The goblin spiders (Araneae, Oonopidae) of the OTONGA Nature Reserve in Ecuador, with the description of seven new species

The goblin spiders (Araneae, Oonopidae) of the Otonga Nature Reserve in the Chocó region of Ecuador are reviewed. A total of 1034 adult specimens were collected in 2014 and 23 morphospecies in eight different genera were identified from these collections. We describe seven new species: one in the genus Niarchos Platnick & Dupérré: Niarchos normani sp. n.; three in Scaphidysderina Platnick & Dupérré: Scaphidysderina chirin sp. n., S. lubanako sp. n., S. tsaran sp. n.; two in Bipoonops Bolzern: Bipoonops lansa sp. n., B. pilan sp. n.; and one in Reductoonops Platnick & Berniker: Reductoonops berun sp. n. The males of Niarchos baehrae Platnick & Dupérré, 2010 and Orchestina yanayacu Izquierdo, 2017 are described here for the first time. Natural history and collecting data are given for all morphospecies collected, including Niarchos barragani Platnick & Dupérré, 2010, Scaphidysderina cotopaxi Platnick & Dupérré, 2011, Scaphidysderina pinocchio Platnick & Dupérré, 2011, Orchestina otonga Izquierdo, 2017, Orchestina santodomingo Izquierdo, 2017, Orchestina truncata Wunderlich, 2004, Reductoonops otonga Platnick & Berniker, 2014, Reductoonops pichincha Platnick & Berniker, 2014, Paradysderina fusiscuta Platnick & Dupérré, 2011, Scaphiella pich Platnick & Dupérré, 2010 and Tinadyserina otonga Platnick et al., 2013. The data show that oonopid spiders are a major element of the arachnofauna present in the Chocó neotropical forests.

Material examined is deposited in the following institutions: AMNH, American Museum of Natural History New York, NY, USA; DTC, Dupérré-Tapia Collection, Fundación OTONGA, Quito, Ecuador; QCAZ, Museum of Invertebrates, Pontificia Universidad Católica, Quito, Ecuador; ZMH, Zoological Museum Hamburg, Universität of Hamburg, Germany. Specimens were examined in 70% ethanol under a SMZ-U Nikon dissection microscope. A Nikon Coolpix 950 digital camera attached to the microscope was used to photograph all the structures to be illustrated. The digital photos were used as a template to draw the structures. Female genitalia were excised using a sharp entomological needle placed on a slide in lactic acid and observed under an AmScope XSG Series T-500 compound microscope. All measurements are expressed in millimeters and were taken using a micrometric ruler on the microscope. Photos were taken with a BK Plus Lab System by Dun, Inc. with a Canon 5DS Macro camera and a Canon 65mm lens. Morphological nomenclature follows: Dupérré (2010b, 2011a); Bolzern (2014); Platnick and Berniker (2014).

Results
A total of 1034 adult specimens were collected which comprised 23 morphospecies in eight different genera. One morphospecies could not be fitted into any of the extisting genera and probably belongs to an undescribed genus. Previous results of this study presented by Dupérré and Tapia (2016) showed that Oonopidae is the second most diverse family after Theridiidae (32 morphospecies), alongside the Tetragnathidae (22 morphospecies) and Linyphiidae (22 morphospecies). Oonopidae is the most abundant family ~27% of the total spider abundance found in the neotropical forests of the Chocó region of Ecuador (Dupérré and Tapia 2016). Among oonopids, results show that Scaphidysderina is the most abundant genus (329 specimens), followed by Niarchos (221 specimens), Tinadysderina (135 specimens), and Paradysderina (115 specimens).
Composition and distribution. Twenty-three species distributed in the Andean regions of Colombia, Ecuador and Peru (Platnick and Dupérré 2010).
Etymology. In honor of Dr. Norman Platnick, curator emeritus at the American Museum of Natural History, for his innovative and comprehensive work on the spider family Oonopidae.
Diagnosis. This species most resemble Niarchos palenque Platnick & Dupérré, 2010, but differs from all species of the genus by the horizontally directed anterior projections of the male endites (Fig. 2) and the general appearance of the bulb (Fig. 1) Description. Male (holotype). Total length: 1.8; carapace length: 1.3; carapace width: 0.9.

Niarchos baehrae
Diagnosis of male. Males are distinguished from most species by the large up-right embolus accompanied by two dark projections (Fig. 3); and from Niarchos loja Platnick & Dupérré, 2010 by the small triangular sclerotization located at the prolateral base of the embolus (Fig. 3).
Description. Male. and ventral scuta pale orange, soft portions of abdomen white. CEPHALOTHORAX: Carapace without any pattern, elongate oval in dorsal view; pars cephalica slightly elevated in lateral view, entire surface with low granulation; fovea absent. Clypeus margin slightly rebordered, vertical in lateral view. Sternum longer than wide, surface finely reticulate, microsculpture covering entire surface. Labium triangular, not fused to sternum, anterior margin indented at middle (Fig. 4). Endites distally not excavated, anteromedian tip with recurved, posteriorly directed triangular projection (Fig. 4). Chelicerae straight; without teeth. EYES: Six; reduced in size, ALE much larger than posterior eyes; all eyes oval; posterior eye row recurved from above; PLE-PME touching; ALE touching; ALE-PLE touching; PME touching.
GENITALIA: Male palp not strongly sclerotized; cymbium ovoid in dorsal view, completely fused with bulb; bulb elongated; embolus dark, accompanied by two dark pointed projections, one basal and one apical (Fig. 3).
Natural history. All specimens were collected between 1625 and 2225m, by pitfall or sifting mosses and litter.
Distribution. Only known from the type locality. Note. In Platnick and Dupérré (2010), Niarchos baehrae was placed in the cotopaxi-group based on the female genitalic features (globose, tentlike anterior receptaculum). In light of the discovery of the male, we proposed that the species belongs in the loja-group instead. The males of Niarchos baehrae does not share the characteristic retroventral projection of the palpal bulb, but an elongated embolus originating distally on the bulb and protruding far beyond the bulb, characteristic of the male of the loja-group.  Natural history. In our study specimens were collected between 1717 and 2225m. Previous records from Platnick and Dupérré (2010) suggest that the species occur from 700m to 2150m.
Composition and distribution. Twenty species distributed across Colombia, Ecuador and Peru.  Tapia Etymology. The specific epithet is a noun in apposition taken from the Tsafi´ki languange, meaning "narrow" for the slender male palpal bulb.
Diagnosis. Males are distinguished by their short, twisted basal process of the embolus (Fig. 5). Females are distinguished by the T-shaped anterior genitalic process ( Fig. 8) and from S. tsaran sp. n. by the more externally positioned apodemic projections (Fig. 8).
Natural history. All specimens were collected between 1209 and 1888m. Most specimens were collected from mid-June to mid-July, and mid-August to early September.
Distribution. Only known from the type locality. Etymology. The specific epithet is a noun in apposition taken from the Tsafi´ki languange, meaning 'red devil'.
Natural history. All specimens were collected between 1209 and 1997m.
Distribution. Only known from the type locality.  Etymology. The specific epithet is a noun in apposition taken from the Tsafi´ki languange, meaning 'beautiful'.
Diagnosis. Males are distinguished from all species by their spine-like embolic basal process (Fig. 13). Females are distinguished by their T-shaped anterior genitalic process (Fig. 17), and from S. chirin sp. n. by the very elongated of apodemic projections (Fig. 17).
Natural history. All specimens except one were collected between 2105-2225m, predominantly from mid-August to the beginning of December.
Distribution. Only known from the type locality. Diagnosis. Clypeus flattened; four spinnerets; often with only two eyes; four pairs of deep grooves on the sides of the sternum, the most anterior pair of which demarcate a short anterior portion of the sternum (Platnick and Berniker 2011: 6).
COLORATION: As in male. CARAPACE, EYES and ABDOMEN: As in male.
Natural history. Specimens were collected by sifting litter or mosses.
Distribution. Only known from the type locality.  Bolzern, 2014. Diagnosis. Carapace with an indistinct, dark spot on the posterior half. Abdomen highly patterned. Dorsal abdominal scutum present in males with dorsal scutum anteromedially fused to the epigastric scutum, absent in females. Leg spines present. Males cymbium and bulb not fused; bipartite conductor. Females differ in having a very short postepigastric scutum surrounding the pedicel, almost as wide as, not fused to the epigastric scutum, and in lacking small lateral sclerites at the epigastric area (Bolzern 2014: 56).

Bipoonops lansa
Etymology. The specific epithet is a noun in apposition taken from the Tsafi´ki languange, meaning 'orange'.
Diagnosis. Males are distinguished from all species by the short, strongly curved embolus and the small waved prolateral extension of the conductor (Fig. 22); from B. baobab Bolzern, 2014 by the large and rounded retrolateral process of the conductor (Fig. 22) which is absent in the latter (Bolzern 2014, fig. 511).
Female. Unknown. Natural history. Only a few specimens were collected but all between 1717 and 1888m.
Distribution. Only known from the type locality.
Natural history. Specimens were collected by sifting litter and by pitfall trap, between 1997 and 2225m.
Distribution. Only known from the type locality.
Natural history. All specimens were collected from mosses.
Natural history. Found in mosses and epiphytes. Distribution. Ecuador: Napo and Cotopaxi Provinces. Note. Male and female a tentatively match as they were found in the same extraction sample, however on several occasions we found up to three Orchestina species in the same sample.

Discussion
As stated by Dupérre and Tapia (2016), there have been few spider biodiversity studies in neotropical premontane-, low evergreen-and cloud forests (Yanoviak et al. 2003, Peckmezian 2009, Maya-Morales et al. 2012) and assessments of arthropod biodiversity in these systems are still rare. In addition, the comparison between studies is difficult, due to the differences in methodology and collecting techniques. Most studies tend to focus on a particular habitat (canopy, forest understory or ground), use only few collecting techniques, and are done over a short period of time. We collected spiders from various habitats except the canopy, used five different collecting techniques, had four sessions of one week sampling week each, and ran pitfall traps for four consecutive months.
Though not complete and mostly qualitative, this biodiversity assessment represents a more complete view of the arachnofauna assemblage found in these types of neotropical forests. The fact that none of the other studies even mentionned the family Oonopidae in their results is an obvious bias due to the choice of their collecting technique, habitat focus and time period. Yanoviak et al. (2003) and Maya-Morales et al. (2012), for example, focussed their studies on spider assemblages found in trees and forest understory, while Peckmezian (2009) studied different habitats and used different techniques but only applied them for a period of six days. As such, it is not surprising that our results are quite different, and reveal for the first time that the family Oonopidae turns out to be a very important component in Neotropical forests. The complementarity of techniques and microhabitats is crucial, for example Dupérré and Tapia (2016) showed that in the family Anyphaenidae, a well known arboreal group, the genus Katissa was almost exclusively collected in moss from trees, while other species were collected by beating or night collecting. In the family Oonopidae, Reductoonops and Orchestina were collected almost exclusively by sifting litter and mosses (only one male of Reductoonops was collected by pitfall); while other genera were practically only collected in pitfall traps, and clearly more abundant in a certain period of the year (Scaphidysderina, Tinadyserina). The importance of the family Oonopidae has been shown in other types of habitats, such as in a fragmented urban Atlantic forest of Brazil. Dias et al., (2005) showed that oonopids constituted over 20% of the adult spiders captured, and over 9% of the total species diversity; ranking second after Salticidae, both in abundance and diversity. Sørensen (2004) and  also showed that oonopids were important in Afrotropical montane forest, rainforests and savannah forest canopies. Sørensen (2004) proposed that at some sites Oonopidae constitute a major element of the arboreal spider fauna in terms of abundance (17%) of all adult spiders collected.  showed that in two lowland rainforests oonopids were second in abundance after Theridiidae, accounting for 10.4%-16.4% and in two savannah sites. Oonopids also ranked second in terms of abundance contributing 14.9%-21.9%.
Our results also show that the genus Scaphidysderina has a very interesting pattern of altitudinal distributions. The three most abundant species Scaphidysderina chirin sp. n., and S. lubanako sp. n., were collected respectively between 1209-1888m and 1209-1997m and are therefore found in the low evergreen montane forest, while S. tsaran sp. n. was only collected between 2105-2225m in the cloud forest, and consequently could represent a cloud forest specialist. A similar pattern was also discovered in the family Ctenidae (Dupérré 2015), four species, Chococtenus cuchilla, C. fantasma, C. neblina, C. waitti and C. kashakara were collected between 1997-2225m, and therefore were hypothesized as cloud forest specialists.
The forests of the Chocó region of Ecuador are under heavy threat from farming and logging (Sierra et al. 2003, Rival 2004. Approximately 46% of South Ecuador's original forest have already been converted into pastures and other anthropogenic land types (Tapia-Armijos et al. 2015). The evaluation of biodiversity of this well-known hotspot is largely based on the study of vertebrate ani-mals and plants and almost nothing is known about arthropod diversity. We present here some preliminary data about the importance and diversity of spiders found in this threathened region, in hope that this gives one more argument to strengthen and ensure the protection of this unique environment.