Systematics of the spider genus Metabus O. P.-Cambridge, 1899 (Araneoidea: Tetragnathidae) with additions to the tetragnathid fauna of Chile and comments on the phylogeny of Tetragnathidae

Abstract

The spider genus Metabus (Tetragnathidae) previously included nine species: the type M. gravidus O. P.-Cambridge, 1899 - junior synonym of Leucauge ocellata (Keyserling) - from Central America and eight species from Chile. In this paper, the classification of the Metabus species-complex is revised, and two new genera, with three new species and five new combinations, are described. Allende gen. nov. is created for four Chilean species not congeneric with the type of Metabus: the type A. puyehuensis sp. nov., A. patagiatus (Simon) comb. nov., A. nigrohumeralis (F. O. P.-Cambridge) comb. nov. and A. longipes (Nicolet) comb. nov. Further additions to the Chilean fauna are under the new genus Mollemeta gen. nov.–created for M. edwardsi (Simon) comb. nov.–and three new species of Chrysometa: C. acino sp. nov., C. lev sp. nov. and C. maitae sp. nov.Metabus now includes four species: M. ocellatus (Keyserling) comb. nov., M. debilis (O. P.-Cambridge) comb. nov., M. ebanover sp. nov. and M. conacyt sp. nov. All of these species were included in a phylogenetic analysis of 38 tetragnathid and 12 orbicularian outgroup terminals scored for 105 morphological and behavioural characters. The results suggest that Metabus as previously circumscribed is polyphyletic. The phylogenetic relationships within tetragnathids are briefly discussed.

morphology, phylogenetics, taxonomy

INTRODUCTION

Tetragnathid spiders are usually found on the centre of their orb-webs or hiding in nearby vegetation. They usually spin their webs along river margins, near ponds or inside the forest. Most tetragnathids build orb-webs with few radii and spirals (see Fig. 10A–D), although webs with many radii and spirals exist such as in the genera Azilia Keyserling, 1881 and Dolichognatha O. P.-Cambridge, 1869 (Levi, 1980: pl. 8; 1981: pl. 2). Tetragnathids are differentiated from other araneoid taxa with similar web architecture by their spinneret and genital anatomy (Hormiga, Eberhard & Coddington, 1995; Griswold et al, 1998). This family has a worldwide distribution and comprises 52 genera and 955 species (Platnick, 2007).

Figure 10

Photos of webs: A, Allende puyehuensis. B, Chrysometa acinosa. C, Mollemeta edwardsi. D, Metabus ebanoverde. Scale lines 10 cm. Photos A–C from Puyehue National Park, Chile. Photo D from Ebano Verde Scientific Reserve, Dominican Republic. (Photos A to C by G. Hormiga).

Tetragnathid and araneoid phylogenetic relationships have been studied previously (Levi, 1980; Coddington, 1990; Hormiga et al, 1995; Scharff & Coddington, 1997; Griswold et al, 1998). Except for Levi (1980), these authors all recovered a monophyletic Tetragnathidae comprising the monophyletic subfamilies Tetragnathinae and Nephilinae and a paraphyletic Metinae. Hormiga et al (1995) studied the phylogenetic relationships within Tetragnathidae in more detail; they included 14 tetragnathid genera and eight orbicularian outgroups in seven families (22 species in total) coded for 60 morphological and behavioural characters. Their analysis found an unstable paraphyletic three-lineage configuration for Metinae; these authors suggested that future studies on tetragnathid phylogenetics should include genera such as Metabus O. P.-Cambridge, 1899 and Tylorida Simon, 1894, among other taxa (Hormiga et al, 1995: 358–359).

The spider genus Metabus formerly included nine nominal species (Platnick, 2007): the type–M. gravidus O. P.-Cambridge, 1899, junior synonym of Leucauge ocellata (Keyserling, 1864) - and eight species of what Levi (1986: 105) termed the ‘Meta porte group’. These Meta C. L.-Koch, 1836 species were transferred to Metabus by Archer (1963: 17) because they have in common a cymbial basal apophysis and lack femoral trichobothria; Meta auroSimon, 1901 and Meta patagiaSimon, 1901 remained in Meta. The species that Archer transferred were: Metabus porteri (Simon, 1900), M. longipes(Nicolet, 1846), M. nigrohumeralis(F. O. P.-Cambridge, 1889), M. fuegianus (Simon, 1902), M. chilensis (Tullgren, 1902), M. cordillera (Tullgren, 1902), M. echinatus (Tullgren, 1902) and M. tortus (Tullgren, 1902).

Metabus ocellatus has received most attention within this genus. These spiders build communal orb-webs over ponds and share a common retreat during the night, a behaviour unique among tetragnathid spiders (Buskirk, 1975a, b, 1986; Uetz & Craig, 1997). Petrunkevitch (1925) studied the anatomy of this species and described the male. Tanikawa (2001) included this species with Okileucauge sasakiiTanikawa, 2001 from Japan and six more tetragnathid genera in a phylogenetic analysis, which showed that M. ocellatus and O. sasakii were not sister taxa. This cladogram optimized the similarities in genital anatomy and the lack of femoral trichobothria between these two species as symplesiomorphic (Tanikawa, 2001: figs 1–8). Metabus and the species of the genus Okileucauge(see also Zhu, Song & Zhang, 2003) share several morphological features with genera such as Leucauge White, 1841 and Tylorida. The other Chilean Metabus species were known only from the original descriptions, many of them without illustrations.

Figure 1

Strict consensus (325 steps CI = 44 RI = 75) of the ten most parsimonious cladograms 322 steps in length (CI = 45 RI = 75). Homoplasious changes are indicated with a white square, non-homoplasious changes with black squares. Discontinuous states are marked as homoplasy. Numbers on the squares refer to the characters in Appendix 1, and state number is indicated beneath the square. Only changes common to all topologies are mapped on polytomies.

Figure 2

Strict consensus of ten most parsimonious cladograms 322 steps in length (CI = 45 RI = 75). Bremer and Jackknife values, respectively, are given beneath each node.

Figure 3

A–E, male genital anatomy of Metabus ocellatus. A, pedipalp ventral view. B, ectal view. C, dorsal view. D, apical view. E, mesal view. F, Leucauge venusta male pedipalp.

Figure 4

A-E, female genital anatomy of Metabus ocellatus. A, epigynum ventral view. B, caudal view. C, spermatheca dorsal view. D, fertilization and copulatory ducts. E, accessory gland openings. F, Leucauge venusta spermatheca dorsal view.

Figure 5

A-E, male genital anatomy of Allende puyehuensis. A, pedipalp ventral view. B, ectal view. C, cymbial processes. D, pedipalp apical view. E, mesal view. F, Chrysometa alajuela pedipalp ectal view.

Figure 6

A, female genital anatomy of Allende patagiatus mating plug. B–C, A. longipes. B, epigynum caudal view. C, dorsal view. D–E, A. nigrohumeralis. D, fertilization and copulatory ducts. E, accessory gland openings. F, Chrysometa saladito spermatheca lateral view.

Figure 7

A-E, male genital anatomy of Mollemeta edwardsi. A, pedipalp ventral view. B, ectal view. C, cymbial basal process. D, mesal view. E, palpal tibia. F, Metellina curtisi pedipalp ectal view.

Figure 8

A-E, female genital anatomy of Mollemeta edwardsi. A, epigynum ventral view. B, caudal view. C, dorsal view. D, spermatheca lateral view. E, accessory gland openings. F, Chrysometa alajuela epigynum anterior view.

The present study was designed to test the monophyly of Metabus as previously circumscribed, to revise the species included within this genus, and to place phylogenetically the Chilean tetragnathids species described here. The results show that Metabus ocellatus is not congeneric with the species of the ‘Meta porte group’ and the new genus Allende is created to accommodate the latter. The new genus Mollemeta and three new species of Chrysometa Simon, 1894 are also described.

While this paper was in revision, a phylogenetic analysis by Kuntner (2006) hypothesized that nephilines should not be grouped with the remaining Tetragnathidae taxa and elevated nephilines to the family rank. In Kuntner's (2006) analysis the possible sister groups of the nephilines were: araneids, the clade with tetragnathids plus araneoid sheet-web builders, or the clade with all remaining araneoids. In a previous analysis nephilines had been suggested as the sister group of Araneidae (Kuntner, 2003; see also Wunderlich, 2004). A more comprehensive analysis (Kuntner, 2005) concluded that nephilines are the sister lineage of all remaining Araneoidea. Furthermore, analysis of DNA sequence data has suggested that the genus Nephiline is more closely related to Araneidae than it is to Tetragnathidae (Pan et al, 2004), but their study used a very shallow taxonomic sample. The taxonomic sample in the present paper is not designed to test the placement of Nephiline and its relatives, but nonetheless the results of the analysis reported here suggest that Nephilinae is the sister to other tetragnathids, as hypothesized by other cladistic studies (Coddington, 1990; Hormiga et al, 1995; Scharff & Coddington, 1997; Griswold et al, 1998; Kuntner & Alvarez-Padilla, 2006). The phylogenetic placement of nephilines within Araneoidea remains controversial and to this date no study has provided a robust hypothesis for the closest relatives of this lineage.

MATERIAL AND METHODS

Taxonomy

General taxonomic methods and description formats follow Hormiga (2002, 2003). Morphological observations and illustrations of external structures were made using Leica MZ APO dissecting and Leica DMRM compound microscopes with a camera lucida. Internal genitalic structures were cleared in methyl salicylate (Holm, 1979) and mounted on temporary slides (Grandjean, 1949; Coddington, 1983). All measurements were taken with a reticule calibrated in millimetres using dissecting and compound microscopes. In addition, an LEO 1430 VP scanning electron microscope (SEM) was used to study the ultrastructure of many morphological internal and external features. Soft tissues surrounding internal structures were effectively digested overnight with the SIGMA Pancreatin P1750 enzyme complex following the concentrations described by Dingerkus & Uhler (1977).

Specimens

The specimen data were managed in Biota: The Biodiversity Data Manager 1.6.1 (Colwell, 1992–2000). A spreadsheet with the specimen information can be downloaded from http://www.gwu.edu/~spiders/taxonomy.htm. Names for districts and provinces not included in the original labels were obtained from GEOnet Names Server (GNS) 2004 and The Global Gazetteer Version 2, 1996–2004. A small label bearing the author's initials (‘FAP’) with the specimen code in the daTabascoe was added to the vials. Size variation is the maximum size range of the taxon. Whether specimens were illustrated, measured or used for SEM vouchers is indicated under the material examined section of the corresponding species. The specimens illustrated were used in the phylogenetic analysis. Additional material is given in Appendix 3.

Taxa

The taxonomic sample analysed here includes 38 tetragnathid species (including all 22 from Hormiga et al, 1995) representing 25 genera and an undescribed Australian tetragnathid species, and six araneoid families (12 species) that served as outgroups (SensuGriswold et al, 1998), rooted with the deinopoid genus Uloborus Latreille, 1806. Each genus is represented by only one species, with the exception of Allende(four), Metabus(four), Leucauge(two), Chrysometa(six) and Orsinome Thorell, 1890 (two). All outgroup families are represented by one species except Araneidae (six). Three Chrysometa species described by Levi (1986) were included to test if Allende rendered Chrysometa paraphyletic, as both genera share many anatomical features. Differences in several anatomical features between Orsinome cf. vet(Hasselt, 1882) (Thorell, 1890: 209) and O. sarasiniBerland, 1924 (Berland, 1924: 210, figs 108–112) suggest that Orsinome may not be monophyletic. The number of araneid species was increased to test whether some of the Chilean species described here grouped within this family or with Tetragnathidae. Araneid species exemplified the two major basal araneid clades Araneinae and the ‘argiopoid’ clade (SensuScharff & Coddington, 1997). Appendix 2 lists species used and the data matrix.

Characters

The 105 characters coded for these taxa include the 60 from Hormiga et al (1995) and 40 new characters, which total 43 male and 14 female genitalic, 35 somatic and 13 behavioural characters (Appendix 1). Some characters from Hormiga et al (1995) required modification to accommodate the morphological diversity found in the new taxa added. For modified or new character state definitions, an exemplar taxon is given in parentheses, and reference to an illustration, if possible. The ‘character state type’ is meant to be taken as a reference for future homology assessments (e.g. Hormiga, 1994: 5). References are provided for the characters not illustrated or discussed in this study. Characters are cited as (character number - state number) throughout the manuscript. Character names are presented as published in the original work to facilitate comparison with other studies, which included 26 ‘presence-absence’ characters. Sometimes this practice makes the homology definition implicit rather than explicit, which can be rectified by providing detailed character definitions and illustrations.

Analyses

Parsimony analyses were performed using TNT 1.0 (Goloboff, Farris & Nixon, 2004) and NONA version 2.0 (Goloboff, 1993). WinClada 1.00.08 (Nixon, 2002) was used to edit the data matrix and to view cladograms. Searches in TNT used seed 1, 500 random additions and TBR. Jackknife values in TNT used seed 1, 100 random additions, TBR and saved 50 trees per addition for 1000 iterations. Bremer support values (Bremer, 1988, 1995) were calculated in NONA using the following commands (h 1000; mult*50; max*; h 1000; sub 1; find*; h 2000; sub 2; find*; h 4000; sub 3; find*; h 6000; sub 4; find*; h 8000; sub 5; find*, bs). All multistate characters were treated as non-additive (Fitch, 1971). Successive character weighting based on the CI (Farris, 1969) was done with the macro SWT.RUN.

Anatomical abbreviations used in text and figures

AC, aciniform gland spigot(s); AG, aggregate gland spigot(s); AI, epigynum apical invaginations; ALE, anterior lateral eyes; ALS, anterior lateral spinnerets; AME, anterior median eyes; BH, basal haematodocha; C, conductor; CB, cymbium; CBP, cymbial basal process; CD, copulatory ducts; CMP, cymbial median process; CO, copulatory opening; CY, cylindrical gland spigot(s); E, embolus; EBA, embolic basal apophysis; F, fundus; FD, fertilization duct; FL, flagelliform gland spigot(s); MAP, major ampullate gland spigot; mAP, minor ampullate gland spigot; MP, epigynum median plate; P, paracymbium; PI, piriform gland spigot(s); PLE, posterior lateral eyes; PLS, posterior lateral spinnerets; PME, posterior median eyes; RPC, ‘reduced piriform clade’; S, spermatheca; ST, subtegulum; T, tegulum; TA, tibial apophysis.

Institutional abbreviations used in text

AMS, The Australian Museum, Sydney; BMNH, The Natural History Museum, London; CAS, California Academy of Sciences, San Francisco; EME, Essig Museum of Entomology, Berkeley; MACN-Ar, Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge; MNHN, Museum National d'Histoire Naturelle, Paris; NRS, Swedish Museum of Natural History, Stockholm; QMB, Queensland Museum, Brisbane; USNM, National Museum of Natural History, Smithsonian Institution, Washington, DC.

RESULTS

Phylogenetics

Heuristic searches under equal weights in TNT and NONA resulted in 10–42 cladograms of length 322 (CI = 45, RI = 75), depending on how zero-length branches were collapsed. The strict consensus for all trees collapsed five nodes and was 325 steps long (Figs 1, 2). Zero-length branches lack support and are problematic (Platnick, Griswold & Coddington, 1991; Coddington & Scharff, 1994), and were filtered using TNT's ‘rule 4’. These branches are produced because for the tree length calculations only cladograms with dichotomous resolutions are considered. Therefore, any taxa permutation that does not add extra steps is stored into memory regardless of node support. Then a set of algorithms is applied to filter the cladograms that contain zero-length branches.

The ten most parsimonious cladograms obtained with ‘rule 4’ resulted from a combination of character conflict and missing data on different parts of the tree (Kearney, 2002). Character conflict affects the phylogenetic relationships of: Tetragnathinae as sister to ‘leucaugines’ (26-1) or ‘metines s.s.’ (64–2); ‘metines s.s.’ sister to ‘leucaugines’ (104–1); Tetragnatha and Agriognatha O. P.-Cambridge, 1896 as sister taxa (2-1); and Agriognatha as sister to all other Tetragnathinae except Diphya Nicolet, 1849 (53-1). The different resolutions for the three new species of Chrysometa are produced by missing data because two of these species are known only by one sex (Fig. 2).

Successive weighting analyses yielded three cladograms of 14 538 weighted steps equivalent to 323 steps under equal weights. They resolved ‘leucaugines’ as sister to Tetragnathinae and Agriognatha as sister to all other Tetragnathinae except Diphya, but did not resolve the Chrysometa species trichotomy. The placement of Dolichognatha, Mollemeta, Meta and Metellina Chamberlin & Ivie, 1941 differed from the equally weighted trees by placing Mollemeta as sister to Meta and Metellina (Fig. 2), but at the cost of an extra step if judged under equal weights.

Bremer support and Jackknife values collapsed all clades in trees of 328 or more steps (numbers below each clade, Fig. 2). Overall clade support was low: of the 42 nodes common to all cladograms, 22 have Jackknife values below 51% and Bremer support values of two or fewer steps. Particularly weak clades are Diphya plus Tetragnathinae, Azilia plus ‘leucaugines’, ‘metines s.s.’ (due to Mollemeta, lines marked with *, Fig. 2) and Metabus(2/53). Clades with relatively strong support were: Gasteracant Sundevall, 1833 plus Micrathena Sundevall, 1833 (4/97), Tetragnathinae excluding Diphya(6/96), Allende(4/96) and ‘leucaugines’ excluding Azilia(4/79).

DISCUSSION

Tetragnathid phylogenetics

Tetragnathidae, Nephilinae and Tetragnathinae were monophyletic. ‘Metines’ (SensuLevi, 1980, 1986; Heimer & Nentwig, 1982; Coddington, 1990; Hormiga et al, 1995) were rearranged into two weakly supported clades, here termed ‘leucaugines’ and ‘metines s.s.’. In this paper, no taxonomic decisions are taken regarding these two clades or the placement of Diphya as a Tetragnathinae because the phylogenetic relationships of these taxa are unresolved, or weakly supported (Fig. 2). Regarding the phylogenetic composition of ‘metines’SensuHormiga et al (1995), the genera Azilia and Dolichognatha are not sister taxa, and they group with ‘leucaugines’ and ‘metines s.s.’, respectively; both taxa represented a third linage of ‘metines’ in that study. This study corroborates the placement of Nephilinae and the overall taxonomic composition of Tetragnathinae found in previous studies (Coddington, 1990; Hormiga et al, 1995; Griswold et al, 1998).

Araneoidea and Tetragnathidae

In this analysis Tetragnathidae is sister to the ‘reduced piriform clade’ (RPC) SensuGriswold et al (1998). Tetragnathidae plus the RPC are sister to Araneidae and all these taxa conform to Araneoidea. The phylogenetic relationships within the RPC are beyond the scope of this paper and were studied in some detail by Griswold et al (1998). Two synapomorphies support the monophyly of the RPC: short bases of the piriform spigots (7-1) and the three-dimensional web structure (105-1). Tetragnathidae and the spiders of the RPC share three synapomorphies: the loss of the aciniform anterior brush of the PMS (9-1), the non-sticky silk spiral and the first sticky silk in contact (102-1), and the use of the iL1 (first leg facing the hub) to measure where to attach the sticky silk spiral (103-1). Two synapomorphies support the monophyly of Araneidae: the grooved booklung cover cuticle (3-1), and the presence of a radix in the embolic division (69-1). The booklung cover cuticle is also grooved in Nephilinae, except Clitaetra Simon, 1889, and in Meta within the ‘metines s.s.’ clade. The phylogenetic relationships of Araneidae have been studied in detail elsewhere (Scharff & Coddington, 1997).

Tetragnathidae

Supporting the monophyly of this family are the loss of the median apophysis (61-1), the conductor wrapping and coiling with the embolus (62-1), the conductor origin on the ventral margin of the tegulum, found in all nephilines, except Phonognatha Simon, 1894 and ‘leucaugines’ (63-1), and the flattened epigynal plate (90-4). The first two synapomorphies were reported and tested in previous studies (Coddington, 1990; Hormiga et al, 1995; Scharff & Coddington, 1997; Griswold et al, 1998), and the last two synapomorphies are new for the family.

Nephilinae

In the present analysis, Nephilinae are represented by the same five genera as in Hormiga et al (1995): Phonognatha, Clitaetra, Nephiline Leach, 1815, Herennia Thorell, 1887 and Nephilengys L. Koch, 1872. These taxa remain monophyletic and sister to the remaining tetragnathids. They are coded as having a conductor and embolus homologous to that of the other tetragnathids, a filiform embolus and the conductor–tegulum membranous attachment. The two last character states are also found among ‘leucaugines’. The monophyly of Nephilinae is supported by six unambiguous synapomorphies, one of them, long and flexible embolus, is new for the subfamily (73-1). Nephilinae spiders have sclerotized spermathecae, with short and relatively straight copulatory and fertilization ducts. The epigynal plate is flat or if protruding no more that one-quarter of its length. The genital opening is located inside a wide atrium (Nephiline) or sometimes separated by a septum (Nephilengys) (Kuntner, 2005). Nephilinae is sister to a clade that includes Tetragnathinae plus ‘leucaugines’ and ‘metines s.s.’. The monophyly of Tetragnathinae, ‘leucaugines’ and ‘metines s.s.’ is supported by the following two synapomorphies, no cheliceral denticles (18-0), and web posture with the legs I and II extended (93-0).

Tetragnathinae

Tetragnathinae are represented by Agriognatha, Tetragnatha, Pachygnatha Sundevall, 1823 and Glenognatha Simon, 1887. The phylogenetic relationships of Agriognatha are either as sister to Tetragnatha or to all other Tetragnathinae. Eight synapomorphies support the monophyly of Agriognatha plus the remaining tetragnathines: male chelicerae enlarged and curving distally more that two times the clypeus width (15-1), the presence of dorsal femoral trichobothria (27-0), enlarged sperm duct (58-1), articulated paracymbium (50-3), loss of epigynum and fertilization ducts (79-1 and 80-1), copulatory openings immersed in a membranous sac (81-2) and epigynal opening spiracle-shaped (90-1). Two synapomorphies group Diphya with Tetragnathinae: the cymbium apical margin constricted (38-1) and the paracymbium with a distal sclerotized triangular projection (51-1).

‘Leucaugines’

The clade informally labelled here as ‘leucaugines’ includes the tetragnathid genera Azilia, Orsinome, Tylorida, Leucauge, Metabus, Mesi Kulczynski, 1911 and Opadometa Archer, 1951. ‘Leucaugines’ are most diverse in the tropics; common species include Leucauge venusta(Walckenaer, 1842) in the Neotropics and Leucauge granulata(Walckenaer, 1842) from the Old World tropics. These two species were illustrated elsewhere (Levi, 1980: figs 44–59; Davies, 1988: fig. 14) and specimens are abundant in museum collections (e.g. AMNH, QMB and AMS). Four synapomorphies support the monophyly of this clade: spermathecae walls weakly sclerotized (55-1) and ventrally enlarged (57-1), sperm duct with a switchback (59-1), and tegulum ectally displaced (85-1). The nodes within this clade have low support values with the exception of the clade formed by Orsinome plus the remaining distal ‘leucaugines’ (4/79) (Fig. 2). A particularly weak node is the phylogenetic placement of Azilia; this taxon can easily behave as a wild card because of a combination of inapplicable characters plus homoplasy. Azilia species lack an important character complex of tetragnathid spiders due to the absence of a conductor, creating instability for the placement of this genus. Characters such as the weakly sclerotized spermathecae (85-1), the ventral enlargement of the tegulum (57-1) and the coils of the sperm duct (59-1) place Azilia as a basal ‘leucaugines’.

‘Metiness.s.’

The overall morphology of ‘metines s.s.’ is similar to that of species such as Meta menardi(Latreille, 1804) or its American relative Meta ovalis(Gertsch, 1933). This clade includes the genera Nanometa Simon, 1908, ‘Metine from Australia’, Mollemeta, Meta, Dolichognatha, Metellina, Allende and Chrysometa, plus the species Orsinome sarasini. Three unambiguous synapomorphies support the monophyly of these taxa: the cymbial median process (46-1), the cylindrical shape of the paracymbium (53-1), and the conductor originating dorsally and running along the tegular margin (63-3). The spermathecae surface is sclerotized and the epigynal plate is roughly flat (but protruding in Meta ovalis, Orsinome sarasini and Allende species). The low support values in most of the nodes within ‘metines s.s.’ are due to character conflict between Mollemeta, Meta, Metellina and Dolichognatha in cladograms one step longer. A reasonably well-supported clade within ‘metines s.s.’ is the one that unites Nanometa, Orsinome sarasini and the undescribed species from Australia (3/63) (Fig. 2). Three synapomorphies support the monophyly of this clade: cymbial basal process with several spines of variable length (44-4), the presence of subdivided spermathecae in two sacs (84-1), one of them membranous and enclosing the copulatory duct (87-1). The genera Nanometa and the species Orsinome lagenifera(Urquhart, 1888) have been referred elsewhere to the Nanometatidae (Forster & Forster, 1999: 167). The undescribed species from Australia is similar to a species illustrated as Metinae sp. (Davies, 1988: 294, fig. 16). Three synapomorphies support the monophyly of Chrysometa, all of them from the female reproductive system: long fertilization ducts (82-1) with a straight path (83-1), originating anteriorly (91-1), and coiling around the spermathecae (92-1). The taxonomy of the genus Chrysometa was revised by Levi (1986).

Tetragnathid genital anatomy

The following discussion of tetragnathid genital anatomy complements the character descriptions given in Appendix 1. These descriptions incorporate information that is not included in this phylogenetic analysis but that was important to establish primary homology hypotheses for these characters.

Male pedipalp

The tetragnathid palpal bulb is characterized by a globular tegulum, an embolus and a conductor for which morphology varies considerably within the family. Tetragnathids lack a median apophysis. In all tetragnathids the conductor encloses almost all of the embolus, coils with it and forms the conductor–embolus complex (Levi, 1980, 1981, 1986; Levi & Eickstedt, 1989; Coddington, 1990; Hormiga et al, 1995; Scharff & Coddington, 1997; Griswold et al, 1998; Tanikawa, 2001). This complex interacts directly with the epigynum during copulation. Teragnathines insert both sclerites into the epigynum (Levi, 1981: 274, pl. 4), perhaps to manipulate sperm, as do some pholcids (Huber, 1999). Eberhard & Huber (1998: 344) documented that the conductor interacted with the epigynal plate in Leucauge mariana(Taczanowski, 1881). Finally, broken parts of the embolus and conductor were found inside the copulatory openings of some Nephilinae (Hormiga et al, 1995: fig. 11G–I). These interactions between the conductor and the epigynum have not been studied for ‘metines s.s.’.

The attachment of the tetragnathid conductor to the tegulum can be sclerotized or membranous. Membranous attachments occur in Nephilinae (Hormiga et al, 1995: fig. 9C) and almost all ‘leucaugines’, and allow the conductor to move during copulation while interacting with the epigynum. The conductor membrane in nephiline and ‘leucaugine’ originates from the tegulum adjacent to the embolic membrane. This membrane proximity has been used to interpret the conductor of nephilines as part of the embolic division rather than a tegular sclerite (M. Kuntner, pers. comm.). Here the nephiline and ‘leucaugine’ conductor are considered homologous to the conductor of other araneoid spiders because they originate from the tegulum. Additional evidence for this homology hypothesis is the similarity in shape of the conductor of these taxa with other tetragnathids. A sclerotized conductor attachment characterizes Tetragnathinae and ‘metines s.s.’ (Fig. 7F).

Figure 9

Spinnerets. A-E, Metabus ocellatus. A, female ALS. B, female PMS. C, female PLS. D, male PLS. G, epiandrous fusules. F-H, Allende longipes. F, female ALS. G, female PMS. H, female PLS. I-J, A. nigrohumeralis. I, male PLS. J, epiandrous fusules. K-O, Mollemeta edwardsi. K, female ALS. L, female PMS. M, female PLS. N, male PLS. O, epiandrous fusules.

The shape and origin of the conductor varies widely across Tetragnathidae. In ‘metines s.s.’ the conductor is a lamellar apophysis that originates dorsally on the tegulum, follows an ecto-ventral path, and partially covers the embolus (Fig. 5D). In tetragnathines the conductor consists of two lamellae that enclose and coil with the embolus, originates centrally on the tegulum and projects apically (Levi, 1981: fig. 19). In Nephila clavipes(Linnaeus, 1767) the conductor originates on the meso-apical tegular margin and projects ventrally (Levi, 1980: figs 21–26). In ‘leucaugines’ the conductor originates ventrally on the tegulum and projects apically. Its tegular dorsal margin is occupied almost completely by the embolic base (Fig. 3D). The conductor apex of almost all tetragnathids varies from hook shaped in Allende species (Fig. 5D–E) and Tetragnatha (Levi, 1981: figs 5–7) to coiled in Nephilengys cruentata(Fabricius, 1775) (Hormiga et al, 1995: fig. 12). Autapomorphic conductor morphologies are found in Mollemeta (Figs 7A, B, D) and Diphya (Tanikawa, 1995: figs 23–25). Azilia is the only tetragnathid known to lack a conductor (Levi, 1980: figs 302–308).

Figure 19

Allende nigrohumeralis. A, female habitus. B, female frontal view. C–G, epigynum. C, caudal view. D, lateral view. E, ventral view. F, caudal view cleared. G, dorsal view cleared. H, male habitus. I, male frontal view. J–P, male pedipalp. J, expanded. K, ventral view. L, dorsal view. M, expanded. N, conductor detail. P, ectal view. Scale bars 0.2 mm, except in habitus 1 mm.

Figure 12

Metabus ocellatus. A, female habitus. B, female frontal view. C, epigynum ventral view. D, caudal view. E, central view cleared. F, dorsal view cleared. G, male habitus. H, male frontal view. I–L, male pedipalp. I, ectal view. J, ventral view. K, mesal view. L, cleared L. Scale bars 0.2 mm, except in habitus 1 mm.

In Tetragnathidae the embolus attaches to the tegulum by means of a membrane, which usually originates apically at the centre of the tegulum. Filiform emboli are found in many ‘leucaugines’ (Fig. 3D) and Nephilinae (Levi, 1980: fig. 25; Hormiga et al, 1995: fig. 10C). The embolic base of ‘leucaugines’ is smooth and rectangular, while in Nephilinae is roughly square and may have tubercles (Hormiga et al, 1995: fig. 12C). Short and rigid emboli are found in Allende (Fig. 5D) and other ‘metines s.s.’ such as Meta, Dolichognatha and Metellina (Hormiga et al, 1995: fig. 13B-F). The embolic base in these taxa often has apophyses (Figs 5D, F, 7F). Long, rigid emboli without apophyses are common in Nanometa, Mollemeta (Fig. 7D) and Tetragnathinae (Levi, 1981: fig. 19) species. Autapomorphic embolic morphologies are found in Azilia (Levi, 1980: figs 302–308) and Diphya (Tanikawa, 1995: fig. 25). The connection of the embolic apophyses to the embolus base can be sclerotized (Metellina and Dolichognatha) or membranous (Allende; Fig. 17A). In many Chrysometa species the embolic apophysis is completely membranous and coils with the embolus (Fig. 5F).

Figure 13

Metabus debilisis. A, female habitus. B, epigynum ventral view. C, caudal view. D, ventral view cleared. E, dorsal view cleared. F-G, male pedipalp. F, ectal view. G, ventral view. H, mesal view. Scale bars 0.2 mm, except in habitus 1 mm.

Figure 17

Allende patagiatus. A, female habitus. B, female frontal view. C–G, epigynum. C, ventral view. D, caudal view. E, lateral view. F, caudal view cleared. G, dorsal view cleared. H, male habitus. I, male frontal view. J–M, male pedipalp. J, ectal view. K, conductor detail. L, dorsal view. M, ventral view. Scale bars 0.2 mm, except in habitus 1 mm.

The tetragnathid tegulum is roughly globular, with the exception of Mollemeta and Phonognatha in which the tegulum is reduced to a ring (Fig. 7A, B). The tegulum is apically orientated in most tetragnathids with the exception of some nephilines. Nephiline spiders (except Phonognatha) have the tegulum ectally displaced (Hormiga et al, 1995: figs 8–11). In Tetragnathinae and ‘metines s.s.’ the tegulum is usually wider than long (Figs 5A, F, 7F) whereas the tegulum in most ‘leucaugines’ is longer than wide and ventrally protruding (Fig. 3A). The subtegulum is usually shell-shaped and located under the tegulum and it is mesally displaced in most ‘leucaugines’ (Fig. 3E–F). The sperm duct can be enlarged and spiral like in most Tetragnathinae (Levi, 1980: fig. 177) or thin with several twists and turns as in Nephilinae and ‘leucaugines’ (Fig. 12L). In ‘metines s.s.’, the sperm duct is narrower and its path less intricate (Fig. 19O; Levi, 1980: figs 98, 125).

The cymbium is usually wide and spoon-shaped. In Diphya and some Tetragnathinae the cymbium is apically constricted (Hormiga et al, 1995: fig. 7). This constriction is restricted to the distal part of the cymbium in Diphya. The cymbium of Glenognatha and Pachygnatha is completely constricted. The cymbial ectal margin can be smooth or bearing two processes, one basal, the other median (Fig. 5B). The cymbial median process is found in ‘metines s.s.’ except in Metellina and Dolichognatha. The cymbial basal process is found in many tetragnathid genera except Nephilinae. These two processes vary considerably in morphology and orientation (Figs 3C, 5C, F, 7C, F; Hormiga et al, 1995: fig. 6A–C labelled as paracymbium secondary process or SP). The paracymbium also varies considerably in shape, and its attachment to the cymbium ranges from membranous (e.g. Tetragnatha) to well sclerotized (e.g. Meta).

Epigynum

The female reproductive system of tetragnathids can be either entelegyne or haplogyne. In those species without fertilization ducts the spermathecae open directly into a membranous chamber located ventrally to the uterus externus. This chamber leads to a wide spiracle-shaped opening displaced posteriorly to the epigastric furrow (Levi, 1980: figs 26, 27). In Tetragnatha, Pachygnatha and Glenognatha this chamber has two sclerotized structures, the spermathecae (Wiehle, 1967: figs 6, 7; Levi, 1980: figs 26, 169, 259; 1981: fig. 92). It was postulated that Meta species lack fertilization ducts and that they have a peculiar fertilization mechanism referred as ‘semi-entelegyne’. Wiehle (1967: 191–92) mentioned that semi-entelegyne spiders fertilize their eggs by pushing out the sperm through the copulatory opening, and then the sperm runs through a sclerotized ridge from the copulatory openings to the uterus externus opening. Levi (1980) has suggested that Meta and Metellina species do indeed have fertilization ducts and that fertilization takes place as in other entelegyne taxa. Detailed study shows that the fertilization ducts and copulatory ducts of Meta and Metellina are adjacent and originate at the posterior end of the spermathecae (Levi, 1980: 3). My observations of cleared epigyna suggest that at least Meta ovalis and Metellina curtisi(McCook, 1894) have flat and short L-shaped fertilization ducts that originate at the dorsal end of the spermathecae and project anteriorly.

Entelegyne tetragnathids have fertilization and copulatory ducts, and their spermathecae walls can be ‘hard’ or ‘soft’ depending on the degree of sclerotization. Hard spermathecae are common among Araneoidea; within Tetragnathidae they are found in ‘metines s.s.’ and Nephilinae. Tetragnathid hard spermathecae are usually associated with short and well-sclerotized copulatory and fertilization ducts (Figs 6D, F, 8D, F). The epigynal plate of tetragnathids with hard spermathecae can be flat as in Metellina (Levi, 1980: figs 102–110), protruding as in Meta and Allende (Fig. 6A, B) or divided by a median septum as in some Nephilinae. These spermathecae are roughly oval and covered with accessory glands grouped in clusters (Figs 6E, 8E, F). Soft spermathecae are found in all ‘leucaugines’ (Fig. 4C–D, F). The epigynal plate of ‘leucaugines’ is usually a flat sclerotized plate with a shallow median depression of variable shape (Fig. 4A, B). The copulatory openings can be located along the sides of this median depression, or as two visible openings on its surface as in some Leucauge species. The fertilization ducts in ‘leucaugines’ are usually well sclerotized, long and coiled; the copulatory ducts are usually straight and weakly sclerotized (Figs 4D, F). The spermatheca surface of these genera is covered with isolated accessory glands (Fig. 4E). Exceptions to this basic pattern are Azilia, whose fertilization ducts are straight, its accessory glands are arranged in clusters and the epigynal plate slightly protruding; and Leucauge argy(Walckenaer, 1842) with an protruding cone-shaped epigynal plate Levi (1980: figs 60, 63, 290, 293).

SYSTEMATICS

Tetragnathidae menge, 1866 Metabus O. P.–Cambridge, 1899 (Figs3A–E, 4A–E, 9A–E, 10D, 11A, 12–15)

Figure 11

Distribution maps: A, Metabus species, Central and northern South America. B, Allende species, Chile. C, Chrysometa species, Chile. D, Mollemeta edwardsi, Chile.

Figure 14

Metabus ebanoverde. A, female habitus. B, epigynum ventral view. C, caudal view. D, ventral view cleared. E, dorsal view cleared. F–G, male pedipalp. F, ectal view. G, ventral view. H, mesal view. Scale bars 0.2 mm, except in habitus 1 mm.

Figure 15

Metabus conacyt. A, female habitus. C, epigynum ventral view. C, caudal view. D, ventral view cleared. E, dorsal view cleared. F–G, male pedipalp. F, ectal view. G, ventral view. H, mesal view. Scale bars 0.2 mm, except in habitus 1 mm.

Metargy F. O. P.-Cambridge, 1903: 444, pl. 42, f. 15-16. New synonymy.

Type species:

Metabus gravidus O. P.–Cambridge, 1899, type fixed by monotypy.

Etymology:

O. P.–Cambridge (1899: 298) did not explain the etymology of Metabus, this genus name means resembling or having the quality of Meta.

Diagnosis:

Metabus species can be distinguished from all other tetragnathid genera by the following combination of characters: epigynum flat, well sclerotized and with a rectangular atrium (Figs 12C, 13B, 15B); male pedipalp conductor longer than wide, and apically projected (Figs 12J, 13–15G); femur IV without trichobothria; and leg I more than four times the body length.

Description

Female:

Medium- to small-sized spiders, total length from 4.7 to 8.1. The smallest specimens belong to M. debilis and the largest to M. ocellatus. Carapace usually light in colour, cephalic region slightly elevated and lighter than thoracic region, except in M. ocellatus, in which the cephalic region is darker. Thoracic region margins delineated in dark brown, fovea shallow and curved. Sternum triangular-shaped, dark in colouration and projecting between coxae IV. AME and PME subequal in size, larger than the lateral eyes. Lateral eyes juxtaposed and on a tubercle. Clypeus height 1.0–1.4 × AME diameter. Secondary eyes with canoe-shaped tapetum. Anterior surface of chelicerae smooth (Fig. 12B), three prolateral and four retrolateral teeth. Labium rebordered, wider than long. Endites rebordered, longer than wide. Leg formula I > II > IV > III, femora without trichobothria, legs I and II considerably longer than the others (2.30 and 2.06 times body length, respectively). Booklung cover smooth. Abdomen oval with silver guanine patches. Spinnerets of M. ocellatus (Fig. 9A–C), ALS with c. 90 piriform spigots. PMS with three aciniform spigots between the cylindrical and the minor ampullate spigots, mAP nubbin present. PLS aciniform field with c. 25 spigots, aggregate spigots embracing the flagelliform, cylindrical spigots peripheral. Female copulatory openings inside grooves that run along the sides of the epigynal atrium (Fig. 4A, B). Copulatory ducts long, weakly sclerotized and entering the spermathecae near the fertilization duct origin region (Fig. 4C, D). Spermathecae also weakly sclerotized, and longer than wide. Fertilization ducts well sclerotized, long and coiled (Fig. 12E, F). Accessory gland openings evenly distributed on the spermathecae surface (Fig. 4E).

Male:

Total length from 2.2 to 7.3. The smallest specimens belong to M. debilis and the largest to M. ocellatus. Abdomen oval, somatic morphology as in female (Fig. 12G). Anterior surface of chelicerae smooth (Fig. 12H). One line of epiandrous fusules anterior to a sclerotized plate, posterior plate margin enlarged (Fig. 9E). Pedipalp patella with one macroseta. Palpal tibia 2.8–3.2 times longer than wide with two mesal rows of trichobothria. Cymbial basal process absent (except in M. ocellatus), distal cymbial macrosetae enlarged. Tegulum oval, longer than wide and ventrally protruding (Fig. 3A). Spermatic ducts coiled several times (Fig. 12L). Subtegulum displaced laterally (Fig. 3E). Embolus base longer than wide, embolus filiform. Conductor longer than wide, originating ventrally (Fig. 3A), attachment with the tegulum membranous.

Natural history:

M. ocellatus builds individually differentiated webs on a communal framework attached to various vegetation points over running water. These spiders share a communal retreat during the night, emerging at day to build individual webs. Prey is not shared among colony members and agonistic behaviour is displayed to defend their webs from other individuals (Buskirk, 1975a, b; Uetz & Craig, 1997). M. ebanover spiders span their webs on open spaces and across small rivers. The largest orb found was 92 cm in diameter (Fig. 10D), the support threads were heavily reinforced and longer than the orb diameter. Some spiders were found under dry leaf clusters with the legs I and II extended and in contact with the support threads. Several individuals were collected above the branches that connected the solitary webs as in some Tetragnatha species, suggesting a possible communal behaviour. The web of M. ebanover has c. 12 radii and a relatively dense sticky spiral (Fig. 10D). The shape of the web is roughly polygonal resembling that of Tetragnatha species.

Composition:

Four species: Metabus ocellatus (Keyserling, 1864), M. debilis(O. P.-Cambridge, 1889), M. ebanover and M. conacyt.

Phylogenetics:

Two unambiguous synapomorphies support the monophyly of Metabus: conductor apex curved apically (66-1) (Fig. 3A), and absence of trichobothria on the fourth femora (27-1). A cladistic analysis that includes Okileucauge species is recommended to explore Metabus generic limits further, and to test hypotheses about the evolution of the fourth femoral trichobothria complex within ‘leucaugines’. Both taxa have similar genitalia and lack trichobothria on the ectal surface of the femora IV (Tanikawa, 2001; Zhu et al, 2003).

Distribution:

Neotropical, from southern Mexico to French Guiana and Dominican Republic (Fig. 11A).

Nomina dubia:

Metabus fuegianus (Simon, 1902: 26) is proposed as a nomen dubium because the holotype (Meta fugianaSimon, 1902) is an immature specimen.

Metabus ocellatus (Keyserling, 1864) comb. nov. (Figs3A–E, 4A–E, 9A–E, 11A, 12)

Tetragnatha ocellataKeyserling, 1864: 149 pl. 7 f. 7–9 (syntypes of in BMNH, examined).

Argyroepeira ocellaKeyserling, 1893: 336 pl. 17 f. 248.

Metabus gravidus O. P.–Cambridge, 1899: 299 pl. 37 f. 7 (female holotype in BMNH, examined). -Archer 1963: 17.–Nentwig, 1993: 122 f. 141a. New synonymy.

Meta gravida(O. P.–Cambridge, 1899): F. O. P.-Cambridge, 1903: 446.–Petrunkevitch, 1925: 109 f. 19–21 (transferred back to Metabus by Archer, 1963).

Leucauge ocellata (Keyserling, 1864): Petrunkevitch, 1911: 356.

Diagnosis:

M. ocellatus can be distinguished from other Metabus species by the following combination of characters: body length between 6.5 and 8.1 mm, cephalothorax margins rebordered with thick dark lines (Fig. 12A, G); epigynum atrium longer than wide, anterior margin slightly curved (Fig. 12C, D); cymbial basal process spine-shaped (Fig. 12I), conductor slightly curved and with a single distal apophysis (Fig. 12I–K, arrow).

Description:

Female from Prusia, Chiapasas, Mexico (FAP0093). Habitus as in Figure 12A. Total length 8.1. Cephalothorax 3.1 long, 2.5 wide, 0.8 high. Sternum 1.1 long, 1.3 wide. Chelicerae surface smooth and same colour as the cephalothorax (Fig. 12B). Abdomen oval, pattern as in Figure 12A. 6.5 long, 4.4 wide, 4.7 high. AME diameter 0.20, ALE 0.20, PME 0.15, PLE 0.15. Clypeus height 1.8 × AME diameter. AME separation 0.8 × AME diameter, AME–ALE separation 1.1 × AME diameter, PME separation 1.3 × PME diameter, PME–PLE separation 1.4 × PME diameter. Leg I length 31.6, leg II 18.2, leg III 8.5, leg IV 12.9, pedipalp 3.9. Epigynum as in Figure 12C, D. Spermathecae anteriorly projected; fertilization ducts with many coils (Fig. 12E).

Male from Prusia, Chiapasas, Mexico (FAP0092). Habitus as in Figure 12G. Chelicerae anterior surface smooth (Fig. 12H). Total length 7.3. Cephalothorax 3.5 long, 2.9 wide, 0.8 high. Sternum 1.4 long, 1.4 wide. Abdomen oval 4.5 long, 2.1 wide, 2.3 high. AME diameter 0.22. ALE 0.18. PME 0.18. PLE 0.16. Clypeus height 1.4 × AME diameter. AME separation 0.6 × AME diameter. AME–ALE separation 1.1 × AME diameter. PME separation 1.5 × PME diameter. PME–PLE separation 1.7 × PME diameter. Leg I length 34.0, leg II 18.1, leg III 7.6, leg IV 12.5, pedipalp. 2.9. Pedipalp tibia 2.8 times longer than wide. Pedipalp as in Figure 12-L.

Variation:

Female total length 6.5–8.1, cephalothorax length 2.8–3.1. Male total length 5.2–7.3, cephalothorax length 25–3.5.

Distribution:

Southern Mexico to Colombia and French Guiana (Fig. 11A). This species can be found from sea level up to 2700 m elevation. Most of the specimens studied were collected at higher altitudes (1100-2700 m).

Material examined (N= 327): COLOMBIA:

Antioquía, La Estrella, Quebrada Grande, 2100 m, 6.iv.1974, P. A. Schneble leg. 1 male and 28 females (MCZ) FAP0013. Ca, Cauca, 1500 m, 5.iii.1974, 1 female (MCZ) FAP0019; Valle de Cali, 1500 m, iii.1973, 1 male and 1 female (MCZ) FAP0017; 1 female (MCZ) FAP0023; 1500 m, viii.1975, 1 female (MCZ) FAP0025; 1300 m, vi.1977, 1 female (MCZ) FAP0018; 3 mi. W. Villavicencio, ii–iii.1955, E. I. Schlinger & E. S. Ross leg. 3 males and 4 females (CAS) FAP0273; New Grenada Bogota, 2 males, 7 females (syntypes of Tetragnatha ocellata BMNH) FAP0272; Valle Saladito, 1700 m, 5.viii, 1 female (MCZ) FAP0027; 1–5.ix.1971, 1 male and 2 females (MCZ) FAP0020. Medellín, Medellín Valley, 1700−1900 m, 1973, A. B. Schneble leg. 5 females (MCZ) FAP0006; Abadia Envigado, 2700 m, 16.xii.1984, M. A. Serna leg. 1 female (MCZ) FAP0021; 1 female (MCZ) FAP0022; 1 female (MCZ) FAP0026; La Estrella, 1720 m, 28.ii.1959, 1 male and 1 female (AMNH) FAP0107; 5.vi.1973, P. A. Schneble leg. 1 female (MCZ) FAP0024. COSTA RICA: Costa Rica, 1983, W. E. Eberhard leg. 1 female (MCZ) FAP0014; Las Nubes de Coronado, 20.vi.1992, M. Blanco leg. 1 female (MCZ) FAP0015. Puntarenas, Monteverde, 1370 m, 17.i.1961, C. W. Palmer leg. 3 females (AMNH) FAP0104; 11.vi.1978, R. Buskirk leg. 3 immatures (MCZ) FAP0030; Río Guacimal, 1550−1850 m, 30.xii.1971, W. & R. Burskirk leg. 2 males, 1 female and 4 immatures (AMNH) FAP0106; 12.i.1998, F. P. Gillette leg. 1 male, 2 females and 2 immatures (MCZ) FAP0001; 1 male, 3 females and 1 immature (MCZ) FAP0002; 1 female (MCZ) FAP0003; 1 male and 1 female (MCZ) FAP0004; 1 female (MCZ) FAP0005; 1 male, 1 female and 1 immature (MCZ) FAP0007; 1 male and 1 female (MCZ) FAP0008 smallest male measured; 1 male and 2 females (MCZ) FAP0009; 2 females (MCZ) FAP0010; 2 females (MCZ) FAP0011; 2 males, 2 females and 1 immature (MCZ) FAP0012; 4 males and 4 females (MCZ) FAP0031; 1 male and 1 female (MCZ) FAP0032; Bosque de la Hoya, 20.vi.1992, M. Blanco leg. 1 female (MCZ) FAP0016. FRENCH GUIANA: St Laurent, 27.iv.1994, 1 female (AMNH) FAP0105. GUATEMALA: Guatemala, 1 female (holotype of Metabus gravidus BMNH) FAP0271. Puntarenasaz, Purulha, 2300 m, 22–24.v.1979, J. A. Coddington leg. 1 female (MCZ) FAP0028 smallest female measured. San Pedro Yepocapa, Yepocapa, iii–iv.1945, H. Elishewitz leg. 1 male and 7 females (AMNH) FAP0101; 7 females and 1 immature (AMNH) FAP0102. MEXICO: Chiapas, Prusia, 1000 m, iv–v.1942, H. Wagner leg. 32 females (AMNH) FAP0091; 17 males and 14 females (AMNH) FAP0092 largest male measured and illustrated, one male specimen used for SEM (SEMFAP036); 14 males, 12 females and 4 immatures (AMNH) FAP0093 largest female measured and illustrated, one female specimen used for SEM (SEMFAP036); Huixtla, Finca Lubeca, 11.i.1945, T. C. Schneirla leg. 2 females (AMNH) FAP0096; 1 male, 4 females and 2 immatures (AMNH) FAP0097; 1 male, 5 females and 8 immatures (AMNH) FAP0098; 5 females (AMNH) FAP0100; La Victoria, 18.i.1945, C. Vargas leg. 5 females (AMNH) FAP0099; Mapastepec, vi–vii.1940, H. Wagner leg. 3 males, 16 females and 1 immature (AMNH) FAP0094; 5 males, 24 females and 1 immature (AMNH) FAP0095. PANAMA: Canal Zone, Soberania NP, 7.viii.1983, R. Raven leg. 1 female (AMNH) FAP0103; Chiriqui, La Fortuna, 1100−1200 m, 5.iv.1984, W. E. Eberhard leg. 1 female (MCZ) FAP0029.

Metabus debilisis (O. P.-Cambridge, 1899) comb. nov. (Figs11A, 13)

Argyroepeira debilis O. P.-Cambridge, 1889: 5, pl. 1, f. 9 (syntypes in BMNH, examined).–Keyserling, 1893: 365, pl. 19, f. 272. - O. P.-Cambridge, 1896: 218, pl. 28, f. 4–5.

Metargyra debilis(O. P.-Cambridge, 1889): F. O. P.-Cambridge, 1903: 444, pl. 42, f. 15–16.

Diagnosis:

This species can be distinguished from the other small Metabus species by the following characters: epigynum atrium wider than long, anterior margin straight; and spermathecae projected laterally (Fig. 13B–D); conductor with two parallel distal apophyses (Fig. 13F, G arrows); and three trichobothria on the palpal tibia mesal surface (Fig. 13F–H).

Description:

Female from Puerto Napo, Ecuador (FAP0385). Habitus as in Figure 13A. Total length 3.9. Cephalothorax 1.5 long, 1.4 wide, 0.6 high. Sternum 0.8 long and wide. Abdomen shape oval 2.4 long, 2.3 wide, 2.4 high, venter with a rectangular patch of silvery guanine spots. AME diameter 0.12. ALE 0.10. PME 0.12. PLE 0.10. Clypeus height 1.2 × AME diameter. AME separation 0.8 × AME diameter. AME–ALE separation 0.6 × AME diameter. PME separation 1.4 × PME diameter. PME–PLE separation 1.4 × PME diameter. Leg I length 16.6 leg II 8.5, leg III 3.9, leg IV 5.5, pedipalp 1.7. Epigynum as in Figure 13B–E.

Male from Finca La Selva, Herediaia, Costa Rica (FAP0387). Total length 2.2. Cephalothorax 1.1 long, 0.1 wide, 0.4 high. Sternum 0.6 long and wide. Abdomen oval with a dorsal dark median longitudinal band, venter as in female. 1.6 long, 0.9 wide, 0.9 high. AME diameter 0.1. ALE 0.1. PME 0.1. PLE 0.1. Clypeus height 1.0 × AME diameter. AME separation 0.7 × AME diameter. AME–ALE separation 0.7 × AME diameter. PME separation 1.1 × PME diameter. PME–PLE separation 0.7 × PME diameter. Leg I length 12.8, leg II 6.7, leg III 3.1, leg IV 4.4, pedipalp 1.2. Pedipalp tibia 3.2 times longer than wide. Pedipalps as in Figure 13F, G.

Variation:

Female total length 3.5–3.9, cephalothorax length 1.5–1.6. Male total length 2.2–2.8, cephalothorax length 1.1–1.3.

Distribution:

Southern Mexico to Ecuador (Fig. 11A).

Material examined (N= 3):

COLOMBIA: 3 miles W Villavicencio, 11.iii.1955, E. I. Schlinger & E. S. Ross leg. 1 male (CAS) FAP0386 largest male measured. COSTA RICA: Heredia, Finca La Selva, 4 km SE Puerto Viejo de Sarapiquí, 15.ix.1981, C. Griswold leg. 1 male (CAS) FAP0387 smallest male measured and illustrated. ECUADOR: Na, Alinahui, 20 Km E Puerto Napo, 1°0′S, 77°25′W, 450 m, ii.1994, E. S. Ross leg. 1 female (CAS) FAP0385 measured and illustrated. MEXICO: Tabasco, Teapa, 2 males and 6 females (syntypes of Argyroepeira debilis BMNH) FAP0399; 2 females (BMNH) FAP0400.

Metabus ebanoverdesp. nov. (Figs10D, 11A, 14)

Types:

Holotype female from Ebano Verde Scientific Reserve, La Vega, Dominican Republic (deposited at MCZ FAP0397). Three male and four female paratypes from the type locality (deposited at MCZ FAP0382).

Etymology:

The specific epithet, a name in apposition, is taken from the Scientific Reserve Ebano Verde (La Vega) where the type series was collected.

Diagnosis:

This species can be distinguished by the following combination of characters: epigynum wider than long, anterior margin discontinuous with two lateral notches (Fig. 14B); conductor with two divergent distal apophyses (Fig. 14G, arrows); and six trichobothria on the palpal tibia mesal surface (Fig. 14F).

Description:

Female from La Vega, Dominican Republic (FAP0381). Habitus as in Figure 14A. Total length 5.0. Cephalothorax 1.8 long, 1.5 wide, 0.5 high. Sternum 0.8 long and wide. Abdomen oval, venter dark grey with a horizontal rectangle formed of guanine spots, 3.7 long, 3.1 wide, 3.1 high. Diameter of all eyes 0.1. Clypeus height 1.4 × AME diameter. AME separation 0.7 × AME diameter. AME–ALE separation 1.4 × AME diameter. PME separation 1.4 × PME diameter. PME–PLE separation 1.4 × PME diameter. Leg I length 21.8, leg II 10.2, leg III 4.7, leg IV 7.4, pedipalp 1.9. Epigynum as in Figure 14B–E.

Male from La Vega, Dominican Republic (FAP0381). Habitus as in female. Total length 3.7. Cephalothorax 1.9 long, 1.4 wide, 0.4 high. Sternum 0.80 long and wide. Abdomen oval venter as in female. 2.5 long, 1.7 wide, 1.7 high. AME diameter 0.11. ALE 0.09. PME 0.11. PLE 0.09. Clypeus height 1.2 × AME diameter. AME separation 0.8 × AME diameter. AME–ALE separation 0.8 × AME diameter. PME separation 1.2 × PME diameter. PME–PLE separation 1.0 × PME diameter. Leg I length 19.3, leg II 9.9, leg III 4.3, leg IV 6.8, pedipalp 1.5. Pedipalp tibia 3.1. times longer than wide. Pedipalp as in Figure 14F–H.

Variation:

Female total length 4.4–5.0, cephalothorax length 1.8-2.0. Male total length 3.1-3.7, cephalothorax length 1.5-1.9.

Distribution:

Dominican Republic and Guatemala (Fig. 11A).

Material examined (N= 15):

DOMINICAN REPUBLIC: Constanza, La Vega, Ebano Verde, Loma La Sal, 18°41′49.4″N, 70°35′23.7″W, 2274 m, 16.iv.2005, F. Álvarez & B. Suresh leg. 1 female (MCZ) FAP0397; 3 males and 1 immature (MCZ) FAP0398; 1 male and 1 female (MCZ) FAP0381 largest male and female measured, male and female illustrated; 3 males, 4 females and 3 immatures (MCZ) FAP0382 smallest male measured; 1 female (MCZ) FAP0383 smallest female measured. 1 female (MCZ) FAP0384. GUATEMALA: Alta Verapaz, Lanquin nr. Gruta, 5.ii.1980, B. & V. Roth leg. 1 female (AMNH) FAP0388.

Metabus conacytsp. nov. (Figs11A, 15)

Types:

Female holotype and one male paratype from Nebaj, Guatemala (AMNH, FAP0392).

Etymology:

A noun in apposition, after the Mexican government agency ‘Consejo Nacional de Ciencia y Tecnología’ (CONACYT).

Diagnosis:

This species can be distinguished from other Metabus species by the following combination of characters: epigynal atrium longer than wide, anterior margin continuous with two lobes under a cuticle flap (Fig. 15B); conductor tip without apophyses, and six trichobothria on the palpal tibia mesal surface (Fig. 15F–H).

Description:

Female, holotype (FAP0392). Habitus as in Figure 15A. Total length 4.9. Cephalothorax 2.1 long, 1.5 wide, 0.6 high. Sternum 0.8 long, 0.9 wide. Abdomen oval, venter of the abdomen with two parallel lines formed of guanine patches. 3.2 long, 2.7 wide, 2.7 high. AME diameter 0.13. ALE 0.1. PME 0.13. PLE 0.1. Clypeus height 1.0 × AME diameter. AME separation 0.7 × AME diameter. AME–ALE separation 0.7 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 1.0 × PME diameter. Leg I length 20.2, leg II 11.3, leg III 4.8, leg IV 6.8, pedipalp 1.9. Epigynum as in Figure 15B-E.

Male from San Luis Potosí, Tamazunchale, Mexico (FAP0390). Habitus as in female. Total length 3.2. Cephalothorax 1.5 long, 1.3 wide, 0.3 high. Sternum 0.8 long, 0.7 wide. Abdomen oval, venter as in female. 2.0 long, 1.1 wide, 1.0 high. AME diameter 0.13. ALE 0.10. PME 0.10. PLE 0.13. Clypeus height 1.0 × AME diameter. AME separation 0.7 × AME diameter. AME–ALE separation 0.7 × AME diameter. PME separation 1.4 × PME diameter. PME–PLE separation 1.0 × PME diameter. Leg I length 21.4, leg II 9.7, leg III 3.8, leg IV 6.2, pedipalp 1.5. Pedipalp tibia 2.9. times longer than wide. Pedipalp as in Figure 15F–H.

Variation:

Female total length 4.2–4.9, cephalothorax length 1.8–2.1. Male total length 3.1–3.2, cephalothorax length 1.4–1.5.

Distribution:

Central Mexico to Guatemala (Fig. 11A).

Material examined (N= 12):

GUATEMALA: Neb, 9.viii.1947, C. ae P. Aurie leg. 1 male and 5 females (AMNH) FAP0392 largest female measured and illustrated. MEXICO: Chiapas, 4 miles S. Simojovel, 18.iii.1953, E. I. Schlinger leg. 1 female (CAS) FAP0389 smallest female measured. San Luis Potos>, Tamazunchale, 20.v.1952, M. Crazier, W. Gertsch, & R. Schrammel leg. 1 male and 2 immatures (AMNH) FAP0390 largest male measured and illustrated. 21°15′N, 98°48′W, 19.iv.1963, W. J. Gertsch & J. W. Ivie leg. 1 male and 1 immature (AMNH) FAP0391 smallest male measured.

Allendegen. nov. (Figs5A–E, 6A–E, 9F–J, 10A, 11B, 16–19)

Figure 16

Allende puyehuensis. A, female habitus. B, female frontal view. C, first leg mesal view. D–H, epigynum. D, caudal view. E, ventral view. F, lateral view. G, caudal view cleared. H, dorsal view cleared. I, male habitus. J, male frontal view. K–N, male pedipalp. K, dorsal view. L, ectal view. M, conductor detail. N, ventral view. Scale bars 0.2 mm, except in habitus 1 mm.

Figure 18

Allende longipes. A, C–E, female habitus. B, female frontal view. F, first leg mesal view. G–K, epigynum. G, ventral view. H, vaudal view. I, lateral view. J, caudal view cleared. K, dorsal view cleared. L, male habitus. M, male frontal view. N–Q, male pedipalp. N, dorsal view. O, ventral view. P, ectal view. Q, conductor detail. Scale bars 0.2 mm, except in habitus 1 mm.

Type species:

Allende puyehuensis sp. nov.

Etymology:

Named after Salvador Allende (1908−73), president of Chile from 1971 to 1973. Allende is a name masculine in gender.

Diagnosis:

Allende species can be distinguished from all other tetragnathid genera by the following combination of characters: epigynum projected and with two apical invaginations (Figs 6B, 19C); cymbial base covered with small denticles, cymbial basal process long and slender, cymbial median process covered with enlarged macrosetae (Figs 17L, 18P, 19L); conductor tip with a distal hook and small basal lobes (Figs 16M, 17K, 18Q, 19N).

Description

Female:

Medium-sized tetragnathid spiders, total length from 5.58 to 13.33. The smallest specimens are those of A. puyehuensis and the largest are those of A. nigrohumeralis. Carapace usually light in colour with the cephalic region darker, thoracic region slightly elevated and with a shallow circular fovea. Sternum triangular-shaped, dark in colouration, decorated with irregular white marks and projecting between coxae IV. Secondary eyes with canoe-shaped tapetum. AME and PME subequal in size, larger than the lateral eyes. Lateral eyes juxtaposed and on a tubercle. Clypeus height 1.0–1.6 × AME diameter. Chelicerae with three prolateral and four retrolateral teeth. Labium rebordered, wider than long. Endites rebordered, longer than wide. Leg formula I > II > IV > III, femora without trichobothria. Booklung cover smooth. Spinnerets: ALS with c. 60 piriform spigots. PMS with three aciniform spigots between the cylindrical and the minor ampullate spigots, minor ampullate nubbin present. PLS with 13 aciniform spigots arranged roughly in two lines, cylindrical spigots peripheral, aggregate spigots embracing the flagelliform. Epigynum protruding and well sclerotized, copulatory openings in the shape of slits located sideways at the base of the median plate (Fig. 6B). Copulatory ducts short and well sclerotized. Fertilization ducts also short, arising behind the copulatory duct insertion area (Fig. 6C, D). Spermathecae surface rugose covered with small openings, accessory gland openings arranged in groups (Fig. 6E).

Male:

Total length from 4.6 to 9.2. Legs longer than the female and without modifications. Abdomen roughly oval shaped, pattern as in female. Epiandrous fusules arranged in clusters inside a depression of the epiandrous plate. PLS triplet reduced to nubbins, cylindrical spigots absent (Fig. 9I). Pedipalp patella with a single macroseta. Palpal tibia 2.4–4.6 times longer than wide, and with dorsal trichobothria at its base. Cymbium distal border sclerotized. Paracymbium cylindrical, longer than wide, with a distal sclerotized outgrowth, covered with macrosetae, attachment to cymbium membranous. Tegulum cylindrical, wider than long, sperm duct spiralled (Fig. 19J, O). Subtegulum displaced dorsally (Fig. 17M). Embolus enlarged basally and with a single apophysis, which is attached to the embolus by an S-shaped sclerotized ridge that inserts to its median portion, the rest of the attachment is membranous (Fig. 19M). Conductor well sclerotized, arising dorsally from the tegulum following its margin and gradually wrapping the embolus, attachment to the tegulum sclerotized (Fig. 19J).

Natural history:

Largely unknown. A. puyehuensis can be found near streams or in relatively open areas inside the forest. They build orb webs with open hubs, few radii and an open sector (Fig. 10A). The median plate of the epigynum was covered with mating plugs in all the individuals examined from all the species (Fig. 6A).

Composition:

Four species: A. longipes (Nicolet, 1849), A. nigrohumeralis(F. O. P.-Cambridge, 1889), A. patagiatus (Simon, 1901) and A. puyehuensis sp. nov.

Phylogenetics:

Six synapomorphies support the monophyly of Allende: PLS aggregate spigots distal part embracing the flagelliform spigot (12-1) (Fig. 9H); cymbial basal process apophysis covered with denticles (44-3) (Fig. 5C) [these denticles permit recognition of the apophysis of the cymbial basal process of Allende as such rather than as a paracymbial apophysis (Figs 17L, 18–19P)]; cymbial mesal process long (47-1) and covered with thicker macrosetae than those of the cymbium (48-2) (Fig. 5C); a distal hook with several lobes at the conductor tip (65-1); and the protruding epigynum plate (90-3). Allende is sister to Chrysometa; this clade is supported by three synapomorphies: rugose male cheliceral anterior cuticle (20-1), the paracymbium in distal position relative to the cymbial basal process (43-1) (Fig. 5C, F), and the articulated paracymbium (50-3). The species of these two genera differ in their spermathecae and epigynum morphology. Allende species have short fertilization ducts that originate posteriorly (Fig. 6C, D), a configuration characteristic of other genera of the ‘metines s.s.’, whereas C hrysome have long and coiled fertilization ducts (82-1, 83-1) that originate anteriorly from the spermatheca (91-1).

Distribution:

Chile: from the IV to the XII Regions, ‘Región Metropolitana de Santiago’, and Juan Fernández Islands. Argentina: Neuquen and Río Negro. All species have overlapping distributions (Fig. 11B).

Allende puyehuensissp. nov. (Figs5A–E, 10A, 11B, 16)

Types:

Female holotype from Aguas Calientes Puyehue National Park, Osorno, Chile deposited at USNM (FAP0219). One male (FAP0213) and one female (FAP0214) paratype from Puyehue, Los Mallines, deposited at USNM.

Etymology:

The species epithet is derived from the type locality.

Diagnosis:

This species can be distinguished from other Allende species by the following combination of characters: femora and tibiae covered with dark dots (Fig. 16C); epigynum longer than wide, dorsal plate folding over apical invaginations (Fig. 16D–F); cymbial median process base enlarged, cymbial basal process apophysis without denticles (Fig. 16L); and conductor tip with a long hook (Fig. 16M).

Description:

Female from Aguas Calientes, Puyehue, Chile (FAP0235). Habitus as in Figure 16A. Total length 6.8. Cephalothorax 2.7 long, 2.0 wide, 0.8 high. Sternum 1.1 long and wide. Chelicerae light in colouration and with a reticulated dark pattern on their bases (Fig. 16B). Abdomen 4.5 long, 2.7 wide, 2.5 high, brown in colouration, caudal region elevated, venter with a dark median mark delineated with white guanine patches. AME diameter 0.15. ALE 0.15. PME 0.18. PLE 0.13. Clypeus height 1.3 × AME diameter. AME separation 0.9 × AME diameter. AME–ALE separation 1.4 × AME diameter. PME separation 0.7 × PME diameter. PME–PLE separation 1.3 × PME diameter. Leg I length 14.9, leg II 10.0, leg III 5.5, leg IV 7.9, pedipalp 3.1. Epigynal median plate with two round protuberances (Fig. 16D, F), spermathecae oval-shaped, longer than wide (Fig. 16G, H).

Male from El Manzano, Región del Bio-Bio, Chile (FAP0228). Habitus as in Figure 16I. Total length 5.3. Cephalothorax 2.7 long, 1.9 wide, 0.9 high. Sternum 1.1 long and wide. Cheliceral cuticle rugose (Fig. 16J). Abdomen oval 3.1 long, 1.5 wide, 1.3 high. AME diameter 0.14. ALE 0.16. PME 0.16. PLE 0.13. Clypeus height 1.1 × AME diameter. AME separation 1.0 × AME diameter. AME–ALE separation 1.4 × AME diameter. PME separation 0.6 × PME diameter. PME–PLE separation 1.4 × PME diameter. Leg I length 21.2, leg II 12.3, leg III 6.0, leg IV 8.6, pedipalp 3.2. Pedipalp tibia 2.4 times longer than wide, embolic basal apophysis short, pedipalp as in Figure 16K–N.

Variation:

Female total length 5.6–7.6, cephalothorax length 2.5–2.9. Male total length 4.9–7.5, cephalothorax length 2.3–3.7.

Distribution:

A. puyehuensis is endemic to Chile (Fig. 11B). Its geographical range covers the following latitudes 32°31′0″S to 42°53′55″S, and altitudes from 50 to 1440 m.

Material examined (N= 82):

CHILE: Región IV del Coquim, Coquimbo, Pichidangui, 32°8′0″S, 71°32′0″W, 12.vii.1996, E. I. Schlinger leg. 1 female (CAS) FAP0220. Region del Maule V, Cauquenes, Reserva Nacional Los Ruiles, NW of Cauquenes, 35°50.028′S, 72°30.613′W, 146 m, 13.xi.2003, L. Prendini, C. I. Mattoni & J. A. Ochoa leg. 1 female (AMNH) FAP0393; Maule, Cauquenes, W of Maule, 350 m, 4.xii.1983, 2 males, 3 females and 11 immatures (AMNH) FAP0236 female leg IV and epigynum illustrated; Talca, 3 km W Vilches, 1070 m, 7.ii.1992. N. I. Platnick, P. A. Goloboff & M. J. Ramírez leg. 1 male (AMNH) FAP0231. Región V de Valparai, Aconcagua, Quebrada el Tigre, 32°31′0″S, 71°26′0″W, 12.viii.1966, E. I. Schlinger & M. Irwin leg. 6 females (CAS) FAP0221. Región VIII del Bio B, Arauco, Contulmo, Monumento Natural, 300 m, 11–12.ii.1992, N. I. Platnick, P. A. Goloboff & M. J. Ramírez leg. 1 male and 2 immatures (AMNH) FAP0233 largest male measured; Bio-Bio, El Manzano, 15.xii.1985, L. E. Peña leg. 1 female (AMNH) FAP0225; 8.xi.1992, T. Cekalovic leg. 1 male, 2 females and 1 immature (AMNH) FAP0228 male illustrated and measured; Concepción, Nonquen Estero, 26.x.1996, T. Cekalovic leg. 1 male, 1 female and 2 immatures (AMNH) FAP0229 largest female measured. Región X de los Lag, Chiloe Is., Pio Pio, 10–12.iii.1987, L. E. Peña leg. 1 female (AMNH) FAP0224 smallest female measured; Osorno, Maicolpue, hills S of 50 m, 19.ii.1992, N. I. Platnick, P. A. Goloboff & M. J. Ramírez leg. 1 male and 1 immature (AMNH) FAP0232; Puyehue NP, 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 female (USNM) FAP0218; Puyehue, Aguas Calientes, 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, G. Hormiga leg. 1 female (USNM) FAP0219; 40°44′0″S, 72°19′0″W, 1440 m, 6–7.xii.1988, V. & H. Roth leg. 1 female (CAS) FAP0222; 480 m, 17–18.ii.1992, N. I. Platnick, P. A. Goloboff & M. J. Ramírez leg. 2 males, 1 females and 5 immatures (AMNH) FAP0223; 40°44′0″S, 71°15′0″W, 550 m, 29.xi.1994, R. Leschen & C. Carlton leg. 2 females (AMNH) FAP0226; 450 m, 27.i.1986, N. I. Platnick & R. T. Schuh leg. 1 male and 5 immatures (AMNH) FAP0230; 500 m, 2–5.i.1982, L. E. Peña leg. 1 female and 2 immatures (AMNH) FAP0235 female measured and illustrated; 450 m, 27.i.1986, N. I. Platnick & R. T. Schuh leg. 1 female and 2 immatures (AMNH) FAP0239; 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 female (USNM) FAP0268; Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 female (USNM) FAP0215; 1 female (USNM) FAP0216; 1 male (USNM) FAP0217 one male specimen used for SEM; Puyehue, Anticura E of, 26.viii.5.xi.1983, L. E. Peña leg. 1 male (AMNH) FAP0238; Puyehue, Los Mallines, 40°46′0″S, 72°17′0″W, 700 m, 12.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 male (USNM) FAP0213 smallest male measured; 1 female (USNM) FAP0214 epigynum cleared illustrated; Repucura, Anticura, 500 m, 6.ii.1985, S. & J. Peck leg. 1 male and 1 immature (AMNH) FAP0227; Palaena, Chaiten, xii.1985, L. E. Peña leg. 1 female (AMNH) FAP0237; Valdivia, Las Trancas, W La Unión, 500 m, 6–10.ii.1988, L. E. Peña leg. 2 males and 4 immatures (AMNH) FAP0234.

Allende patagiatus (Simon, 1901) comb. nov.

Meta patagiaSimon, 1901: 18 (female holotype in MNHN, examined).

Meta echinaTullgren, 1902: 19, pl. 2, f. 1 (male holotype in NRS, examined). New synonymy.

Metabus echinatus (Tullgren, 1902): Archer, 1963: 17.

Diagnosis:

This species can be distinguished from other Allende species by the following combination of characters: epigynum wider than long, and small apical invaginations covered by the dorsal plate (Fig. 17C–E); cymbial median process straight and covered with short macrosetae (Fig. 17J), cymbial basal process apophyses with two apical denticles (Fig. 17L); and the shape of the conductor tip (Fig. 17K).

Description:

Female from Nahuelbuta, sendero Piedra del Aguila, Chile (FAP0243). Habitus as in Figure 17A. Total length 12.3. Cephalothorax 4.0 long, 3.3 wide, 1.1 high. Sternum 1.8 long, 1.7 wide. Chelicerae smooth and light in colouration (Fig. 17B). Abdomen oval 10.0 long, 5.8 wide, 6.3 high, with a caudal protuberance, venter with a median dark brown rectangular area delineated by two thin white lines. AME diameter 0.19. ALE 0.19. PME 0.20. PLE 0.15. Clypeus height 1.3 × AME diameter. AME separation 1.1 × AME diameter. AME–ALE separation 1.7 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 1.5 × PME diameter. Leg I length 19.3, leg II 17.3, leg III 8.7, leg IV 13.3, pedipalp 4.8. Epigynal median plate with two round protuberances (Fig. 17D, E), spermathecae oval-shaped longer than wide (Fig. 17F, G).

Male from Puyehue, Anticura, Chile (FAP0261). Habitus as in Figure 17H. Total length 7.8. Cephalothorax 3.3 long, 2.5 wide, 0.9 high. Sternum 1.6 long, 1.5 wide. Chelicerae pattern as in Figure 17I. Abdomen oval, venter as in female. 4.9 long, 3.9 wide, 2.5 high. AME diameter 0.17. ALE 0.14. PME 0.17. PLE 0.13. Clypeus height 1.1 × AME diameter. AME separation 0.9 × AME diameter. AME–ALE separation 1.4 × AME diameter. PME separation 0.9 × PME diameter. PME–PLE separation 1.4 × PME diameter. Leg I length 29.4, leg II 18.3, leg III 8.8, leg IV 12.7, pedipalp 6.1. Pedipalp tibia 3.1 times longer than wide. Embolic basal apophysis short, pedipalp as in Figure 17J–M.

Variation:

The colouration of the dorsal surface of the abdomen varies from almost white to dark brown, and no geographical pattern correlates with this colour variation. Female total length 7.1–12.7, cephalothorax length 2.9–4.0. Male total length 6.2–9.2, cephalothorax length 3.3–4.2.

Distribution:

A. patagiatus is found in Chile and Argentina (Fig. 11B). Its geographical range covers the following latitudes 31°50′0″S to 42°53′55″S, and altitudes from 40 to 1100 m.

Material examined (N= 83):

ARGENTINA: Río Neg, El Bolsón, 41°58′0″S, 71°31′0″W, 1.ii.1961, A. Kovacs leg. 1 female (AMNH) FAP0258. CHILE: Región IV del Coquim, Los Vilos, 21.x.1994, L. E. Peña leg. 1 male (AMNH) FAP0252 largest male measured. Región V de Valparai, Valparaiso, Cerro Molle, 1 female (holotype of Meta patagia MNHN) FAP0284. Región VII del Mau, Maule, Cauquenes W of Maule, 350 m, 4.x.1983, 1 male (AMNH) FAP0261 male illustrated and measured; 400 m, 24–27.i.1981, L. E. Peña leg. 1 female (AMNH) FAP0251; Tregualemu, 24.i.1976, G. Moreno leg. 1 female (AMNH) FAP0260; Talca, Bramadora, 600 m, 21–22.i.1993, L. E. Peña leg. 6 females and 2 immatures (AMNH) FAP0249. Región VIII del Bio B, Concepción, Nonquen Estero, 11.xii.1977, T. Cekalovic leg. 1 female (AMNH) FAP0259; Ñuble, Las Trancas, iii–iv.1979, L. E. Peña leg. 2 females and 5 immatures (AMNH) FAP0250. Región IX de la Araucar, Malleco, Curacautín, 16 km N of, 800 m, 15.ii.1992, N. I. Platnick, P. A. Goloboff & M. J. Ramírez leg. 1 male (AMNH) FAP0263; Nahuelbuta, sendero Piedra del Aguila, 37°49′45″S, 73°0′30″W, 1450 m, 4–9.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 6 females (USNM) FAP0243 largest female measured epigynum cleared illustrated. Región X de los Lag, Chiloe, Huequetrumao Is. de Chiloe, 27.xii.1981, L. E. Peña leg. 2 females (AMNH) FAP0255; Lago Coluco, 26.i.1981, L. E. Peña leg. 1 female (AMNH) FAP0256; Puente Murror, Ancud, 16.xii.1982, L. E. Peña leg. 6 males, 1 female and 7 immatures (AMNH) FAP0257; Llanquihue, Alerce Andino, Sector Sargazo, 41°30′30″S, 72°37′0″W, 350 m, 20–27.xii.2000, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 3 females and 3 immatures (USNM) FAP0245; Osorno, Pucatrihue coast, 10–24.ii.1985, L. E. Peña leg.1 female and 6 immatures (AMNH) FAP0253; 1–10.ii.1980, L. E. Peña leg. 2 females (AMNH) FAP0254; Puyehue, Aguas Calientes, 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, G. Hormiga leg. 2 females (USNM) FAP0244; Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 2 males (USNM) FAP0240 one female specimen used for SEM; 1 female and 2 immatures (USNM) FAP0241 smallest female measured epigynum illustrated; 3 females and 2 immatures (USNM) FAP0242; Palena, Chaiten, 25–27 km N of, 40 m, 16.i.1986, N. I. Platnick, P. A. Goloboff & R. T. Schuh leg. 2 females and 1 immature (AMNH) FAP0262; Valdivia, Purolon, 10.i.1985, L. E. Peña leg. 1 female (AMNH) FAP0248; Valdivia, 25.xii.1979, E. Krahmer leg. 2 females (AMNH) FAP0247; Valdivia, 30 km S of, 13.i.1951, Ross & Michelbacher leg. 1 female (CAS) FAP0246. Región XI del Ais, Aisen, Río Aisen Valley, i.1897. P. Dusen leg. 1 male (holotype of Meta echina NRS) FAP0275.

Allende longipes (Nicolet, 1849) comb. nov. (Figs6b, C, 9F–H, 11B, 18)

Epeira longipesNicolet, 1849: 491 (type presumably deposited at MNHN. Species identifications made with Dr Levi's illustrations of the type material).

Meta auroSimon, 1901: 19 (female holotype MNHN, examined). New synonymy.

Meta chilensisTullgren, 1902: 23 pl. 1 f. 7 (male holotype in NSR, examined). New synonymy.

Meta cordilleraTullgren, 1902: 23 pl. 2 f. 3 (female lectotype in NSR, examined). New synonymy.

Meta torTullgren, 1902: 21 pl. 2 f. 2 (female holotype in NSR, examined). New synonymy.

Meta longipes (Nicolet, 1849): Simon, 1900: 51.

Metabus longipes (Tullgren, 1902): Archer, 1963: 17.

Metabus chilensis (Tullgren, 1902): Archer, 1963: 17.

Metabus cordillera (Tullgren, 1902): Archer, 1963: 17.

Metabus tortus (Tullgren, 1902): Archer, 1963: 17.

Diagnosis:

A. longipes differs from other Allende species by the following combination of characters: female metatarsus I and II mesal surface with stout macrosetae (Fig. 18F); epigynum apical invaginations exposed and delineated by narrow ridges (Fig. 18H); cymbial median process globular-shaped (Fig. 18N); and shape of the conductor tip (Fig. 18Q).

Description:

Female from Cautín, Chacamo, Chile (FAP0090). Habitus as in Figure 18A, C–E. Total length 8.8. Cephalothorax 3.7 long, 2.8 wide, 1.1 high. Sternum 1.6 long, 1.5 wide. Chelicerae orange, darker than the cephalothorax (Fig. 18B). Abdomen oval, 5.8 long, 4.0 wide, 4.2 high, venter with a median white rectangular area and two white spots around the spinnerets. AME diameter 0.16. ALE 0.14. PME 0.16. PLE 0.14. Clypeus height 1.3 × AME diameter. AME separation 1.1 × AME diameter. AME–ALE separation 2.3 × AME diameter. PME separation 1.1 × PME diameter. PME–PLE separation 2.6 × PME diameter. Leg I length 17.5, leg II 13.0, leg III 7.4, leg IV 10.8, pedipalp 4.06. Spermathecae almost spherical, epigynum as in Figure 18G–K.

Male from Nahuelbuta, Chile (FAP0080). Habitus as in Figure 18L. Total length 5.3. Cephalothorax 2.5 long, 1.9 wide, 0.9 high. Sternum 1.2 long, 1.1 wide. Chelicerae same colour as the cephalothorax (Fig. 18M). Abdomen oval 3.1 long, 2.0 wide, 1.9 high. AME diameter 0.11. ALE 0.09. PME 0.11. PLE 0.09. Clypeus height 1.3 × AME diameter. AME separation 1.3 × AME diameter. AME–ALE separation 2.1 × AME diameter. PME separation 1.3 × PME diameter. PME–PLE separation 2.2 × PME diameter. Leg I length 17.1, leg II 11.9, leg III 5.9, leg IV 8.6, pedipalp 4.6. Pedipalp tibia 3.7 times longer than wide. Embolic apophysis long, pedipalp as in Figure 18N–Q.

Variation:

Four different variations of dorsal patterns were identified (Fig. 18A, C, D). These differences correspond to three different species described by Tullgren (1902) and synonymized here under A. longipes. These species are synonymized because the diagnostic pedipalp and epigynal characters presented no variation among dorsal patterns. Furthermore, a handful of specimens were found with intermediate abdominal patterns, suggesting that this could be a case of polymorphism for A. longipes; however, the locality data provided was insufficient to test if these abdominal variations were segregated at a population level. The only exceptions are the number of denticles on the cymbial basal process projection (which varies from three to five, even within individuals) and the epigynal median plate distal constriction width. Female total length 5.6–8.8, cephalothorax length 2.5–3.7. Male total length 4.6–6.3, cephalothorax length 2.3-3.2.

Distribution:

A. longipes is found in Chile and Argentina (Fig. 11B). Its geographical range covers the following latitudes 25°47′0″S to 53°40′0″S, and altitudes from 30 to 1450 m.

Material examined (N= 232):

ARGENTINA: Neuqu, Pucara, 1 male (CAS) FAP0040. CHILE: Región Metropolitana de Santia, Santiago, C. Joseph leg. 2 females (AMNH) FAP0200. Región V de Valparai, Valparaiso, Cerro Molle, C. Porter leg. 1 female. (holotype of Meta auro MNHN) FAP0282. Región VII del Mau, Curico, Las Tablas, ii.1985, L. E. Peña leg. 1 female (AMNH) FAP0071. Región VIII del Bio B, Guallali, Lago El Barco, 1200 m, 21–28.ii.1981, L. E. Peña leg. 1 male (AMNH) FAP0077 largest male measured; Ñuble, Atacalco, Chillan area, 17.iii.1983, L. E. Peña leg. 1 female (AMNH) FAP0083; Las Cabras, 26-28.xii.1986, L. Umaña leg. 1 female and 1 immature (AMNH) FAP0049; Las Trancas, 18.iii.1983, L. E. Peña leg. 2 females (AMNH) FAP0062; Las Trancas, El Recinto, 1100 m, ii.1987, L. E. Peña leg. 1 male (AMNH) FAP0073; Los Lleques, 5–20.xii.1985, L. Umaña leg. 4 females (AMNH) FAP0055. Región IX de la Araucar, Cautín, Chacamo, 600–700 m, 16–24.ii.1984, L. E. Peña leg. 2 males and 2 females (AMNH) FAP0088; 17–23.ii.1981, L. E. Peña leg. 1 female (AMNH) FAP0090 abdominal morpho 1 (Fig. 18A, L) largest female measured and illustrated; Toltan, 27.ii.1979, L. E. Peña leg. 1 male and 1 female (AMNH) FAP0072; Villarrica, 27–28.ii.1979, L. E. Peña leg. 32 males, 17 females and 4 immatures. (AMNH) FAP0065 smallest female measured, one male and one female specimens used for SEM; L. E. Peña leg. 3 males and 3 females (AMNH) FAP0067; Malleco, Angol, Nahuelbuta, 14–24.ii.1977, G. Moreno leg. 3 females (AMNH) FAP0050; Contulmo Natur., 350 m, 11.xii.1984−13.ii.1985, S. & W. Peck leg. 1 male (AMNH) FAP0058; Malalcahuello, 9–15.xii.1985, L. E. Peña leg. 1 male and 2 immatures (AMNH) FAP0087; Nahuelbuta Cord., 1300−1400 m, 6–12.i.1982, L. E. Peña leg. 1 male (AMNH) FAP0089; Nahuelbuta, 1300 m, 6–12.i.1982, L. E. Peña leg. 7 males and 9 immatures (AMNH) FAP0047; 1120 m, 13.ii.1992, N. I. Platnick, P. A. Goloboff & M. J. Ramírez leg. 3 males (AMNH) FAP0048; 1300 m, 1–6.ii.1979, L. E. Peña leg. 1 male, 3 females and 3 immatures (AMNH) FAP0069; L. E. Peña leg. 1 male and 4 females (AMNH) FAP0075; L. E. Peña leg. 5 males and 1 female (AMNH) FAP0080 male measured and illustrated; Nahuelbuta, sendero Piedra del Aguila, 37°49′30″S, 73°2′15″W, 1450 m, 8.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 2 males (USNM) FAP0039; Tolhuaca, 15–23.iii.1986, L. E. Peña leg. 1 female (AMNH) FAP0085. Región X de los Lag, Chiloe Is., Cerros del Cucao, Cord de Piuche, 1.iii.1981, L. E. Peña leg. 1 male and 5 females (AMNH) FAP0079; Continental, Río Ventisquero, Lago Velcho, 5–9.xii.1985, L. E. Peña leg. 1 male and 4 immatures (AMNH) FAP0086; Llanquehue, Los Muermos, 41°24′0″S, 73°29′0″W, 13.xi.1966, E. I. Schlinger & M. Irwin leg. 1 female (CAS) FAP0043; Calbuco, 21–28.ii.1967, A. F. Archer leg. 2 males and 2 immatures (AMNH) FAP0056; 3 males and 1 immature (AMNH) FAP0264 smallest male measured; Lago Chapo, 7-10.ii.1985, L. E. Peña leg. 1 male (AMNH) FAP0068; Osorno, La Pelada Chica, 1–2.iii.1987, L. E. Peña leg. 1 female (AMNH) FAP0070; La Picada, El Refugio, 41°4′0″S, 72°26′0″W, 6.ii.1967, E. I. Schlinger leg. 1 male (AMNH) FAP0053; La Unión, 600 m, 25–28.iii.1987, L. E. Peña leg. 4 females (AMNH) FAP0076; 900 m, 1–2.iii.1987, L. E. Peña leg. 5 females and 1 immature (AMNH) FAP0066; Puyehue NP, 40°46′30″S, 72°12′0″W, 700 m, 12.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 male (USNM) FAP0038; 24.i.1951, Ross & Michelbacher leg. 1 male (CAS) FAP0041; 26.i.1951, Ross & Michelbacher leg. 1 female and 4 immatures (CAS) FAP0042; Puyehue, Aguas Calientes, 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, G. Hormiga leg. 2 males and 1 immature (USNM) FAP0035; 13.xii.2000−1.i.2001, G. Hormiga leg. 1 male (USNM) FAP0036; 1 female (USNM) FAP0037; 40°44′0″S, 72°18′0″W, 500 m, 16.i.1995, N. I. Platnick, K. Catley & D. Silva leg. 6 females (AMNH) FAP0046; 600 m, 18.xii.1984−8.i.1985, S. & J. Peck leg. 1 female (AMNH) FAP0054; 1 female (AMNH) FAP0059; 425 m, 31.i.1985, N. I. Platnick & O. Francke leg. 2 males and 1 female (AMNH) FAP0060; 500 m, 2–5.i.1982, L. E. Peña leg. 2 males and 1 immature (AMNH) FAP0081; 600 m, 12–20.ii.1979, L. E. Peña leg. 1 male and 4 females (AMNH) FAP0082; 480 m, 17–18.ii.1992, N. I. Platnick, P. A. Goloboff & M. J. Ramírez leg. 1 female (AMNH) FAP0197 female morpho 3 illustrated (Fig. 18C); Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 2 males and 1 female (USNM) FAP0033; Puyehue, Los Mallines, 40°46′0″S, 72°17′0″W, 700 m, 12.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 male (USNM) FAP0034; Palena, Chaiten, 40 m, 17.i.1986, N. I. Platnick, P. A. Goloboff & R. T. Schuh leg. 1 female (AMNH) FAP0209; Chaiten, 70 km S of, 500 m, 16.i.1986, N. I. Platnick, P. A. Goloboff & R. T. Schuh leg. 1 female (AMNH) FAP0196; Valdivia, Las Lajas, 9–13.i.1990, L. E. Peña leg. 1 male, 3 females and 3 immatures (AMNH) FAP0064; Las Trancas, W La Unión, 500 m, 6–10.ii.1988, L. E. Peña leg. 2 females (AMNH) FAP0084; Neltume, ii.1987, L. E. Peña leg. 3 females (AMNH) FAP0045 female morpho 2 illustrated (Fig. 18E); Santo Domingo, Valdivia, 1.xi.1976, E. Krahmer leg. 1 female (AMNH) FAP0051. Región XI del Ais, Aisen, Cisnes Medio, 200 m, 30.xii.1984−28.i.1985, S. & J. Peck leg. 1 female (AMNH) FAP0057; Cocharne, 1–3.ii.1990, L. E. Peña leg. 1 male and 2 females (AMNH) FAP0074; Cocharne, Confl. Backer & Nef rivers, 180 m, 5–6.ii.1990, L. E. Peña leg. 1 male (AMNH) FAP0078; Cocharne, Río Backer, 180 m, 30.i.1990, L. E. Peña leg. 1 female (AMNH) FAP0063. Quelat, PN nr. Pto. Cisnes, 500 m, 6.ii.1985, N. I. Platnick & O. Francke leg. 2 females (AMNH) FAP0198 female morpho 4 illustrated (Fig. 18D); Río Aisen, Valley, i. 1897, P. Dusen leg. 1 male (holotype of Meta tor NRS) FAP0274; 1 female (holotype Meta chilensis NRS) FAP0276; 3 females (paralectotypes Meta cordillera NRS) FAP0277; 1 female (lectotype Meta cordillera designated by Levi NRS) FAP0278; 2 females (NRS) FAP0279. Región XII de Magallan, Magallanes, Chorrilla de la Piedra, 3.ii.1990, T. Cekalovic leg. 1 female (AMNH) FAP0052; Tierra del Fuego, Cameron, 30 m, 2.xii.1966, E. I. Schlinger & M. Irwin leg. 2 males (CAS) FAP0044.

Allende nigrohumeralis (F. O. P.–Cambridge, 1889) comb. nov. (Figs6d, E, 9I–J, 11B, 19)

Meta nigrohumeralis F. O. P.-Cambridge, 1899: 18 pl. 2 f. 4–5 (female holotype in BMNH, examined). New synonymy.

Meta porteSimon, 1900: 51 (female holotype in MNHN, examined). New synonymy.

Metabus nigrohumeralis(F. O. P.-Cambridge, 1889): Archer (1963): 17.

Metabus porteri (Simon, 1900): Archer, 1963: 17.

Diagnosis:

A. nigrohumeralis differs from other Allende species by the following combination of characters: two anterior dark patches on the abdomen (Fig. 19A); epigynum apical invaginations delineated by thick ridges (Fig. 19C); cymbial median process curved apically (Fig. 19L), cymbial basal process projection covered with small denticles (Fig. 19P); and the shape of the conductor tip (Fig. 19N).

Description:

Female from Valparaiso, Cerro Las Vizcachas, Chile (FAP0142). Habitus as in Figure 19A. Total length 10.4. Cephalothorax 4.4 long, 3.3 wide, 1.3 high. Sternum 1.9 long, 1.8 wide. Chelicerae anterior surface covered with dark dots (Fig. 19B). Abdomen oval, 6.5 long, 5.2 wide, 4.9 high, venter with a median white rectangular area delineated by two thin dark lines. AME diameter 0.19. ALE 0.18. PME 0.19. PLE 0.18. Clypeus height 1.5 × AME diameter. AME separation 1.1 × AME diameter. AME–ALE separation 2.8 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 3.3 × PME diameter. Legs light brown, femur I dark brown. Leg I length 25.7, leg II 19.1, leg III 9.8, leg IV 16.7, pedipalp 5.4. Spermathecae spherical, epigynum as in Figure 19C–G.

Male from Valparaiso, Cerro Las Vizcachas, Chile (FAP0142). Habitus as in Figure 19H. Total length 7.3. Cephalothorax 3.7 long, 2.9 wide, 1.1 high. Sternum 1.7 long, 1.5 wide. Chelicerae pattern as in Figure 19I. Abdomen oval, pattern as in female 4.5 long, 2.8 wide, 2.4 high. AME diameter 0.16. ALE 0.14. PME 0.16. PLE 0.14. Clypeus height 1.6 × AME diameter. AME separation 1.0 × AME diameter. AME–ALE separation 2.3 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 2.5 × PME diameter. Femur I dark brown. Leg I length 28.6, leg II 19.6, leg III 9.6, leg IV 13.1, pedipalp 5.9. Pedipalp tibia 3.5 times longer than wide. Embolic apophysis long, pedipalp morphology as in Figure 19J–P.

Variation:

Female total length 7.6–13.3, cephalothorax length 3.0–5.9. The epigynal median plate distal constriction varies in width. Male total length 5.4–8.9, cephalothorax length 2.6–4.3. The number of denticles on the cymbial basal process projection also varies; nevertheless, the margin is always completely covered by them.

Distribution:

A. nigrohumeralis is found in Chile and Argentina (Fig. 11B). Its geographical range covers the following latitudes 24°18′0″S to 36°56′0″S, and altitudes from 40 to 1300 m.

Material examined (N= 335):

ARGENTINA: Chub, Epuyén, 17.x.1966, A. Kovacs leg. 1 female (AMNH) FAP0194 smallest female measured. Río Neg, El Bolson, 41°58′0″S, 71°31′0″W, 23.x.1961, A. Kovacs leg. 1 male (AMNH) FAP0189; 13.iii.1961, 1 female (AMNH) FAP0190; 1.ii.1961, 1 female (AMNH) FAP0191; 3 females (AMNH) FAP0192; 21.ii.1963, 1 female (AMNH) FAP0193; Ñorquinco, 20.vi.1966, A. Kovacs leg. 1 male (AMNH) FAP0195. CHILE: Chile, 2 females and 1 immature (MNHN) FAP0290; 25 females and 2 males (MNHN) FAP0291; 1 male (MNHN) FAP0292. Juan Fernández I, Cumberland Bay, 11-12.iii.1962. B. Malkin leg. 1 female (AMNH) FAP0178; 1 female (AMNH) FAP0179; 7 females and 2 immatures (AMNH) FAP0180; Galpon, Valle Villagra, Mas a tierra, 23-24.iv.1962, B. Malkin leg. 6 females and 7 immatures (AMNH) FAP0183; Juan Fernández, 8.xii.1926, W. L. Schmith leg. 1 male, 2 females and 1 immature (USNM) FAP0206; Ghe, 1 female (holotype of Meta nigrohumeralis BMNH) FAP0270; Juan Fernández, Caracoles, viii.1943, Rosa leg. 1 male (CAS) FAP0112; La Piña, 33°40′6″S, 78°48′0±W, 280–400 m, 31.xii.2002, S. Lew, K. Will & S. Ocare leg. 1 male and 1 female (EME) FAP0208; Plazoleto de Yunque, Mas a tierra, 23–24.iv.1962, B. Malkin leg. 3 females and 2 immatures (AMNH) FAP0182; San Juan Bautista, 33°38′9″S. 78°49′54″W, 28.xii.2002, S. Lew, K. Will & W. Ocare leg. 7 females (EME) FAP0207. Región Metropolitana de Santia, Aculeo, El Paragual, 5–8.xii.1983, L. Irarrazaval leg. 1 female and 2 immatures (AMNH) FAP0164; Melpilla, La Viluma, SE Melpilla, 350 m, 15.xii.1987, L. E. Peña leg. 1 female (AMNH) FAP0203; San Manuel, 13–14.v.1980, L. E. Peña leg. 1 female (AMNH) FAP0133; 6–8.xii.1980, L. E. Peña leg. 1 male and 2 females (AMNH) FAP0135; Santiago, Cuesta La Dormida, 800–1300 m, 13–18.xi.1982, Peña leg. 2 males, 2 females and 2 immatures (AMNH) FAP0149; El Golf, 9.iv.1961, A. F. Archer leg. 3 females (AMNH) FAP0185; 3 females and 2 immatures (AMNH) FAP0267 largest female measured, one male specimen used for SEM; Quilicura, vs. 1979, L. E. Peña leg. 1 female (AMNH) FAP0140; L. E. Peña leg. 1 female (AMNH) FAP0205; Los Lirios, 1 female (MNHN) FAP0288; Mnt. Bell, 914 m, 17.xii.1950, Ross & Michelbacher leg. 1 female (CAS) FAP0117; Mnt. Bell, 914 m, 17.xii.1950, Ross & Michelbacher leg. 1 male (CAS) FAP0119. Región IV del Coquim, Choapa, Los Vilos Km 26 Rt 5, 40 m, 31°50′0″S, 71°31′0″W, 13.xii.1993, N. I. Platnick, K. Catley, M. J. Ramírez & T. Allende leg. 1 male (AMNH) FAP0175; Coquimbo, Fray Jorge NP, 11.xii.1950, Ross & Michelbacher leg. 1 female (CAS) FAP0120; Pichidangui, 32°8′0″S, 71°32′0″W, 12.viii.1996, E. I. Schlinger leg. 1 female (AMNH) FAP0199; Quilican, 16 km E la Serena, 2.x.1961, R. Wagnerknetch leg. 1 female (AMNH) FAP0265; Los Vilos, 12.x.1986, L. E. Peña leg. 1 male, 1 female and 1 immature (AMNH) FAP0141; Ovaile, 13.viii.1950, Ross & Michelbacher leg. 1 female (CAS) FAP0115. Region V de Valparai, Aconcagua, C. Porter leg. 1 female (MNHN) FAP0289; Cachagua, 14.xii.1980, L. E. Peña leg. 1 male, 4 females and 1 immature (AMNH) FAP0129; Las Palmas, 1 female (holotype of Meta porte MNHN) FAP0283; Petorca, 8.x.1986, L. E. Peña leg. 1 male 4 immatures (AMNH) FAP0128; Concon, 16.viii.1950, Ross & Michelbacher leg. 1 male (CAS) FAP0114; Valparaiso, Cerro Las Vizcachas, 1800−2200 m, 1–12.xii.1982, L. E. Peña leg. 2 males and 1 female (AMNH) FAP0142 male and female illustrated and measured; Cordoba, Santiago Coast, 15–20.ii.1979, L. E. Peña leg. 1 female (AMNH) FAP0201; Cuesta El Melón, 10-12.x.1986, L. E. Peña leg. 1 male, 3 females and 15 immatures (AMNH) FAP0145; 15.xii.1985, L. E. Peña leg. 1 female and 3 immatures (AMNH) FAP0147; Quintero, 12.xii.1980, L. E. Peña leg. 3 males (AMNH) FAP0132; Valparaiso Central Coast, 31.x.1982, 1 male (AMNH) FAP0173; 1 male (AMNH) FAP0187; Viña del Mar, vi.1979, A. Tobar leg. 2 females (AMNH) FAP0168; xii.1978, A. Tobar leg. 1 female (AMNH) FAP0169; i–ii.1978, A. Tobar leg. 3 females (AMNH) FAP0202. Región VII del Mau, Cauquenes, Reserva Nacional Los Ruiles, NW of Cauquenes, 35°50.028′S, 72°30.613′W, 146 m, 13.xi.2003, L. Prendini, C. I. Mattoni & J. A. Ochoa leg. 1 female (AMNH) FAP0394; 1 female (AMNH) FAP0395; Curico, Las Tablas, ii.1985, L. E. Peña leg. 1 female (AMNH) FAP0148; L. E. Peña leg. 1 female and 2 immatures (AMNH) FAP0150; Linares, Bullileo, Parral, 5–8.xii.1990, L. E. Peña leg. 1 male 2 immatures (AMNH) FAP0127; Maule, Cayurranquil, 400 m, 24–27.i.1981, L. E. Peña leg. 1 male (AMNH) FAP0155; El Parrón, 30.i.1981, L. E. Peña leg. 1 female (AMNH) FAP0136; Tregualemu, 4–5.xi.1993, L. E. Peña leg. 3 males and 5 immatures (AMNH) FAP0126; Talca, Bramadora, 600 m, 21–22.i.1993, L. E. Peña leg. 1 female (AMNH) FAP0210 cleared epigynum illustrated; Gil de Vilches NP, 35°36′0″S, 71°4′0″W, 1200 m, 14.i.1995, N. I. Platnick, K. Catley & D. Silva leg. 1 female (AMNH) FAP0174. Región VIII del Bio B, Bio-Bio, El Manzano, 15.xii.1985, L. E. Peña leg. 2 males, 3 females and 2 immatures (AMNH) FAP0124; 5 males, 4 females and 6 immatures (AMNH) FAP0146 smallest male measured; 1 female (AMNH) FAP0204; Carampangue, 4.3 km E Las Corrientes, campsite 5 km W, 37°15.547′S, 73°11.508′W, 116 m 13.xi.2003, L. Prendini, C. I. Mattoni & J. A. Ochoa leg. 2 females (AMNH) FAP0396; 8.xii.1992, T. Cekalovic leg. 1 male 5 immatures (AMNH); Concepción, Chiguayante, 23.xi.1975, T. Cekalovic leg. 2 females (AMNH) FAP0061; Escuadrón, 24.iii.1989, T. Cekalovic leg. 1 female (AMNH) FAP0157; Escuadrón, Laguna la Posada, 17.i.1976, T. Cekalovic leg. 1 female (AMNH) FAP0165; Estación Cosmito, 17.iv.1977, G. Muñoz leg. 1 female (AMNH) FAP0186; Nonquen Estero, 23.i.1987, T. Cekalovic leg. 1 female (AMNH) FAP0166; 3.ii.1985, T. Cekalovic leg. 2 females (AMNH) FAP0181; 11.xii.1977, T. Cekalovic leg. 1 female (AMNH) FAP0211; Nonquen Valle, 6.xii.1981, T. Cekalovic leg. 1 female (AMNH) FAP0162; Penco, 22.xii.1979, T. Cekalovic leg. 1 female (AMNH) FAP0158; Periquillo, 21.xii.1996, T. Cekalovic leg. 1 male and 1 immature (AMNH) FAP0163; Sector Escuadrón, 27.xii.1987, T. Cekalovic leg. 1 female (AMNH) FAP0156; Tome, 8.x.1983, T. Cekalovic leg. 1 female and 8 immatures (AMNH) FAP0159; 1.i.1992, T. Cekalovic leg. 2 females (AMNH) FAP0160; Ñuble, Chillan, 31.xii.1975, G. Moreno leg. 1 female (AMNH) FAP0176; Coloquecura, 8–9.xi.1993, L. E. Peña leg. 2 males (AMNH) FAP0125 one female specimen used for SEM; Las Cabras, 26–28.xii.1986, L. Umaña leg. 1 male (AMNH) FAP0171; Las Trancas, iii–iv.1979, L. E. Peña leg. 3 females and 14 immatures (AMNH) FAP0143; Los Lleques, 5-20.xii.1985, L. Umaña leg. 4 males (AMNH) FAP0170; Pinto 4 km E road to, 4.i.1976, G. Moreno leg. 1 female (AMNH) FAP0177; W. Ralco, Santa Barbara, 400 m, 22–23.xi.1994, L. E. Peña leg. 1 female (AMNH) FAP0154. Región IX de la Araucar, Cautín, Chacamo, 600–700 m, 17–23.ii.1981, L. E. Peña leg. 1 male and 1 female (AMNH) FAP0144; Pucón, 15.xi.2.xii.1989, S. A. Marshall leg. 1 male (AMNH) FAP0184 largest male measured; Villarrica, 27–28.ii.1979, L. E. Peña leg. 1 female (AMNH) FAP0131; 12.xii.1961, A. F. Archer leg. 1 female and 2 immatures (AMNH) FAP0266; Malleco, Curacautín, W Selva Oscura, 27.i.1985, L. E. Peña leg. (AMNH) 1 female FAP0152; Malleco, xi.1979, L. E. Peña leg. 1 female (AMNH) FAP0153; Nahuelbuta, Pehuenco, 37°49′45″S, 73°0′30″W, 1100 m, 4–9.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 3 females (USNM) FAP0269; J. Miller, F. Alvarez, J. A. Coddington & G. Hormiga leg. 1 female (USNM) FAP0108. Región X de los Lag, Chiloe, Lago Coluco, 26.i.1981, L. E. Peña leg. 1 female (AMNH) FAP0139; Puntra, 21–23.xii.1981, L. E. Peña leg. 1 female (AMNH) FAP0137; Pio Pio, 10–12.iii.1987, L. E. Peña leg. 2 females (AMNH) FAP0151; Llanquihue, Petrohue, 41°8′0″S, 72°25′0″W, 2700 m, 15.xi.1966, E. I. Schlinger & M. Irwin leg. 1 female (CAS) FAP0118; Pt. Montt, 20.iii.1991, T. Cekalovic leg. 2 females (AMNH) FAP0161; 17.iii.1991, T. Cekalovic leg. 1 female (AMNH) FAP0167; Osorno, La Picada, El Refugio, 41°4′0″S, 72°26′0″W, 6.ii.6197, E. I. Schlinger leg. 1 female (CAS) FAP0113; Puyehue NP, 600 m, 12–22.ii.1979, L. E. Peña leg. 1 female (AMNH) FAP0134; Puyehue, Aguas Calientes, 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, G. Hormiga leg. 3 females (USNM) FAP0110; xii.1981, L. E. Peña leg. 1 female (AMNH) FAP0138; 425 m, 31.i.1985, N. I. Platnick & O. Francke leg. 2 females (AMNH) FAP0172; Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 female (USNM) FAP0109; 2 females (USNM) FAP0111; Pt. Montt, Lepihue, 21.i.1951, Ross & Michelbacher leg. 1 female (CAS) FAP0121; Valdivia, Neltume, 2. 39°48′0″S, 71°57′0″W, 23.xi.1988, V. & B. Roth leg. 1 female (CAS) FAP0122; Purolon, 10.i.1985, L. E. Peña leg. 1 male, 5 females and 3 immatures (AMNH) FAP0130; Río Bueno, 15.i.1951, Ross & Michelbacher leg. 1 female (CAS) FAP0116; Ross & Michelbacher leg. 1 female (CAS) FAP0123; Valdivia, 25.xii.1979, E. Krahmer leg. 1 female (AMNH) FAP0188.

Mollemetagen. nov. (Figs7A–E, 8A–E, 9K–O, 10C, 11D, 20)

Figure 20

Mollemeta edwardsi. A, female habitus. B, epigynum ventral view. C, caudal view. D, ventral view cleared. E, dorsal view cleared. F–J, male pedipalp. F, cymbium (bulb dissected). G, expanded. H, dorsal view. I, palpal tibia. J, ventral view. Scale bars 0.2 mm, except in habitus 1 mm.

Type species:

Landana edwardsiSimon, 1904.

Etymology:

The genus name is derived from the Mapuche word for tree (Molle) and thus Mollemeta means ‘The Meta that builds orb webs on trees.’Mollemeta is a name feminine in gender.

Diagnosis:

Mollemeta can be distinguished from other tetragnathids by the following combination of characters: epigynum with a spherical small scape, genital openings situated in deep curved groves (Fig. 8A, B); tegulum reduced to a narrow ring (Fig. 20G); membranous conductor, embolus without basal apophyses (Fig. 20J).

Description:

Female from Llanquihue, P. N. Alerce Andino, Chile (FAP0332). Habitus as in Figure 20A. Total length 9.2. Cephalothorax 3.8 long, 2.9 wide, 1.2 high. Sternum 1.8 long, 1.5 wide. Chelicerae with a large boss. Abdomen oval 6.14 long, 5.27 wide, 4.46 high, venter with two parallel lines of white guanine patches. AME diameter 0.21. ALE 0.2. PME 0.22. PLE 0.19. Clypeus height 0.9 × AME diameter. AME separation 0.9 × AME diameter. AME–ALE separation 0.4 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 1.0 × PME diameter. Spinnerets (Fig. 9K–M). ALS with several piriform spigots. PMS with three aciniform spigots between the cylindrical spigot and the minor ampullate, nubbin present. PLS with a field of c. 20 aciniform spigots, aggregate spigots embracing the flagelliform and cylindrical spigots peripheral. Leg I length 17.6, leg II 13.8, leg III 9.4, leg IV 11.3, pedipalp 4.3. Epigynum as in Figure 20B–E. Copulatory ducts curved and almost as thick as the spermathecae (Fig. 8C). Fertilization ducts short and well sclerotized (Fig. 8D). Spermathecae well sclerotized, accessory gland openings arranged in clusters (Fig. 8E).

Male from Aguas Calientes, Puyehue, Chile (FAP0328). Habitus as in female. Total length 8.2. Cephalothorax 4.3 long, 3.1 wide, 1.3 high. Sternum 3.9 long, 3.2 wide. Abdomen oval longer than wide, venter as in female. 4.4 long, 3.1 wide, 2.6 high. Epiandrous fusules on a plate and concentrated in clusters (Fig. 9O). PLS and PMS cylindrical and aggregate spigots reduced to nubbins (Fig. 9N). AME diameter 0.19. ALE 0.22. PME 0.22. PLE 0.2. Clypeus height 1.3 × AME diameter. AME separation 1.3 × AME diameter. AME–ALE separation 0.8 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 1.1 × PME diameter. Leg I length 32.6, leg II 23.0, leg III 11.6, leg IV 14.8, pedipalp 6.5. Pedipalp tibia 3.2 times longer than wide (Fig. 7E); pedipalp as in Figure 20F–J.

Variation:

Female total length 7.1–13.6, cephalothorax length 3.7–5.9. Male total length 5.8–8.4, cephalothorax length 2.7–4.5.

Natural history:

Mollemeta edwardsi builds vertical orb webs on tree trunks. The web is longer than wide with c. 48 radii, and several spirals of the sticky silk (more than 50). The temporary spiral is removed and the hub is closed (Fig. 10C).

Phylogenetics:

The monotypy of this genus is based on the cladistic placement of M. edwardsi as sister to a clade that contains the genera Meta, Metellina, Dolichognatha, Allende and Chrysometa (Figs 1, 2). Autapomorphies of Mollemeta are the ring-shaped cymbium, and the unique shape of the conductor and embolus (Fig. 7A, B).

Composition:

Monotypic.

Distribution:

This species is endemic to Chile (Fig. 11D).

Mollemeta edwardsi (Simon, 1904) comb. nov.

Landana edwardsiSimon, 1904: 94 (female holotype in MNHN, examined).

Dolichognatha edwardsi (Simon, 1904): Levi, 1981: 277.

Diagnosis:

See above under genus description.

Material examined (N= 75):

CHILE: Chili (MNHN) FAP0311. Región IX de la Araucan, Cautín, P.N. Nietol nr. Temuco, site 652, pyrethrin fogging tree bark, 250 m, 14–30.xii.1982, A. Newton & M. Thayer leg. 1 male (AMNH) FAP0351; Malleco, Puren Contulmo Natur. Mon, 350 m, 11.xii.1984−13.ii.1985, S. & J. Peck leg. 1 male (AMNH) FAP0350. Región X de los Lag, Llanquihue, P. N. Alerce Andino, Sector Sargazo, 41°30′30″S, 72°37′0″W, 350 m, 20–27.xii.2000, J. Miller, I. Agnarsson, F. Álvarez, J. A. Coddington & G. Hormiga leg. 2 females (USNM) FAP0330 one female specimen used for SEM; 2 females (USNM) FAP0331; 3 females (USNM) FAP0332 female illustrated; 2 females (USNM) FAP0333; Petrohue, 23.i.1951, Ross & Michelbacher leg. 1 female (CAS) FAP0340; Osorno, Aguas Calientes, P. N. Puyehue, 480 m, 17–18.ii.1992, N. I. Platnick, P. A. Goloboff, & M. I. Ramírez leg. 2 males, 1 female and 5 immatures (AMNH) FAP0342; 600–700 m, 28.i.1986, N. I. Platnick, & R. T. Schuh leg. 1 female (AMNH) FAP0344; 450 m, 27.i.1986, N. I. Platnick, P. A. Goloboff & R. T. Schuh leg. 9 females and 4 immatures (AMNH) FAP0345; 5 females and 4 immatures (AMNH) FAP0346 largest female measured; 40°44′0′S, 72°18′0′W, 480 m, 21.xi.1993, N. I. Platnick, K. Catley, M. I. Ramírez & T. Allende leg. 3 males and 3 females (AMNH) FAP0347 largest male measured; 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 2 females (USNM) FAP0325; 1 female (USNM) FAP0326; 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000−2.i.2001, G. Hormiga leg. 1 male and 1 female (USNM) FAP0327 one male specimen used for SEM; 1 male and 1 female (USNM) FAP0328 male illustrated; 1 male and 1 female (USNM) FAP0329; 1 female (USNM) FAP0379; 13-17.xii.1998, M. J. Ramírez, M. Compagnucci, C. Grismado & L. Lopardo leg. 1 male (MACN-Ar) FAP0338; P. N. Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga, 1 female (USNM) FAP0334; 1 female (USNM) FAP0337; 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000−2.i.2001, G. Hormiga leg. 1 male (USNM) FAP0335; 1 female (USNM) FAP0336; 1 female (USNM) FAP0380; Puyehue, 10 km E of, 24.i.1951, Ross & Michelbacher leg. 1 male (CAS) FAP0339; Osorno, xii.1978, A. Tobar leg. 1 female (AMNH) FAP0349; Palena, Chaiten, 10 m, 16.i.1986, N. I. Platnick, P. A. Goloboff, & R. T. Schuh leg. 1 female (AMNH) FAP0343; 2 males and 4 females (AMNH) FAP0348 smallest female and largest male measured; Valdivia, La Herradura, 6 km E Niebla, 20 m, 16.ii.1992, N. I. Platnick, P. A. Goloboff, & M. I. Ramírez leg. 1 female (AMNH) FAP0341.

Chrysometa Simon, 1895Chrysometa acinosasp. nov. (Figs10b, 11C, 21)

Types:

Female holotype from Osorno, Puyehue National Park, Chile (FAP0373), two male and three female paratypes, all deposited at the USNM (FAP0374).

Etymology:

Acinos means like grapes and refers to the spermathecae shape of this species.

Diagnosis:

Females of C. acino can be distinguished from other Chrysometa species by the grape-cluster-shaped spermathecae (Fig. 21D), and from C. butamLevi, 1986 by the constricted median plate (Fig. 21C). Males are distinguished by a massive process on the tibial dorsal margin (Fig. 21G, H).

Figure 21

Chrysometa acinos A, female habitus. B, epigynum ventral view. C, caudal view. D, ventral view cleared.E, dorsal view cleared. F-H, male pedipalp. F, ventral view. G, mesal view. H, ectal view. Scale bars 0.2 mm; except A 1 mm.

Description:

Female from Aguas Calientes, Puyehue, Chile (FAP0377). Habitus as in Figure 21A. Total length 6.3. Cephalothorax 2.7 long, 2.1 wide, 0.9 high. Sternum 1.1 long and wide. Chelicerae same colour as the cephalothorax. Abdomen oval 4.4 long, 4.1 wide, 3.2 high, venter with two parallel lines of silver guanine patches. AME diameter 0.15. ALE 0.19. PME 0.18. PLE 0.18. Clypeus height 1.3 × AME diameter. AME separation 1.0 × AME diameter. AME–ALE separation 1.1 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 1.1 × PME diameter. Leg I length 13.7, leg II 10.2, leg III 5.7, leg IV 8.0, pedipalp 3.0. Epigynum as in Figure 21B–E.

Male from Aguas Calientes, Puyehue, Chile (FAP0377). Habitus as in female. Total length 5.4. Cephalothorax 2.5 long, 2.1 wide, 0.9 high. Sternum 1.2 long, 1.1 wide. Cheliceral anterior surface covered with scales. Abdomen oval venter as in female. 3.2 long, 2.1 wide, 2.0 high. AME diameter 0.16. ALE 0.16. PME 0.19. PLE 0.16. Clypeus height 1.2 × AME diameter. AME separation 0.9 × AME diameter. AME–ALE separation 1.0 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 1.2 × PME diameter. Leg I length 17.5, leg II 11.6, leg III 6.3, leg IV 8.7, pedipalp 3.2. Pedipalp tibia 2.0 times longer than wide. Cymbial median process globular and covered with small openings, cymbial basal process longer than wide and membranous (Fig. 21H). Embolic basal apophysis short (Fig. 21F).

Variation:

Female total length 5.3–6.3, cephalothorax length 2.1–2.8. Male total length 3.1–5.4, cephalothorax length 1.8-2.5.

Distribution:

Endemic to Chile (Fig. 11C).

Material examined (N= 15):

CHILE: Región VIII del Bio B, Arauco, Contulmo, Monumento Natural, 300 m, 11.ii.1992, Platnick, Goloboff, & Ramírez, 2 males (AMNH) FAP0375 smallest male measured. Región IX de la Araucar, Malleco, Nahuelbuta, Pehuenco, 37°49′45″S, 73°0′30″W, 100 m, 4.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 2 males and 3 females (paratype USNM) FAP0374 smallest female measured. Región X de los Lag, Osorno, Puyehue, Aguas Calientes, 40°44′0″S, 72°18′45″W, 450 m, 12.xii.2000, G. Hormiga leg. 1 female (USNM) FAP0373 largest female measured; 1 male and 1 female (USNM) FAP0377 largest male measured, both specimens illustrated; 1 female (USNM) FAP0378; Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 2 males and 2 females (USNM) FAP0376.

Chrysometa leviisp. nov. (Figs11c, 22F–I)

Figure 22

A-E, Chrysometa maitae. A, female habitus. B, epigynum ventral view cleared. C, dorsal view cleared. D, central view. E, caudal view. F–I, Chrysometa lev. F, male habitus. G, male pedipalp ectal view. H, dorsal view. I, mesal view. Scale bars 0.2 mm; except A 1 mm.

Types:

Male holotype and one male paratype from Osorno, Puyehue National Park, Chile (FAP0370) deposited at USNM (FAP0371).

Etymology:

Named in honor of Dr Herbert W. Levi in recognition to his monumental contributions to the taxonomy of araneoid spiders.

Diagnosis:

This species can be distinguished from other Chrysometa species by the G-shaped embolus (Fig. 22I), and by the distally constricted cymbium (Fig. 22H).

Description:

Male from Aguas Calientes, Puyehue, Chile (FAP0368). Habitus as in Figure 22F. Total length 3.7. Cephalothorax 1.9 long, 1.5 wide, 0.4 high. Sternum 0.9 long, 0.8 wide. Anterior surface of chelicerae covered with scales. Abdomen oval 2.15 long, 1.50 wide, 1.24 high, venter without guanine patches. AME diameter 0.11. ALE 0.12. PME 0.12. PLE 0.11. Clypeus height 0.8 × AME diameter. AME separation 1.1 × AME diameter. AME–ALE separation 1.3 × AME diameter. PME separation 1.1 × PME diameter. PME–PLE separation 1.3 × PME diameter. Leg I length 9.5, leg II 7.3, leg III 3.6, leg IV 4.7, pedipalp 1.5. Palpal tibia 3.6 times longer than wide, pedipalps as in Figure 22G–I.

Variation:

Male total length 3.0–3.7, cephalothorax length 1.6–1.9.

Distribution:

Endemic to Chile (Fig. 11C).

Material examined (N= 25):

CHILE: Región IX de la Araucar, Malleco, Nahuelbuta, Pehuenco, 37°49′45″S, 73°0′30″W, 1100 m, 4.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 male (USNM) FAP0371 smallest male measured; 4.i.2001, G. Hormiga leg. 1 male (USNM) FAP0372. Región X de los Lag, Chiloe Is., Pio Pio, 10.iii.1987, L. E. Peña leg. 1 male (AMNH) FAP0369; Llanquihue, Calbuco, 21.ii.1967, A. F. Archer leg. 1 male and 2 immatures (AMNH) FAP0367; Osorno, 1 km E. Termas de Puyehue, 305 m, 31.i.1985, N. I. Platnick & O. Francke leg. 2 males (AMNH) FAP0366; Aguas Calientes, P. N. Puyehue, 600 m, 18.xii.1984−8.ii.1985, S. & W. Peck leg. 2 males (AMNH) FAP0360; Puyehue, Aguas Calientes, 450 m, 27.i.1986, N. I. Platnick & R. T. Schuh leg. 1 male (AMNH) FAP0368 largest male measured and illustrated; Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 male (USNM) FAP0370; Palena, 37 km SE Chaiten, 60 m, 28.xii.1984−30.i.1985, S. & W. Peck leg. 1 male (AMNH) FAP0364; Valdivia, 34 Km WNW La Unión, 700 m, 17.xii.1984−7.ii.1985, S. & W. Peck leg. 1 male (AMNH) FAP0362; Neltume, 17.xi.2003, L. E. Peña leg. 3 males and 3 immatures (AMNH) FAP0365. Región IX del Ais, Aisen, 35 km E Pto. Aisen, Río Simpson NP, 70 m, 31.xii.1984−26.i.1985, S. & W. Peck leg. 1 male (AMNH) FAP0363; Cisnes Medio, 16 km NW Río Grande, 200 m, 30.xii.1984, S. & W. Peck leg. 2 males (AMNH) FAP0361; Las Juntas, 15 km W. 30 km N. Puyuhuapi, 100 m, 30.xii.1984−29.i.1985, S. & W. Peck leg. 2 males (AMNH) FAP0359.

Chrysometa maitaesp. nov. (Figs11c, 22A–E)

Types:

Female holotype (FAP0352) and one female paratype (FAP0353) from Anticura, Puyehue National Park, Chile, deposited at USNM. Another female paratype from Antillanca, Chile, deposited at AMNH (FAP0358).

Etymology:

The species epithet ‘maitae’ is a derivation of the name of my niece, Maite.

Diagnosis:

This species can be distinguished from other Chrysometa species by the abdominal tubercles (Fig. 22A); the epigynum shape, and the posterior origin of the fertilization ducts (Fig. 22B–E).

Description:

Female from Anticura, Puyehue National Park, Chile (FAP0354). Habitus as in Figure 22A. Total length 4.1. Cephalothorax 1.7 long, 1.3 wide, 0.7 high. Sternum 0.8 long and wide. Chelicerae same colour as cephalothorax. Abdomen oval with two dorsal tubercles 2.82 long, 2.97 wide, 1.88 high, venter with two parallel lines of guanine silver patches. AME diameter 0.11. ALE 0.11. PME 0.13. PLE 0.11. Clypeus height 0.7 × AME diameter. AME separation 1.0 × AME diameter. AME–ALE separation 1.3 × AME diameter. PME separation 1.0 × PME diameter. PME–PLE separation 1.2 × PME diameter. Leg I length 6.7, leg II 5.0, leg III 2.9, leg IV 4.2, pedipalp 1.6. Epigynum as in Figure 22B–E.

Variation:

Female total length 3.3–4.1, cephalothorax length 1.6–1.7.

Distribution:

Endemic of to Chile (Fig. 11C).

Material examined (N= 12):

CHILE: Región X de los Lag, Osorno, 4.1 Km E of Anticura, trap site 662, 19–26.xii.1982. A. Newton & M. Thayer leg. 1 female (AMNH) FAP0357; Aguas Calientes, P. N. Puyehue, 500 m, 2–5.i.1982, L. E. Peña leg. 1 female and 5 immatures (AMNH) FAP0354 largest female measured; 450 m, 27.i.1986, N. I. Platnick, & R. T. Schuh leg. 1 female (AMNH) FAP0356; Anticura, E. of Puyehue, 26-31.viii.1–5.xi.1983, L. E. Peña leg. 1 female (AMNH) FAP0355; Antillanca rd., 470–720 m, 18–24.xii.1982, A. Newton & M. Thayer leg.1 female (AMNH) FAP0358; P. N. Puyehue, Anticura, 40°40′0″S, 72°10′30″W, 350 m, 13.xii.2000−2.i.2001, J. Miller, F. Álvarez, J. A. Coddington & G. Hormiga leg. 1 female (USNM) FAP0352 smallest female measured; G. Hormiga leg. 1 female (USNM) FAP0353.

ACKNOWLEDGEMENTS

Special thanks to G. Hormiga for helping in many ways and for his valuable comments on the manuscript. I would like to thank Nikolaj Scharff, Petra Sierwald and Bernhard Huber for the critical revision of this manuscript and their valuable comments. L. Lopardo, J. Coddington, D. Dimitrov and S. Benjamin also provided valuable comments and corrections on early drafts of the manuscript. I would like to thank Herbert W. Levi for sending copies of his unpublished illustrations of Simon's Meta type specimens deposited at the MNHN in Paris, and to G. Hormiga for providing unpublished data and photographs of the webs of A. puyehuensis, C. acino and M. edwardsi. Thanks to R. Clouse and J. Coddington for revising the English. I also would like to thank the following curators and museums for the loan of specimens: J. Beccaloni (BMNH), J. Coddington (USNM), G. Giribet and L. Leibensperger (MCZ), M. Gray and G. Milledge (AMS), C. Griswold (CAS), T. Kronestedt (NRS), S. Lou (EME), N. Platnick and L. Sorkin (AMNH), M. J. Ramírez and Cristina Scioscia (MACN-Ar), R. Raven and V. Davies (QMB), and C. Rollard (MNHN). I also would like to thank the following people for their valuable help during fieldwork: R. Raven, V. Davies, G. Hormiga, N. Scharff, S. Larsen and M. Kuntner (Australia); G. Hormiga, J. Coddington and J. Miller (Chile); and K. Guerrero, G. Hormiga, S. Benjamin and the staff of the scientific reserves ‘Ebano Verde’ and ‘Loma Quita Espuela’ (Dominican Republic). SEM facilities were provided by the Department of Biological Sciences at the George Washington University. Support for this project was provided by an NSF-PEET grant to Hormiga and Giribet (DEB-0328644), a scholarship from the Consejo Nacional de Ciencia y Tecnología of México (CONACYT), a REF grant from The George Washington University (to G. Hormiga), and collection study grants from the American Museum of Natural History and the California Academy of Sciences.

Appendices

APPENDIX 1

Character descriptions

64 of the 105 characters had been used in previous phylogenetic studies; in that case, a code at the end of each character indicates these references. The following codes are used: Levi, 1980=L80; Coddington, 1990=C90; Hormiga et al, 1995=H95; Scharff & Coddington, 1997=S97; Griswold et al (1998)=G98; Tanikawa, 2001=T01. New characters are marked with an asterisk after the character number. Citation does not necessarily mean character authorship, many of these characters have been used and described in spider systematics since the 19th century.

1. Male dorsal scutum: (0) absent; (1) present (H95, G98).

2. Abdominal dorsal surface: (0) without guanine silver patches (Meta ovalis); (1) with guanine silver patches (Leucauge venusta) (T01). These guanine crystals may help to prevent overheating by reflecting light (Oxford & Gillespie, 1998).

3. Booklung cover: (0) smooth; (1) grooved (H95, S97, G98, T01).

4. Tracheal spiracle: (0) posterior; (1) advanced (H95).

5. Gasteracanthinae abdominal spines: (0) absent; (1) present (S97).

6. Sclerotized ring around spinnerets: (0) absent; (1) present (S97).

7. ALS piriform bases: (0) normal; (1) reduced (H95, S97, G98).

8. PMS nubbin: (0) present; (1) absent (H95, S97, G98).

9. PMS aciniform gland spigots: (0) extensive; (1) reduced (H95, S97, G98).

10. PLS mesal cylindrical spigot base: (0) same size of the other PLS cylindrical spigots; (1) enlarged (H95).

11. PLS mesal cylindrical spigot position: (0) central; (1) peripheral (H95, G98).

12. PLS aggregate flagelliform triad: (0) aggregate spigots apart from flagelliform spigot; (1) distal end of the aggregate spigots embrace the distal end of the flagelliform spigot (H95).

13. PLS aggregate gland spigots: (0) normal; (1) enlarged (H95, G98).

14. Cheliceral stridulatory striae: (0) absent; (1) present (H95, S97, G98).

15. Male cheliceral distal curvature: (0) fewer than two times clypeus width (Leucauge venustaLevi, 1980: fig 54); (1) more than two times clypeus width (Tetragnatha versicolLevi, 1981: fig. 102). The chelicerae curvature is measured as the distance between the distal ectal surfaces of the chelicera divided by the clypeus width.

16. Cheliceral boss: (0) smooth; (1) striated (H95, S97, G98).

17. Cheliceral boss striae: (0) few (< 20); (1) numerous (> 25) (H95).

18. Cheliceral denticles: (0) absent; (1) present (H95).

19. Male cheliceral length: (0) proportionally the same size of the female (Leucauge venustaLevi, 1980: figs 47, 54); (1) distally elongated (Dolichognatha pentagoLevi, 1981: figs 3, 11).

20. Male cheliceral anterior cuticle: (0) like the clypeal area (Tetragnatha versicolLevi, 1981: fig 102); (1) covered with scales (Allende patagiatus, Fig. 17I); (2) covered with small protuberances (Dolichognatha pentagoLevi, 1981: fig. 11). The cheliceral cuticle with scales is a synapomorphy of the Allende plus Chrysometa clade; in Chrysometa, these scales are reduced to a ridge on the cheliceral margin.

21. Cheliceral dorso-mesal macrosetae: (0) like other macrosetae in the chelicerae (Azilia affin); (1) thicker and with a base larger than the other cheliceral macrosetae (Leucauge argy).

22. Lateral eyes of the male: (0) separate; (1) touching (C90, H95, S97, G98, T01).

23. PME tapeta: (0) absent; (1) present (L80, C90, H95, S97, G98).

24. PLE tapeta: (0) absent; (1) present (L80, C90, H95, S97, G98).

25. Clypeus height: (0) smaller than one AME diameter; (1) equal or larger than one AME diameter; (2) larger than two AME diameter (L80, H95, S97, G98, T01). The last character state was added to account for the similarities of clypeal length found in the ‘reduced piriform clade’.

26. Posterior gut caeca: (0) absent; (1) present (L80, H95, G98).

27. Dorsal femoral trichobothria: (0) present; (1) absent (L80, H95, S97, G98, T01).

28. Femur IV trichobothria arrangement: (0) one irregular line (Tetragnatha versicol as in Tetragnatha labourioLevi, 1981: fig. 17); (1) two parallel rows (Mesida argentiopunctaDavies, 1988: fig. 15).

29. Trichobothria morphology: (0) annulated (Uloborus glomos); (1) smooth (Tetragnatha versicol as in Tetragnatha labourioLevi, 1981: fig. 17); (2) branched (Mesida argentiopunctaDavies, 1988: fig. 15).

30. Femur IV trichobothria extension: (0) less than 1/3 of the femur length (Tetragnatha versicol as in Tetragnatha labourioLevi, 1981: fig. 17); (1) more than 1/3 on the femur length (Leucauge venustaLevi, 1980: fig. 51) (T01).

31. Tibia IV: (0) without tufts of setae on the distal end (Leucauge venustaLevi, 1980: fig. 50); (1) with tufts of setae on the distal end (Nephila clavipesLevi, 1980: fig. 31).

32. Patella-tibia autospasy: (0) absent; (1) present (H95, S97, G98).

33. Sustentaculum: (0) absent; (1) present (H95, S97, G98).

34. Tarsus IV theridiid comb(34): (0) absent; (1) present (C90, H95, G98).

35. Size of the male: (0) > 0.5 female; (1) < 0.4 female (H95, S97, G98).

36. Palpal patella macrosetae of the male: (0) one (1) none (2) two (H95, S97, T01).

37. Male palpal tibial distal margin: (0) smooth (Leucauge venustaLevi, 1980: figs 56–59); (1) with apophyses (Chrysometa acinosaFig. 21G, H).

38. Cymbium apical margin: (0) entire; (1) constricted (H95, G98).

39. Cymbium orientation: (0) dorsal; (1) mesal (H95, S97, G98).

40. Cymbial distal margin: (0) smooth (Leucauge venustaLevi, 1980: figs 55, 57); (1) with a process (Metine from Australia Davies, 1988: fig. 16).

41. Cymbial base: (0) smooth (Leucauge venustaLevi, 1980: figs 56–59); (1) with a process (Metellina curtisiFig. 7F). This character would correspond to the paracymbium secondary process of Hormiga et al (1995; Fig. 6C). The primary homology statement of this character was modified because this process originates at the base of the cymbium and not from the paracymbium (Kuntner & Alvarez-Padilla, 2006). This primary homology hypothesis has other possible interpretations, for example Levi (1980: fig. 124) considered this process and the paracymbium as a single unit, the paracymbium.

42. Cymbial basal process shape; (0) massive (Mollemeta edwardsiFig. 7C); (1) long spine with cuticle different from cymbium (Allende puyehuensisFig. 3I); (2) small roughly triangular (Chrysometa alajuelaFig. 5F); (3) covered with denticles (Allende puyehuensisFig. 5C); (4) slightly protruding and with several spines (Nanometa sp. Davies, 1988: fig. 17); (5) short spine with the base enlarged (Tylorida striaDavies, 1988: fig. 13); (6) long triangular spine (Metabus ocellatusFig. 3C); (7) short spine on a ridge (Diphya palliTanikawa, 1995: fig. 23).

43. Cymbial basal process and paracymbium relative position: (0) basal (Meta ovalisLevi, 1980: figs 112–127); (1) distal (Allende puyehuensisFig. 5D); (2) paracymbium not originating at the cymbial basal process (Metabus ocellatusFig. 3B, C).

44. Cymbial basal process modifications: (0) cymbial base slightly swollen (Tylorida striaDavies, 1988: fig. 13); (1) short protuberances (Metellina curtisiFig. 7F); (2) one long spine (Mollemeta edwardsiFig. 7C); (3) long apophysis covered with denticles (Allende longipes species Fig. 18P); (4) several spines of variable length (Nanometa sp. Davies, 1988: fig. 17).

45. Cymbial basal process position: (0) dorsal (Metabus ocellatusFig. 3B, C); (1) ectal (Metellina curtisiFig. 7F).

46. Cymbial ectal edge median portion: (0) smooth (Metellina curtisiFig. 7F); (1) with a process (Allende puyehuensisFig. 5B).

47. Cymbial median process shape: (0) short (Mollemeta edwardsiFig. 7C); (1) long (Allende puyehuensisFig. 5B); (2) globular (Allende longipesFig. 18N).

48. Cymbial median process macrosetae: (0) none (Mollemeta edwardsiFig. 7C); (1) like the macrosetae on the cymbium (Chrysometa alajuelaFig. 5F); (2) enlarged (Allende puyehuensisFig. 5C).

49. Cymbial median process base diameter: (0) gradually increasing diameter towards its base (Chrysometa alajuelaFig. 5F); (1) abruptly enlarged (Allende puyehuensisFig. 5C).

50. Paracymbium: (0) absent; (1) integral; (2) intersegmental; (3) articulated (L80, C90, H95, S97, G98).

51. Paracymbium distal margin: (0) smooth (Metabus ocellatusFig. 3B); (1) with a triangular sclerotized growth (Allende longipesFig. 18P).

52. Degree of sclerotization of the paracymbium base: (0) sclerotized like cymbium; (1) less sclerotized (H95).

53. Paracymbium morphology: (0) short basal structure, hook-shaped; (1) longer than wide and finger-like; (2) flat and roughly rectangular; (3) U-shaped; (4) long projection of the basal half of the cymbium continuous with the retrolateral margin; (5) flat and roughly triangular; (6) very large and broader than width of cymbium (H95, G98).

54. Paracymbium apophysis: (0) absent; (1) anterior; (2) folding of margin; (3) basal; (4) several apophyses (H95).

55. Subtegulum and tegulum relative position: (0) subtegulum located under the tegulum; (1) ectally displaced. The character state one is found only in ‘leucaugines’ (Fig. 3E, F).

56. Tegular sclerites: (0) subterminal; (1) apical (H95).

57. Tegulum ventral surface: (0) not enlarged; (1) ventrally enlarged. The tegulum in many ‘leucaugines’ is ventrally enlarged; the best example is Tylorida stria (Davies, 1988: fig. 13). The tegulum of M. ocellatus is less swollen (Fig. 3A).

58. Sperm reservoir (sperm duct): (0) normal; (1) enlarged (C90, H95, G98).

59. Sperm duct: (0) with fewer than four turns; (1) more than four turns. In many ‘leucaugines’ including Azilia (Levi, 1980: figs 301–303) the sperm duct makes several coils and turns. One of these coils may correspond with the switchback of C90; this character codes all the extra turns. This character is coded for Metabus (Fig. 12L), and some Nephilines (M. Kuntner, pers. comm.).

60. Reservoir course (sperm duct): (0) spiraled; (1) with a switchback (C90, H95, T01).

61. Median apophysis: (0) present; (1) absent (C90, H95, S97, G98, T01).

62. Conductor and embolus: (0) separate; (1) conductor wraps embolus (C90, H95, S97, G98, T01).

63. Conductor origin: (0) near embolus (Phonognatha graeffHormiga et al, 1995: fig. 8G); (1) ventral margin of the tegulum (Metabus ocellatusFig. 3A); (2) centre of the tegulum (Tetragnatha versicol as in Tetragnatha labourioLevi, 1981: fig. 19); (3) dorsal margin of the tegulum (Allende nigrohumeralisFig. 19J); (4) tegular rim (Mangora gibberoLevi, 1975: figs 126, 127).

64. Conductor embolus coverage: (0) fewer than 1/3 of the embolus (Mollemeta edwardsiFig. 7D); (1) entire (Metabus ocellatusFig. 3A, D); (2) between 1/3 and 2/3 of the embolus (Metellina curtisiLevi, 1980: figs 84–86).

65. Conductor tip: (0) lamelliform and without ornamentations (Mangora gibberoLevi (1975): figs 126, 127); (1) with several lobes and a distal hook (Allende patagiatusFig. 17K).

66. Conductor apex: (0) straight (Mollemeta edwardsiFig. 7A); (1) curved apically (Metabus ocellatusFig. 3A).

67. Sigmoid distal end of the conductor: (0) absent; (1) present (H95, G98).

68. Embolus tegulum orientation: (0) parallel; (1) 90 ° (H95, G98).

69. Araneid radix: (0) absent; (1) present (C90, H95, S97, G98).

70. Stipes: (0) absent; (1) present (H95, S97).

71. Distal haematodocha: (0) absent; (1) present (H95, S97).

72. Metine embolic apophysis: (0) absent; (1) present (C90, H95, S97, G98, T01).

73. Embolus length: (0) fewer than 2 times the embolus base length (Metellina curtisiHormiga et al, 1995,Fig. 13E); (1) more than 2 times the embolus base length (Nephila clavipesHormiga et al, 1995Fig. 10C).

74. Embolic basal apophysis attachment: (0) membranous (Allende nigrohumeralisFig. 19J); (1) sclerotized (Metellina curtisiLevi, 1980: figs 97, 98).

75. Embolic basal apophysis shape: (0) several apophyses (Mangora gibberoLevi (1975): fig. 129); (1) single massive sclerite (Metellina curtisiFig. 7F); (2) long and slender (Dolichognatha pentagoLevi, 1981: fig. 13); (3) hook shaped (Allende nigrohumeralisFig. 19M); (4) coiling with embolus (Chrysometa alajuelaFig. 5F); (5) short spine (Chrysometa acinosaFig. 21F).

76. Embolic apophysis apex: (0) straight (Dolichognatha pentagoLevi, 1981: fig 13); (1) curved (Allende nigrohumeralisFig. 19M).

77. Embolus tegulum membrane: (0) absent; (1) present (H95, G98).

78. Theridiid tegular apophysis: (0) absent; (1) present (C90, H95, G98).

79. Epigynum: (0) present; (1) absent (H95).

80. Fertilization ducts: (0) present; (1) absent (L80, C90, H95, S97, G98).

81. Copulatory openings: (0) divided by a plate (Allende longipesFig. 6B); (1) within a sclerotized atrium (Metabus ocellatusFig. 4A); (2) within a membranous sack (Tetragnatha versicolLevi, 1981: figs 92, 93).

82. Fertilization ducts length: (0) less than half the spermatheca length (Allende longipesFig. 6C); (1) more than half the spermathecae length (Metabus ocellatusFig. 12E).

83. Fertilization ducts path: (0) straight (Allende longipesFig. 6C); (1) coiled (Metabus ocellatusFig. 12E).

84. Spermathecae: (0) oval (Allende longipesFig. 6C); (1) subdivided in two lobes (Nanometa sp. Davies, 1988: fig. 17).

85. Spermatheca surface: (0) sclerotized (Allende longipesFig. 6C); (1) membranous (Metabus ocellatusFig. 4C).

86. Epigynum ventral plate: (0) truncate to gently rounded (Metabus ocellatusFig. 4C).; (1) with a scape (Mangora gibberoLevi (1975): fig. 120).

87. Copulatory ducts: (0) free between the spermathecae (Metabus ocellatusFig. 4C); (1) within a membranous sack (NanometaDavies, 1988: fig. 17).

88. Epigynum tip: (0) smooth (Meta ovalis as Meta menardiLevi, 1980: fig. 117); (1) with two invaginations (Allende longipesFig. 6B); (2) invaginations rebordered (Allende patagiatusFig. 17C, D).

89. Epigynal caudal plate: (0) smooth (Allende longipesFig. 18H); (1) with distal outgrowths (Allende patagiatusFig. 17C–E). The epigyna of Allende species is formed by two plates. A dorsal plate that covers the anterior part of the spermathecae, and a caudal plate that supports the spermathecae and bears the genital openings.

90. Epigynal opening shape: (0) like a spiracle (Glenognatha foLevi, 1980: fig. 274); (1) excavated longer than wide (Metabus ocellatusFig. 4A); (2) longer than wide with the anterior edges rounded (Metabus conacytFig. 15B); (3) protruding (Leucauge argyLevi, 1980: fig. 63); (4) flat (Metellina curtisiLevi, 1980: fig. 78); (5) longitudinal atria (Nephila clavipesLevi, 1980: figs 17, 18); (6) excavated wider than long (Metabus debilisisFig. 13B).

91. Fertilization ducts origin: (0) posterior (Mollemeta edwardsiFig. 8D); (1) anterior (Chrysometa alajuelaFig. 8F).

92. Fertilization ducts and spermathecae: (0) separated (Mollemeta edwardsiFig. 8D); (1) coiling around it (Chrysometa saladitoFig. 6F).

93. Web posture: (0) extended legs 1 and 2; (1) flexed. (L80, C90, H95, S97, G98).

94. Web architecture: (0) orb; (1) sheet; (2) gum foot. (C90, H95, S97, G98).

95. Hub against substrate: (0) absent; (1) present. (H95).

96. Hub bite-out: (0) absent; (1) present. (C90, H95, G98).

97. Hub: (0) closed; (1) open. (L80, C90, H95, G98).

98. Hub loop-no sticky spiral shift: (0) gradual; (1) abrupt. (H95).

99. Radii construction: (0) radii singly attached; (1) radii attached twice. (C90, H95, G98).

100. Radii construction: (0) radii not cut and reeled; (1) radii cut and reeled. (C90, H95, G98).

101. Non sticky spiral: (0) removed from finished web; (1) remains in finished web. (H95, S97, G98).

102. Non sticky contact in 1st sticky spiral construction: (0) present; (1) absent. (C90, H95).

103. Sticky spiral location: (0) oL1; (1) iL1; (2) oL4. (L80, C90, H95, S97; G98).

104. Wrap-bite attack: (0) present; (1) absent. (C90, H95, G98).

105. Web frame; (0) 2D; (1) 3D. (C90, G98).

APPENDIX 2

Data matrix used for cladistic analysis

Uloborus glomos(Walckenaer, 1842)	0000000100	0–000––100	00000?0000	0000000000
Araneus diademat Clerck, 1757	0010000000	010000–100	0111001–––	0010020010
Argiope argenta(Fabricius, 1775)	0010000000	010000–100	0111101–––	0010100010
Mecynogea lemnisca(Walckenaer, 1842)	0010000000	010000–100	01110?1–––	00100000?0
Gasteracantha cancriform(Linnaeus, 1758)	0010110000	110?00–102	01110?1–––	00001100?0
Micrathena gracil(Walckenaer, 1805)	001011000?	???000–100	01110?1–––	00001000?0
Mangora gibbero(Hentz, 1847)	001000000?	???000–100	01110?1–––	00100000?0
Linyphia triangular(Clerck, 1757)	0000001111	100110–000	0111201–––	0100000000
Pimoa altiocula(Keyserling, 1886)	0000001111	10010––000	01112?1–––	0100000000
Steatoda america(Emerton, 1882)	0000001110	101000–002	01112?1–––	0001000000
Nesticus cellulan(Clerck, 1757)	0010001010	101000-102	01112?1–––	0001000000
Epeirotypus brevip O. P.-Cambridge, 1894	0000001010	10000––100	01111?1–––	0000010000
Phonognatha graeff(Keyserling, 1865)	0010000010	100000–100	01110?1–––	0000000000
Clitaetra episinoid Simon, 1889	0000000010	1100010100	00000?1–––	0000000000
Nephila clavipes(Linnaeus, 1767)	1010000010	1100011100	01111?1–––	1000100000
Herennia ornatissi(Doleschall, 1859)	1010000010	1100011100	01101?1–––	0000100000
Nephilengys malabarens(Walckenaer, 1842)	1010000010	1100011100	01101?1–	0000100000
Tetragnatha versicol(Walckenaer, 1842)	0100000010	100010–000	0000110010	0000000100
Glenognatha fo(McCook, 1894)	0001000011	100010–001	01012?0010	0000010100
Pachygnathaha autumnal Marx, 1884	0001000111	100010–000	0101210010	0000010100
Agriognatha sp.	0100000010	1?0010–001	01??1?0021	0000000100
Azilia affin O. P.-Cambridge, 1893	0000000000	100000–010	00000?1–––	0000000000
Leucauge venusta(Walckenaer, 1842)	0100000010	100000–000	1111110121	0000000000
L. argy(Walckenaer, 1842)	010000??1?	1?0000–000	11111?0121	00000000?0
Orsinome nr. vet(Hasselt, 1882)	010000????	1?0010–000	01111?0010	00000000?0
Metabus ocellatus(Keyserling, 1864)	0100000010	100000–000	01111?1–––	00000000?0
M. debilis(O. P.-Cambridge, 1899)	010000??1?	1?0000–000	01111?1–––	00000000?0
M. ebanover	010000??1?	1?0000–000	01111?1–––	00000000?0
M. conacyt	010000??1?	1?0000–000	01111?1–––	00000000?0
Mesida argentiopuncta(Rainbow, 1916)	010000??1?	1?0000–012	01111?0121	00000000?0
Opadometa sp.	010000??1?	1?0000–010	01111?0121	10001100?0
Tylorida stria(Thorell, 1877)	010000??1?	1?0000–000	01111?0010	00000100?0
Meta menardi(Latreille, 1804)	0010000000	100000–000	0111101–––	0000000000
Mollemeta edwardsi(Simon, 1904)	0000000010	110000–000	01111?1–––	00000000?0
Metellina curtisi(McCook, 1894)	0000000010	100010–000	01111?1–––	0000000000
Dolichognatha pentago(Hentz, 1850)	0000000010	100000–012	00000?1–––	0000010000
C. alajue Levi, 1986	0100000010	100000–001	01111?1–––	00000000?0
C. nubo Levi, 1986	0100000010	100000–001	01111?1–––	00000000?0
C. saladi Levi, 1986	0100000010	100000–001	01111?1–––	00000000?0
A. nigrohumeralis(F. O. P.-Cambridge, 1899)	0000000010	110000–001	01111?1–––	00000000?0
Allende puyehuensis	0000000010	110000–001	01111?1–––	00000000?0
A. patagiatus(Simon, 1901)	0000000010	110000–001	01111?1–––	00000000?0
A. longipes(Nicolet, 1849)	0000000010	110000–001	01111?1–––	00000000?0
Nanometa sp.	010000??1?	1?0000–000	01111?1–––	00000000?0
Metine from Australia	000000??1?	1?0000–001	01111?1–––	00000000?1
Orsinome sarasini Berland, 1924	000000??1?	1?0000–001	01111?1–––	00000000?1
Chrysometa acinosa	000000??1?	1?0000–001	01111?1–––	00000011?0
C. lev	0??000????	???0?0–0?1	01111?1–––	???0?001?0
C. maitae	?00000??1?	1?0??0–0??	0???1?1–––	0?00??????
Diphya sp.	000000??1?	1?0000–002	00002?1–––	00000011?0
0–––-0–––0	–––-000000	0040000000	000–––000	0011010000	3000000--0	000000
0–––-0–––1	1000000000	0040000011	110000100	0000001000	3??1001000	100000
0–––-0–––1	0000000000	0040000011	000–––100	0000000000	3001001000	100000
0–––-0–––1	0000000000	0040000011	101–––100	0100000000	3??100?00?	?1???1
0–––-0–––1	0001000000	0040000010	000–––100	0000001000	300100?01?	10?0?0
0–––-0–––1	0001000000	0040000010	000–––100	0000001000	3??100?11?	10?0?0
0–––-0–––1	1000000000	0040000011	110000100	0000001000	3001001000	1000?0
0–––-0–––2	0-30010000	1–––00–000	010000100	0111000000	30011–––--	–––-11
0–––-1–––1	0050000001	00?0000000	000–––000	0000000000	31011–––--	–––-11
–––--0–––0	–––-000000	0041000000	000–––010	0000100000	40012–––--	–––-11
1–––-0–––1	0064000001	0040000000	000–––010	0000000100	31012–––--	–––-?1
0–––-0–––1	0000000001	0031000000	000–––000	0100000000	3??1001010	101111
0–––-0–––1	0042000000	1101000000	001–––100	0000100000	4001010000	?10210
0–––-0–––1	0122010011	1111000100	001–––100	0000000000	400?01??1?	?????0
0–––-0–––1	0122010011	1111000100	001–––100	0000000000	5001000001	01?210
0–––-0–––1	0122010011	1111001100	001–––100	0000000000	4001010001	01?210
0–––-0–––1	0122010011	1111001100	001–––100	0000000000	4001010001	011210
0–––-0–––3	0-11010100	1121000000	000–––101	12--10–0--	0--0001110	101110
0–––-0–––3	1-11010100	1121000000	000–––101	12--00–0--	0--0000-?0	10-1?0
0–––-0–––3	1-11010100	1121000000	000–––101	12--00–0--	0--?0011??	?????0
0–––-0–––3	1000010100	1122000000	000–––101	12–––--0--	0--?00?1??	?0???0
0–––-0–––1	0000111011	1–––-0–000	000–––100	0000010000	4000001010	10?1?0
0–––-0–––1	1000111011	1111000000	001–––100	0111010000	1000001110	101100
162000–––1	1000111001	1111000000	001–––100	0000010000	300?00?1??	?????0
172000–––1	10001110?1	1111000000	001–––100	0100010000	400?00?1??	?0???0
162000–––1	0000111011	1111010000	001–––100	0111010000	100?00????	?????0
0–––-0–––1	0000111011	1111010000	001–––100	0111010000	600???????	??????
0–––-0–––1	0000111011	1111010000	001–––100	0111010000	600?00?1??	?0???0
0–––-0–––1	0000111011	1111010000	001–––100	0111010000	100???????	??????
152000–––1	0000111011	1111000000	001–––100	0111010000	400?00?1??	?0???0
152000–––1	1000111001	1111001000	001–––100	0110010000	400?00?1??	?0???0
152000–––1	0000111011	1111000000	001–––100	0111010000	400?00?1??	?0???0
1001110001	0010010001	1132000000	010110100	0000000000	3000001110	101100
1002110001	0010010000	1130000000	000–––100	0000000000	400?01?00?	?0???0
100110–––1	0010010000	1132000000	010110100	0100100000	400000111?	?0???0
101210–––1	0010010001	1132000000	010120100	0110000000	5001001010	1011?0
1210111103	?-64010000	1132000000	010040100	0011000000	411?00????	?0???0
1210111103	?-64010000	1132000000	010040100	0011000000	411?00????	?????0
1210110003	?-64010000	1132000000	011040100	0011000000	411?00?1??	?0???0
1313111203	1-10010000	1132100000	010031100	0000000010	300?00????	??????
1112111213	1-10010000	1132100000	010030100	0000000021	300?00?1??	?0????
1313111203	1-10010000	1132100000	010030100	0000000021	300?00????	??????
1313112213	1-10010000	1132100000	010031100	0000000010	300?00????	??????
1424110001	?010010000	1132000000	000–––100	0100100100	400?00?1??	?0???0
1424110001	?000010000	1132000000	000–––100	0000100100	400?00?1??	?0???0
1424110001	?010010000	1132000000	000–––100	0000100100	300???????	??????
1110112003	1-10010000	1132000000	010150100	0011000000	410?00?1??	?0???0
1210110003	1-10010000	1132000000	01004010?	??????????	??????????	??????
??????????	??????????	?????0????	????????0	0011000000	410?00?1??	?0???0
172000–––1	10010100–0	1122000000	000–––100	0000?00000	410???????	??????

APPENDIX 3

Additional material examined

This appendix includes the specimens used in this study and not mentioned within the Material examined section of each species description. Specimens are listed alphabetically by family, genus and species. Localities are given as shown in museum labels.

Araneidae

Araneus diademat Clerck, 1757

USA: Oregon, Colburg, 2.ix.1968 (USNM).

USA: Massachusetts, Co. Quisset Barnstable, 27.viii.1983, Edwards leg. (USNM).

Argiope argenta(Fabricius, 1775)

MEXICO: Tamaulipas, Industrial area near Altamira, Ejido F. Magon, 13.v.1995, Carlow leg. (USNM).

TRINIDAD: St. George, 2 km E of Blanchisseuse, 21.iii.1985, G. F. & J. F. Hevel leg. (USNM).

Gasteracantha cancriform(Linnaeus, 1758)

DOMINICA: Roseau.x.1967, Krauss leg. (USNM).

PANAMA: Canal Zone, Colon, 2–14.vii.1979, Humid forest Canopy fogging, Broadhead et al leg. (USNM).

Mangora gibbero(Hentz, 1847)

USA: Virginia, Fairfax Co., Bull Run Occoquan Regional Park. vii-ix 2004, Álvarez-Padilla leg. (MCZ).

Mecynogea lemnisca(Walckenaer, 1842)

USA: South Carolina, Oconee. Co., Chattooga River landing just below Rt. 28 bridge, 1580′ elev, 22.vii.1988 (USNM).

USA: South Carolina, Oconee. Co., Yellow Branch Picnic Araea on Rt. 28, 2.9 mi S intersec with Rt. 107, 1300′ elev, 22.vii.1988. Smith leg. (USNM).

Micrathena gracil(Walckenaer, 1805)

USA: Virginia, Fairfax Co., Bull Run Occoquan Regional Park. vii-ix 2004, Álvarez-Padilla leg. (MCZ).

Linyphiidae

Linyphia triangular(Clerck, 1757)

SWEDEN: 14–15.viii.1995, Hormiga leg. (GWU).

Nesticidae

Nesticus cellulan(Clerck, 1757)

USA: North Carolina, Macon Co., Glenn Falls (nr. Highlands), Mixed pine hardwood rhododendron forest, 35°01′N 83°14′W, 22.vi.1991, Hormiga leg. (USNM).

Pimoidae

Pimoa altiocula(Keyserling, 1886)

USA: Washington, Seattle/King Co., Ravenna Park, 100′ elev, beaten from woodland understory, 44.637°N 122.307°W, 10.xi1987, Crawford leg. (USNM).

Theridiidae

Steatoda america(Emerton, 1882)

USA: West Virginia, Berkeley Co., Sleepy Creek Hunt & Fish Area, Third Hill Mtn. Oak-Pine forest, 20–27.vi.1986, Martinat leg. (USNM).

USA: West Virginia, Berkeley Co., Sleepy Creek Hunt & Fish Area, Third Hill Mtn. Oak-Pine forest, 9–17.vii.1986, Martinat leg. (USNM).

Theridiosomatidae

Epeirotypus brevip O. P.-Cambridge, 1894

COSTA RICA: Puntarenasas, Monteverde Cloud Forest, 1500 m, 9.ii.1979, Coddington leg. (USNM).

Tetragnathidae

Agriognatha sp.

COSTA RICA: Puntarenasas, P. N. (ACLA) Estacion Pittier, 9°01′-N 82°5′-W, 1800 m, 8–11.vi.1995, Hormiga leg. (GWU).

Azilia affin O. P.-Cambridge, 1893

USA: Florida, Alachura Co., Dudley Caves, 18.iii.1938 (AMNH).

Chrysometa alajuela Levi, 1986

COSTA RICA: Puntarenasas, P. N. (ACLA) Estacion Pittier, 9°01′-N 82°5′-W, 1800 m, 8–11.vi.1995, Hormiga leg. (GWU). (voucher specimen SEMFAP 046).

Chrysometa nubo Levi, 1986

COSTA RICA: Puntarenasas, P. N. (ACLA) Estacion Pittier, 9°01′-N 82°5′-W, 1800 m, 8–11.vi.1995, Hormiga leg. (GWU).

Chrysometa saladito Levi, 1986

COSTA RICA: Puntarenasas, P. N. (ACLA) Estacion Pittier, 9°01′-N 82°5′-W, 1800 m, 8–11.vi.1995, Hormiga leg. (GWU).

Clitaetra episinoid Simon, 1889

SOUTH AFRICA: Faine's Is., 40–50 m, 31.iii.2001, Hormiga leg. (GWU).

Diphya sp.

CHILE: Región VII Bio-Bio, Ñuble, E. Recinto las Trancas, 1100 m, ii.1987, Peña leg. (AMNH).

CHILE: Región IX de la Araucaria, Caut'n, Chacamo NW of Nueva Imperial W. of Temuco, 16–24.ii.1981, Peña leg. (AMNH).

Dolichognatha pentago(Hentz, 1850)

USA: Mississippi, Forrest Co., Camp Shelby, 1945−46, Archer leg. (AMNH).

Glenognatha fo(McCook, 1894)

USA: Louisiana, Hamburg Co., 29.iv.1963 (AMNH).

Herennia ornatissi(Doleschall, 1859)

THAILAND: Surat Thani Prov., Khao Sok NP, Wing Hin Waterfall trail, N 8°55′0.4′; E 98°31′40.9′, 300 m 19–20.x.2003. ATOL Expedition 2003 leg. (USNM).

Leucauge venusta(Walckenaer, 1842)

USA: Virginia, Fairfax Co., Bull Run Occoquan Regional Park. vii-ix 2004, Álvarez-Padilla leg. (MCZ). (voucher specimen SEMFAP033).

Mesida argentiopuncta(Rainbow, 1916)

AUSTRALIA: Queensland, Bellender Ker Range, NQ Cable Tower 7, 500 m, 1–7.xi.1981, EARTHWATCH/QLD. MUSEUM QMB 27777 (QMB).

Meta menardi(Latreille, 1804)

USA: Connecticut, Litchfield from a cellar, 5.i.1935, Woodbury leg. (USNM).

Metellina curtisi(McCook, 1894)

USA: Washington: Snohomish Co., Crystal Creek, from moss on trees, logs and stumps, 300′, 47.831°N 121.652°E, 24.x.1986, Crawford leg. (USNM).

USA: Oregon: Corvallis (USNM).

USA: California, Sequoia N. P., 4.vi.2002, Álvarez-Padilla leg. (voucher specimen SEMFAP063).

Metine from Australia

AUSTRALIA: Queensland, Lamington NP, tracks near O'Reilly's Rain forest, 28°14′05′′S 153°08′13′′E, 920 m, 13–17.iv.2002, Hormiga, Kuntner & Álvarez leg. (USNM).

Nanometa sp.

AUSTRALIA: Queensland, Bellender Ker Range, NQ Cable Tower 3, 1054 m, x.17–24, 1981. EARTHWATCH/QLD. MUSEUM (QMB).

Nephila clavipes(Linnaeus, 1767)

USA: Florida, Hillsborough Co., Tampa, 1.ix.1984, Larcher leg. (USNM).

Nephilengys malabarens(Walckenaer, 1842)

THAILAND: Surat Thani Prov., Khao Sok NP, Wing Hin Waterfall trail, N 8°55′0.4′; E 98°31′40.9′, 300 m 19–20.x.2003, ATOL Expedition 2003 leg. (USNM).

Opadometa sp.

PAPUA NEW GUINEA: Louisiade Arch. Misima Is, north slopes of Mt. Misima, 350 m, Camp 7. 16–30.vii.1956, Fifth Archbold Expedition, Brass leg. (AMNH).

SOUTH-WEST PACIFIC: 1986−87, Krauss leg. (AMNH).

Orsinome sarasini Berland, 1924

TASMANIA: Scotts Peak, Unnamed cave Cliffline cave site 4, 29.x.1988, Clarke leg. KS21369(AUM).

TASMANIA: Florentine Valley, Punishment Pot JF 373, 1988, Eberhard leg. KS20172 (AUM).

Orsinome nr. vet(Hasselt, 1882)

PAPUA NEW GUINEA: Louisiade Arch. Rossel Is., south slopes of Mt. Rossel, 400 m, Camp 13, 12–20.x.1956, Fifth Archbold Expedition, Brass leg. (AMNH).

Pachygnathaha autumnal Marx, 1884

USA: Massachusetts, Quisset Co., 24.iii.1989, Edwards leg. (USNM).

Phonognatha graeff(Keyserling, 1865)

AUSTRALIA: Queensland, Tamborine NP, Witches Falls Rainforest, 27°56′27′′S 153°10′48′′E, 17.iv.2002, Hormiga, Kuntner, & Álvarez (USNM).

Leucauge argy(Walckenaer, 1842)

USA: Florida, Highlands Co., Archbold Biol. Sta., 11.x.1964, Arnaud Jr. leg. (CAS).

MEXICO: Nayarit, Highway 15 near Acaponeta 7 mi N junction of Highways 15 and 23, 16.x.1973, Williams, Blair & Mullinex leg. (CAS).

Tetragnatha versicol(Walckenaer, 1842)

USA: Virginia, Fairfax Co., Bull Run Occoquan Regional Park. vii-ix 2004, Álvarez-Padilla leg. (MCZ).

Tylorida stria(Thorell, 1877)

NIGERIA: Abeokouta Prov., Ilugun, 11.xii.1948 (AMNH).

PAPUA NEW GUINEA: Kairiru Is., 03°20′-S 143°33′-E, 28.x.1981, Borrell leg. KS66865 (AUM).

Uloboridae

Uloborus glomos(Walckenaer 1842)

USA: Virginia, Fairfax Co., Bull Run Occoquan Regional Park. vii-ix.2004, Álvarez-Padilla leg. (MCZ).