Article. Five new species of lapsiine jumping spiders from Ecuador (Araneae: Salticidae)

Zootaxa 3424: 51 65 (2012) www.mapress.com/zootaxa/ Copyright 2012 Magnolia Press Article ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) Five new species of lapsiine jumping spiders from Ecuador (Araneae: Salticidae) WAYNE P. MADDISON Departments of Zoology and Botany and Beaty Biodiversity Museum, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia, V6T 1Z4, Canada. E-mail: wmaddisn@interchange.ubc.ca Abstract Five new species of lapsiine jumping spiders from Ecuador are described, including the first Lapsias Simon from outside Venezuela. Lapsias lorax, sp. nov. is known from a cloud forest just west of Quito. A new species from the slopes of Volcan Sumaco is tentatively assigned to Lapsias, Lapsias guamani sp. nov. Lapsias canandea, sp. nov. is the first lapsiine described from the lowlands west of the Andes. The genus Lapsias is poorly defined, and some of these new species may merit separate genera when the group's phylogeny is better known. Two new species of Thrandina Maddison are described from about 2000 m elevation, Thrandina cosanga sp. nov. from the eastern slopes of the Andes, and Thrandina bellavista sp. nov. from the western slopes. New illustrations are provided for the already-described Thrandina parocula Maddison. Photographs of living individuals are presented for all species. Key words: Araneae, Salticidae, lapsiines, jumping spider Introduction Lapsiine jumping spiders are distinctive within the neotropical fauna, falling outside the major salticid clade Salticoida and having several ancestral traits such as a median apophysis on the male palp and a tarsal claw on the female palp (Maddison, 2006; Maddison & Needham, 2006; Ruiz & Maddison, 2012). Only fourteen species in four genera have been described: four species of Lapsias Simon from Venezuela (Platnick, 2011), one of Thrandina Maddison and two of Galianora Maddison from Ecuador (Maddison, 2006), and seven species of Soesiladeepakius Makhan from Brazil and Surinam (Ruiz & Maddison, 2012). I here describe five new lapsiine species from two recent expeditions to Ecuador: three species of Lapsias, the first from outside Venezuela, and two species of Thrandina. Material and methods Figure 1 shows the localities from which the specimens here described were sampled. All specimens are deposited in the Spencer Entomological Museum of the University of British Columbia (UBC-SEM) or the Museum of Zoology, Pontificia Universidad Católica, Quito, Ecuador (QCAZ). Photographs of living specimens were taken with a Pentax Optio 33WR digital camera with a small lens glued to it for macro capability; photographs of bodies in alcohol were compiled by Helicon Focus from photographs of different focal planes taken with a Nikon D7000 camera. Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope. Terminology is standard for Araneae. All measurements are given in millimeters. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; PLE, posterior lateral eyes; PME, posterior median eyes (the "small eyes"). Accepted by T. Szuts: 22 Jun. 2012; published: 15 Aug. 2012 51

FIGURE 1. Ecuador, with collecting localities for lapsiine species described here. The southern locality for Thrandina parocula was reported by Maddison (2006). Map based on http://en.wikipedia.org/wiki/file:ecuador_topography.png, in the public domain. Taxonomy Genus Lapsias Simon The four described species of Lapsias (Simon, 1900, 1901; Galiano, 1963; Platnick, 2011) share a relatively short embolus that arises distally on the prolateral side, a delicate hook-like median apophysis, an elongate tegulum, small PME's, and body proportions typical of most salticids (i.e. a reasonably high carapace and short legs). These features distinguish Lapsias from Thrandina (which has large PME's and an expanded median apophyses), Galianora (which has a round and rotated tegulum with adpressed embolus) and Soesiladeepakius (which is flatter-bodied and whose median apophysis is fused to the tegulum or perhaps absent). However, all of the listed features of Lapsias are arguably plesiomorphies or widespread enough to be uninformative. For instance, the median apophyses of G. sacha Maddison and the Old World hisponines are also delicate hooks. Although the described Lapsias species are quite similar to one another in palpi and body form, as yet no clear synapomorphies have been proposed for the genus. Because of the lack of definition of Lapsias, I place three new species there with some hesitation. Lapsias lorax has proposed synapomorphies with two of the Lapsias species, L. tovarensis Simon and the type species L. estebanensis Simon (see below). While Lapsias could have been restricted to include just those three species, this would leave L. ciliatus Simon and L. cyrboides Simon without a place. The other two new species are difficult to place. L. canandea is quite distinctive, having a flattened body, a long embolus and a long hook-like conductor. It shares a delicate hook-like median apophysis with Lapsias, and it has no features that would link it to any of the 52 Zootaxa 3424 2012 Magnolia Press MADDISON

other lapsiine genera. L. guamani is of unclear relationships, known only from a single female. While new genera could have been erected for L. canandea and L. guamani, it seems more prudent to expand an existing mess (Lapsias) rather than generate more nomenclatorial clutter, at least until the relationships are understood. With L. canandea added to Lapsias, the genus can be characterized by the hook-like median apophysis and lack of a round bulb with adpressed embolus. Lapsias lorax, sp. nov. (Figs 2 9) Type material. Holotype male in QCAZ, temporarily held at the UBC-SEM, with data: "ECUADOR: Pichincha: Bellavista Cloud Forest Reserve, Discovery Trail. S 0.01462 W 78.68326. 2240 m elev. 9 November 2010. W Maddison. WPM#10-067", "Male W624 Photo'd 9 Nov #ECU2010-3221", "UBC-SEM AR00194" Etymology. Named after Theodor Geisel's fictional character The Lorax, a small creature with yellow mustache who advocated for the preservation of trees. This name was suggested by Tristan Long to refer to the bars of yellow setae on the chelicerae (appearing as a mustache) and to reflect on the importance of preserving forests such as those where this spider was found. Diagnosis. The placement of the embolus, conductor and median apophysis toward the distal tip of the palp is more extreme than in other Lapsias. Accordingly, the straight portion of the sperm duct along the prolateral margin of the tegulum stands out as being particularly long, before the duct turns retrolaterally to meet the median apophysis. The yellow bars of setae across the chelicerae appear to be unique among Lapsias, although Simon's specimens of the other species are old and there is the possibility that the setae may have been lost (type series of L. estebanensis, L. tovarensis, L. ciliatus, and L. cyrboides, in MNHN Paris examined). Notes. Two possible synapomorphies with the type species, Lapsias estebanensis, could be proposed as justification for placing this new species in the genus. The first is the very short embolus (shared also with L. tovarensis, but also with the genus Soesiladeepakius); the second is the migration of the retrolateral tibial apophysis dorsally to approach a second dorsal apophysis (Figs. 9 10). In L. estebanensis, L. tovarensis, and L. lorax, a dorsal view of the palp tibia shows a V-shaped cleft framed by two ridges, the lateral representing the retrolateral tibial apophysis, and the medial representing a dorsal apophysis. This cleft is not apparent in L. ciliatus or L. cyrboides. In body form and markings, L. lorax is quite similar to L. tovarensis. Description. Male (holotype). Carapace length 2.0; abdomen length 2.1. PME small. Chelicerae not particularly enlarged, with 4 promarginal and 2 retromarginal teeth, closely placed (more or less fissident). Palpus (Figs 7 9) with short terminal embolus and small, thin median apophysis which is hooked at the tip. Accompanying the median apophysis is a membranous fold that probably is the conductor. Tibial apophysis short, shifted dorsally. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 4 prolateral and three retrolateral macrosetae. Colour (Figs 2 6): honey-coloured (in alcohol, honey-cream) with dark brown markings. Thoracic area dark except two pale side bars and a central pale stripe; ocular quadrangle brown. Clypeus pale, with white setae. Chelicerae black in front, with a distinct oblique yellow band of setae just below the clypeus (Fig. 2). Femur of palp mostly pale; patella, tibia and cymbium dark. Femora of legs pale except dark spots dorsally. Patellae and tibiae orange-brown. Metatarsi and tarsi with dark termini, especially on legs 3 and 4. Abdomen medium brown, with chevrons. Natural history. The single male was found by shaking lianas climbing within moss covering a tree trunk in cloud forest. Extensive subsequent searching for one day in this microhabitat yielded many Thrandina bellavista (4 adults and 23 juveniles), but no more specimens of Lapsias lorax. This may indicate the species is uncommon, or that the location where it was found was not its core habitat it may, for instance, live primarily in the canopy or at higher elevation. When walking, the second pair of legs is waved periodically, as seen in Thrandina and many other basal salticids (Maddison, 2006, 2009). A video of the living holotype is available at http://www.youtube.com/watch?v=tgf2jcdr_88. NEW SPECIES OF LAPSIINE JUMPING SPIDERS FROM ECUADOR Zootaxa 3424 2012 Magnolia Press 53

FIGURES 2 8. Lapsias lorax, sp. n. 2 6 holotype male 7 left male palp, ventral view 8 left male palp, retrolateral view. Scale bar = 0.1 mm. Figures 2 8 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. 54 Zootaxa 3424 2012 Magnolia Press MADDISON

FIGURES 9 10. Male palp tibia of Lapsias species, dorsal view. 9 Lapsias lorax, sp. n. holotype 10 Lapsias estebanensis Simon, type. Scale bar = 0.1 mm. Figures 9 10 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. Lapsias canandea, sp. nov. (Figs 11 20) Type material. Holotype male in QCAZ with data: "ECUADOR: ESMERALDAS: Reserva Canandé, nr Cuckoo Trail. N 0.5210 W 79.2081 to N 0.5204 W 79.2067, 460 500 m el. 23.Aug.2011 W. MaddisonWPM#11-188", "Photo'd 25 Aug #ECU2011-9737", "UBC-SEM AR00191" Etymology. Derived from the type locality, Reserva Canandé. To be treated as an arbitrary combination of letters, and hence without the need to agree in gender with the genus. Diagnosis. The palp with prolaterally-arising embolus and two hooked apophyses is distinctive (Fig 11), as are the fringed legs of the male (Fig 17). No other lapsiine has such a large, sclerotized conductor. The body is somewhat elongate and flat, resembling a large Soesiladeepakius, or a darker Galianora sacha. The anterior guide of the epigynum of L. canandea distinguishes it from most lapsiines except some females in the type series of Lapsias cyrboides (see Galiano 1963: Plate XXV, figure 12; other females in the type series are of a different species, without anterior guide; which females pair with the holotype male is unclear; in MNHN Paris, examined). Description. Male (holotype). Carapace length 2.1; abdomen length 2.5. Carapace: Fig. 16. PME small. Chelicerae not noticeably enlarged compared to female, with 3 small promarginal and 2 small retromarginal teeth. Palpus (Figs 11 12) with bulb rotated so that embolus arises retrolaterally and revolves about 300 around the tegulum. Two hook-shaped processes arise from the tegulum, a larger one that arises almost in the middle (interpreted as the conductor), and a smaller one behind it that arises along the retrolateral distal edge (interpreted as the median apophysis). The larger apophysis has no visible separation from the tegulum, but the smaller is clearly separated from the tegulum by a membrane, like the median apophyses of many other lapsiines and cocalodines (Maddison, 2006, 2009). Following the sperm duct of L. canandea as it approaches the embolus, the first of the two apophyses encountered is the smaller; later, as the sperm duct loops inward, it approaches the larger apophysis. This also suggests that the smaller apophysis is the median apophysis, because in other lapsiines and cocalodines NEW SPECIES OF LAPSIINE JUMPING SPIDERS FROM ECUADOR Zootaxa 3424 2012 Magnolia Press 55

the sequence following the sperm duct is median apophysis, conductor, embolus (Maddison, 2006: Figs. 1, 2; Maddison, 2009: Figs. 26, 37, 57). Cymbium with retrolateral side excavate. Retrolateral tibial apophysis projecting as a small keel that might be mistaken for a paracymbium (Figs 11, 12, rta). Two small dorsal tibial apophyses. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 3 pairs. Colour (Figs 15 20): Body medium to dark brown, with orange tones. Thorax slightly paler in band along lower margin and central longitudinal band. Clypeus dark brown, glabrous except for a few setae. Chelicerae dark brown to black. Abdomen with trace of chevrons. Segments of palp paler except for cymbium, which is dark brown to black. White scales on patella and tibia. First legs darkest, with femur, patella and tibia dark brown to black. Patella and tibia with small dorsal and longer ventral fringe of black setae. Legs 2 4 honey-coloured in alcohol, with indistinct darker areas at the ends of the tibiae and metatarsi. FIGURES 11 14. Lapsias canandea, sp. n. 11 male palp, ventral view (rta = retrolateral tibial apophysis) 12 male palp, retrolateral view 13 epigynum, ventral view 14 epigynum, cleared, dorsal view. Scale bar = 0.1 mm. Figures 11 14 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. Female (paratype in UBC-SEM, #UBC SEM AR00192, from ECUADOR: ESMERALDAS: Reserva Canandé, Quail Trail. N 0.5255 W 79.2069 to N 0.5265 W 79.2118, ~330 m el. 17.Aug.2011 W. Maddison WPM#11-135). Carapace length 2.0; abdomen length 2.1. Carapace shape as in male. PME small. Chelicerae with 3 promarginal and 2 retromarginal teeth. Palp with tarsal claw. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 3 pairs. Epigynum (Fig. 13 14) with a distinctive guide anteriorly. Paired openings hidden behind dark flaps, about half way between hood and posterior margin (arrow, Fig. 13). Internally the ducts proceed posteriorly, then laterally, then loop back to return medially and then anteriorly to the fertilization duct. The lateralmedial loop is twisted into a spiral. Colour (Figs 19 20): much like the male, though the brown is duller (with gray rather than orange tones), and the first legs lack the fringe. Some females are paler and grayish. 56 Zootaxa 3424 2012 Magnolia Press MADDISON

FIGURES 15 20. Lapsias canandea, sp. n. (15 18) male 15, 17 male from Quail Trail (locality WPM#11-131) 16, 18 holotype male (19 20) female from Quail Trail (locality WPM#11-129). Figures 15 20 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. NEW SPECIES OF LAPSIINE JUMPING SPIDERS FROM ECUADOR Zootaxa 3424 2012 Magnolia Press 57

Additional material examined. 16 males, 9 females and 40 juveniles, in UBC-SEM, all from ECUADOR and in UBC-SEM: ESMERALDAS: Reserva Canandé, 17 23 August 2011. Details: N 0.5254 W 79.2080 to N 0.5252 W 79.2076, ~300 m el., WPM#11-129 (1 female); N 0.5253 W 79.2073 to N 0.5255 W 79.2069, ~330 m el., WPM#11-131 (1 male); N 0.5261 W 79.2119, ~330 m el., WPM#11-132 (1 juv.); N 0.5261 W 79.2119, ~320 m el., WPM#11-133 (2 juv.); N 0.5220 W 79.1974 to N 0.5221 W 79.1969, ~580 m el., WPM#11-142 (2 male, 1 juv.); N 0.5222 W 79.1961 to N 0.5223 W 79.1958, ~580 m el., WPM#11-144 (1 female, 2 juv.); N 0.5220 W 79.1954 to N 0.5217 W 79.1951, ~580 m el., WPM#11-146 (1 male, 1 female, 2 juv.; male is designated as paratype, with additional data "Photo'd 18 Aug #ECU2011-8164"); N 0.5279 W 79.2036 to N 0.5274 W 79.2088, ~300 m el., WPM#11-153 (1 male, 3 juv.); N 0.5274 W 79.2088 to N 0.5270 W 79.2089, ~300 m el., WPM#11-154 (2 juv.); N 0.5283 W 79.2059 to N 0.5286 W 79.2062, ~280 m el., WPM#11-158 (1 juv.); N 0.5171 W 79.1935 to N 0.5175 W 79.1935, ~580 m el., WPM#11-160 (4 male, 6 juv.); N 0.5175 W 79.1935 to N 0.5179 W 79.1936, ~590 m el., WPM#11-161 (2 females, 1 juv.); N 0.5179 W 79.1937 to N 0.5184 W 79.1939, ~560 m el., WPM#11-162 (2 females); N 0.5184 W 79.1939 to N 0.5187 W 79.1942, ~580 m el., WPM#11-163 (1 male); N 0.5167 W 79.1934, ~580 m el., WPM#11-165 (1 male); N 0.5217 W 79.1951, ~580 m el., WPM#11-166 (3 male 1 female 4 juv.); N 0.5193 W 79.1946, ~550 m el., WPM#11-168 (1 male, 9 juv.); N 0.5214 W 79.2033 to N 0.5214 W 79.2036, ~530 m el., WPM#11-174 (1 male); N 0.5214 W 79.2036 to N 0.5216 W 79.2040, ~540 m el., WPM#11-175 (2 juv.); N 0.5202 W 79.2076 to N 0.5198 W 79.2077, ~480 m el., WPM#11-184 (1 female, 4 juv.). Natural history. This species was relatively common in the leaf litter at Reserva Canandé. It was rarely seen walking on top of the litter, but was usually collected by quickly moving leaf litter onto a beating sheet, on which the specimens would be seen running quickly toward the edge. They could be found commonly in leaf litter not only on the ground throughout the forest, but also in litter suspended up to 2 m above ground, especially in tree fall clearings. It appeared that when on the ground, to house L. canandea the litter needed to be loose and well-drained. Their breadth of habitat is unusual; in my experience salticid species typically live either in suspended litter, or on ground litter, but not both. Indeed, L. canandea females and juveniles were occasionally found beating vegetation, though it is possible in all such cases the vegetation held some suspended litter. It has previously been noted that many basal salticids have an unusual gait that includes waving of the second pair of legs (see notes under other Lapsias species here, and W. Maddison, 2009). L. canandea was never seen to wave the second pair of legs, but nonetheless it does have the fluid gait typical of basal salticids, lacking the pulsed gait of salticoids. Indeed, the first specimen of L. canandea found was seen walking on leaf litter, and it was recognized immediately as a basal salticid based on its gait. In this regard, L. canandea is like Galianora sacha, which also has the fluid gait but has not been seen to wave the second legs (W. Maddison, unpublished observations). Videos of living L. canandea specimens are available: male, http://www.youtube.com/watch?v=pywzwquhivu and http://www.youtube.com/watch?v=zu81v8ttmng; female, http://www.youtube.com/watch?v=t8em-lbezvq. Lapsias guamani, sp. nov. (Figs 21 24) Type material. Holotype female in QCAZ, temporarily held at the UBC-SEM, with data: "ECUADOR: Napo: Río Guamani on Jondachi-Loreto road. S 0.7223 W 77.6408. 1050 m elev. 31 October 2010. D Maddison. WPM#10-038", "Photo'd 31 Oct #ECU2010-1627", "UBC-SEM AR00193" Etymology. The name refers to the type locality. Diagnosis. The epigynum with an overhanging projection resembling a human epiglottis is distinctive. This is also the smallest Lapsias known. Notes. This species is placed provisionally in Lapsias, as its body form is unremarkable (like Lapsias) and there is no other genus where it is more justified. Phylogenetic studies may very well revise its placement. Description. Female (holotype). Carapace length 1.3; abdomen length 1.5. Carapace relatively square, high. PME small. Chelicerae with 2 3 promarginal and 2 retromarginal teeth. Palp with tarsal claw. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 3 pairs. Epigynum (Fig. 24) with a central depression, over which is an epiglottis-shaped projection. Colour (Figs 21 23): Dark brown; abdomen with faint chevrons. Natural history. The one specimen was collected from a small patch of moist forest. The forest was somewhat disturbed less than 10 m from where the specimen was found was a highway; on the other side was a river 58 Zootaxa 3424 2012 Magnolia Press MADDISON

within 50 m. Although the exact location and microhabitat where the specimen was found is not known, the primary microhabitats searched were leaf litter and mossy tree trunks. When walking, the second pair of legs are waved periodically, as seen in Thrandina and many other basal salticids (Maddison, 2006, 2009). A video of the living holotype is available at http://www.youtube.com/watch?v=egfizph1lou. FIGURES 21 24. Lapsias guamani, sp. n. 21 24 holotype female 24 epigynum, ventral view. Scale bar = 0.1 mm. Figures 21 24 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. Genus Thrandina Maddison The type species of Thrandina, T. parocula Maddison, and the two described here are unusual among lapsiines, indeed salticids, in the large size of the PMEs. The palp has a robust median apophysis. T. parocula was described from 1000 m elevation in southeastern Ecuador. A new record of T. parocula from further north (Fig. 1) is also at about 1000m. The two new species, similar in appearance to T. parocula, are from higher elevation, approximately 2000m. Of the two subadult males mentioned by Maddison (2006) as possibly NEW SPECIES OF LAPSIINE JUMPING SPIDERS FROM ECUADOR Zootaxa 3424 2012 Magnolia Press 59

belonging to T. parocula, the one from Yanayacu is almost certainly T. cosanga, and the one from Morona Santiago may also be T. cosanga, as it was found above 2000 m elevation. All three known Thrandina species are found on mossy tree trunks and branches in wet forests, especially where the moss is green and moist and yet exposed to some sun. Living male specimens of the three species can be distinguished relatively easily: T. cosanga and T. bellavista have the cymbium almost entirely clothed in white hairs, while T. parocula has less extensive white, with the prolateral third of the cymbium black. The first two species also have much more distinct white patches at the base of the first leg metatarsus. T. bellavista is unusual in having two white spots on the posterior of the abdomen. These differences are easily seen in the photographs and the videos (T. cosanga, http://www.youtube.com/watch?v=-8xx8rn5iku; T. bellavista, http://www.youtube.com/watch?v=gbuwsax2xbm; T. parocula, http://www.youtube.com/watch?v=2rlt7_0q9zs). Thrandina cosanga, sp. nov. (Figs 25 29, 37 42) Type material. Holotype male in QCAZ with data: "ECUADOR: Napo: Vinillos, near Cosanga. S 0.6025 W 77.8508. 2080 m elev. 29 30 October 2010. W & D Maddison, M Vega, M Reyes. WPM#10-036", "Male W606", UBC-SEM AR00195 Etymology. Named after the type locality. Diagnosis. Closely similar to T. bellavista, from which it can be distinguished by the narrower tip of embolus (Fig. 25), the more robust median apophysis (Fig. 27), and the lack of two large pale spots on the back of the abdomen. Although there are few records, T. cosanga and T. bellavista may be allopatric, with T. cosanga on the eastern slopes of the Andes. Like T. bellavista, T. cosanga differs in many respects from T. parocula, including the smaller median apophysis, embolus with a single curve, cymbium clothed entirely in white setae (compare Figs 39 and 51), paler base of the first leg metatarsus (compare Figs 38 and 48), and epigynal opening more anteriorly placed. Description. Male (holotype). Carapace length 2.5; abdomen length 2.5. PME large. Chelicerae not particularly enlarged, with 3 promarginal and 2 retromarginal teeth. Palpus (Figs 25 27): curved embolus arises prolaterally. Median apophysis large, shaped almost like a human ear. Conductor pale, triangular, between embolus and median apophysis. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 3 prolateral and 2 retrolateral macrosetae. Colour (Figs 37 40): In general dark brown. Thorax with pale medial longitudinal stripe. Palpus dark but with white setae covering almost the entire cymbium, dorsum of the tibia, and parts of the patella. The extreme prolateral edge of the cymbium has dark setae. Legs medium to dark brown, the first legs especially dark, with paler patella and pale annulae at the bases of the metatarsi and tarsi. Abdomen with paler chevrons. Female (paratype in UBC-SEM, #UBC SEM AR00196, from: ECUADOR: Napo: Cosanga, Yanayacu Biological Station, forest. S 0.600-1 W 77.888-890. 2100 m elev. 7 Nov. 2010. W & D Maddison, M Vega, M Reyes. WPM#10-058). Carapace length 2.0; abdomen length 2.6. PME large. Chelicerae with 3 promarginal and 2 retromarginal teeth. Palp with tarsal claw. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 3 prolateral and 2 retrolateral setae. Epigynum (Fig. 28 29) with openings in single round anterior cavity. Colour (Figs 41 42): as in male except first legs not so dark. Additional material examined. 7 males, 8 females, in UBC-SEM, from: ECUADOR: Napo: Reserva Ecologica Antisana, Sendero Jumandy. S 0.624-5 W 77.840-2. 2260 m elev. 29 Oct. 2010. W & D Maddison, M Reyes. WPM#10-035 (1 male); ECUADOR: Napo: Vinillos, near Cosanga. S 0.6025 W 77.8508. 2080 m elev. 29 30 October 2010. W & D Maddison, M Vega, M Reyes. WPM#10-036 (3 male and 1 female paratypes, and an additional 2 males and 3 females); ECUADOR: Napo: Cosanga, Yanayacu Biological Station, forest. S 0.600-1 W 77.888-890. 2100 m elev. 7 Nov. 2010. W & D Maddison, M Vega, M Reyes. WPM#10-058 (1 male paratype, 1 female paratype, and 3 additional females). Natural history. All specimens were found in moist forests, usually on mossy tree trunks and branches. Partial courtship behaviour was observed once (Fig. 38, video http://www.youtube.com/watch?v=ztb4g4fxh0k). The first legs were held down and forward, and the palpi angled outward over the femora of the first legs. The male flickered the palpi and first legs occasionally. Another video of a living male is available: http://www.youtube.com/watch?v=-8xx8rn5iku. 60 Zootaxa 3424 2012 Magnolia Press MADDISON

FIGURES 25 36. Thrandina species, genitalia. (25 29) Thrandina cosanga, sp. n., holotype male 25 tip of embolus, oblique view between ventral and prolateral 26 male palp, ventral view 27 male palp, retrolateral view 28 epigynum, ventral view 29 epigynum, cleared, dorsal view (30 33) Thrandina bellavista, sp. n., 30 tip of embolus, oblique view between ventral and prolateral 31 male palp, ventral view 32 male palp, retrolateral view 33 epigynum, ventral view (34 36) Thrandina parocula Maddison. 34 epigynum, ventral view 35 epigynum, cleared, dorsal view 36 male palp, ventral view. (25 27, 30 32, 36) holotypes. (28 29, 33) paratypes. Scale bar = 0.1 mm. Figures 25 36 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. NEW SPECIES OF LAPSIINE JUMPING SPIDERS FROM ECUADOR Zootaxa 3424 2012 Magnolia Press 61

FIGURES 37 42. Thrandina cosanga, sp. n. 37, 39 male from Vinillos (locality WPM#10-036) 38 holotype male, courtship pose 40 male from Yanayacu (locality WPM#10-058) 41 female from Vinillos (locality WPM#10-036) 42 female from Yanayacu (locality WPM#10-058). Figures 37 42 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. Thrandina bellavista, sp. nov. (Figs 30 33, 43 47) Type material. Holotype male in QCAZ with data: "ECUADOR: Pichincha: Bellavista Cloud Forest Reserve. S 0.0125-0.127 W 78.680. 2070 m elev. 10 November 2010. D. Maddison, M. Reyes & M. Vega. WPM#10-068", "Photo'd 11 Nov #ECU2010-3261", "UBC-SEM AR00197" 62 Zootaxa 3424 2012 Magnolia Press MADDISON

FIGURES 43 47. Thrandina bellavista, sp. n. (43, 44, 46) holotype male (45, 47) female from type locality (WPM#11-068), same female as Fig. 33. Figures 43 47 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. Etymology. Refers to the type locality. Diagnosis. Known from the west slopes of the Andes, in contrast to the similar T. cosanga, which is known from the east. The palp can be distinguished from that of T. cosanga by a wider tip of embolus (Fig. 30) and slightly more delicate median apophysis (Fig. 32). Unlike both T. cosanga and T. parocula, the abdomen has a pair of pale spots in the distal third (Figs. 46 47). Like T. cosanga, T. bellavista differs in many respects from T. parocula, including the smaller median apophysis, embolus with a single curve, cymbium clothed entirely in white setae (compare Fig. 43 and 51), paler base of the first leg metatarsus (compare Fig. 43 and 48), and epigynal opening more anteriorly placed. NEW SPECIES OF LAPSIINE JUMPING SPIDERS FROM ECUADOR Zootaxa 3424 2012 Magnolia Press 63

FIGURES 48 52. Thrandina parocula Maddison. (48, 49, 51) male from Río Guamani (locality WPM#10-038) 50 female from Río Guamani (locality WPM#10-053) 52 female from Río Guamani (locality WPM#10-038). Figures 48 52 are copyright 2012 W. P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. Description. Male (holotype). Carapace length 2.4; abdomen length 2.6. PME large. Chelicerae not particularly enlarged, with 3 promarginal and 2 retromarginal teeth. Palpus (Figs 30 32): as in T. cosanga, though tip of embolus wider and median apophysis less robust. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 3 prolateral and 2 retrolateral setae. Colour (Figs 43, 44, 46): In general dark brown, as in T. cosanga. First legs darkest. The bases of the first leg metatarsus and tarsus are especially pale in contrast with the rest of the leg. The abdomen has, in addition to faint chevrons, two lateral pale spots in the posteriormost third. Female (paratype in UBC-SEM, #UBC SEM AR00198, from ECUADOR: Pichincha: Bellavista Cloud Forest 64 Zootaxa 3424 2012 Magnolia Press MADDISON

Reserve. S 0.0125 0.127 W 78.680. 2070 m elev. 10 November 2010. D. Maddison, M. Reyes & M. Vega. WPM#10-068). Carapace length 2.0; abdomen length 2.6. PME large. Chelicerae: 3 promarginal and 2 retromarginal teeth. Palp with tarsal claw. Tibia of first leg with 3 pairs of ventral macrosetae; first metatarsus with 3 prolateral and 2 retrolateral macrosetae. Epigynum (Fig. 33) with openings in single round anterior cavity. Colour (Figs 45, 47): As in T. cosanga. The abdomen has, in addition to faint chevrons, two lateral pale spots in the posteriormost third. Additional material examined. Paratypes from: ECUADOR: Pichincha: Bellavista Cloud Forest Reserve. S 0.012 016 W 78.682. 2050 2240 m elev. 9 10 November 2010. W & D Maddison, M Vega, M Reyes. WPM#10-065 (1 female). ECUADOR: Pichincha: Bellavista Cloud Forest Reserve. S 0.0125 0.127 W 78.680. 2070 m elev. 10 November 2010. D. Maddison, M. Reyes & M. Vega. WPM#10-068 (1 male). Natural history. All specimens were found in a moist forest, usually on mossy tree trunks and branches. A video of a living male is available: http://www.youtube.com/watch?v=gbuwsax2xbm. Thrandina parocula Maddison (Figs 34 36, 48 52) Diagnosis. T. parocula can be distinguished from both T. cosanga and T. bellavista by its larger median apophysis with a long spur, sinuate embolus, prolateral side of cymbium clothed in black (see Fig. 51), darker base of the metatarsus of leg 1 in males, and epigynal opening more posteriorly placed. New material examined. 2 males and 7 females in UBC-SEM from: ECUADOR: Napo: Río Guamani on Jondachi-Loreto road. S 0.7223 W 77.6408. 1050 m elev. W & D Maddison, M Vega, M Reyes. (1 male 2 females from 31 October 2010, WPM#10-038; 1 males 5 females from 6 November 2010, WPM#10-053) Natural history. A video of a living male is available: http://www.youtube.com/watch?v=2rlt7_0q9zs Acknowledgments Mauricio Vega provided critical assistance in arranging both expeditions to Ecuador. David Maddison, Mauricio Vega, Edyta Piascik and Marco Reyes assisted with collecting. The Ecuadorian Ministry of the Environment and the Museum of Zoology of the Pontificia Universidad Católica de Ecuador assisted with permits for our research. Funding was provided primarily from an NSERC Discovery grant to the author, with some funding for the 2010 field work supplied by David Maddison from the Harold E. and Leona M. Rice Endowment at Oregon State University. Literature cited Galiano, M.E. (1963) Las especies americanas de arañas de la familia Salticidae descriptas por Eugène Simon: Redescripciones basadas en los ejemplares típicos. Physis, Buenos Aires, 23, 273 470. Maddison, W.P. (2006) New lapsiine jumping spiders from Ecuador (Araneae: Salticidae). Zootaxa, 1255, 17 28. Maddison, W.P. (2009) New cocalodine jumping spiders from Papua New Guinea (Araneae: Salticidae: Cocalodinae). Zootaxa, 2021, 1 22. Maddison, W.P. & Needham, K. (2006) Lapsiines and hisponines as phylogenetically basal salticid spiders (Araneae: Salticidae). Zootaxa, 1255, 37 55. Platnick, N.I. (2011) World spider catalog, version 12.0. Available from http://research.amnh.org/iz/spiders/catalog/saltici- DAE.html. (accessed 16 October 2011) Ruiz, G. & Maddison, W.P. (2012) DNA sequences corroborate Soesiladeepakius as a non-salticoid genus of jumping spiders: placement with lapsiines, phylogeny and description of six new species (Araneae, Salticidae). Zoological Journal of the Linnean Society, 165, 274 295 Simon, E. (1900) Descriptions d'arachnides nouveaux de la famille des Attidae. Annales de la Société entomologique belge, 44, 381 407. Simon, E. (1901) Histoire naturelle des araignées. Paris, 2, 381 668. NEW SPECIES OF LAPSIINE JUMPING SPIDERS FROM ECUADOR Zootaxa 3424 2012 Magnolia Press 65