–28.II.1961, L. Peña, 3♂, 2♂ 3♀ 1 immature, 1♂ 2 immatures (IRSN IG 23077); Río Cisnes Medio, Río Grande, 30.XII.1984–28.I.1985, 200 m, FIT, S. and J. Peck, 1♂ (AMNH); Río Simpson, 37 km W Cohiaique, 20 m, wet forest, 20.I.1986, N. Platnick, P. Goloboff, R. Schuh, 1♂ (AMNH). Región XII (Magallanes y Antártica): Ultima Esperanza: Rio Pérez, Seno Skyring, 10.X.1952, R. Rodríguez González, 1♂ 1♀ (AMNH), 1♀ penultimate (AMNH); Torres del Paine Natl. Park: near Refugio Chileno, 50°56′45″S, 72°55′0″W, 400–600 m, 8–9.XII.2000, J. Miller, I. Agnarsson, 1♀ (USNM), same, steppe, under rock, 1♀. Magallanes: Camerón, 14–17.XI.1960, L. Peña, 4♀ 1 immature (MCZ); Cueva del Milodón, 28.I.1976, T. Cekalovic, 1♀ (AMNH); Isla Daly, I.1962, L. Peña, 4 immatures (IRSN IG 23077); Dos Lagunas, 28.I.1976, T. Cekalovic, 2♀ (AMNH); Estancia La Vicuña, 1956, 1♀ 1 immature, no. 13, 4.III.1957, 3♀, 4 immatures, no. 14, 4.III.1957, 2 immatures, no. 15, 4.III.1957, 1 immature, no. 17, 15.II.1959, 2♀, J. Vellard (MACN-Ar); Estancia La Vicuña, SE Camerón, 1–6.XII.1960, L. Peña, 1♀ (MCZ); 35 km SW Camerón, Nothofagus assoc., 2.XII.1966, E. Schlinger and M. Irwin, 2♀, 2♀ penultimates (CAS); Península Hardy, Isla Hoste, Bahía Orange, 9.III.1961, B. Malkin, 2 immatures, (AMNH); Isla Navarino, Puerto Toro, 8.II.1896, Svenska expeditionen till Magleansläderna, 2♀ 2 immatures (NRS), 16–17.III.1961, B. Malkin, 3♀ 3 immatures (AMNH), 19.XII.1992, Michelsen, 1♀ 2 immatures (ZMH 178); Isla Navarino, Puerto Williams, 22.VIII.1966, T. Cekalovic, 1♀ (UC), 14.VIII.1976, T. Cekalovic, 1♀, 1 immature (AMNH); XII.1962–I.1963, P. Darlington, 3♀ 2♀ penultimates (MCZ); Isla Picton, Svenska expeditionen till Magleansläderna, 1♀ (NRS); no. 14, María Virginia, J. Vellard, 1♀ (MACN-Ar); 30 km Natales, road to Magallanes, 21.III.1948, M. Birabén, 5♀, 1♀, 2 immatures (MLP); Monte Alto, 7.III.1960, T. Cekalovic, 1♀ (UC), 17.III.1971, T. Cekalovic, 1♀ (AMNH); 7–11.III.1969, L. Peña, 1♀, 4♀ (MCZ); Paine, 10.X.1952, R. Rodríguez González, 1♂ (AMNH); Torres del Paine Natl. Park, 150 m, scrub, 10.II.1985, N. Platnick and O. Francke, 2♀ (AMNH); Península Brunswick, Tres Brazos, 9.III.1961, T. Cekalovic, 1♀ (UC); Port Famine, 10.I.1977, T. Cekalovic, 2♂ (AMNH); Puerto Hambre, II.1956, J. Vellard, 1♀ (MACN-Ar); Punta Arenas, IX.1892, Michelsen, 1 immature (ZMH); (La Turba), 23.II.1960, T. Cekalovic, 1♂ (MACN-Ar); dry forest remnant, 10.II.1985, N. Platnick and O. Francke, 2♀ (AMNH); 102.2 km NNW Punta Arenas, 430 m, Nothofagus assoc., 6.XII.1966, E. Schlinger and M. Irwin, 1♀ 1 immature, 2♀ 1 immature (CAS); Punta Arenas, Puerto del Hambre, 500 m, 8.II.1979, Misión Científica Danesa, 1♂ (ZMK); Cerro Castillo Natl. Res., 500–600 m, dry forest, 7.II.1985, N.I. Platnick and O.F. Francke, 2♀ (AMNH); Aserradero Río Bueno, no. 13, J. Vellard, 1 immature (MACN-Ar); Río Chico, 1956, J. Vellard, 5 immatures (MACN-Ar); Río San Juan, 25.I.1976, T. Cekalovic, 1♀ (AMNH); Río Rubens, ca. 52°S, 30.XI–3.XII.1962, P. Darlington, 2♀ 5 immatures (MCZ); Río Santa María, 25.I.1976, T. Cekalovic, 1♂ (AMNH); 8.I.1977, T. Cekalovic, 1♀ (UC); Rubens, 22.III.1948, M. Birabén, 3 immatures (MLP), 10.X.1952, R. Rodríguez González, 1♂ (AMNH), 13.XII.1960, L. Peña, 1♀ (MCZ); Rusffin, no. 11, III.1957, 1♀, 3 immatures, 5 immatures, 1♀, no. 12, 1.III.1957, 2♀, Silla del Diablo, 28.I.1976, T. Cekalovic, 1♂ 1♀ (AMNH); Spanger, 27.IV.1899, E, Nordenskiöld, 1♀ (NRS); Chorrillo Tres Puentes, IV.1969, T. Cekalovic, 1♀ (MACN-Ar); Tres Vientos, Puerto Arturo, 53°34′S, 73°23′W, 25–28.XI.1960, L. Peña, 2♀ (MCZ); Aserradero Yendegaia, no. 1, 12.II.1957, 1♂ 1♀ 2 immatures, 3♀, no. 2, 12.II.1957, 1♂ 2♀, no. 3, 13.II.1957, 1♀, 2 immatures, no. 4, 14.II.1957, 4♀, no. 5, 14.II.1957, 2 immatures, 10♀ 5 immatures, 1♀, no. 6, 15.II.1957, 3♀, 1♂ 2♀ 1 immature, 18.II.1957, 5 immatures, J. Vellard (MACN-Ar). No Specific Locality: South Chile, 190708, Skottsberg, 1♂ (NRS). Mistaken Locality: Santiago Prov., Malleco, XI.1979, L. Peña, 1♂ 2♀ 1♀ penultimate (AMNH) (see Ramírez, 1995b: 83).

    Morphological Remarks:

    Some details of the abdominal musculature, tracheae, and claw tufts were described in Ramírez (1995a).

    Tomopisthes varius Simon Figures 104B, 105C, D, 108

  • Tomopisthes varius Simon, 1884: 134 (male and female syntypes, from Chile, Cabo de Hornos, in MHNP 6670, examined), 1887: E30, 1897b: 107, 1902: 31, 1904: 102. Ramírez, 1995a: 368.

  • Gayenna varia: Tullgren, 1901: 233, 259, 1902: 59.

  • Diagnosis:

    Resembles T. horrendus in having a relatively large (but usually smaller) size, distinguished by having a simpler epigyne, with copulatory openings close to the epigastric furrow, and a short paramedian apophysis with several cusps. Immatures are distinguished from those of T. horrendus by lacking lateral spines on metatarsus I.

    Female (syntype, measurements of specimen from Quetrihué):

    Total length 11.30. Carapace length 5.05, width 3.65, wider on legs II–III. Length of tibia/metatarsus: I, 3.27/2.77; II, 3.17/2.77; III, 2.60/2.67; IV, 3.40/3.86. Palpal tarsus length 1.67. Chelicerae unmodified, with two teeth on retromargin. Sternum length 2.63, width 1.97. Spines: leg I, femur d 1–1–1, p 0-d1-(1-d1), r 0-d1-d1; tibia v 2–2–2; metatarsus v 2bas. II, femur = I; tibia v r1–2–2; metatarsus v 2bas, p d1–0. III, femur d 1–1–1, p and r 0-d1-d1; patella r d1; tibia v p1–2–2, p and r d1–1, d r1bas; metatarsus v 2–2–2, p and r d1–1–1, d 0-p1–2. IV, femur d 1–1–1, p 0-d1-d1, r d1ap; patella, tibia and metatarsus = III. Abdomen length 6.52, width 3.85, spiracle–epigastrium 2.67, spiracle–spinnerets 0.65. Color (figs. 104B, 108A): carapace grayish with dark spots, reddish to cephalic area. Legs pale grayish with dark spots. Sternum brown, darker on margins. Abdomen yellow with dark grayish pattern, venter paler, with small spots, mainly on median band. Epigyne (fig. 108E, F): median field ample in posterior view, weakly sclerotized, rugose. Copulatory ducts short. Ducts of accessory bulbs slightly converging.

    Male (syntype, measurements of specimen from Quetrihué):

    Total length 8.40. Carapace length 5.05, width 2.70. Length of tibia/metatarsus: I, 3.20/2.70; II, 2.80/2.50; III, 2.23/2.27; IV, 2.83/3.17. Chelicerae slightly longer and narrower than those of female, teeth on retromargin more separated, fang long, thick. Sternum length 1.90, width 1.50. Spines as in female, except: leg II, tibia v 2–2–2. Abdomen length 4.80, width 2.40, spiracle–epigastrium 2.17, spiracle–spinnerets 0.60. Color as in female (fig. 108B). Palp (figs. 105C, D, 108C, D): tibia long, width/length 0.49. Cymbium relatively narrow. Embolus short, basal process lobate, thick, separated by ample, weakly sclerotized area (fig. 105C). Median apophysis short, relatively wide. Paramedian apophysis with complex base, flattened tip. Primary conductor small, triangular. Secondary conductor small, complex, quite modified.

    Morphological Remarks:

    Tracheal system and claw tufts described in Ramírez (1995a).

    Variability:

    Size is quite variable. Male Spines: tibiae III, IV v 2–2–2.

    Natural History:

    A very opportunistic species that builds retreats on foliage, under bark, in crevices in rotten logs, and occasionally under stones or between flat stones in ravines.

    Distribution:

    Southern forests in Argentina and Chile, from the relict forest in Fray Jorge to Cabo de Hornos.

    Other Material Examined:

    ARGENTINA: Neuquén: Estancia San Ramón, Rincón Chico, Río Limay, X–XII.1962, Havrylenko, 1♂ (MACN-Ar); Lago Nompehuén, NW Aluminé, 12.I.1985, E. Maury and A. Toth, 2♀ 1 immature (MACN-Ar); Lago Aluminé, II.1974, E. Maury, 3♀ (MACN-Ar); Lago Carrhué Chico, 15–16.I.1983, E. Maury, 2♀ (MACN-Ar); Lanín Natl. Park: Lago Lácar, Pucará, I.1961, M.E. Galiano, 2♀ (MACN-Ar 5287); I–II.1971, S. Schajovskoy, 1♀ (MACN-Ar), 10.XI.1978, Misión Científica Danesa, 1♂ (ZMK); 650 m, 28–29.XI.1981, Nielsen and Karsholt, 4♂ (ZMK); San Martín de los Andes, 40°10′S, 71°21′W, 20–21.XI.1988, V. and B. Roth, 1♂, 1♀ (CAS); Nahuel Huapi Natl. Park: Isla Victoria, Piedras Blancas, 1964, Contreras, 1♂ (MACN-Ar); Isla Victoria, Playa del Toro, VI.1984, M. Ramírez, 1♀ (MACN-Ar); Río Frías superior, I.1990, M. Ramírez, 1♀ (MACN-Ar); Lago Nahuel Huapi, Península Quetrihué, 16–25.XII.1984, 1♂ (MACN-Ar); II.1986, M. Ramírez, 1♂ (MACN-Ar); 23.I.1985, M. Ramírez, 3♀ 1 immature (MACN-Ar); Península Quetrihué, “arrayan” forest, stony beach, 24.II.1996, M. Ramírez, 1♂ (MACN-Ar); Puerto Blest, II.1986, M. Ramírez, 1♂ (MACN-Ar); 770 m, 4.XII.1978, 2♀, 25–27.X.1981, 1♂ 1♀, Nielsen and Karsholt (ZMK), 10.I.1998, M. Ramírez, 1♂ (MACN-Ar); 7–20.I.2000, L. Lopardo and A. Quaglino, 9 immatures (MACN-Ar); Puerto Blest, Lago Ortiz Basualdo, I.1990, M. Ramírez, 2♀ (MACN-Ar); San Carlos de Bariloche, II.1954, M.E. Galiano, 2♀ (MACN-Ar), 1964, Monrós, 1♀ (MACN-Ar); San Carlos de Bariloche, Colonia Suiza, 810 m, 20.XI.1978, 1♀, 5.XII.1978, 1♀, Misión Científica Danesa (ZMK); Nahuel Huapi, 1950, Havrylenko, 2♀ (MACN-Ar 5507). Chubut: El Maitén, 29.VII.1961, A. Kovács, 2 immatures (AMNH); Los Cipreses, XI.1982, M. Ramírez, 2♀ penultimates 2 immatures (MACN-Ar); Los Alerces Natl. Park, I.1982, P. Goloboff, 1♂ (MACN-Ar); Los Alerces Natl. Park: Lago Futalaufquen, I.1990, M.J. Ramírez, 1♀ 1 immature, 1♀ (MACN-Ar); Lago Menéndez, I.1990, M. Ramírez, 1♂ (MACN-Ar); Lago Verde, II.1985, M. Ramírez, 1♀, 1♀ 1 immature (MACN-Ar). Santa Cruz: Lago Frías, no date, E. Maury, 1♂ 1♀ (MACN-Ar); Los Glaciares Natl. Park, II.1977, no collector, 1♂ (MACN-Ar), 18.I.1980, P. Goloboff, 1♂ 1♀ 1 immature (MACN-Ar); no date, E.R. Fernández, 1♂ (MACN-Ar); Los Glaciares Natl. Park, Península Magallanes, in front of Glaciar Moreno, II.1977, D. Pepe and M. Rumboll, 1♂, 1♀ (MACN-Ar); Ventisquero Moreno, 18–24.I.1971, J. Vellard, 2♂ (MACN-Ar); 28.II.1971, J. Vellard, 1♂ (MACN-Ar). Tierra del Fuego: Lago Fagnano, XI.1984, 1♀ (MACN-Ar); Lago Roca, Nothofagus antarctica forest, II.1967, Williner, 1♀ (MACN-Ar); 27.I.1971, J. Vellard, 1♀ 1 immature (MACN-Ar); Lapataia, I.1948, J.M. Viana, 1♂ 1♀ (MACN-Ar); II.1963, E. Maury, 4♀, 1♂ (MACN-Ar); Río Grande, XI.1973, M. Rumboll, 1♀ (MACN-Ar); Ushuaia, Playa Larga, 13.XII.1967, M.E. Galiano, 1♀, 1♀ (MACN-Ar); Punta Remolino 24, 24.II.1959, J. Vellard, 1♂ 1♀ penultimate (MACN-Ar); Río Pipo, XII.1989, A. González, 1♀ (MLP); Aserradero Yendegaya, no. 1, 12.II.1957, 1♂ 1♀; no. 3, 13.II.1957, 1♀, no. 4, 14.II.1957, 1♂, no. 5, 14.II.1957, 1♀ 2 immatures, 1♀, no. 7, 16.II.1957, 1♀ 1 immature, no. 9, 17.II.1957, 2♀, J. Vellard (MACN-Ar). CHILE: Región IV (Coquimbo): Limarí: Fray Jorge Natl. Park, 21.X.1966, E. Schlinger, M. Irwin, 1♀ (CAS); pitfalls in relict Valdivian forest, 16.II.19??, 1♀, 17.IV.19??, 1♀, 19.IV.19??, 1♀, 19.VIII.19??, 1♀, 17.X.19??, 2♂ 1♀, R. Calderón G. (AMNH), 27.IX.1970, 1♂ 1♀ (UC); 580 m, relict valdivian forest, 5.I.1985, N. Platnick and O. Francke, 1♀ 1 immature (AMNH); 9.I.1984, A. Roig, 1♀ (MACN-Ar); 560 m, relict Valdivian fog forest, under rocks, 8.II.1986, N. Platnick and R. Schuh, 2♀ (AMNH); 560 m, 3.X.1992, N. Platnick, P. Goloboff, K. Catley, 1♂ 4 immatures (AMNH); elev. 580 m, 10.XI.1993, 30°40′S, 71°41′W, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH); Fray Jorge, Rancho, 10.XII.1950, Ross and Michelbacher, 1♀ (CAS); Fray Jorge, forest, 11.XII.1950, Ross and Michelbacher, 1♀ 1 immature (CAS). Choapa: Los Vilos, 12.X.1986, L. Peña, 1♀ (AMNH); Quereo, Los Vilos, 6.XI.1988, P. Goloboff, E. Maury, and C. Szumik, 1♀ (MACN-Ar). Región V (Valparaíso): Los Vilos, 6.XI.1988, P. Goloboff, E. Maury, and C. Szumik, 1♀ (MACN-Ar). Petorca: Los Molles, Ovalle, 1600 m, 17.X.1994, L. Peña, 1♀ (AMNH); E La Ligua, relict forest, 27.IX.1980, L. Peña, 1♀ (AMNH). Quillota: Cuesta El Melón, nr. La Calera, 15.XI.1985, L. Peña, 2♀ 3 immatures (AMNH). Valparaíso: Quintero, pitfalls in relict forest, 12.VIII.1968, 3♂, 2.X.1968, 1♂, 3♂, R. Calderón G. (AMNH); 2.X.1968, 1♀ (UC); Quintero, Barber VII.I, 12.VIII.1968, 1♂, 1♂ (UC). Región Metropolitana (Santiago): Santiago: El Canelo, 950 m, 33°37′S, 71°35′W, M. Irwin and E. Schlinger, 5♀ (CAS); Pan de Azúcar, 2700 m, road to Cerro Tupungato, I.1984, A. Mann Z., 1 vial (number of specimens not recorded) (MHNS 867). Región VII (Maule): Curicó: Cajón del Río Claro, Cordillera Curicó, 9.X.1966, E. Schlinger, 2♀ (CAS); Las Tablas, E Curicó, II.1985, L. Peña, 3♀ (AMNH); Los Queñes I.1984, P. Goloboff, 1♀ 1 immature (MACN-Ar), 14.I.1984, P. Goloboff, 1♀ 1 immature (MACN-Ar). Talca: Gil de Vilches, 1200 m, I.1984, P. Goloboff and E. Maury, 3♀, 2♀ (MACN-Ar); 7.I.1989, M. Ramírez, 1♀ (MACN-Ar, photo MJR 10); 7–8.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH). Cauquenes: Tregualemu, 520 m, 6–7.XI.1993, L. Peña and A. Ugarte, 1♀ (AMNH). Región VIII (Biobío): Ñuble: 60 km SE Chillán, Termas Road, beech forest, FIT, 1300 m, 7.XII.1984–19.II.1985, S. and J. Peck, 2♂ (ANMH); 22.7 km ESE Recinto, 1330 m, Nothofagus forest, site 646, window trap, 10.XII.1983, A. Newton and M. Thayer, 1♀ (AMNH). Concepción: Bosque de Ramuntcho, Cerro Caracol, Concepción, elev. 200 m, 36°51′S, 73°02′W, 17.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1 vial (number of specimens not recorded) (AMNH); Lomas Colorada, 24.XI.1996, T. Cekalovic, 1♀ (AMNH); Periquillo, 8.XII.1994, T. Cekalovic 1 vial (number of specimens not recorded) (AMNH); San Pedro, 22.X.1974, T. Cekalovic 1♀ (UC); Tomé, 1.I.1992, T. Cekalovic, 1♀ (AMNH); Lenga, Teta Norte, 12.III.1979, M. Casanueva, 1♀ (UC). Biobío: W Ralco, Santa Bárbara, 400 m, 22–23.XI.1994, L. Peña, 1♀ (AMNH). Región IX (Araucanía): Malleco: 17 km W Angol, 800 m, FIT, mixed Nothofagus, 8.XII.1984–16.II.1985, S. and J. Peck, 9♂ 3♀, 2♂ 4♀ 9 immatures (AMNH); 40 km W Angol, Nahuelbuta Natl. Park, FITS, 1200–1500 m, Nothofagus/Araucaria forest, 9.XII.1984–17.II.1985, S. and J. Peck, 1♂ 1♀, 1♀ (AMNH); 20 km W Curacautín, 1000 m, FIT, 1000 m, Nothofagus forest, 12.XII.1984–16.II.1985, S. and J. Peck, 1♀, 1♀ (AMNH), 1500 m, Malaise, Nothofagus-Araucaria forest, 16.II.1985, S. and J. Peck, 3♀ (AMNH); 40 km W Curacautín, Nothofagus/Araucaria, 12.XII.1984–16.II.1985, S. and J. Peck, 1♀, 1♀ (AMNH); Fundo María Ester, 15 km W Victoria, 14.I.1989, M. Ramírez, 1♀ (MACN-Ar); Malalcahuello, 9–15.XII.1985, L. Peña, 2♀ (AMNH); 6.5 km E Malalcahuello, 1080 m, trap site 651, widow trap, Nothofagus dombeyi with Chusquea, 13–31.XII.1982, A. Newton and M. Thayer, 2♀ (AMNH); Monumento Natural Contulmo, 12.I.1989, M. Ramírez, 5♀ (MACN-Ar); mixed evergreen forest, 11.XII.1984–13.II.1985, S. and J. Peck, 1♀, 3♂ (AMNH); Nahuelbuta Natl. Park, 1230 m, dry forest, 1.II.1986, N. Platnick and T. Schuh, 2♂ (AMNH); Tolhuaca, 15–23.III.1986, L. Peña, 1♂ 6♀ 6 immatures (AMNH), 15.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1 immature (MACN-Ar). Cautín: Bellavista, N shore Lago Villarrica, 310 m, site 655, window trap, valdivian rainforest, 15–30.XII.1982, A. Newton and M. Thayer, 1♂ (AMNH); undisturbed forest, 260 m, 30.I.1986, N. Platnick, T. Schuh, 1♀ (AMNH); 39°12′S, 72°08′W, 240 m, 20.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♂ (AMNH); Chacamó, NW Nueva Imperial, W Temuco, 16–24.II.1981, L. Peña, 1♀ (AMNH); Cuesta Lascarría, 23.IX.1968, H. Prats, 1♂ (UC); Lago Caburgua, 39°08′S, 71°46′W, 10.XII.1988, V. and B. Roth, 2♀ (CAS); Conguillio Natl. Park, 23.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH); 10 km S Pucón, Volcán Villarrica Natl. Park, FIT, 900 m, Nothofagus grove on ash, 15.XII.1984–10.II.1985, S. and J. Peck, 1♂ (AMNH); 21 km NE Pucón, Lago Caburga, FIT, 600 m, mixed forest remnant, 15.XII.1984–10.II.1985, S. and J. Peck, 4♂ (AMNH); Pucón, FIT (toxic), hilltop beech (peninsula), 8–13.XI.1989, S.A. Marshall, 1♂ (AMNH); FIT, mature forest, 15.XI–1.XII.1989, S.A. Marshall, 3♂ (AMNH); 17 km E Pucón, 23.XI.1983, N. Platnick, T. Schuh, 2♀ (AMNH); Termas de Palguin, Salto Puma, 725 m, 39°22′S, 71°50′W, fogging fungusy logs, 24.XI.1994, R. Leschen and D. Carlton no. 157, 1♀ (AMNH); Tolten (coastal town), 27.II.1979, L. Peña, 1♀ 1 immature (AMNH); 14 km N Villarrica, elev. 250 m, 39°10′S, 72°12′W, 20.XI.1993, N. Platnick, K. Catley, M. Ramírez, T. Allen, 1♀ (AMNH); 15 km NE Villarrica, Flor del Lago, 300 m, 2 FITS, Nothofagus forest, 14.XII.1984–10.II.1985, S. and J. Peck, 6♂ (AMNH). Región X (Los Lagos): Valdivia: Las Lajas, W La Unión, 19–20.XI.1990, L. Peña, 1♂ 2♀ (AMNH); Peulla, 200 m, 25.III.1965, H. Levi, 1 immature (MCZ); I.1990, M. Ramírez, 3♀ (MACN-Ar); Purolón, NW Panguipulli, 10.I.1985, L.E. Peña, 1♀ (AMNH); Valdivia, 12.X.1976, E. Krahmer, 2♀ (AMNH), 1983, 1♀ (MHNS 822), 1♀ (MHNS 829), XI–XII.1982, 1♀ (MHNS 700), E. Krahmer; 30 km S Valdivia, 11.II.1981, T. Cekalovic, 1♀ (AMHN). Osorno: Maicolpué, 64 km W Osorno, 19.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1♀ (AMNH), 1♀ (MACN-Ar); Puyehue Natl. Park: Aguas Calientes, FIT, Derrumbes forest trail, 20.XII.1984–8.II.1985, S. and J. Peck, 4♂ 1♀ 2 immatures (AMNH), 18.II.1992, M. Ramírez, N. Platnick, P. Goloboff, 1 immature (MACN-Ar); 40°44′S, 72°19′W, 1440 m, 5–7.XII.1988, V. and B. Roth, 1♂ 2♀ (CAS), 13–17.XII.1998, M. Ramírez, L. Compagnucci, C. Grismado, L. Lopardo, 1♂ (MACN-Ar 15.XII.98/8, frame 20); 600 m, Malaise, Nothofagus forest, 18.XII.1984–8.II.1985, S. and J. Peck, 2♂ 1♀ (AMNH); Anticura, 19–29.X.1985, L. Peña, 1♂ 1♀ (AMNH); Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.XII.2000–2.I.2001, forest, J. Miller, Alvarez, J. Coddington, G. Hormiga, 1♂ (USNM); 4.1 km W Anticura, 270 m, trap site 663, window trap, valdivian rainforest, 19–25.XII.1982, A. Newton and M. Thayer, 3♂ (AMNH); Osorno, VIII.1977, A. Tobar, 1♂ (AMNH); 34.5 km E Osorno, 40°38′S, 71°42′W, 200 m, fogging fungusy log, 1.XII.1994, R. Leschen and C. Carlton no. 191, 1 immature (AMNH); 48.5 km W Osorno, 40°37′S, 73°45′W, 75 m, sifting leaf litter, 28.XI.1994, R. Leschen and C. Carlton no. 191, 1♀ (AMNH); Termas de Puyehue, 180 m, forest, 28.XI.1981, N. Platnick and R. Schuh, 1♂ 1♀ (AMNH); 460 m, litter from moss forest floor, 25.XI.1981, N. Platnick and R. Schuh, 1♂ 2♀ (AMNH); Puyehue, 700 m, 9.XII.1994, L. Peña, 1♂ 2♀ (AMNH); Pucatrihue, 25–31.I.1978, L. Peña, 1♀ (AMNH). Llanquihue: Caleta La Arena, on Marcypospermum, 30.I.1991, M. Ramírez, 1 immature (MACN-Ar); Correntoso, XII.1969, L. Peña, 1♀ (MCZ); N Correntoso, NE Puerto Montt, VIII.IX.1989, L. Peña, 3♂ 1 immature (AMNH); NW Shore, Lago Chapo, 250 m, 41°27′S, 72°30′W, 13.XI.1966, M. Irwin and E. Schlinger, 1♂ 1♀ 7 immatures (CAS); 2–3 km NW Ensenada, 18.III.1965, H. Levi, 1♀ (MCZ); Los Muermos, S. Chile, forest, 19.I.1951, Ross and Michelbacher, 1♀ 7 immatures (CAS); Vicente Pérez Rosales Natl. Park, Salto Petrohué, mixed most forest, 150 m, FIT, 23.XII.1984–4.II.1985, S. and J. Peck, 1♂ (AMNH); Petrohué, 41°08′S, 72°25′W, 100 m, 15.XI.1966, E.I. Schlinger and M.E. Irwin, 1♀ (CAS); 8 mi W Puerto Varas, 18.I.1951, Ross and Michelbacher, 1♀ (CAS). Chiloé: Isla de Chiloé: Cucao, tepual, 12.II.1990, M. Ramírez, 2♀ (MACN-Ar); Chepu, 17 m, 29.XI.1981, N. Platnick and R. Schuh, 1♀ 1 immature (AMNH); 3.II.1991, M. Ramírez, 3♀ (MACN-Ar); 15 km S Chepu, 3.II.1991, M. Ramírez, 1♀ (MACN-Ar); Pid-Pid, 17.II.1997, T. Cekalovic, 1♀ (AMNH); “Port San Pedro, Chiloes, leg. Keyserling, unentwickeltes (imm.)”, 1 immature (MCZ); 5 km N Quellón, 105 m, modified forest, floor litter and moss, concentrate Berlese, 1.XII.1981, N. Platnick and R. Schuh, 1♀ (AMNH), 107 m, 1.XII.1981, N. Platnick and R. Schuh, 1♀ (AMNH); Terao, S Chonchi, 10–20.III.1988, L. Peña, 1♂ (AMNH). Región XI (Ibáñez del Campo): Aisén: Cochrane, Río Backer, 180 m, 30.I.1990, L. Peña, 1♂ (AMNH); Río Simpson Natl. Park, 33 km Puerto Aisen, selectively cut forest, 31.XII.1984–26.I.1985, S. and J. Peck, 1♂, 1♀ (AMNH); Puerto Aisén, 24–26.I.1962, L. Peña, 2 immatures (IRSN IG23077); Río Cisnes, 1–28.II.1961, L. Peña, 1♂ (IRSN IG23077); Río Cisnes Medio, Río Grande, 30.XII.1984–28.I.1985, 200 m, FIT, S. and J. Peck, 1♀ (AMNH); Valle del Río Aisen, I.1897, P. Dusen, 1♂ 3♀ 8 immatures (NRS). Región XII (Magallanes y Antártica): Ultima Esperanza: Rio Pérez, Seno Skyring, 10.X.1952, R. Rodríguez González, 1♂ (AMNH); Ultima Esperanza, 3.IV.1896, 1♂ (NRS). Magallanes: Camerón, 14–17.XI.1960, L. Peña, 1♀ (MCZ); Cueva del Milodón, 28.I.1976, T. Cekalovic, 1♂ 3♀ (UC), 3♂ 4♀ (AMNH); Estancia Gazy Harbour, 10.II.1990, T. Cekalovic, 1♀ (AMNH); Isla Daly, I.1962, L. Peña, 1♀ 3 immatures (IRSN IG 23077); Isla Navarino, Puerto Toro, 1896, Svenska expeditionen till Magleansläderna, 1♀ (NRS); XI.1892, Delfin, 2♀ (ZMH 179); Isla Nueva, 4.II.1896, 1♀ 4 immatures (NRS); Torres del Paine Natl. Park, 150 m, scrub, 10.II.1985, N. Platnick and O. Francke, 2♀ (AMNH); Península Brunswick, Barranco Amarillo, 27.I.1976, T. Cekalovic, 2♀ (AMNH); Port Famine, 10.I.1977, T. Cekalovic, 1♂ (AMNH); Puerto Bridges, 10.I.1893, Michelsen, 2♀ 1 immature (ZMH 165); Puerto Hambre, 25.III.1991, T. Cekalovic, 1♀ 2 immatures (AMNH); Punta Arenas, IX.1892, Michelsen, 1♂ 1♀ (ZMH 75); Punta Arenas (La Turba), 27.VIII.1976, T. Cekalovic, 1♀ (AMNH); Chorrillo Tres Puentes, 7.II.1971, T. Cekalovic, 1♀ (AMNH); no date, T. Cekalovic, 1♀ (MACN-Ar); Tres Vientos, Puerto Arturo, 53°34′S, 73°23′W, 25–28.XI.1960, L. Peña, 3♀, 8♀ (MCZ); Puerto Bulnes, 25.III.1991, T. Cekalovic, 1♀ (AMNH); Tweedie, Sierra del Toro, 19.III.1896, E. Nordenskjöld, Svenska expeditionen till Magleansläderna, 1♀ (NRS); Aserradero Yendegaia, no. 3, 13.II.1957, J. Vellard, 1♀ (MACN-Ar). No Locality: E67–2–12, 1♀ (CAS). Locality Not Found: Puerto Anchorena (presumably Argentina), 1.XI.1969, Contreras, 1♂ (MACN-Ar). Mistaken Locality: Santiago Prov., Malleco, XI.1979,