From the Lowlands to the Highlands of Ecuador, a Study of the Genus Masteria (Araneae, Mygalomorphae, Dipluridae) with Description of Seven New Species

Zootaxa 5005 (4): 538–568 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2021 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.5005.4.4 http://zoobank.org/urn:lsid:zoobank.org:pub:43AB6083-4E39-47DD-819E-8EC21F3B3C90

From the lowlands to the highlands of Ecuador, a study of the genus Masteria (Araneae, Mygalomorphae, Dipluridae) with description of seven new species

NADINE DUPÉRRÉ1,2*, ELICIO TAPIA3, DIETMAR QUANDT4,5, VERÓNICA CRESPO-PÉREZ6 & DANILO HARMS7 1Zoological Museum Hamburg, Leibniz-Institute for the Analysis of Biodiversity Change (LIB), Center for Taxonomy and Morphology, Martin-Luther-King-Platz 3, 20146, Hamburg, Germany. 2Research Associate, American Museum of Natural History, New York, USA. 3Fundación OTONGA, Calle Rither N° 20-10 y Bolivia, Quito, Ecuador. https://orcid.org/0000-0001-9005-5328 4Nees Institute for Plant Biodiversity, University of Bonn, Meckenheimer Allee 170, 53115 Bonn, Germany. 5Leibniz Institute of Plant Genetics and Crop Plant Research (IPK), Corrensstraße 3, 06466 Gatersleben, Germany https://orcid.org/0000-0003-4304-6028 6Laboratorio de Entomología, Museo de Zoología QCAZ, Escuela de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Avenida 10 de Octubre 1076, 170143, Quito, Ecuador. https://orcid.org/0000-0002-8811-1965 7Zoological Museum, Center of Natural History, Universität Hamburg, Martin-Luther-King-Platz 3, 20146, Hamburg, Germany. https://orcid.org/0000-0002-7189-5345 *Corresponding author. �[email protected]; https://orcid.org/0000-0003-2195-878X

Abstract

Dipluridae represent a small Mygalomorphae family of South American origin, the family includes two subfamilies Diplurinae and Masteriinae although the placement of the latter in Dipluridae is still under debate. The family has a predominantly South American distribution although the genus Masteria L. Koch, 1873 presents an interesting distribution with representatives found in Fiji, Micronesia, New Caledonia, New Guinea and Australia. This genus is diverse at the species level in tropical South America and the Caribbean but no species have been described from Ecuador to date. Ongoing field work as part of the BIO-GEEC Project—a consortium established by several Ecuadorian and German institutions—has resulted in the discovery of several new species from both lowlands and highland habitats in Ecuador. Herein we described seven new species of Masteria from Ecuador: M. jatunsacha n. sp. (male); M. machay n. sp. (female); M. chalupas n. sp. (male); M. papallacta n. sp. (male and female); M. pasochoa n. sp. (male and female); M. lasdamas n. sp. (male); and M. otongachi n. sp. (male). The type species of the genus, Masteria hirsuta L. Koch, 1873 from Fiji, is redescribed and re-illustrated, from the original type specimen.

Key words: New species, DNA-barcoding, taxonomy

Introduction

The BIO-GEEC (German-Ecuadorian Biodiversity Consortium) project was launched in 2019 with the mandate to develop a platform that disseminates Ecuadorian biodiversity data through a combination of targeted field work, taxonomy, DNA barcoding and WebApp assessment. The consortium led by four Ecuadorian universities and four German partner institutions in collaboration with the Ministerio del Ambiente y Agua (MAAE) and the Instituto Nacional de Biodiversidad (INABIO) is developing pipelines for the rapid characterisation of diverse organismic groups such as seed plants (tissue, seeds & pollen), pollinators (in particularly insects), arachnids, and soil mi- croorganisms. The Web App will include relevant morphological and genetic and genetic information, as well as high-resolution images, thereby facilitating fast species-level identification, ideally by non-arachnologists. Barcod- ing these and additional samples from Ecuador and providing baseline data in EcBioDiv (Ecuadorian Biodiversity Database) will help to dissociate the growing knowledge on Ecuador’s impressive Arachnid fauna, establish a stable

538 Accepted by J. Bond: 9 Jul. 2021; published: 28 Jul. 2021 baseline for future species discoveries. Furthermore, these pipelines shall be applied to other taxa and handed over to government agencies and industry. Ecuador’s distinctive geography and topography represents the foremost unique, highly diverse study system. In fact, all ecoregions of Ecuador belong to one of the prime global biodiversity hotspots with extreme levels of species endemism and genetic richness (e.g., Mutke & Barthlott 2005, Myers et al. 2000). The Andes separate the entire country longitudinally and both sides of the Andes are covered with tropical rainforests at low elevation up to the Páramo or alpine tundra (6,000 m), though with different ecological parameters. One aim of the BIO-GEEC project is to discover, describe and characterize the morphological and genetic biodiversity of spiders found along an altitudinal gradient on the eastern side of the Andes. In February–March 2020 a series of pitfall traps were in- stalled along a transect near Tena (1,048 m) up to Papallacta (4,020 m). Additional sampling was also done on both sides of the Andes at different elevations. The samples collected led to the discovery of seven new species belonging to the mygalomorph spider family Dipluridae Simon, 1889. The family was recently redelimited by Opatova et al. (2020) but the limits are not fully resolved, and the family remains non-monophyletic (Opatova et al. 2020). The family currently includes two subfamilies, the Diplurinae Simon, 1889 and the Masteriinae Simon, 1889, but the latter may in future be elevated to family level (Opatova et al. 2020). As of present, Masteriinae includes three recent genera next to the fossil Edwa Raven, Jell & Knezour, 2015: Masteria C.L. Koch, 1873 (32 valid species) and occurs in the Americas and Oceania including New Caledonia and New Guinea and Australia, while Siremata Passanha & Brescovit, 2018 (3 species) is endemic to Brazil and Striamea Raven, 1981 (2 species) is endemic to Colombia (WSC 2021). Masteria is a genus of small (1.97–6.06 mm), 6 or 8 eyed mygalomorph spiders. The American fauna has been revised by Passanha & Brescovit (2018), Raven (1991) reviewed the New Caledonia fauna, but the rest of the genus is still pending revision. In the present paper we described seven new species of Masteria from Ecuador’s lowland forest systems (430 m) up to the Páramo (4,020 m), on both sides of the Andes. The type species of the genus, Mas- teria hirsuta L. Koch 1873, from Fiji is redescribed and the female internal genitalia is imaged for the first time. A distribution map of all Ecuadorian species is presented.

Materials and methods

Specimens were initially collected and examined in 90% ethanol under a Leica M125 dissection microscope. Female genitalia were excised using a sharp entomological needle, washed in 80% alcohol and digested with Pancreatin solution following Álvarez-Padilla & Hormiga (2007), then placed on a slide in lactic acid for imaging. Imaging was achieved using a custom-made BK Plus lab System by Dun, Inc. with integrated Canon camera, macro lens (65mm) and the Zerene stacking software (Zerene Systems LLC 2018). All measurements are in millimeters and were made using a Leica M205A with Leica Application Suite X. Descriptions of specimens colours are based on specimens conserved in ethanol, in specimens recently collected (e.g. M. chalupas n. sp., M. jatunsacha n. sp.) the colouration is likely to be more authentic. Collection abbreviations are as follows: QCAZ-I, Museum of Invertebrates, Pontificia Universidad Católica, Quito (Ecuador); ZMH, Zoological Museum Hamburg, Hamburg (Germany). Morphological nomenclature follows Passanha & Brescovit (2018). Abbreviations in text and figures are as follows: ALE, anterior lateral eyes; AME, anterior median eyes; BS, basal spine; EcL, ectal lobe; EnL, ental lobe; GR, glandular region; P1, Process 1; P2, Process 2; P3, Process 3; PLE, posterior lateral eyes; PLS, posterior lateral spinnerets; PME, posterior median eyes; PMS, posterior median spinnerets; plv, prolatero-ventral; rlv, retrolatero-ventral; SB, spermathecae base.

Taxonomy

Family Dipluridae Simon, 1889

Genus Masteria L. Koch, 1873

Type species. Masteria hirsuta L. Koch, 1873

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 539 Diagnosis. See Passanha & Brescovit 2018: 8. Composition. For complete list of species see WSC 2021. M. chalupas n. sp., M. jatunsacha n. sp., M. lasdamas n. sp., M. machay n. sp., M. otongachi n. sp., M. papallacta n. sp., M. pasochoa n. sp. Distribution in the Americas. Costa Rica, Panama, Cuba, Puerto Rico, Martinique, Dominican Republic, St. Vincent, Jamaica, Trinidad, Tobago, Venezuela, Guyana, Colombia, Ecuador, Brazil and Peru.

Masteria hirsuta L. Koch, 1873 Figs 1A–C, 20A.

Masteria hirsuta L. Koch, 1873: 458, pl. 35, f. 5 (Description female from Ovalau). Masteria hirsuta Simon, 1892a: 190, f. 131. Masteria hirsuta Roewer, 1963a: 109, f. 2a–d (Description male from Ponape). Masteria hirsuta Raven, 1979: 627, f. 5–7. (Illustration of holotype female). Masteria hirsuta Passanha & Brescovit, 2018: 13, f. 3B, 6A, 7A–B (Illustration female from Ponape, images of male from Ponape per Roewer).

Type material. Ein Exemplar im Museum Godeffroy. Von Ovalau. (Koch 1873: 458). Holotype female in Zoolo- gisches Museum Hamburg (ZMH-A0003083). Examined. Diagnosis. Masteria hirsuta is distinguished from all species by the combination of the following characters: six eyes with strongly recurved posterior row (Figs 1A, 20A), internal genitalia with unilobed spermathecae, spermathecae bases wide and short, oval ectal lobe (EcL) situated externally on spermathecae bases (Fig. 1C). Description. Female (holotype): Total length: 5.40; carapace length: 2.64; carapace width: 2.18; abdomen length: 2.76. Carapace yellow-orange (Fig. 1A). Chelicerae yellow; promargin with 12 teeth and ~15 smaller mesobasal teeth. Endites whitish without cuspules. Labium whitish without cuspules. Sternum whitish, covered with long setae (Fig. 1B). Eyes: six eyes grouped; AME absent; ALE and PLE almost equal; PME rounded: posterior row strongly recurved (Fig. 20A). Abdomen elongated, uniformly beige covered with long setae (Fig. 1A). Spinnerets: PLS 2.66 (0.91/0.88/0.87). Legs coloration uniformly pale yellow. Leg measurements: I, femur, 2.26; II, missing; III, femur 1.5; IV, femur 2.7, patella 1.07 and tibia 1.9. Leg spination: legs damaged. Palpal claw with ~ 21 teeth. Genitalia: Unilobed spermathecae with large, oval ectal lobes (EcL) situated externally on spermathecae bases; spermathecae bases wide and short; glandular region wide, with glands large and widely interspaced (Fig. 1C). Note. The type species of the genus Masteria hirsuta was described by L. Koch in 1873, based on a female specimen from Ovalau (Fiji Islands). Roewer (1963) described the male based on a specimen from Pohnpei (or Ponape) in Micronesia and examined several juvenile females from Marina Islands (Tinian), Palau (Babelthuap, Koror, Peleliu), Chuuk (or Truk), Kosrae Is. (or Kusaie) next to adult females specimens from Yap and the Marshall Island (Eniwetok). Roewer concluded that all the specimens he examined represented a single widespread species in Micronesia, even though the type came from Ovalau, about 1000km away. The internal genitalia of the female holotype (and type species of the genus) is illustrated here for the first time (Fig. 1C). The internal genitalia of the female holotype differ from the illustration given for a specimen identified as M. hirsuta by Passanha & Brescovit 2018 from Ponape (see Passanha & Brescovit 2018: fig. 6A). The differences observed are as such: the spermathecae bases are much shorter and broader in the type specimen and the ectal lobes are situated externally on the spermathecae bases while in M. hirsuta from Ponape the spermathecae bases are long and thin and the ectal lobes are situated medially on the spermathecae bases. The female internal genitalia of Masteria are fragile, the differences observed could be due to distortion, variation, or just a different angle of illustration. At this point, a re-examination of fresh specimens from Ovalau, and Ponape is necessary in order to conclude if these specimens belong to the same or represent separate species. Bearing in mind the high diversity of the genus (WSC 2021), it would not be surprising that the species from Ovalau and Ponape are two different species and under these circumstances the male described by Roewer (1963) would not be male of M. hirsuta. Interestingly, Raven (1979) redescribed the species based on the type specimen and illustrated the carapace but the information was overlooked in the World Spider Catalog (2021).

540 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. FIGURES 1A–C. Masteria hirsuta L. Koch, 1873 (holotype). A. Female habitus, dorsal view. B. Female habitus, ventral view. C. Internal genitalia, dorsal view. Scale bars: 0.5mm.

FIGURES 2A, B. Masteria jatunsacha n. sp. (holotype). A. Male habitus, dorsal view. B. Male habitus, ventral view. Scale bars: 0.5mm.

542 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. Type material. Male holotype from Napo Province, Ecuador, Jatun Sacha Natural Reserve (-01.666464 -77.617192) 400–430m, 10–15 Mar. 2020, pitfall, E.E. Tapia, N. Dupérré, A.A. Tapia (QCAZ). Paratypes: 2♂ same data as the holotype (ZMH-A0014697, QCAZ). Etymology. The specific name is a noun in apposition taken from the type locality Jatun Sacha Natural Reserve, Napo, Ecuador.

FIGURES 3A–D. Masteria jatunsacha n. sp. (holotype). A. Male palp, prolateral view. B. Male palp, retrolateral view. C. Male palpal bulb, prolateral view. D. Male leg I, prolateral view. Scale bars: 0.1mm.

Diagnosis. Males most resemble M. chalupas n. sp. and M. papallacta n. sp. due to the presence of spines on the retrolateral side of the male palpal tibia but can be distinguished from M. papallacta n. sp. by the grouped spines

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 543 positioned on the apical half (Fig. 3B, C), whilst they are positioned on the basal half in the latter species (Fig. 9B, C); from M. chalupas n. sp. by the presence of 21 grouped spines (Fig. 3C) vs. 14 grouped spines in the latter species (Fig. 7B, C) and basal spine of metatarsus I shorter (Fig. 22A), clearly longer in M. chalupas n. sp. (Fig. 22B). Description. Male (holotype): Total length: 2.79; carapace length: 1.37; carapace width: 1.07; abdomen length: 1.42. Carapace yellow with reticulation in the cephalic groove and pars thoracica, covered with long setae (Fig. 2A). Chelicerae yellow; promargin with 11 teeth and ~10 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow, covered with long setae (Fig. 2B). Eyes: six eyes grouped; AME absent; ALE the largest; PLE oval; PME small and oval; posterior row slightly recurved (Figs 2A, 20B). Abdo- men elongated, uniformly beige covered with long setae (Fig. 2B). Spinnerets: PLS 1.15 (0.39/0.41/0.35). Legs coloration uniformly pale yellow. Leg formula 4123; leg measurements: I 4.44 (1.27/0.63/1.03/0.80/0.72); II 3.54 (1.02/0.43/0.81/0.69/0.59); III 3.38 (0.92/0.35/0.82/0.72/0.57); IV 4.55 (1.24/0.29/1.12/1.09/0.61). Leg spination: I: femur d1-1-1-1-1; tibia v1, p-1ap; metatarsus v1-1; II: femur d1-1-1-1-1, p1; patella v1, p1; tibia v1-1-1ap, p1; metatarsus v1-1-1; III: femur d1-1-1-1, p1; patella p1; tibia d1-1, v1-1-1-3ap, p1-1, r1-1; metatarsus d1-2, v2-2-3ap, p1, r1-1; IV: femur d1-1-1-1-1; patella v-1ap, r1; tibia d1, v2-1ap, p1-1-1, r1-1, metatarsus d1, v2-2-2ap, p1-1, r1-1- 1. Tibia I: P1 absent; P2 with an apical, narrow strong spine; P3 with two apical spines (Fig. 22A). Metatarsus I with small basal spine (Fig. 22A). Palp: palpal tibia 2x the length of cymbium, 2.6x longer than wide, with 21 grouped long spines on retrolateral side, positioned on apical half (Fig. 3B, C); cymbium 2x longer than wide, with four apical spines; bulb rounded, with laminar twisted embolus (Fig. 3C). Female: unknown. Distribution. Ecuador: only found at the type locality. Natural History. Specimens were collected by pitfall between 400–430m in an evergreen lowland forest of the Napo-Curaray region (Guevara et al. 2013b).

Masteria machay Dupérré & Tapia, new species Figs 4A, B, 5A, 20C, map.1.

Type material. Female holotype from Ecuador, Napo, Miraflores community (-00.958370 -77.645851) 854m, 24 Feb. 2018, hand collected 10m in cave, E.E. Tapia (QCAZ). Paratype: one female, same data as holotype (ZMH- A0002513). Other material examined. Ecuador: Napo Province: Miraflores community (-00.958370 -77.645851) 854m, 24 Feb. 2018, 3 juv., hand collected 10m in cave, E.E. Tapia (ZMH-A0002511, QCAZ). Etymology. The specific name is a noun in apposition taken from the Kichwa language “mach’ay” meaning cave. Diagnosis. Females most resemble M. tayrona Passanha & Brescovit, 2018 but are distinguished from all species by their reduced eyes (Figs 4A, 20C) and their internal genitalia composed of multilobed (8 lobes) spermathecae (Fig. 5A); whilst M. tayrona female multilobed spermathecae is composed of 4–5 lobes (Passanha & Brescovit 2018; fig. 22F). Description. Female (holotype): Total length: 4.57; carapace length: 1.85; carapace width: 1.43; abdomen length: 2.72. Carapace light yellow, smooth, covered with long setae (Figs 4A, 20C). Chelicerae yellow; promargin with 10 teeth and ~20 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow, covered with long setae (Fig. 4B). Eyes: six reduced eyes grouped; AME absent; ALE and PLE almost equal; PME the smallest, rounded: posterior row slightly recurved (Figs 4A, 20C). Abdomen elongated, uniformly beige covered with long setae (Fig. 4A). Spinnerets: PLS 1.75 (0.61/0.57/0.57). Legs coloration uniformly pale yellow. Leg formula 4123; leg measurements: I 6.25 (1.71/0.79/1.52/1.27/0.96); II 5.16 (1.47/0.74/1.19/0.92/ 0.84); III 4.97 (1.39/0.53/1.11/1.04/0.90); IV 6.28 (1.67/0.63/1.58/1.42/0.98). Leg spination: I: femur d1-1-1-1-1; tibia v1-1; metatarsus v1-1; II: femur d1-1-1-1-1, p1; tibia v1-1ap, p1; metatarsus v1-1-1; III: femur d1-1-1-1, p1; patella p1; tibia d1-1, v1-1-1-3ap, p1-1, r1-1; metatarsus d1-2, v2-2-3ap, p1, r1-1; IV: femur d1-1-1-1-1; patella v-1ap; tibia d1, v2-1ap, p-1-1, r1-1, metatarsus d1, v2-2-2ap, p1-1, r1-1-1. Palpal claw with ~14 teeth. Genitalia: multilobed (8 lobes) spermathecae with large, oval ectal lobes (EcL) on elongated stems; ental lobes (EnL) triangular; spermathecae bases long; glandular region with small glands close together at the base of lobes and large glands widely interspaced on the spermathecae bases (Fig. 5A).

Male: unknown. Distribution. Ecuador: only found at the type locality. Natural History. Specimens were found in an open karstic cave under rocks, about 6–10m from the entrance. The cave floor was covered with residues (fruits) from fruit bats.

Masteria chalupas Dupérré & Tapia, new species Figs 6A, B, 7A–D, 20D, 22B, map 1.

Type material. Male holotype from Ecuador, Napo, Tena, Colonso Chalupas Biological Reserve (-00.9014 - 77.88588) 1048m, 9–15 Mar. 2020, pitfall trap, E.E. Tapia, N. Dupérré & A.A. Tapia (QCAZ). Paratypes. 2♂ same data as holotype (QCAZ, ZMH-A0013145); one 1♂ from Ecuador, Napo, Tena, Colonso Chalupas Biological Reserve (-00.9019 -77.88588) 1044m, 9–15 Mar. 2020, pitfall trap, E.E. Tapia, N. Dupérré & A.A. Tapia (ZMH- A0013146). Etymology. The specific name is a noun in apposition taken from the type locality Colonso Chalupas Biological Reserve, Napo, Ecuador. Diagnosis. Males most resemble M. jatunsacha n. sp., and M. papallacta n. sp. but can be distinguished as such: from M. papallacta n. sp. by their twisted embolus (Fig. 7B, D) while straight in M. papallacta n. sp. (Fig. 9B, C); from M. jatunsacha n. sp. by the presence of 14 grouped spines on the retrolateral side of the male palpal tibia (Fig. 7B, C), while 21 grouped spines are present in the latter species (Fig. 3C), and the longer basal spine of metatarsus (Fig. 22B), shorter in M. jatunsacha n. sp. (Fig. 22A). Description. Male (holotype): Total length: 1.97; carapace length: 1.04; carapace width: 0.74; abdomen length: 0.93. Carapace light yellow with reticulation in the cephalic groove and pars thoracica, covered with long setae (Fig. 6A). Chelicerae yellow; promargin with 10 teeth and ~9 smaller mesobasal teeth. Endites whitish without cuspules. Labium whitish without cuspules. Sternum whitish, covered with long setae (Fig. 6B). Eyes: six eyes grouped; AME absent; ALE the largest; PLE oval; PME the smallest, oval; posterior row recurved (Figs 6A, 20D). Abdomen elongated, uniformly beige covered with elongated setae (Fig. 6B). Spinnerets: PLS 0.69 (0.24/0.19/0.26). Legs coloration uniformly pale yellow. Leg formula 4123; leg measurements: I 2.64 (0.73/0.46/0.62/0.42/0.41); II missing;

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 547 III 1.95 (0.56/0.27/0.37/0.41/0.34); IV 2.72 (0.79/0.26/0.64/0.60/0.43). Leg spination: I: femur d1-1-1-1-1; metatarsus v1; II missing; III: femur d1-1-1-1, p1; patella p1; tibia d1-1, v1-1-1-3ap, p1-1, r1-1; metatarsus d1-2, v2-2ap, p1, r1-1; IV: femur d1-1-1-1-1; patella v-1ap, r1; tibia v2-1ap, p1-1-1, r1-1, metatarsus v1-1-2ap, p1-1. Tibia I: P1 absent; P2 with an apical, narrow strong spine; P3 with two apical spines (Fig. 22B). Metatarsus I with elongated basal spine (Fig. 22B). Palp: palpal tibia 1.6x the length of cymbium, 2.6x longer than wide, with 14 grouped long spines on retrolateral side, positioned on apical half (Fig. 7A-C); cymbium 2x longer than wide, with four apical spines; bulb constricted, with laminar twisted embolus (Fig. 7C). Female: unknown. Distribution. Ecuador: only found at the type locality.

FIGURES 7A–D. Masteria chalupas n. sp. (holotype). A. Male palp, prolateral view. B. Male palp, retrolateral view. C. Male palpal bulb, prolateral view. D. Male leg I, prolateral view. Scale bars: 0.1mm.

548 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. Natural History. Specimens were collected by pitfall trap between 1044–1048m in an evergreen piemontane forest of the northern part of the oriental Andean mountain range (Guevara et al. 2013a).

Masteria papallacta Dupérré & Tapia, new species Figs 8A, B, 9A–D, 10 A, B, 11A, 20E, F, 22C. map. 1.

Type material. Male holotype and female allotype from Napo Province, Ecuador, Páramo de Papallacta, site 3 (- 00.281881 -78.137040) 4020m, 1–8 Mar. 2020, pitfall, E.E. Tapia, N. Dupérré, A.A. Tapia (QCAZ). Paratypes: 4♀ same data as the holotype (ZMH-A00014698, A00014699, QCAZ). Etymology. The specific name is a noun in apposition taken from the type locality Páramo de Papallacta. Diagnosis. Males resemble M. jatunsacha n. sp., and M. chalupas n. sp. but can be distinguished from both species by the straight embolus (Fig. 9C), that is twisted in the two other species (Figs 3C, 7C). Females are distinguished from all species by their small, tortuous ental lobes (Fig. 11A). Description. Male (holotype): Total length: 2.74; carapace length: 1.35; carapace width: 1.08; abdomen length: 1.39. Carapace light brown without obvious reticulation in the cephalic groove and pars thoracica, covered with long setae (Fig. 8A). Chelicerae yellow; promargin with 12 teeth and ~12 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow, covered with setae (Fig. 8B). Eyes: six eyes grouped; AME absent; ALE oval; PLE the largest, oval; PME the smallest, oval; posterior row recurved (Figs 8A, 20E). Abdomen elongated, grayish covered with long setae (Fig. 8A). Spinnerets: PLS 0.97 (0.36/0.29/0.32). Legs coloration uniformly pale yellow. Leg formula 4123; leg measurements: I 3.65 (1.02/0.56/0.79/0.67/0.61); II 2.85 (0.81/0.48/0.55/0.55/0.46); III 2.69 (0.82/0.33/0.53/0.49/0.52); IV 3.84 (1.07/0.43/0.90/0.79/0.65). Leg spination: I: no spines observed; II: femur d1-1; patella v1, p1; tibia v1-1, p1; metatarsus v1-1; III: femur d1-1-1; patella p1, v2ap; tibia d1-1, v2-2, p1-1, r1-1; metatarsus d1-2, v2-3ap, p1, r1-1; IV: femur d1-1-1; patella p1, v1ap, r1; tibia d1, v1-2ap, p1-1-1, r1-1, metatarsus d1, v2-2-2ap, p1-1, r1-1-1. Tibia I: P1 absent; P2 with an apical strong, curved spine; P3 with two apical spines (Fig. 22C). Metatarsus I with small pointed basal spine (Fig. 22C). Palpal tibia 1.7x the length of cymbium, 2.3x longer than wide, with 28 spines grouped in U on retrolateral side, (Fig. 9B, C); cymbium 2x longer than wide, with four apical spines; palpal bulb elongated-oval, embolus short, straight, strongly sclerotized (Fig. 9C). Female (allotype): Total length: 4.47; carapace length: 1.96; carapace width: 1.57; abdomen length: 2.51. Coloration as in male (Fig. 10A, B). Chelicerae yellow; promargin with 10 teeth and ~16 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow, covered with long setae (Fig. 10B). Eyes: six eyes grouped; AME absent; ALE oval; PLE the largest, oval; PME the smallest, oval; posterior almost straight (Fig. 20F). Abdomen elongated, uniformly beige, covered with setae (Fig. 10B). Spinnerets: PLS 0.97 (0.56/0.43/0.50). Legs coloration uniformly pale yellow. Leg formula 4123; leg measurements: I 4.44 (1.28/0. 74/1.03/0.76/0.63); II 3.63 (1.05/0.59/0.69/0.71/0.59); III 3.73 (1.17/0.44/0.70/0.76/0.66); IV 5.16 (1.47/0.61/1.24/ 1.06/0.78). Leg spination: I: d1-1-1-1-1; patella v1, tibiae v1-1; II: femur d1-1-1-1; patella v2ap, p1; tibia v1-1, p1; metatarsus v1-1-1, p1; III: femur d1-1-1; patella d1-2, p1, v1ap; tibia d1-1, v2-2, p1-1, r1-1; metatarsus d1-2, v1-1- 2ap, p1, r1-1; IV: femur d1-1-1; patella pd1, v2ap, r1; tibia d1-1, v1-2-2ap, p1-1-1, r1-1, metatarsus d1, v2-1-1ap, p1-1, r1-1-1. Palpal claw with ~20 teeth. Genitalia: unilobed spermathecae with short, oval ectal lobes (EcL), ental lobes tortuous (EnL); spermathecae bases wide; glandular region with large glands closely spaced on the spermathecae bases (Fig. 11A). Distribution. Ecuador: only found at the type locality. Natural History. Specimens were collected by pitfall at high elevation (4020m) in an evergreen shrubland and grassland of the Andean Páramo (Salgado et al. 2013).

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 549 FIGURES 8A, 8B. Masteria papallacta n. sp. (holotype). A. Male habitus, dorsal view. B. Male habitus, ventral view. Scale bars: 0.5mm.

550 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. FIGURES 9A–9D. Masteria papallacta n. sp. (holotype). A. Male palp, prolateral view. B. Male palp, retrolateral view. C. Male palpal bulb, prolateral view. D. Male leg I, prolateral view. Scale bars: 0.1mm.

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 551 FIGURES 10A, B. Masteria papallacta n. sp. (allotype). A. Female habitus, dorsal view. B. Female habitus, ventral view. Scale bars: 0.5mm.

Masteria pasochoa Dupérré & Tapia, new species Figs 12A, B, 13A–D, 14A, B, 15A, 20G, H, 23A, map 1.

Type material. Male holotype from Ecuador, Pichincha Province, Valle Chillos, Pasochoa Forest Reserve (- 0.443585 -78.507632) 3080m, June 2013, E.E. Tapia, N. Dupérré (QCAZ). Female allotype: Pichincha Province, Pululahua Geobotanical Reserve (00.059720–78.509230), 2128m, 2 Au- gust 2015, E.E. Tapia, N. Dupérré, A.A. Tapia (QCAZ). Etymology. The specific name is a noun in apposition taken from the type locality the Pasochoa Reserve. Diagnosis. Males most resembles M. colombiensis, M. lewisi and M. amarumayu but can be distinguished as such: from M. colombiensis by the presence of six eyes (Fig. 12A), eight in the latter species (Passanha & Brescovit 2018: fig. 13A); from M. lewisi by the absence of 4–6 short strong spines on palpal tibia (Fig. 13B) that are present in the latter (Passanha & Brescovit 2018: fig. 15B); and from M. amarumayu by the presence of P1 (Fig. 23A) that is absent in the latter (Passanha & Brescovit 2018: fig. 16C). Females are distinguished from all species by their unique internal genitalia with unilobed spermathecae, and large, triangular spermathecae bases (Fig. 15A). Description. Male (holotype): Total length: 4.10; carapace length: 1.95; carapace width: 1.38; abdomen length: 2.15. Carapace light brown with reticulation in the cephalic groove, radiating lines and pars thoracica; pars thoracica with arched row of long strong macrosetae (Fig. 12A). Chelicerae yellow; promargin with nine teeth and ~15 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow (Fig. 12B). Eyes: six eyes grouped; AME absent, ALE oval; PLE the largest; PME the smallest, oval, posterior row recurved (Figs 12A, 20G). Abdomen elongated, whitish-gray covered with long setae (Fig. 12B). Spinnerets: PLS 1.55 (0.58/0.43/0.54). Leg formula 4123; leg measurements: I 5.28 (1.45/0.71/1.35/0.94/083); II 4.37 (1.24/0.62/0. 88/0.89/074); III 4.13 (1.17/0.55/0.78/0.94/0.69); IV 5.53 (1.55/0.68/1.23/1.21/0.86). Leg spination: I: d1-1-1-1-1; tibiae v1-1; II: femur d1-1-1-1; patella v2ap, p1; tibia v1-2-1, p1; metatarsus v1-1-1, p1; III: femur d1-1-1; patella d2-2, p1, v2ap; tibia d1-1, v2-2, p2-2, r2-2; metatarsus d1-2, v1-1-2ap, p1, r1-1; IV: femur d1-1-1; patella pd1, v2ap; tibia d1-1-1, v1-1-2ap, p1-1-1, r1-1-1, metatarsus d1, v2-2-1ap, p1-1, r1-1-1. Tibia I: P1 triangular, P2 with an apical strong spine; P3 with a with two apical spines (Fig. 23A). Metatarsus I with large, rounded basal spine (Fig. 23A). Palp: palpal tibia 2x the size of cymbium, 3x longer than wide (Fig. 13A); cymbium 2x longer than wide, with four apical spines (Fig. 13A, B); palpal bulb elongated-oval, embolus short, straight with strongly bend tip (Fig. 13C). Female (allotype): Total length: 5.04; carapace length: 2.35; carapace width: 1.94; abdomen length: 2.69. Carapace light brown with reticulation in the cephalic groove, radiating lines and pars thoracica; pars thoracica with arched row of long strong macrosetae (Fig. 14A). Chelicerae light brown; promargin with 10 teeth and ~18 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow, without

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 553 FIGURES 12A–D. Masteria pasochoa n. sp. (holotype). A. Male habitus, dorsal view. B. Male habitus, ventral view. Scale bars: 0.5mm.

554 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. sigilla (Fig. 14B). Eyes: six eyes grouped; AME absent, ALE and PLE almost equal; PME the smallest, oval; posterior row strongly recurved (Figs 14A, 20H). Abdomen elongated, grayish covered with long setae (Fig. 14A). Spinnerets: PLS 1.81 (0.66/0.38/0.77). Legs coloration uniformly pale yellow. Leg measurements: I missing; II missing; III 4.96 (1.48/0.68/0.95/0.98/0.87); IV 6.69 (1.73/0.80/1.70/1.48/0.98). Leg spination: I and II missing; III: femur d1-1-1-1; patella d1-2, p1, v1ap; tibia d1-1, v2-2, p1-1, r1-1; metatarsus d1-2, v1-1-2ap, p1, r1-1; IV: femur d1-1-1-1; patella p1, v2ap, r1; tibia d1-1, v1-2-2ap, p1-1-1, r1-1, metatarsus d1, v2-1-1ap, p1-1, r1-1. Palpal claw with ~15 teeth.

FIGURES 13A–D. Masteria pasochoa n. sp. (holotype). A. Male palp, prolateral view. B. Male palp, retrolateral view. C. Male palpal bulb, prolateral view. D. Male leg I, prolateral view. Scale bars: 0.1mm.

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 555 FIGURES 14A, B. Masteria pasochoa n. sp. (allotype). A. Female habitus, dorsal view. B. Female habitus, ventral view. Scale bars: 0.5mm.

556 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. Genitalia: unilobed spermathecae with large, oval ectal lobes (EcL), ental lobes (EnL) small and triangular; spermathecae bases wide, triangular; glandular region with large glands widely interspaced on the spermathecae bases (Fig. 15A). Distribution. Known from two localities in Pichincha Province. Natural History. The male specimen was hand collected at 3080m in an inter-Andean humid montane forest (Valencia & Jørgensen 2008) and the female was hand collected at 2128m, in an evergreen mountain shrubland of the northern Ecuadorian Andes (Josse & Aguirre 2013). Note. Even though the male and female were not collected at the same locality (~50km), the specimens were matched based on eyes configuration, size, and altitudinal range.

FIGURES 15A. Masteria pasochoa n. sp. (allotype), female internal genitalia, dorsal view.

Masteria lasdamas Dupérré & Tapia, new species Figs 16A, B, 17A–D, 21A, 23B, map 1.

Type material. Male holotype from Santo Domingo de los Tsachilás, OTONGA Biological Reserve, Las Damas (-00.39506 -78.98100) 1290m, 05–16 August 2014, pitfall trap, E. Tapia, I. Tapia, N. Dupérré (QCAZ). Paratypes: same locality as holotype: 1♂, 12–23 July 2014, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (ZMH-A0015054); 1♂, 28 June–12 July 2014, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (QCAZ); 1♂, 23 Jul.–5 Aug. 2014, pitfall trap, E. Tapia, C. Tapia & N. Dupérré (QCAZ). Etymology. The specific name is a noun in apposition taken from the type locality Las Damas in the OTONGA Biological Reserve. Diagnosis. Male most resemble M. otongachi n. sp. but can be distinguished by their much smaller size (2.69 vs. 4.41); prolateral process of leg I, P1 inconspicuous (Fig. 23B) vs. P1 well sclerotized and triangular in M. otongachi n. sp. (Fig. 23 C). Description. Male (holotype): Total length: 2.69; carapace length: 1.32; carapace width: 1.01; abdomen length: 1.37. Carapace light brown; pars thoracica with long setae (Fig. 16A). Chelicerae yellow; promargin with 10 teeth and ~15 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow (Fig. 16B). Eyes: six eyes grouped; AME absent, ALE the largest; PLE oval; PME small, rounded; posterior eye row recurved (Figs 16A, 21A). Abdomen elongated, whitish-gray, covered with long setae (Fig. 16A). Spin- nerets: missing. Leg formula 4123; leg measurements: I 4.59 (1.17/0.60/1.09/0.93/0.80); II 3.53 (0.95/0.45/0.74 /0.71/0.68); III 2.32 (0.92/0.41/0.68/0.68/0.53); IV 4.44 (1.23/0.41/0.99/1.09/0.72). Leg spination: I: d1-1-1-1-; tibiae v1; II: femur d1-1-1; tibia v1-1, p1; metatarsus v1-1, p1; III: femur d1-1-1-1; patella d1, p1; tibia d1, v1-1-2, p2-2, r2-2; metatarsus d1-2, v1-2ap, p1, r1-1; IV: femur d1-1-1; patella p1; tibia d1-1-1, v1-1-1ap, p1-1-1, r1-1-1,

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 557 FIGURES 16A, B. Masteria lasdamas n. sp. (holotype). A. Male habitus, dorsal view. B. Male habitus, ventral view. Scale bars: 0.5mm.

558 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. FIGURES 17A–D. Masteria lasdamas n. sp. (holotype). A. Male palp, prolateral view. B. Male palp, retrolateral view. C. Male palpal bulb, prolateral view. D. Male leg I, prolateral view. Scale bars: 0.1mm.

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 559 metatarsus d1, v1-1, p1-1, r1-1. Tibia I: P1 inconspicuous; P2 with an apical strong spine; P3 with a with two apical spines, one slightly curved apically (Fig. 23B). Metatarsus I with rounded basal spine (Fig. 23B). Palp: palpal tibia 2x the size of cymbium, 3.5x longer than wide (Fig. 17A); cymbium 2.5x longer than wide, with four apical spines (Fig. 17A, B); palpal bulb elongated-oval, embolus short, straight with bend tip (Fig. 17C). Female: unknown. Distribution. Ecuador: only found at the type locality. Natural History. Specimens were collected by pitfall trap at 1290m in an evergreen foothill forest of the occidental Andean mountain range (Guevara & Morales 2013).

Masteria otongachi Dupérré & Tapia, new species Figs 18A, B, 19A–D, 21B, 23C, map 1.

Type material. Male holotype from Ecuador, Cotopaxi, La Union del Toachi, Otongachi Natural Reserve (- 00.330510 -78.934420) 25 Feb. 2020, 1087m, pitfall, N. Dupérré, E.E. Tapia, I.G. Tapia (QCAZ). Paratypes: Ec- uador: Cotopaxi Province: La Union del Toachi, Otongachi Natural Reserve (-00.330510 -78.934420) 18 Feb. 2020, 1087m, pitfall, N. Dupérré, E.E. Tapia, I.G. Tapia (QCAZ). OTONGA Biological Reserve, Rio Esmeraldas (-00.41941 -78.99607) 1717m, 24 May-8 June 2014, 1♂, E.E. Tapia, C. Tapia & N. Dupérré (ZMH-A0015057); 5-19 Sept. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (QCAZ); (-00.41433 -79.00035) 1888m, 3- 16 Aug. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (ZMH-A0015055); 5-19 Sept. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (QCAZ); 19 Sept.–2 Oct. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (QCAZ); 25 Nov.–8 Dec. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (QCAZ); (-00.41994 -79.99623) 1997m, 21 June–2 July 2014, 1♂ (QCAZ), 5–19 Sept. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (ZMH-A0015056); (-00.41564 -79.00452) 2105m, 16 Aug.–5 Sept. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (ZMH-A0015058); (-00.42180 -79.01325) 2225m, 16 Aug.–5 Sept. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (QCAZ). Other material examined. Cotopaxi Province: OTONGA Biological Reserve, Rio Esmeraldas (-00.41941 -78.99607) 1717m, 25 Nov.–8 Dec. 2014, 2♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (DTC); (-00.41994 -79.99623) 1997m, 25 Nov.–8 Dec. 2014, 1♂, pitfall trap, E.E. Tapia, C. Tapia & N. Dupérré (QCAZ). Etymology. The specific name is a noun in apposition taken from the type locality Otongachi Natural Re- serve. Diagnosis. Males most resemble M. lasdamas n. sp., but can be distinguished by their much larger size (4.41 vs. 2.69); prolateral process of leg I, P1 well sclerotized and triangular (Fig. 23 C) whilst P1 inconspicuous in M. lasdamas n. sp. (Fig. 23B). Description. Male (holotype): Total length: 4.41; carapace length: 2.02; carapace width: 1.57; abdomen length: 2.39. Carapace light brown with reticulation in the cephalic groove and pars thoracica, covered with long setae (Fig. 18A). Chelicerae yellow; promargin with 11 teeth and ~15 smaller mesobasal teeth. Endites yellow without cuspules. Labium yellow without cuspules. Sternum yellow (Fig. 18B). Eyes: six eyes grouped; AME absent, ALE the largest; PLE oval; PME the smallest, oval; posterior row recurved (Fig. 21B). Abdomen elongated, whitish-gray covered with long setae (Fig. 18A). Spinnerets: PLS 1.78 (0.54/0.59/0.65). Leg formula 4123; leg measurements: I 6.56 (1.77/0.85/1.63/1.22/1.09); II 5.03 (1.39/0.67/1.13/1.00/0.84); III 4.20 (1.05/0.53/0.97/0.89/0.76); IV 6.72 (1.78/0.81/1.51/1.55/1.07). Leg spination: I: d1-1-1; tibiae v1-1, p1; metatarsus v1; II: femur d1-1-1-1; patella v2; tibia v1-1-1ap, p1; metatarsus v1-1, p1; III: femur d1-1-1; patella d1, r1-1; tibia d1-2, v2-1-1, p2-2, r2-2; metatarsus d1-1-2, v1-2ap, p1, r1-1; IV: femur d1-1-1; patella d1, p1; tibia d1-1-1, v1-1-1-2ap, p1-1-1, r1-1-1, metatarsus d1, v1-1-1, p1-1-1, r1-1-1. Tibia I: P1 triangular; P2 with an apical strong spine; P3 with two apical spines (Fig. 23C). Metatarsus I with large rounded basal spine (Fig. 23C). Palp: palpal tibia 2x size of cymbium, 4x longer than wide (Fig. 19A, B); cymbium 2x longer than wide, with four apical spines; palpal bulb elongated-oval, embolus short, sinuous, bent apically (Fig. 19C). Female: unknown. Distribution. Ecuador: found at two localities in the Cotopaxi Province. Natural History. Specimens were collected between 1290–2225m in evergreen foothill (Guevara & Morales 2013) to montane forest (incl. cloud forest) of the occidental Andean mountain range (Aguirre et al. 2013).

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 561 FIGURES 19A–D. Masteria otongachi n. sp. (holotype). A. Male palp, prolateral view. B. Male palp, retrolateral view. C. Male palpal bulb, prolateral view. D. Male leg I, prolateral view. Scale bars: 0.1mm.

562 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. FIGURES 20A–H. Cephalothorax, dorsal view. A. Masteria hirsuta L. Koch, 1873 (female holotype). B. Masteria jatunsacha n. sp. (male holotype). C. Masteria machay n. sp. (female holotype). D. Masteria chalupas n. sp. (male holotype). E. Masteria papallacta n. sp. (male holotype). F. Masteria papallacta n. sp. (female allotype). G. Masteria pasochoa n. sp. (male holotype). H. Masteria pasochoa n. sp. (female allotype).

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 563 FIGURES 21A, B. Cephalothorax, dorsal view. A. Masteria lasdamas n. sp. (male holotype). B. Masteria otongachi n. sp. (male holotype).

564 · Zootaxa 5005 (4) © 2021 Magnolia Press Dupérré et al. FIGURES 22A–C. Prolateral processes of leg I, male. A. Masteria jatunsacha n. sp. (holotype). B. Masteria chalupas n. sp. (holotype). C. Masteria papallacta n. sp.

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 565 FIGURES 23A–C. Prolateral processes of leg I, male. A. Masteria pasochoa n. sp. (holotype). B. Masteria lasdamas n. sp. (holotype). C. Masteria otongachi n. sp. (holotype).

Acknowledgement

The authors sincerely thank Alvaro Barragán and María Fernanda Salazar from the Museum of Invertebrates, Pon- tificia Universidad Católica del Ecuador, Quito (QCAZ) for their support. Special thanks to Dr. Giovanni Onore of OTONGA Foundation and Dr. Luis Coloma of Jambatu Foundation, Centro de Investigación y Conservacioìn de Anfibios for their friendship and technical support. To Italo G. Tapia, César A. Tapia, Carmen Caisaguano, Irène Tapia and Anabelle A. Tapia for their help in collecting specimens in the field. Sincere thanks to all our BIO-GEEC partners Maria Cristina Peñuela Mora, Maria Claudia Segovia Salcedo and Santiago Zarate Baca for their support. Special thanks to Marianela Mariño for her work in the permits process and logistics in field collection. The collection of specimens was done under the permits (No005-17 IC-FAU-DNB/MA, No003-18 IC-FAU-DNB/MA, MAE-DNB-CM-2020-0130) and the exportation of specimens was done under exportation authorization (MAE- DNB-CM-2020-0130) of the Ministerio de Ambiente, Quito, Ecuador.

References

Aguirre, Z., Toasa, G. & Gálvez, J. (2013) Región Andes. In: Galeas, R., Guevara, J.E., Medina-Torres, B., Chinchero, M.A. & Herrera, X. (Eds.), Sistema de Clasificación de Ecosistemas del Ecuador Continental. Ministerio del Ambiente del Ecua- dor, Quito, pp. 88–90. Álvarez-Padilla, F. & Hormiga, G. (2007) A protocol for digesting internal soft tissues and mounting spiders for scanning elec- tron microscopy. Journal of Arachnology, 35, 538–542.

Masteria from lowlands to highlands of Ecuador Zootaxa 5005 (4) © 2021 Magnolia Press · 567 https://doi.org/10.1636/Sh06-55.1 Guevara, J. & Morales, C. (2013) Región Andes. In: Galeas, R., Guevara, J.E., Medina-Torres, B., Chinchero, M.A. & Herrera, X. (Eds.), Sistema de Clasificación de Ecosistemas del Ecuador Continental. Ministerio del Ambiente del Ecuador, Quito, pp. 83–84. Guevara, J., Mogollón, H., Céron, C. & Josse, C. (2013a) Región Andes. In: Galeas, R., Guevara, J.E., Medina-Torres, B., Chinchero, M.A. & Herrera, X. (Eds.), Sistema de Clasificación de Ecosistemas del Ecuador Continental. Ministerio del Ambiente del Ecuador, Quito, pp. 108–110. Guevara, J., Pitmann, N., Mogollón, H., Céron, C. & Palacios, W. (2013b) Región Andes. In: Galeas, R., Guevara, J.E., Me- dina-Torres, B., Chinchero, M.A. & Herrera, X. (Eds.), Sistema de Clasificación de Ecosistemas del Ecuador Continental. Ministerio del Ambiente del Ecuador, Quito, pp. 178–180. Josse, C. & Aguirre, Z. (2013) Región Andes. In: Galeas, R., Guevara, J.E., Medina-Torres, B., Chinchero, M.A. & Herrera, X. (Eds.), Sistema de Clasificación de Ecosistemas del Ecuador Continental. Ministerio del Ambiente del Ecuador, Quito, pp. 76–77. Koch, L. (1873) In: Die Arachniden Australiens, nach der Natur beschrieben und abgebildet Bauer & Raspe, Nürnberg, pp. 369–472, pls. 28–36. https://doi.org/10.5962/bhl.title.121660 Mutke, J. & Barthlott, W. (2005) Patterns of vascular plant diversity at continental to global scales. Biologiske Skrifter, 55, 521–531. Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858. https://doi.org/10.1038/35002501 Opatova, V., Hamilton, C.A., Hedin, M., Montes de Oca, L., Král, J. & Bond, J.E. (2020) Phylogenetic systematics and evolu- tion of the spider infraorder Mygalomorphae using genomic scale data. Systematic Biology, 69 (4), 671–707. https://doi.org/10.1093/sysbio/syz064 Passanha, V. & Brescovit, A.D. (2018) On the Neotropical spider subfamily Masteriinae (Araneae, Dipluridae). Zootaxa, 4463 (1), 1–73. https://doi.org/10.11646/zootaxa.4463.1.1 Raven, R.J. (1979) Systematics of the mygalomorph spider genus Masteria (Masteriinae: Dipluridae: Arachnida). Australian Journal of Zoology, 27 (4), 623–636. https://doi.org/10.1071/ZO9790623 Raven, R.J. (1991) A revision of the mygalomorph spider family Dipluridae in New Caledonia (Araneae). In Chazeau, J. & Tillier, S. (Eds.), Zoologia Neocaledonica, Vol. 2. Mémoires du Muséum National d’Histoire Naturelle de Paris, (A), 149, pp. 87–117. Raven, R.J., Jell, P.A. & Knezour, R.A. (2015) Edwa maryae gen. et sp. nov. in the Norian Blackstone Formation of the Ipswich Basin- the first Triassic spider (Mygalomorphae) from Australia. Alcheringa: An Australasian Journal of Palaeontology, 39 (2), 259–263. https://doi.org/10.1080/03115518.2015.993300 Roewer, C.F. (1963a) Araneina: Orthognatha, Labidognatha. Insects of Micronesia, 3 (4), 105–132. Salgado, S., Cuesta, F., Báez, S., Josse, C. & Menina-Torres, B. (2013) Región Andes. In: Galeas, R., Guevara, J.E., Medina- Torres, B., Chinchero, M.A. & Herrera, X. (Eds.), Sistema de Clasificación de Ecosistemas del Ecuador Continental. Min- isterio del Ambiente del Ecuador, Quito, pp. 134–135. Simon, E. (1892a) Histoire naturelle des araignées. Deuxième édition, tome premier. Roret, Paris, 256 pp. https://doi.org/10.5962/bhl.title.51973 Simon, E. (1892c) Arachnides. In: Raffrey, A., Bolivar, I. & Simon, E., Études cavernicoles de l’île Luzon. Voyage de M. E. Simon aux l’îles Phillipines. 4e Mémoire. Annales de la Société Entomologique de France, 61, pp. 35–52. Valencia, R. & P.M. Jørgensen (2008) Composition and Structure of a Humid Montane Forest on the Pasochoa Volcano, Ecua- dor. Nordic Journal of Botany, 12 (2), 239–247. https://doi.org/10.1111/j.1756-1051.1992.tb01301.x World Spider Catalog (2021) World Spider Catalog. Version 22.0. Natural History Museum Bern, Bern. Available from: http// wsc.nmbe.ch (accessed 2 January 2021)