First Record of the Family Hahniidae in Ecuador with Description of Thirteen New Species and Three New Genera (Araneae: Hahniidae) ^†

Abstract

The Nearctic family Hahniidae is seldom found in South America; only 20 species occur on the continent. Herein, we present the first record of the family in mainland Ecuador, with the description of thirteen new species in five different genera. In Amaloxenops: A. minimalista n. sp. (female); in Kasha n. gen.: Kasha patpa n. sp. (male, female); in Neohahnia: Neohahnia catleyi (female) n. sp., N. piemontana n. sp. (male, female), N. pristirana n. sp. (male, female), N. freibergi n. sp. (male, female), N. paramo n. sp. (male, female), and N. chalupas n. sp. (male); in Paramito n. gen.: Paramito papallacta n. sp. and P. oyacachi n. sp.; and in Pristirana n. gen.: Pristirana barthlotti n. sp. (male, female), P. niederi n. sp. (female), and P. nowickii n. sp. (male, female). Distribution maps are presented for all species, as well as a key to the South American Hahniidae genera.

1. Introduction

The small spider family Hahniidae contains 354 species belonging to 23 genera and is distributed worldwide [1], but is more diverse in the Holarctic bioregion. South American Hahniidae diversity is relatively low, with only eight genera and 20 species occurring on the continent [1]; however, this region has a high degree of specific endemism (95%). In South America, Hahniidae is found in Argentina, Brazil, Chile, Colombia, the Falkland Islands, and Venezuela [1], but the bulk of the diversity (45%) is found in Argentina.

Most South American Hahniid species were initially studied by Simon [2], who described the genus Cybaeolus Simon, 1884 [2], alongside five species [1]. Later on, in the 1900s, Mello-Leitão [3,4] described two genera (Neohahnia Mello-Leitão, 1917 [3] and Austrohahnia Mello-Leitão, 1942 [4]), followed by Schiapelli and Gerschman [5], Roth [6], and Brignoli [7], each describing a monotypic genus, Amaloxenops Schiapelli & Gerschman, 1958 [5], Harmiella Brignoli, 1979 [7], and Lizarba Roth, 1967 [6], respectively. Since 1979, no new genera have been added to the South American fauna, and only two species, Neohahnia chibcha Heimer & Müller, 1988 [8], and Austrohahnia catleyi Rubio et al., 2014 [9] have been described.

Two subfamilies occur in South America, Hahniinae Bertkau, 1878 [10] and Cybaeolinae Lehtinen, 1967 [11]. Hahniinae are found worldwide; in South America, the genera Amaloxenops, Austrohahnia, and Intihuatana Lehtinen, 1967 [11] are endemic to Argentina; Lizarba Roth, 1967 [6] and Harmiella Brignoli, 1979 [7] are endemic to Brazil, and Neohahnia is more widely distributed and occurs in Colombia, Venezuela, and Brazil; finally, the genus Hahnia C. L. Koch, 1841 [12] is found in Argentina, Chile, Venezuela, and the Falkland Is., but the generic placement of the South American species is considered doubtful [1]. The Cybaeolinae subfamily is endemic to the Neotropical bioregion, and the genus Cybaeolus Simon, 1884 [2] occurs in Chile and Argentina.

Hahniidae are small (1.3–4.9) six or eight-eyed entelegyne ecribellate spiders [13,14]. They make small meshy webs in leaf litter, mosses, or trees [13,14], and are often collected while sifting mosses and litter or using pitfall traps (pers. obs.). Traditionally, Hahniidae were morphologically characterized through their spinnerets being aligned in row, but with recent taxonomic changes, this characteristic is no longer found in all members of the family (e.g., Chorizomma, Cicurina, Cybaeolus, and Lizarba). Interestingly, some members of the family also present a reduction in the eyes (AME reduced), have completely lost the AME, or have no eyes at all [15,16].

While conducting a biodiversity survey in the Chocó region of Ecuador, we came across a surprising number of specimens of Hahniidae from mosses, epiphytes, and litter samples. Later on, we collected specimens in pitfall traps in the Andean Paramó, and also in the Amazonian region, revealing incredible, unknown biodiversity. After detailed examination and comparison, we established that half of the species did not belong to any known genera, and all species were new to science. In this paper, 13 new species of Hahniidae are described, as well as 3 new genera.

2. Materials and Methods

Specimen imaging was conducted using a custom-made BK Plus lab System by Dun, Inc. (Palmyra, PA, USA) with an integrated Canon camera, a macro lens (65 mm), and Zerene stacking software (Zerene Systems LLC 2018, Richland, WA, USA). The female genitalia were excised utilizing a sharp entomological needle, washed in 80% alcohol, and digested with Pancreatin solution following Álvarez-Padilla and Hormiga [17]. All measurements are in millimeters and were made using a Leica M205A stereomicroscope (Wetzlar, Germany) with Leica Application Suite X. Specimens were prepared for SEM imaging via dehydration using an ethanol solution from 70% to 100%, and then transferred to hexamethyldisilazane (HMDS 99%) for 3 h. Specimens were mounted on an SEM stub, and images were obtained using a Hitachi tabletop microscope TM4000 plus (Tokyo, Japan). Schemes were made with Google Earth Pro software (version 7.3.6.9345). Scheme 1 represents the three provinces in which Hahniidae specimens were collected. The acronym ECFN, found in the text and on the labels, refers to the Ecuador Field Number and is unique to every specimen.

Abbreviations

Somatic Morphology
ac	aciniform gland spigot
cy	cylindrical gland spigot
ALE	anterior lateral eye
AME	anterior median eye
ALS	anterior lateral spinneret
mAm	minor ampullate gland spigot
MAm	major ampullate gland spigot
PLE	posterior lateral eye
PLS	posterior lateral spinneret
PMS	posterior median spinneret
PME	posterior median eye
PMS	posterior median spinneret
Genitalia (female)
at	atrium
cd	copulatory ducts
co	copulatory openings
h	hood
fd	fertilization ducts
s	spermathecae
ss	secondary spermathecae
Genitalia (male)
cf	cymbial furrow
e	embolus
ma	median apophysis
dta	dorsal tibial apophysis
rta	retrolateral tibial apophysis
pta	patellar apophysis
vl	ventral lamella
Material examined is deposited in the following institutions:
AMNH	American Museum of Natural History, New York, USA.
DTC	Dupérré-Tapia Collection, Quito, Ecuador.
QCAZ	Museum of Invertebrates, Pontificia Universidad Católica, Quito, Ecuador.
USNM	Smithsonian National Museum of Natural History, Washington D.C., USA.
ZIMG	Zoologisches Institut und Museum Greifswald, Germany.
ZMH	Zoological Museum Hamburg, Hamburg, Germany.

3. Results

3.1. Taxonomy

Family Hahniidae Bertkau, 1878 [10].

Genus Amaloxenops Schiapelli & Gerschman, 1958 [5].

Type species. Amaloxenops vianai Schiapelli & Gerschman, 1958 [5].

Diagnosis. Male palpal patellar apophysis absent ([5]; fig 19), present in all other genera (e.g., Figure 5B, Figure 11B, Figure 25B, and Figure 31B); female spermathecae elongated, without secondary spermathecae (Figure 1C; [5]; fig. 13, 14) while spermathecae are not elongated in all other genera (e.g., Figure 5D, Figure 9, Figure 25D, and Figure 29C).

Composition. Amaloxenops minimalista n. sp., A. palmarum Schiapelli & Gerschman, 1958, and A. vianai Schiapelli & Gerschman, 1958.

Distribution. Argentina and Ecuador.

3.1.1. Amaloxenops minimalista New Species

Figure 1A–C, Figure 2A–F, and Figure 3A–H, Scheme 2.

Type Material

Holotype: ♀ from Ecuador: Cotopaxi Province, Pristirana Reserve, sector Rio Esmeraldas (-00.42414°-78.95719°) 1485 m, 6 Sept. 2015, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10882 (QCAZ). Paratypes: 1♀ same data as holotype, ECFN 11263 (QCAZ); Pristirana Reserve (-00.42414°-78.95719°), 1♀, June–August 2014, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10883 (ZMH-A0020679); 1♀, Nov. 2014, ECFN 10880 (ZMH-A0020682); 2♀, 6 May 2014, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10884 (ZMH-A0020680); 2♀, Oct.–Nov. 2014, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10885 (ZIMG); 2♀, 7 Nov. 2015, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10886 (QCAZ); 1♀, 8 May 2017, sifting litter from epiphytes, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10887 (ZMH-A0020681); 2♀, Dec. 2014, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10888 (USNM); 1♀, Nov. 2014, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10889 (QCAZ); 17♀, Oct.–Nov. 2014, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10890 (AMNH).

Etymology. The specific epithet is a noun in apposition taken from the Latin, referring to the very simple form of the female genitalia.

Diagnosis. Females closely resemble A. palmarum (Schiapelli & Gerschman [5]) but can be differentiated by their copulatory ducts being close together (Figure 1C) while the copulatory ducts are widely separated in the latter ([5]; fig. 19).

Description. Female (holotype). Total length: 1.73; carapace length: 0.72; carapace width: 0.56; abdomen length: 1.01.

Cephalothorax. Carapace pear-shaped, light yellow suffused with dark brown medially and submarginally (Figure 1A), smooth dorsally with granulation on radiating lines; fovea absent (Figure 1A and Figure 2A,B); pars cephalica flat, pars thoracica incline. Sternum as long as wide, light yellow lightly suffused with dark brown marginally (Figure 1A); truncated (Figure 2C); labium and endites light brown (Figure 1A); serrula present with 34 teeth (Figure 2D). Clypeus: 0.06. Chelicerae light yellow; promargin, retromargin with two teeth. Eight eyes surrounded by black pigmentation; AME: 0.02, ALE 0.05, PLE: 0.05, PME: 0.04 (Figure 1A and Figure 2A).

Abdomen. Oval, whitish with pattern of four–five broken chevrons, covered with long black setae (Figure 1A); ventral side with light beige; tracheal spiracle recurved, 6× further from the epigastric groove than the spinnerets (Figure 1A and Figure 2F); spinnerets aligned (Figure 3A); PMS beige, separated by half their width; ALS basal segment elongated, dark brown, apical segment short, whitish; PLS basal segment elongated, dark brown, apical segment elongated and whitish (Figure 1A); PMS with one minor ampullate, one cylindrical and five aciniform gland spigots (Figure 3E,F); ALS with one major ampullate (Figure 3G), and PLS with one cylindrical and four–five aciniform gland spigots (Figure 3H).

Legs. Light yellow–orange with two dark ventral marks on femurs I–III; one dark ventral mark on femur IV, tibia, patellae, and metatarsi; tarsi uniformly light yellow–orange; palpal femur and patellae light yellow–orange suffused lightly with black; tibiae and tarsi dark orange–brown suffused with black; palpal tarsal claw present; leg measurements: I 1.78 (0.54/0.20/0.41/0.35/0.28); II 1.76 (0.53/0.20/0.40/0.35/0.28); III 1.57 (0.45/0.18/0.37/0.33/0.27); IV 1.92 (0.53/0.18/0.45/0.44/0.32); leg formula: 4123. Macrosetae present on tibia and patellae I–IV dorsally; ventrally on metatarsi and tarsi III–IV; trichobothria present on tibia, metatarsi, and tarsi (Figure 3C,D); tarsal organ rounded located in concavity (Figure 3C,D, white arrow); tarsal claw uniseriate with six–seven teeth, lower claw with two teeth (Figure 3B).

Genitalia. Epigynal plate lightly sclerotized; with single medial atria, with double hood (Figure 1B); copulatory openings close together (Figure 1C). Internal genitalia copulatory ducts, short and sinuous; spermathecae elongated; fertilization duct wide, positioned basally (Figure 1C).

Male. Unknown.

Distribution. Only known from the type locality in Cotopaxi province.

Natural History. All specimens were collected in a low evergreen mountain forest (BsBn04) of the Western Cordillera [18] at 1485 m where it lives in sympatry with Kasha patpa n. sp., Neohahnia pristirana n. sp., and Pristirana niederi n. sp.

Figure 1. Amaloxenops minimalista new species, female holotype. (A) Habitus, dorsal and ventral view (arrow points to tracheal spiracle). (B) Epigynum, ventral view. (C) Internal genitalia, dorsal view. Scale bar: 0.5 mm.

Figure 1. Amaloxenops minimalista new species, female holotype. (A) Habitus, dorsal and ventral view (arrow points to tracheal spiracle). (B) Epigynum, ventral view. (C) Internal genitalia, dorsal view. Scale bar: 0.5 mm.

Figure 2. Amaloxenops minimalista new species, female paratype (ECFN 10890), SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Sternum, ventral view. (D) Endites and labium, ventral view. (E) Epigynal plate, ventral view. (F) Abdomen, ventral view (arrow points to tracheal spiracle).

Figure 2. Amaloxenops minimalista new species, female paratype (ECFN 10890), SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Sternum, ventral view. (D) Endites and labium, ventral view. (E) Epigynal plate, ventral view. (F) Abdomen, ventral view (arrow points to tracheal spiracle).

Figure 3. Amaloxenops minimalista new species, female paratype (ECFN 10890) SEM. (A) Spinnerets, ventral view. (B) Tarsal claw II, lateral view. (C) Tarsus I, dorsal view (arrow points to tarsal organ). (D) Tarsus II, dorsal view (arrow points to tarsal organ). (E) PMS, apical view. (F) PMS, apical view. (G) ALS, apical view. (H) PLS, apical view.

Figure 3. Amaloxenops minimalista new species, female paratype (ECFN 10890) SEM. (A) Spinnerets, ventral view. (B) Tarsal claw II, lateral view. (C) Tarsus I, dorsal view (arrow points to tarsal organ). (D) Tarsus II, dorsal view (arrow points to tarsal organ). (E) PMS, apical view. (F) PMS, apical view. (G) ALS, apical view. (H) PLS, apical view.

3.1.2. Kasha New Genus

Type species. Kasha patpa n. sp.

Etymology. The generic epithet is taken from the Kichwa language “Kasha” meaning “spiny” referring to the elongated spines found on the species. The gender is feminine.

Diagnosis. The genus is distinguished from all other Hahniidae genera by the presence of white plumose setae on the dorsal surface of abdomen (Figure 4A and Figure 6F) and all patellae with two extremely long macrosetae (Figure 4A and Figure 6H arrows).

Description. Small spider, total length 1.97–2.2.

Cephalothorax. Carapace pear-shaped, longer than wide; smooth; pars cephalica flat, pars thoracica inclined; fovea absent (Figure 4A and Figure 6A,B). Sternum heart-shaped, wider than long; smooth; truncated basally (Figure 4B and Figure 6D). Labium hexagonal, endites with serrula (Figure 6D). Cheliceral promargin and retromargin with two–three teeth. Eight eyes surrounded with black pigmentation, AME minute 1/4 the size of PME; ALE, PLE, and PME of almost equal size (Figure 4A and Figure 6A,B).

Abdomen. Oval, dorsally covered with short white plumose setae and long grayish setae (Figure 4A and Figure 6E,F); ventrally with plumose setae (Figure 7C,D).

Spinnerets aligned; PMS long, almost as long as ALS; ALS, PLS long, PLS apical segment as long as basal one (Figure 4B and Figure 7B,C).

Legs. Macrosetae present on femur, tibia, and metatarsi III, IV; patellae with two elongated macrosetae (Figure 4A and Figure 6H arrows); trichobothria present on tibia, metatarsi, and tarsi; leg formula: 4123; three claws, first part enlarged (only observed for claw IV) (Figure 7A); tarsal organ rounded set in a concavity (Figure 7E). Female palpal claw present; epigynum with large copulatory openings (Figure 5D and Figure 6G); male palpal patellar apophysis short, bidentate; tibia with two apophyses (Figure 5B); palpal cymbium pointed, with small furrow; embolus short, starting prolatero-basally, median apophysis absent (Figure 5A).

Composition. Kasha patpa n. sp.

Distribution. Only found in the Chocó region of Ecuador.

3.1.3. Kasha patpa New Species

Figure 4A,B, Figure 5A–D, Figure 6A–H and Figure 7A–E, Scheme 2.

Type Material

Holotype: ♀ from Ecuador, Cotopaxi Province, San Francisco de las Pampas, Pristirana Reserve (-00.42414°-78.95719°) 1485 m, XI. 2013, E. Tapia, N. Dupérré, C. Tapia, ECFN 10893 (QCAZ). Paratypes: 1♂, same data as holotype, ECFN 10894 (QCAZ); 1 ♀, San Francisco de las Pampas, Pristirana Reserve (-00.42414°-78.95719°) 1480 m, 24 Nov. 2019, E. Tapia, Faml. Tapia Caisaguano, ECFN 4035 (QCAZ); 2♀, San Francisco de las Pampas, Pristirana Reserve (-00.424920°-78.957080°) 1449 m, beating epiphytes, Faml. Tapia Caisaguano, ECFN 1755 (ZMH-A0020683).

Etymology. The specific epithet is a noun in apposition taken from the Kichwa language meaning “feather” in reference to the feathery setae found on the abdomen of the species.

Diagnosis. Males and females are distinguished from all other Hahniidae genera by their abdomen covered by white plumose setae dorsally (Figure 5A), plumose setae absent in all other genera (e.g., Figure 1A, Figure 17A, and Figure 30A).

Description. Female (holotype). Total length: 2.22; carapace length: 0.83; carapace width: 0.58; abdomen length: 1.39.

Cephalothorax. Carapace pear-shaped, light yellow suffused with black medially and marginally, smooth; fovea absent (Figure 4A and Figure 6A,B); pars cephalica flat, pars thoracica inclined. Clypeus: 0.12. Sternum wider than long, uniformly light yellow, smooth, truncated posteriorly (Figure 4B and Figure 6D); labium and endites light yellow, serrula present with ~37 teeth (Figure 4B and Figure 6D). Chelicerae yellow; promargin with two teeth, retromargin with three teeth. Eight eyes surrounded by black pigmentation; AME 0.02, ALE 0.07, PLE 0.08, PME 0.08.

Abdomen. Oval, light grayish dorsally suffused with dark gray medially and darker gray laterally, covered with whitish short, plumose setae, and grayish elongated setae (Figure 4A); ventral side uniformly beige with semi-plumose setae (Figure 4B and Figure 7C); tracheal spiracle recurved, situated 8× further from the epigastric groove than the spinnerets; spinnerets aligned (Figure 4B and Figure 7C); PMS long whitish, touching; ALS basal segment whitish basally, apically suffused with dark ring, apical segment short, brownish; PLS long, basal and apical segments whitish (Figure 7B,C).

Legs. Light yellow, femur I–IV with two ventral dark marks, tibia and metatarsi with one ventral dark mark (Figure 4A,B); macrosetae present on all femur, tibiae, and on metatarsi III and IV, absent on tarsi; two extremely elongated macrosetae on all patellae (Figure 4A,B and Figure 6H); upper tarsal claw wide, with five–six teeth (Figure 7A); tarsal organ rounded in deep concavity (Figure 7E). Leg measurements: I 2.06 (0.54/0.23/0.47/0.46/0.36); II 1.94 (0.49/0.22/0.43/0.46/0.33); III 1.91 (0.41/0.21/0.42/0.56/0.31); IV 2.41 (0.61/0.21/0.62/0.59/0.38); leg formula: 4123.

Genitalia. Epigynal plate flat, lightly sclerotized; copulatory openings large, C-shaped; atrium deep (Figure 5C,D). Internal genitalia with short, wide copulatory ducts; secondary spermathecae drop-shaped; spermathecae rounded with antero-externally positioned small knob-like extrusion; fertilization ducts short, externo-basally positioned (Figure 5D).

Male (paratype). Total length: 1.97; carapace length: 0.87; carapace width: 0.60; abdomen length: 1.10.

Cephalothorax. Carapace and sternum as in female. Clypeus: 0.11. Chelicerae promargin, retromargin with three teeth. Eight eyes surrounded by black pigmentation; AME minute, 0.02, ALE, PLE, and PME 0.08.

Abdomen. As in female, slightly paler.

Legs. As in female. Leg measurements: I 2.73 (0.78/0.26/0.66/0.61/0.42); II 2.58 (0.77/0.23/0.61/0.55/0.42); III 2.36 (0.70/0.21/0.52/0.54/0.39); IV 2.88 (0.76/0.27/0.66/0.75/0.44); leg formula: 4123.

Genitalia. Palpal patellar apophysis short, bidentate basally positioned; tibia with two apophyses, one large retrolateral, bidentate apophysis, and a small dorsal pointed one; (Figure 5B); two retrolateral and two dorsal trichobothria (Figure 5B); cymbium pointed, cymbial furrow small, with small lamella basally (Figure 5A,B); embolus short starting prolatero-basally; median apophysis absent (Figure 5A).

Distribution. Only found at the type locality in Cotopaxi Province.

Natural History. All specimens were collected in a low evergreen mountain forest (BsBn04) of the Western Cordillera [18] between 1449 and 1485 m, where it lives in sympatry with Amaloxenops minimalista n. sp., Neohahnia pristirana n. sp., and Pristirana niederi n. sp.

Figure 4. Kasha patpa new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 4. Kasha patpa new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 5. Kasha patpa new species. (A) Male palp, ventral view. (B) Male palp, retrolateral view. (C) Epigynum, ventral view. (D) Internal genitalia, dorsal view.

Figure 5. Kasha patpa new species. (A) Male palp, ventral view. (B) Male palp, retrolateral view. (C) Epigynum, ventral view. (D) Internal genitalia, dorsal view.

Figure 6. Kasha patpa new species, female paratype (ECFN 1755) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Carapace, frontal view. (D) Sternum and endites, ventral view. (E) Abdominal plumose setae and macrosetae, ventral view (arrow points to macrosetae). (F) Abdominal plumose setae, ventral view details. (G) Epigynal plate, ventral view. (H) Patella I, dorsal view (arrows point to macrosetae).

Figure 6. Kasha patpa new species, female paratype (ECFN 1755) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Carapace, frontal view. (D) Sternum and endites, ventral view. (E) Abdominal plumose setae and macrosetae, ventral view (arrow points to macrosetae). (F) Abdominal plumose setae, ventral view details. (G) Epigynal plate, ventral view. (H) Patella I, dorsal view (arrows point to macrosetae).

Figure 7. Kasha patpa new species, female paratype (ECFN 1755) SEM. (A) Tarsal claw IV, apical view. (B) Spinnerets, ventral view. (C) Tracheal spiracle, ventral view (arrow points to tracheal spiracle). (D) Tracheal spiracle, details. (E) Tarsus III, dorsal view (arrow points to tarsal organ).

Figure 7. Kasha patpa new species, female paratype (ECFN 1755) SEM. (A) Tarsal claw IV, apical view. (B) Spinnerets, ventral view. (C) Tracheal spiracle, ventral view (arrow points to tracheal spiracle). (D) Tracheal spiracle, details. (E) Tarsus III, dorsal view (arrow points to tarsal organ).

3.1.4. Genus Neohahnia Mello-Leitão 1917

Type species. Neohahnia sylviae Mello-Leitão, 1917.

Composition. Neohahnia catleyi n. sp., N. chalupas n. sp., N. chibcha Heimer and Müller, 1988, N. ernsti (Simon, 1897), N. freibergi n. sp., N. palmicola Mello-Leitão, 1917, N. paramo n. sp., N. piemontana n. sp., N. pristirana n. sp., N. sylviae Mello-Leitão, 1917.

Distribution. St. Vincent, Venezuela, Colombia, Ecuador, and Brazil.

Diagnosis. Males are distinguished from all other genera by their palp with embolus long, 1–2× encircling the tegulum (Figure 11A and Figure 18A); whereas all other genera have the embolus shorter, not encircling the tegulum ([5]; fig. 14; Figure 5A, Figure 25A, and Figure 35A); female internal genitalia with long copulatory ducts with two or more loops (Figure 11D, Figure 18D, and Figure 20D), all other genera with copulatory ducts short without loops (e.g., Figure 1C, Figure 5D, Figure 26D, and Figure 29C).

3.1.5. Neohania catleyi New Species

Figure 8A–D, Scheme 2.

Type Material

Holotype: ♀ from Ecuador: Santo Domingo de los Tsáchilas province, Alluriquin, La Florida (-00.25254°-79.03043°) 884 m, 26 Sept. 2014, hand collected in mosses from Lauraceae tree 20m high, N. Dupérré, E. Tapia, ECFN 10919 (QCAZ). Paratype: 1 ♀, same data as holotype, ECFN 10918 (QCAZ).

Etymology. The specific epithet is a patronym in honor of Dr. Kefyn M. Catley in recognition for his work on Austral South American Hahniidae.

Diagnosis. Females most resemble Neohania pristirana n. sp. in overall coloration (compare Figure 8A,B and Figure 17A,B) but can be distinguished by their oval spermathecae and elongated with small knob-like extrusion secondary spermathecae (Figure 8D), while the latter have bean-shaped spermathecae and rounded secondary spermathecae (Figure 18D).

Description. Female (holotype). Total length: 1.14; carapace length: 0.54; carapace width: 0.40; abdomen length: 0.60.

Cephalothorax. Carapace pear-shaped, light yellow–brown strongly suffused with dark pattern, shiny; fovea absent (Figure 8A); pars cephalica flat, pars thoracica with strong incline. Sternum longer than wide, light yellow medially, strongly suffused with black submarginally and marginally; truncated (Figure 8A); labium and endites light yellow–brown suffused with black; serrula present. Clypeus: 0.04. Chelicerae light yellow–brown suffused with black; promargin and retromargin not observed. Six eyes surrounded by black pigmentation; AME absent, ALE 0.05, PLE 0.06, PME 0.05.

Abdomen. Oval; dorsally light beige suffused with black, with pattern of three–four complete chevrons (Figure 8A); ventrally light beige suffused with dark gray in mottled pattern (Figure 8A); tracheal spiracle recurved, 11× further from the epigastric groove than the spinnerets; spinnerets aligned; PMS whitish, almost touching; ALS basal segment suffused with black, apical segment white; PLS basal segment suffused with black basally, apical segment white (Figure 8A).

Legs. Light yellow–brown with two ventral black marks on femurs, yellow–brown with one ventral black mark on tibia and metatarsi; tarsi uniformly light yellow–brown; leg measurements: I 1.33 (0.40/0.14/0.30/0.24/0.25); II 1.15 (0.35/0.13/0.24/0.22/0.21); III 1.02 (0.27/0.13/0.19/0.22/0.21); IV 1.37 (0.35/0.16/0.32/0.30/0.24); leg formula: 4123. Macrosetae present on patellae and tibia I–IV dorsally; trichobothria present on tibia, metatarsi, and tarsi.

Genitalia. Epigynal plate flat, lightly sclerotized, copulatory opening as rugged slit; situated medially (Figure 8C). Internal genitalia with long, semi-transparent coiled copulatory ducts; secondary spermathecae elongated; spermathecae oval; fertilization ducts long and thick (Figure 8D).

Distribution. Only known from the type locality in Santo Domingo de los Tsáchilas province.

Natural History. Specimens were collected in tree mosses, in an evergreen foothill forest (BsPn01) of the Western Cordillera [19] at 884 m where it lives in sympatry with Neohahnia piemontana n. sp.

Figure 8. Neohahnia catleyi new species, female holotype. (A) Habitus, dorsal and ventral view (arrow points to tracheal spiracle). (B) Habitus, frontal view. (C) Epigynum, ventral view. (D) Internal genitalia, dorsal view. Scale bars: 0.5 mm.

Figure 8. Neohahnia catleyi new species, female holotype. (A) Habitus, dorsal and ventral view (arrow points to tracheal spiracle). (B) Habitus, frontal view. (C) Epigynum, ventral view. (D) Internal genitalia, dorsal view. Scale bars: 0.5 mm.

3.1.6. Neohahnia piemontana New Species

Figure 9, Figure 10A,B, Figure 11A–F, Figure 12A–F, Figure 13A–H, Figure 14A–F, and Figure 15A–H, Scheme 2.

Type Material

Holotype: ♀ from Ecuador, Pichincha province, Otongachi reserve (-00.321295°-78.95163°) 900 m, 27.xii.2012, sifting litter, Berlese, N. Dupérré & E. Tapia, ECFN 10900 (QCAZ). Paratypes: 3♂6♀, same data as holotype, ECFN 11264 (QCAZ); Otongachi reserve (-00.321295°-78.95163°) 900 m, 23.xii.2013, sifting litter, N. Dupérré & E. Tapia, ECFN 10892 (ZIMG), 1♂, ECFN 10891 (ZMH-A0020689); La Unión del Toachi, Otongachi Natural Reserve (-00.330510°-78.934420°) 1087 m, 13 Feb. 2020, 2♂1♀, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 6784 (USNM); 3♂3♀, ECFN 6253 (ZIMG); 2♂2♀, ECFN 6757 (AMNH); 27 Feb. 2020, 2♂, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 4814 (QCAZ); 13 Feb. 2020, 1♂1♀, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 6771 (ZMH-A0020684); 20 Feb. 2020, 5♂2♀, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 6722 (QCAZ); 16 Feb. 2020, 5♂2♀, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 6743 (ZMH- A0020685).

Other material examined: Ecuador: Pichincha province: La Unión del Toachi, Otongachi Natural Reserve (-00.321295°-78.95163°) 900 m xii.2012, 1♂8♀, sifting litter, N. Dupérré & E. Tapia, ECFN 10899 (ZMH-A0020691); (-00.330510°-78.934420°) 1087 m, 7 Feb. 2020, 1♂, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 6665 (DTC); 13 Feb. 2020, 4♂1♀, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 6223, ECFN 6247 (AMNH); 1♂2♀, ECFN 6647 (DTC); 1♂, ECFN 6737 (DTC); 3♂, ECFN 5622, 6220, 6734 (ZMH-A0013180, A0013186, A0020686); 18 Feb. 2020, 1♂, pitfall traps, I.G. Tapia, N. Dupérré & E. Tapia, ECFN 7450 (ZMH-A0020686); 2♂, ECFN 5650 (DTC); 20 Feb. 2020, 2♂, pitfall traps, I.G. Tapia, N. Dupérré, E. Tapia, ECFN 6209 (ZIMG); 3♂1♀, ECFN 6697 (ZIMG); ECFN 6200 (ZMH-A0013185); 25 Feb. 2020, 1♂, pitfall traps, I.G. Tapia, N. Dupérré, E. Tapia, ECFN 4826 (DTC); ECFN 5613 (ZMH-A0013179); 27 Feb. 2020, 2♂2♀, pitfall traps, I.G. Tapia, N. Dupérré, E. Tapia, ECFN 6685 (USNM); 3♂1♀, ECFN 6666 (ZMH-A0020687); 3♂, ECFN 6635 (ZIMG); 1♂1♀, ECFN 6609 (ZIMG); ECFN 6633, 6634 (ZMH-A0013087, A0013088). Santo Domingo de los Tsáchilas: La Florida (-00.248529°-79.026887°) 868 m, 6 Feb. 2020, 1♂, ECFN 6892 (DTC); 1♂, ECFN 6908 (DTC); 13 Feb. 2020, 3♂, ECFN 6817 (QCAZ); 20 Feb. 2020, 2♂1♀, pitfall traps, I. Tapia, N. Dupérré, E. Tapia, ECFN 6878 (DTC); 1♂, ECFN 6868 (QCAZ); 27 Feb. 2020, 1♂1♀, pitfall traps, I. Tapia, N. Dupérré, E. Tapia, ECFN 6841 (ZMH-A0020690); OTONGA Reserve, Las Damas (-00.39506°-78. 98100°) 1290 m, 23 March–5 April 2014, 1♀, pitfall traps, I. Tapia, N. Dupérré, E. Tapia, ECFN 10898 (DTC); 5–15 April 2014, 1♂, ECFN 10897 (DTC).

Etymology. The specific epithet is a noun in apposition taken from the Spanish language, referring to the region where the species occur.

Diagnosis. Males and females are distinguished from most species by the absence of abdominal chevrons (Figure 9 and Figure 10A). From similar species N. ernsti Simon, 1898 [20] and N. freibergi n. sp. by the female epigynum with copulatory openings separated by 1.5× their diameter (Figure 11C), while in N. ernsti the copulatory openings are separated by 1× their diameter ([11]: p. 370) and joined in N. freibergi n. sp. (Figure 20C). Males are separated from both species by their bi-dentate patellar apophysis (Figure 11B), while tri-pointed in N. ernsti ([11]: fig. 366) and single pointed in N. freibergi n. sp. (Figure 20B).

Description. Female (holotype). Total length: 1.6; carapace length: 0.8; carapace width: 0.5; abdomen length: 0.8.

Cephalothorax. Carapace pear-shaped, light yellow with light gray pattern, shiny; pars cephalica flat, pars thoracica with strong incline (Figure 10A and Figure 12A). Sternum light yellow, margin sometimes suffused with gray; heart-shaped, truncated (Figure 10B and Figure 13A); labium light yellow; endites light yellow, serrula present with 34 teeth (Figure 12F). Clypeus short: 0.06. Chelicerae light yellow; promargin and retromargin with two teeth. Eight eyes surrounded by black pigmentation; AME 0.03, ALE 0.03, PLE 0.03, PME 0.04.

Abdomen. Oval; light to dark gray, molted, covered with long black setae (Figure 9 and Figure 10A); ventral side uniformly light gray; tracheal spiracle straight, 1.5× further from the epigastric groove than the spinnerets (Figure 10B and Figure 12C, arrow); spinnerets aligned; uniformly light brown; PMS separated, with minor ampullate, two cylindrical and four aciniform glans spigots (Figure 10B and Figure 13D,E); ALS apical segment short, with major ampullate gland spigots (Figure 13F); PLS with segments of equal size, with one cylindrical and seven aciniform gland spigots (Figure 10B and Figure 13G).

Legs. Femur light yellow, tibia to tarsi light brown; leg measurements: I 1.39 (0.43/0.14/0.31/0.26/0.25); II 1.31 (0.40/0.14/0.27/0.23/0.27); III 1.22 (0.34/0.13/0.30/0.21/0.24); IV 1.49 (0.39/0.16/0.33/0.32/0.29); leg formula: 4123, palpal tarsal claw present (Figure 13H). Macrosetae present on patellae and tibia I–IV dorsally; trichobothria present on tibia, metatarsi and tarsi; tarsal organ rounded located in concavity (Figure 13B,C).

Genitalia. Epigynal plate flat, lightly sclerotized; large, oval copulatory openings, deep atria, positioned antero-laterally (Figure 11C and Figure 12E). Internal genitalia with long and coiled copulatory ducts; secondary spermathecae large; spermathecae rounded; fertilization duct thin, positioned externally (Figure 11D).

Male (paratype). Total length: 1.3; carapace length: 0.5; carapace width: 0.4.

Cephalothorax. Carapace as in female (Figure 10A and Figure 14B). Chelicerae, sternum, labium, and endites as in female (Figure 10B and Figure 14C,D); serrula with 37 teeth (Figure 14E). Clypeus short, 0.07. Eyes: AME 0.03, ALE 0.03, PLE 0.03, PME 0.04.

Abdomen. As in female (Figure 10B and Figure 14A). Spinnerets aligned; uniformly light brown; PMS separated, with minor ampullate, and three aciniform glans spigots (Figure 15F); ALS apical segment short, with major ampullate gland spigot (Figure 15G); PLS with segments of equal size, with one cylindrical, and four aciniform gland spigots (Figure 15H).

Legs. As in female (Figure 10A,B).

Leg measurements: I 1.34 (0.39/0.15/0.31/0.25/0.24); II 1.19 (0.33/0.13/0.26/0.23/0.24); III 1.16 (0.31/0.13/0.23/0.24/0.25); IV 1.46 (0.37/0.15/0.33/0.32/0.29); leg formula: 4123.

Genitalia. Palpal patella with long basal spur, bi-pointed apically (Figure 11B and Figure 15E); tibia with large, curved and rugose retrolateral tibial apophysis (Figure 11B and Figure 15C,D); cymbium rounded apically; with retrolateral cymbial furrow (Figure 11A,B); embolus originating basally, long and thin; 2.5× circling the tegulum; not protruding (Figure 11A,B and Figure 15F).

Distribution. Only known from the type locality in Pichincha Province.

Natural history. All specimens were collected in an evergreen foothill forest (BsPn01) of the Western Cordillera [19] between 868 and 1290 m. At the locality, La Florida, the species lives in sympatry with Neohania catleyi n. sp. and in OTONGA Reserve, Las Damas with Neohania pristirana n. sp.

Figure 9. Neohahnia piemontana new species, female. Habitus, dorsal view.

Figure 9. Neohahnia piemontana new species, female. Habitus, dorsal view.

Figure 10. Neohahnia piemontana new species. (A) Male paratype and female holotype, habitus dorsal view. (B) Male paratype and female holotype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 10. Neohahnia piemontana new species. (A) Male paratype and female holotype, habitus dorsal view. (B) Male paratype and female holotype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 11. Neohahnia piemontana new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female paratype, internal genitalia dorsal view. (E) Male chelicerae, posterior view. (F) Female spinnerets, ventral view.

Figure 11. Neohahnia piemontana new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female paratype, internal genitalia dorsal view. (E) Male chelicerae, posterior view. (F) Female spinnerets, ventral view.

Figure 12. Neohahnia piemontana new species, female (ECFN 6878) SEM. (A) Carapace, dorsal view. (B) Abdomen, dorsal view. (C) Abdomen, ventral view (arrow point to tracheal spiracle). (D) Epigastric region, ventral view. (E) Epigynum, ventral view. (F) Serrula, ventral view.

Figure 12. Neohahnia piemontana new species, female (ECFN 6878) SEM. (A) Carapace, dorsal view. (B) Abdomen, dorsal view. (C) Abdomen, ventral view (arrow point to tracheal spiracle). (D) Epigastric region, ventral view. (E) Epigynum, ventral view. (F) Serrula, ventral view.

Figure 13. Neohahnia piemontana new species, female (ECFN 6878) SEM. (A) Sternum, ventral view. (B) Tarsus II, dorsal view (arrow points to tarsal organ). (C) Tarsus IV, dorsal view (arrow points to tarsal organ). (D) Spinnerets, ventral view. (E) PMS, ventral view. (F) ALS, apical view. (G) PLS, apical view. (H) Palpal claw, lateral view.

Figure 13. Neohahnia piemontana new species, female (ECFN 6878) SEM. (A) Sternum, ventral view. (B) Tarsus II, dorsal view (arrow points to tarsal organ). (C) Tarsus IV, dorsal view (arrow points to tarsal organ). (D) Spinnerets, ventral view. (E) PMS, ventral view. (F) ALS, apical view. (G) PLS, apical view. (H) Palpal claw, lateral view.

Figure 14. Neohahnia piemontana new species, male (ECFN 6878) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Endites, ventral view. (D) Sternum, ventral view. (E) Serrula, ventral view. (F) Epigastric region, ventral view.

Figure 14. Neohahnia piemontana new species, male (ECFN 6878) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Endites, ventral view. (D) Sternum, ventral view. (E) Serrula, ventral view. (F) Epigastric region, ventral view.

Figure 15. Neohahnia piemontana new species, male (ECFN 6878) SEM. (A) Tibia IV, dorsal view. (B) Palp, retrolateral view. (C) Palpal tibia, retrolateral view. (D) Palpal patella retrolateral view. (E) Palp, ventral view. (F) PMS, apical view. (G) ALS, apical view. (H) PLS, apical view.

Figure 15. Neohahnia piemontana new species, male (ECFN 6878) SEM. (A) Tibia IV, dorsal view. (B) Palp, retrolateral view. (C) Palpal tibia, retrolateral view. (D) Palpal patella retrolateral view. (E) Palp, ventral view. (F) PMS, apical view. (G) ALS, apical view. (H) PLS, apical view.

3.1.7. Neohahnia pristirana New Species

Figure 16, Figure 17A,B, and Figure 18A–D, Scheme 2.

Type Material

Holotype: ♀ from Ecuador, Cotopaxi Province, San Francisco de Las Pampas, Pristirana Reserve (-00.42414°-78.95719°) 1485 m, June–August 2013, sifting litter from epiphytes, N. Dupérré, E.E. Tapia, C. Tapia, ECFN 10903 (QCAZ). Paratypes: 1♂2♀1juvenile, same data as holotype (QCAZ) ECFN 11255; Cotopaxi Province, San Francisco de la Pampas, Pristirana Reserve (-00.424920°-78.957080°) 1449 m, 27 Feb. 2019, 1♀, hand collecting, Faml. Tapia Caisaguano, ECFN 2138 (ZMH-A0020673); 15 Feb. 2019, 1♂, beating epiphytes, Faml. Tapia Caisaguano ECFN 1738 (ZMH-A0020674); 27 Feb. 2019, 1♀, hand collecting, Faml. Tapia Caisaguano ECFN 2229 (QCAZ); San Francisco de la Pampas, Pristirana Reserve, sector Rio Esmeraldas (-00.424414°-78.95719°) 1485 m, 8 May 2017, 1♀, sifting litter from epiphytes, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10902 (QCAZ); 7 Nov. 2015, 1♀ (ZMH-A0020675).

Other material examined. ECUADOR: Cotopaxi province: San Francisco de la Pampas, OTONGA Natural Reserve (-00.41994°-79.00623°), 1997 m, 4–7 Sept. 2014, 6♀, sifting litter, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10912, 10915 (AMNH); 13–15 Nov. 2014, 4♀, sifting litter from epiphytes, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10907 (ZIMG); 13 Nov. 2014, 1♂, sifting litter from epiphytes, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10914 (ZIMG); 8–21 June 2014, 1♀, sifting mosses, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10906 (DTC); 24 May–8 June 2014, 2♀, pitfall traps, N. Dupérré, E. E. Tapia, C. A. Tapia, 10913 (DTC); 24–30 May 2014, 2♂3♀, beating trees, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10909 (AMNH); 24–30 May 2014, 1♀, sifting mosses, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10908 (ZMH-A0020676); 8 Dec. 2014, 1♀, sifting litter from base of epiphytes, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10910 (QCAZ); San Francisco de la Pampas, OTONGA reserve (-00.42180°-79.01325°), 2225 m, 25 Nov.–8 Dec. 2014, 1♂, pitfall traps, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10911 (DTC). Santo Domingo de los Tsáchilas: OTONGA reserve, sector las Damas (-00.40855°-78.99681°) 1582 m, 4–7 Sept. 2014, 3♂10♀, beating branches, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10905 (USNM).

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

Diagnosis. Females and males are easily distinguished from other species by the contrasting coloration of their carapace and abdominal pattern (Figure 16 and Figure 17A). Furthermore, females are distinguished from Neohania catleyi n. sp. by their bean-shaped spermathecae and rounded secondary spermathecae (Figure 18D), while the latter have oval spermathecae and are elongated with small knob-like extrusion secondary spermathecae (Figure 8D). Males most resemble Neohania chibcha Heimer & Müller, 1988 but are distinguished by their wide, angular retrolateral tibial apophysis (Figure 18B), while in the latter, the retrolateral tibial apophysis is thin and curved ([8]: fig. 3).

Description. Female (holotype). Total length: 1.9; carapace length: 0.7; carapace width: 0.6; abdomen length: 1.2.

Cephalothorax. Carapace pear-shaped, light yellow with dark gray pattern (Figure 17A); pars cephalica flat, pars thoracica with strong incline. Sternum light yellow with dark wide lateral band; labium and endites light yellow (Figure 17B). Clypeus: 0.06. Chelicerae light yellow; promargin with three teeth, retromargin with two teeth. Six eyes of equal size, surrounded by black pigmentation; AME absent; ALE, PLE, PME: 0.08.

Abdomen. Oval, whitish with dark gray pattern of four–five chevrons, covered with long black setae (Figure 17A); ventral side with dark gray triangular mark medially below the epigastric groove reaching the tracheal spiracle (Figure 17B); tracheal spiracle 5× further from the epigastric groove than the spinnerets (Figure 17B, arrow); spinnerets aligned; PMS whitish, short, separated by half their width; ALS basal segment dark gray, apical segment yellowish; PLS basal segment medially dark gray, apical segment medially suffused with dark gray, basal segment slightly shorter than ALS (Figure 17B).

Legs. Light yellow with dark ventral mark on trochanters; two dark ventral marks on femurs and tibia; one ventral mark on patellae; one dorsal apical mark on metatarsi; macrosetae present on all segments except femur, metatarsi, and tarsi; trichobothria present on all segments except femur and patellae. Leg measurements: I 2.29 (0.64/0.20/0.58/0.49/0.38); II 1.98 (0.57/0.20/0.43/0.43/0.34); III 1.80 (0.57/0.17/0.41/0.37/0.28); IV 2.41 (0.67/0.17/0.58/0.58/0.41); leg formula: 4123; palpal tarsal claw present.

Genitalia: Epigynal plate flat, slightly sclerotized with dark pigmentation pattern; single copulatory opening, slit-like, atrium shallow (Figure 18C). Internal genitalia with long and coiled copulatory ducts; secondary spermathecae rounded; spermathecae bean-shaped; fertilization ducts wide, positioned externally (Figure 18D).

Male (paratype). Total length: 1.6; carapace length: 0.5; carapace width: 0.6; abdomen length 1.1.

Cephalothorax. As in female (Figure 17A); pars cephalica rounded. Sternum as long as wide, medially light yellow with wide, light gray lateral bands (Figure 17B); labium and endites as in female. Clypeus: 0.1. Chelicerae light yellow; promargin with three teeth, retromargin two teeth. Six eyes of equal size, surrounded by black pigmentation, AME absent; ALE, PLE, PME: 0.08.

Abdomen. As in female; spinnerets as in female, except for PLS apical segment uniformly whitish; tracheal spiracle position as in female (Figure 17B, arrow).

Legs. As in female. Leg measurements: I 2.21 (0.56/0.20/0.56/0.48/0.41); II 1.98 (0.54/0.21/0.47/0.28/0.38); III 1.61 (0.40/0.17/0.37/0.38/0.29); IV 2.06 (0.55/0.19/0.48/0.46/0.38); leg formula: 4123.

Genitalia. Palpal patella with basal, pointed spur; tibia with wide, angular, and rugose retrolateral tibial apophysis (Figure 18B); cymbium oval; cymbial furrow present (Figure 18B); embolus originating apically, long and thin, 2× circling the tegulum; tegulum not protruding (Figure 18A,B).

Distribution. Ecuador, Cotopaxi, and Santo Domingo de los Tsáchilas provinces.

Natural History. Specimens were collected in a low evergreen mountain forest (BsBn04) of the Western Cordillera between 1449 and 1997 m and in an evergreen mountain forest (cloud forest) at 2225 m of the Western Cordillera (BsMn03) [18]. At the locality Las Damas, the species is found in sympatry with Neohahnia piemontana n. sp., in the OTONGA reserve with Neohahnia freibergi n. sp., Pristirana nowickii n. sp., and Pristirana barthlotti n. sp. and in Pristirana reserve it occurs with Amaloxenops minimalista n. sp., Kasha patpa n. sp., and Pristirana niederi n. sp.

Figure 16. Neohania pristirana new species, female dorsal view.

Figure 16. Neohania pristirana new species, female dorsal view.

Figure 17. Neohania pristirana new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 17. Neohania pristirana new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 18. Neohania pristirana new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

Figure 18. Neohania pristirana new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

3.1.8. Neohahnia freibergi New Species

Figure 19A,B and Figure 20A–D, Scheme 2.

Type Material

Holotype: ♀ from Ecuador, Cotopaxi Province, San Francisco de las Pampas, OTONGA Reserve (-00.41994°-79.00623°) 1997 m, 24–30 May 2014, hand collecting, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10876 (QCAZ). Paratype: 1♂, same data as holotype ECFN 10878 (QCAZ); 1♀, 4–7 Sept. 2014, beating trees, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10877 (QCAZ).

Etymology. The specific name is a patronym in honor of the German botanist Dr. Martin Freiberg for his contribution to the knowledge of the OTONGA reserve plant biodiversity.

Diagnosis. Females and males are distinguished from other species by their coloration, their dark gray abdomen without pattern and smaller eyes 0.02–0.03 (Figure 19A). Females are further distinguished from eight eye species, Neohahnia piemontana n. sp., by their elongated and joined copulatory openings, with small triangular hood (Figure 20C,D), while atria are separated and hoods are absent in the latter (Figure 11C). Males are separated from Neohahnia piemontana n. sp. by their palpal patellar apophysis subterminal and angular retrolateral tibial apophysis (Figure 20A,B), whereas the patellar apophyses are terminal and the retrolateral tibial apophyses are curved in the latter species (Figure 11A,B).

Description. Female (holotype). Total length: 1.53; carapace length: 0.57; carapace width: 0.44; abdomen length: 096.

Cephalothorax. Carapace pear-shaped, dark brown with dark gray pattern (Figure 19A); pars cephalica slightly elevated, pars thoracica with strong incline. Sternum dark brown, as long as wide, suffused with dark gray pattern; labium and endites light brown (Figure 19B), serrula present. Clypeus: 0.06. Chelicerae light yellow-brown; promarginal teeth not observed. Eight eyes surrounded by black pigmentation; AME 0.02, ALE 0.03, PLE 0.03, PME 0.03.

Abdomen. Oval, dark gray without pattern (Figure 19A); ventral side dark gray; tracheal spiracle 4× further the epigastric groove than the spinnerets (Figure 19B, arrows); spinnerets aligned; light brown; PMS short, touching; ALS apical segment short; PLS apical segment slightly shorter than basal segment (Figure 19B).

Legs. Femur and patellae light yellow, tibia to tarsi light orange–brown; macrosetae present on all segments, except on femur and tarsi; trichobothria present on all segments except femur. Leg measurements: I 1.27 (0.38/0.12/0.29/0.24/0.24); II 1.12 (0.36/0.12/0.23/0.21/0.20); III 1.04 (0.30/0.11/0.23/0.21/0.19); IV 1.29 (0.41/0.12/0.30/0.26/0.20); leg formula: 4123; palpal tarsal claw present.

Genitalia. Epigynal plate flat, with dark pigmentation pattern; long slit-like copulatory opening, with hood, shallow atria, positioned anteriorly (Figure 20C). Internal genitalia with long and coiled copulatory ducts, secondary spermathecae rounded; spermathecae oval; fertilization duct narrow, positioned externally (Figure 20D).

Male (paratype). Total length: 1.26; carapace length: 0.59; carapace width: 0.48; abdomen length: 0.67.

Cephalothorax. As in female (Figure 19A). Sternum, labium, and endites as in female (Figure 19B). Clypeus: 0.09. Chelicerae as in female. Eight eyes surrounded by black pigmentation; AME 0.01, ALE 0.03, PLE 0.03, PME 0.04.

Abdomen. As in female (Figure 19A); spinnerets as in female, except for PLS apical segment uniformly whitish; tracheal spiracle position as in female (Figure 19B).

Legs. As in female (Figure 19A,B). Leg measurements: I 1.49 (0.43/0.14/0.37/0.26/0.29); II 1.26 (0.36/0.12/0.27/0.26/0.25); III 1.02 (0.32/0.12/0.24/0.25/0.21); IV 1.58 (0.46/0.12/0.37/0.33/0.30); leg formula: 4123.

Genitalia. Palpal patella with sub-basal pointed spur; tibia with narrow, angular, and rugose retrolateral tibial apophysis (Figure 20B); cymbium rounded apically; with wide cymbial furrow (Figure 20A,B); embolus originating prolatero-apically, long and thin, 2× circling the tegulum; tegulum not protruding (Figure 20A,B).

Distribution. Only known from the type locality in Cotopaxi province.

Natural History. All specimens were collected beating trees in the low evergreen mountain forest of the of the Western Cordillera, at 1997 m (BsBn04) [18]. The species occurs in sympatry with N. pristirana n. sp., Pristirana nowickii n. sp., and Pristirana barthlotti n. sp.

Figure 19. Neohania freibergi new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 19. Neohania freibergi new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 20. Neohania freibergi new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

Figure 20. Neohania freibergi new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

Scheme 2. Distribution map of Hahniidae species occurring on the Western Cordillera. (Amaloxenops minimalista n. sp. (purple circle), Kasha papta n. sp. (turquoise star), Neohahnia catleyi n. sp., (orange circle), N. piemontana n. sp. (yellow lozenge), N. pristirana n. sp. (green inverse triangle), N. freibergi n. sp. (black circle), P. niederi n. sp. (red cross), P. nowickii n. sp. (red inverse triangle), and Pristirana barthlotti n. sp. (yellow cross)).

Scheme 2. Distribution map of Hahniidae species occurring on the Western Cordillera. (Amaloxenops minimalista n. sp. (purple circle), Kasha papta n. sp. (turquoise star), Neohahnia catleyi n. sp., (orange circle), N. piemontana n. sp. (yellow lozenge), N. pristirana n. sp. (green inverse triangle), N. freibergi n. sp. (black circle), P. niederi n. sp. (red cross), P. nowickii n. sp. (red inverse triangle), and Pristirana barthlotti n. sp. (yellow cross)).

3.1.9. Neohahnia paramo New Species

Figure 21A,B and Figure 22A–D, Scheme 3.

Type Material

Holotype: ♀ from Ecuador, Napo Province, Páramo de Papallacta (-00.281913°-78.137172°) 4016 m, 21 March 2021, hand collected, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 7194 (QCAZ). Paratypes: same data as holotype; 1♂1♀, ECFN 11256 (QCAZ); 1♀, ECFN 5913 (USNM); 2♀ ECFN 5902, 5903 (ZMH-A0013171, A0013172); 1♀ ECFN 5914 (AMNH).

Etymology. The specific epithet is a noun in apposition in regard to the ecoregion where the species is found.

Diagnosis. Females and males most resemble N. pristirana n. sp. and N. chibcha Heimer & Müller, 1988 [8] but are distinguished from N. pristirana n. sp. in having eight eyes (Figure 21A); while the latter have only six (Figure 17A); from N. chibcha by the embolus starting apically (Figure 22A), while retrolaterally in the latter ([8]; fig. 4).

Description. Female (holotype): Total length: 2.30; carapace length: 0.93; carapace width: 0.73; abdomen length: 1.37.

Cephalothorax. Carapace pear-shaped, light brown suffused with black (Figure 21A); pars cephalica flat, pars thoracica inclined. Sternum as long as wide, brown strongly suffused with black (Figure 21B); labium and endites light brown, serrula present. Clypeus: 0.1. Chelicerae yellow; promargin with three teeth, retromargin with two teeth. Eight eyes surrounded by black pigmentation; AME 0.02, ALE 0.05, PLE 0.05, PME 0.06.

Abdomen. Oval, dark gray with four–five chevrons (Figure 21A); ventral side dark gray with two whitish longitudinal bands; tracheal spiracle 5× further from the epigastric groove than the spinnerets (Figure 21B, arrow); spinnerets aligned; PMS whitish, touching; ALS basal segment dark gray, apical segment whitish; PLS basal segment suffused gray apically, apical segment whitish (Figure 21B).

Legs. Light yellow with two dark ventral marks on femurs and tibia; one ventral mark on patellae; one dorsal apical mark on metatarsi; macrosetae present on all segments except femur, metatarsi and tarsi; trichobothria present on all segments except femur and patellae. Leg measurements: I 2.70 (0.76/0.26/0.64/0.59/0.45); II 2.40 (0.68/0.26/0.52/0.53/0.41); III 2.28 (0.63/0.24/0.51/0.50/0.40); IV 2.98 (0.86/0.29/0.73/0.63/0.47); leg formula: 4123; palpal tarsal claw present.

Genitalia. Epigynal plate flat, well sclerotized; single copulatory opening, atrium deep, positioned antero-medially (Figure 22C). Internal genitalia with long and coiled copulatory ducts; secondary spermathecae oval with dorsal knob; spermathecae rounded; fertilization duct short and wide, positioned baso-externally (Figure 22D).

Male (paratype). Total length: 2.01; carapace length: 0.94; carapace width: 0.72; abdomen length: 1.07.

Cephalothorax. Carapace light brown suffused with black, and shiny (Figure 21A); pars cephalica rounded, pars thoracica inclined. Sternum as long as wide, brown suffused with black, medially light yellow (Figure 21B); labium and endites as in female. Clypeus: 0.15. Chelicerae as in female. Eight eyes surrounded by black pigmentation; AME 0.02, ALE 0.05, PLE 0.05, PME 0.06.

Abdomen. Oval, dark gray with four–five chevrons (Figure 21A); ventral side as in female; spinnerets as in female; tracheal spiracle position as in female (Figure 21B, arrow).

Legs. As in female (Figure 21A,B). Leg measurements: I 2.37 (0.73/0.28/0.62/0.59/0.45); II 2.40 (0.69/0.28/0.51/0.50/0.42); III 2.23 (0.65/0.26/0.50/0.43/0.39); IV 2.82 (0.79/0.26/0.66/0.62/0.49); leg formula: 4123.

Genitalia. Palpal patella with basal pointed spur (Figure 22B); tibia with thin, angular, and rugose retrolateral tibial apophysis (Figure 22B); cymbium oval apically; with cymbial furrow (Figure 22A); embolus originating prolatero-apically, long and thin, circling 2× around tegulum; tegulum not protruding (Figure 22A,B).

Distribution. Only known from the type locality in Napo Province.

Natural History. Specimens were collected using pitfall traps in the Paramó grassland (HsSn02) at 4016 m [21]. The species lives in sympatry with Paramito papallacta n. sp.

Figure 21. Neohania paramo new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 21. Neohania paramo new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 22. Neohania paramo new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

Figure 22. Neohania paramo new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

3.1.10. Neohahnia chalupas New Species

Figure 23, Scheme 3.

Type Material

Holotype: ♂ from Ecuador, Napo Province, Tena, Colonso Chalupas Natural Reserve (-00.914889°-77.877944°) 782 m, 2–9 March 2020, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 4691 (QCAZ).

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

Diagnosis. Males most resemble N. piemontana n. sp. and N. freibergi n. sp., but can be distinguished from both species by their tegulum strongly protruding (Figure 23D), and not protruding in the latter (Figure 11A,B and Figure 20A,B).

Description. Male (holotype). Total length: 1.42; carapace length: 0.62; carapace width: 0.64; abdomen length: 0.80.

Cephalothorax. Carapace light brown suffused with black (Figure 23A); pars cephalica flat, pars thoracica inclined. Sternum as long as wide, light brown; labium and endites light brown (Figure 23B), serrula present. Clypeus: 0.09. Chelicerae yellow; teeth not observed. Eight eyes surrounded by black pigmentation; AME 0.08, ALE 0.07, PLE 0.05, PME 0.06 (Figure 23A).

Abdomen. Oval, dark gray with chevron patterns (Figure 23A); ventral side light gray; tracheal spiracle position not observed; spinneret positions not observed.

Legs. Uniformly light yellow–brown; macrosetae only observed on patellae and tibiae; present on all segments except femur. Leg measurements: I 1.98 (0.56/0.19/0.43/0.37/0.43); II 1.80 (0.55/0.18/0.39/0.36/0.32); III 1.70 (0.52/0.17/0.33/0.36/0.32); IV 2.20 (0.59/0.17/0.52/0.50/0.42); leg formula: 4123.

Genitalia. Palpal patella with elongate bifid sub-basal apophysis (Figure 23D); tibia with elongate curved, retrolateral tibial apophysis, with some asperity (Figure 23D); cymbium rounded apically; with large cymbial furrow (Figure 23C,D); embolus long, originating retrolaterally, long and thin, circling 1× the tegulum (Figure 23C,D); tegulum protruding ventrally (Figure 23D).

Distribution. Only found at the type locality in Napo Province.

Natural History. The only male was collected with a pitfall trap at 782 m in Amazonian region, in the Napo-Curaray lowland evergreen (BsTa02) [22].

Figure 23. Neohania chalupas new species, male holotype. (A) Habitus, dorsal view. (B) Habitus, ventral view. (C) Male palp, ventral view. (D) Male palp, retrolateral view. Scale bars: 0.5 mm.

Figure 23. Neohania chalupas new species, male holotype. (A) Habitus, dorsal view. (B) Habitus, ventral view. (C) Male palp, ventral view. (D) Male palp, retrolateral view. Scale bars: 0.5 mm.

3.1.11. Paramito New Genus

Type species. Paramito papallacta n. sp.

Etymology. The generic epithet is derived from the Spanish language; the noun “Páramo” refers to the alpine tundra ecosystem where the type species was collected. The gender is masculine.

Diagnosis. Males and female are distinguished from all other genera by their carapace with well-marked longitudinal fovea (Figure 24A and Figure 28A), absent in all other genera (e.g., Figure 1A, Figure 9 and Figure 30A); male chelicerae with patch of elongated setae and excavation internally (Figure 26H, arrow) absent in all other genera; spinnerets not aligned (Figure 24A) except in Lizarba Roth, 1967 [6] (but separated by the absence of white spatulate setae, present in the latter [6]; p. 306), spinnerets aligned in all other genera (e.g., Figure 1A and Figure 4B).

Description. Small size spider, females (1.90–2.77), males (2.44).

Cephalothorax. Carapace elongated pear-shaped, longer than wide, pars cephalica flat, pars thoracica inclined, fovea marked by a longitudinal line (Figure 24A, Figure 26B, and Figure 28A). Sternum longer than wide, not truncated posteriorly (Figure 26D and Figure 28A). Labium wider than long, endites straight with serrula (Figure 26F). Chelicerae promargin and retromargin with two–three teeth; male with patch of elongated setae and excavation internally (Figure 26H). Eight eyes surrounded by black pigmentation; AME minute 1/4 the size of PME; ALE, PLE, and PME of almost equal size.

Abdomen. Oval, with chevron patterns (Figure 24A and Figure 28A), covered with long setae (Figure 26A). Spinnerets not aligned; PME separated by half to their diameter, PLS behind ALS, PLS apical segment short (Figure 24B and Figure 28A).

Legs. Legs uniformly colored; macrosetae presents on all segments, except on tarsi; trichobothria present on all segments except femur; tarsal organ rounded situated in concavity (Figure 27F,H); three claws (Figure 27G); female palpal tarsal claw present; leg formula: 4123.

Genitalia. Palpal patellar apophysis bifid; tibial apophysis elongated, smooth (Figure 25B and Figure 27B); cymbium pointed, with small retrolateral furrow, with ventral lamella (Figure 25A,B and Figure 27A); embolus elongated starting retrolaterally, with basal membrane (Figure 27A, arrow), median apophysis present (Figure 25A and Figure 27A).

Epigynum with copulatory openings, vertical slit-like (Figure 25C and Figure 28B).

Internal genitalia with short copulatory ducts; secondary spermathecae absent; spermathecae rounded to oval; fertilization ducts short (Figure 25D and Figure 28C).

Composition. Paramito papallacta n. sp. and P. oyacachi n. sp.

Distribution. Ecuador.

3.1.12. Paramito papallacta New Species

Figure 24, Figure 25, Figure 26 and Figure 27, Scheme 3.

Type Material

Holotype: ♂ from Ecuador, Napo Province, Páramo de Papallacta (-00.281614°-78.136955°) 4018 m, 8–21 March 2020, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 6561 (QCAZ). Paratypes: Páramo de Papallacta (-00.30022° -78.133618°) 3983 m, 1–8 March 2020, 1♀, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 6531 (QCAZ); Páramo de Papallacta (-00.281913°-78.137172°) 4016 m, 21 March 2020, 2♂, hand collected, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 5916, 5922 (USNM); Páramo de Papallacta (-00.300167°-78.133806°) 3984 m, 8–21 March 2020, 1♂, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 5031 (AMNH); Páramo de Papallacta (-00.281614°-78.136955°) 4018 m, 8–21 March 2020, 2♂, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 6022 (ZMH-A0013747); (-00.295778°-78.12208°) 3830 m, 1♂, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 4726 (ZMH-A0013745); Páramo de Papallacta (-00.30022°-78.133618°) 3988 m, 8–21 March 2020, 2♂, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 6012, 6013 (ZMH-A0013744, A0013746); Páramo de Papallacta (-00.296278°-78.121250°) 3784 m, 1 March 2020, 1♂, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 5732 (ZMH-A0013170); (-00.300194°-78.133694°) 3986 m, 1–8 March 2020, 1♀, pitfall traps, N. Dupérré, E. E. Tapia, A. A. Tapia, ECFN 6015 (ZMH-A00013184).

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

Diagnosis. Males are distinguished by their pointed cymbium with ventral basal lamella (Figure 25A and Figure 27A); palpal patella elongated and bifid; median apophysis present (Figure 25B). Females most resemble P. oyacachi n. sp. but are distinguished by their longer copulatory ducts, and small rounded spermathecae (Figure 25D), while the latter have short copulatory ducts, and large oval spermathecae (Figure 28C).

Description. Male (holotype). Total length: 2.44; carapace length: 1.22; carapace width: 0.76; abdomen length: 1.22.

Cephalothorax. Carapace elongated, pear-shaped, light yellow-orange (Figure 24A); pars cephalica flat, pars thoracica inclined; fovea mark by longitudinal line (Figure 24A). Clypeus: 0.09. Sternum wider than long (Figure 26D), light yellow–orange, lightly suffused with black, darker marginally, not truncated (Figure 24B and Figure 26D); labium and endites light yellow–orange, serrula with ~40 teeth (Figure 26E). Chelicerae orange; promargin with three teeth, retromargin with two teeth; retromargin with patch of elongated setae and excavation (Figure 26H, arrow). Eight eyes surrounded by black pigmentation (Figure 25A); AME minute 0.015 (Figure 26G); ALE 0.06, PLE 0.07, PME 0.07.

Abdomen. Oval, beige with suffused with grayish–brown markings of four–five chevrons, covered with elongated grayish setae (Figure 24A); ventrally beige medially, gray laterally (Figure 24B); tracheal spiracle slightly recurved, situated 10× further from the epigastric groove than the spinnerets (Figure 24B and Figure 26E, arrow); spinnerets not aligned; PMS light brown, separated by half their diameter, with minor ampullate and one cylindrical gland spigot (Figure 27C); ALS basal and apical segments light brown, apical segment short, with major ampullate and one cylindrical? gland spigot (Figure 27D); PLS basal and apical segments light brown, apical segment short (Figure 24B), with one cylindrical and two aciniform gland spigots (Figure 27E).

Legs. Uniformly light brown (Figure 24A,B); macrosetae present on all segments, except on tarsi; trichobothria present on all segments except femur and patellae; three claws, paired claw with ten teeth, unpaired claw with teeth (Figure 27G); tarsal organ rounded in deep concavity (Figure 27F,H, arrows). Leg measurements: I 2.84 (0.82/0.28/0.65/0.60/0.49); II 2.51 (0.73/0.28/0.52/0.54/0.44); III 2.37 (0.66/0.27/0.49/0.52/0.43); IV 2.86 (0.78/0.25/0.65/0.67/0.51); leg formula: 4123.

Genitalia. Palpal patella with bifid, sub-basal patellar apophysis; tibial apophysis elongated (Figure 25B and Figure 27B); cymbium pointed, with small retrolateral furrow, and ventral lamella (Figure 25A,B and Figure 27A); embolus elongated starting retrolaterally, with basal membrane (Figure 27A, arrow), median apophysis present (Figure 25A and Figure 27A).

Female (paratype). Total length: 1.90; carapace length: 0.93; carapace width: 0.58; abdomen length: 0.97.

Cephalothorax. As in male (Figure 24A). Clypeus: 0.12. Sternum longer then wide, light yellow–orange (Figure 24B); labium and endites as in male. Chelicerae as in male. Eight eyes surrounded by black pigmentation; AME 0.15, ALE 0.06, PLE 0.06, PME 0.05.

Abdomen. Oval, light beige dorsally suffused with pale chevron patterns, covered with elongated grayish elongate setae (Figure 24A); ventrally beige (Figure 24B); tracheal spiracle not observed, abdomen shrunk; spinnerets not aligned; PMS light brown, separated by half their diameter; ALS basal segment and apical segments light brown, apical segment short; PLS, basal long, light brown, apical whitish, short (Figure 24B).

Legs. As in male. Leg measurements: I 2.25 (0.67/0.26/0.49/0.47/0.36); II 2.06 (0.61/0.24/0.42/0.43/0.36); III 1.75 (0.48/0.25/0.34/0.40/0.28); IV 2.36 (0.64/0.28/0.53/0.56/0.35); leg formula: 4123; palpal tarsal claw.

Genitalia. Epigynal plate flat, strongly sclerotized; copulatory openings vertical, slit-like (Figure 25C). Internal genitalia with wide copulatory ducts; secondary spermathecae absent; spermathecae rounded; fertilization ducts short, internally directed (Figure 25D).

Distribution. Only found at the type locality in Napo province.

Natural History. Specimens were collected by pitfall trap in the Paramó grassland (HsSn02) between 3983 and 4018 m [21]. The species lives in sympatry with Neohahnia paramo n. sp.

Figure 24. Paramito papallacta new species. (A) Male holotype and female paratype, habitus dorsal view. (B) Male holotype and female paratype, habitus ventral view (arrow points to tracheal spiracle). Scale bars: 0.5 mm.

Figure 24. Paramito papallacta new species. (A) Male holotype and female paratype, habitus dorsal view. (B) Male holotype and female paratype, habitus ventral view (arrow points to tracheal spiracle). Scale bars: 0.5 mm.

Figure 25. Paramito papallacta new species. (A) Male holotype, palp ventral view. (B) Male holotype, palp retrolateral view. (C) Female paratype, epigynum ventral view (ECFN 6531). (D) Female paratype, internal genitalia dorsal view (ECFN 6015). Scale bars: 0.5 mm.

Figure 25. Paramito papallacta new species. (A) Male holotype, palp ventral view. (B) Male holotype, palp retrolateral view. (C) Female paratype, epigynum ventral view (ECFN 6531). (D) Female paratype, internal genitalia dorsal view (ECFN 6015). Scale bars: 0.5 mm.

Figure 26. Paramito papallacta new species, male (ECFN 4726) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Habitus, ventral view. (D) Sternum, ventral view. (E) Abdomen, ventral view (arrow points to tracheal spiracle). (F) Labium and endites, ventral view. (G) Carapace, frontal view. (H) Chelicerae, posterior view (arrow points to excavation).

Figure 26. Paramito papallacta new species, male (ECFN 4726) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Habitus, ventral view. (D) Sternum, ventral view. (E) Abdomen, ventral view (arrow points to tracheal spiracle). (F) Labium and endites, ventral view. (G) Carapace, frontal view. (H) Chelicerae, posterior view (arrow points to excavation).

Figure 27. Paramito papallacta new species, male (ECFN 11247) SEM. (A) Palp, ventral view (arrow point to membrane). (B) Palp, retrolateral view. (C) PMS, apical view. (D) ALS, apical view. (E) PLS, apical view. (F) Tarsi I, dorsal view (arrow points to tarsal organ). (G) Tarsal claw I, lateral view (arrow points to third claw). (H) Tarsi IV, dorsal view (arrow points to tarsal organ).

Figure 27. Paramito papallacta new species, male (ECFN 11247) SEM. (A) Palp, ventral view (arrow point to membrane). (B) Palp, retrolateral view. (C) PMS, apical view. (D) ALS, apical view. (E) PLS, apical view. (F) Tarsi I, dorsal view (arrow points to tarsal organ). (G) Tarsal claw I, lateral view (arrow points to third claw). (H) Tarsi IV, dorsal view (arrow points to tarsal organ).

3.1.13. Paramito oyacachi New Species

Figure 28, Scheme 3.

Type Material

Holotype: ♀ from Ecuador, Pichincha Province, Cayambe, Via Cangahua, Oyacachi (-00.181166°-78.067601°) 3860 m, 25 August 2013, sifting mosses on ground, Berlese traps, E. Tapia, N. Dupérré, ECFN 10916 (QCAZ). Paratypes: 3♀, same data as holotype, ECFN 11258 (QCAZ).

Etymology. The specific epithet in a noun in apposition, taken from the nearby town, Oyacachi where the specimens were collected.

Diagnosis. Females most resemble P. papallacta n. sp. but are distinguished by their internal genitalia with shorter copulatory ducts and large, oval spermathecae (Figure 28C), while the latter have longer copulatory ducts and small, rounded spermathecae (Figure 25D).

Description. Female (holotype). Total length: 2.77; carapace length: 1.13; carapace width: 0.75; abdomen length: 1.64.

Cephalothorax. Carapace elongated pear-shaped, light brown–orange, lightly suffused with black medially, along radiating lines and marginally (Figure 28A); pars cephalica flat, pars thoracica inclined, fovea marked by longitudinal line (Figure 28A). Clypeus: 0.10. Sternum longer then wide, light brown–orange, lightly suffused with black medially, strongly suffused with black laterally, not truncated (Figure 28A); labium brown–orange suffused with black, endites light brown suffused with black laterally (Figure 28A); serrula present. Chelicerae light brown; promargin with two teeth, retromargin with three teeth. Eight eyes surrounded by black pigmentation; AME minute, 0.02, ALE 0.08, PLE 0.06, PME 0.06.

Abdomen. Oval, light grayish dorsally suffused with some dark gray, with four chevrons, covered with elongated grayish elongated setae (Figure 24A); ventrally light gray medially and dark gray laterally (Figure 24A); tracheal spiracle slightly procurved, situated 6× further from the epigastric groove than the spinnerets (Figure 24A, arrow); spinnerets not aligned; PMS long, light brown, separated by diameter; ALS basal segment and apical segments light brown; PLS, basal long, light brown, and apical segments basally whitish, apically brownish, short (Figure 24A).

Legs. Uniformly light brown (Figure 24A); macrosetae present on all segments, except on tarsi; trichobothria present on all segments except femur. Leg measurements: I 2.84 (0.81/0.31/0.66/0.61/0.45); II 2.48 (0.72/0.26/0.54/0.53/0.43); III 2.17 (0.54/0.30/0.47/0.47/0.39); IV 2.86 (0.76/0.30/0.72/0.65/0.43); leg formula: 4123; tarsal palpal claw present.

Genitalia. Epigynal plate flat, strongly sclerotized; copulatory openings vertical, slit-like (Figure 28B). Internal genitalia with short, thin copulatory ducts; spermathecae oval; fertilization ducts short (Figure 28C).

Male. Unknown.

Distribution. Only found at the type locality in Pichincha Province.

Natural History. All specimens were collected by sifting mosses in Paramó grassland (HsSn02) at 3860 m [21].

Figure 28. Paramito oyacachi new species, female holotype (ECFN 10916) (A) Habitus, dorsal and ventral view (arrow points to tracheal spiracle). (B) Epigynum, ventral view. (C) Internal genitalia, dorsal view. Scale bars: 0.5 mm.

Figure 28. Paramito oyacachi new species, female holotype (ECFN 10916) (A) Habitus, dorsal and ventral view (arrow points to tracheal spiracle). (B) Epigynum, ventral view. (C) Internal genitalia, dorsal view. Scale bars: 0.5 mm.

Scheme 3. Distribution map of species occurring on the Eastern Cordillera and Amazonian regions (Paramito papallacta n. sp. (inversed yellow triangle), Paramito oyacachi n. sp. (turquoise circle), Neohahnia paramo n. sp. (green star), and N. chalupas n. sp. (red circle)).

Scheme 3. Distribution map of species occurring on the Eastern Cordillera and Amazonian regions (Paramito papallacta n. sp. (inversed yellow triangle), Paramito oyacachi n. sp. (turquoise circle), Neohahnia paramo n. sp. (green star), and N. chalupas n. sp. (red circle)).

3.1.14. Pristirana New Genus

Type species: Pristirana niederi n. sp.

Diagnosis. Members of the genus are distinguished from all other Hahniidae by the male palpal cymbium blunt apically (Figure 31A and Figure 35A) while pointed (e.g., Figure 5A and Figure 25A) to rounded (e.g., Figure 11A and Figure 20A) in other genera; female internal genitalia with spermathecae on an elongated base (Figure 29C, Figure 31D, and Figure 35D); elongated bases not found in all other genera (e.g., Figure 1C, Figure 5D, and Figure 11D).

Etymology. The generic name is feminine and taken from Pristirana Reserve where the type species is found. The gender is feminine.

Description. Small size spider; females (1.14–1.74), males (1.15–1.74).

Cephalothorax. Carapace pear-shaped, longer than wide, pars cephalica flat, pars thoracica inclined, fovea absent (Figure 29A and Figure 34A). Sternum wider than long, or as wide as long, truncated posteriorly (Figure 30B and Figure 32C). Labium wider than long, endites slightly converging with serrula (Figure 32D). Chelicerae promargin and retromargin with two–three teeth. Eight eyes surrounded by black pigmentation, AME minute 1/3 the size of PME; ALE, PLE, and PME of almost equal size.

Abdomen. Oval, with (Figure 29A and Figure 34A) or without chevron patterns (Figure 30A), covered with long setae; tracheal spiracle situated 7–13× further from the epigastric groove than spinnerets. Spinnerets aligned; ALS longer than first segment of PLS (Figure 29A and Figure 34B).

Legs. Macrosetae and trichobothria present on tibia, metatarsi, and tarsi; macrosetae present on femur, tibia, and sometimes metatarsi; tarsal organ rounded situated in concavity (Figure 33B); three claws (Figure 33C); leg formula: 4123; female palpal claw present.

Genitalia. Epigynum with small, externally positioned copulatory openings (Figure 29B and Figure 31C); male palpal patella apophysis sub-apical; palpal tibia recurved, smooth; palp cymbium truncated, without retrolateral fold; embolus filiform; median apophysis absent (Figure 31A and Figure 35A).

Composition. Pristirana niederi n. sp., P. nowickii n. sp., and P. barthlotti n. sp.

Distribution. Found only in the Chocó region of Ecuador.

3.1.15. Pristirana niederi New Species

Figure 29, Scheme 2.

Type Material

Holotype: ♀ from Ecuador, Cotopaxi Province, San Francisco de las Pampas, Pristirana Reserve (-00.42414°-78.95719°) 1485 m, 08.vi.2017, N. Dupérré, E. E. Tapia, C. Tapia, ECFN 10881 (QCAZ). Paratypes: 1♀ same data as holotype; 1♀, San Francisco de las Pampas, Pristirana Reserve (-00.42414°-78.95719°) 1480 m, 15 Feb. 2019, E. Tapia, Faml. Tapia, ECFN 1512 (QCAZ).

Etymology. The specific epithet is a patronym in honor of the German botanist Dr. Jürgen Nieder, for his contribution to the knowledge of the OTONGA reserve’s epiphyte biodiversity.

Diagnosis. Females most resembles P. nowickii n. sp., and P. barthlotti n. sp. but are differentiated by their light color pattern with five chevrons (Figure 29A) absent in P. nowickii n. sp. (Figure 30A), and dark gray pattern with chevrons in P. barthlotti n. sp. (Figure 34A); furthermore, females are differentiated by their spermathecae with copulatory ducts attached ventrally (Figure 29B), apically in P. nowickii n. sp. (Figure 31D), and from P. barthlotti n. sp. by their spermathecae elongated oval (Figure 29C), rounded in the latter (Figure 35D).

Description. Female (holotype): Total length: 1.65; carapace length: 0.72; carapace width: 0.54; abdomen length: 0.93.

Cephalothorax. Carapace pear-shaped, light yellow-brown suffused with dark brown pattern (Figure 29A); pars cephalica flat, pars thoracica incline. Sternum light yellow suffused with dark brown, wider than long, truncated apically (Figure 29A); labium and endites light brown (Figure 29A); serrula present. Clypeus: 0.06. Chelicerae light yellow–brown; promargin, retromargin with two teeth. Eight eyes surrounded by black pigmentation; AME 0.02, ALE, PLE, and PME 0.07.

Abdomen. Oval, beige with dark gray pattern of four–five chevrons (Figure 29A); ventrally beige medially, laterally dark gray; tracheal spiracle slightly recurved, situated 7× further from the epigastric groove than the spinnerets (Figure 29B, arrow); spinnerets aligned; PMS white with brown band, almost touching; ALS basal segment white basally, dark brown apically, as long as PLS basal segment; apical segment light brown; PLS basal segment white basally, dark brown apically, apical segment elongated light brown basally, whitish apically, both segment of almost equal size (Figure 29B).

Legs. Light yellow–brown with two dark ventral marks on femurs; one ventral mark on tibia I-II and patellae, and two ventral marks on tibia III–IV; metatarsi and tarsi uniformly light yellow–brown; palpal femur and patellae light yellow–brown, tibia with one dark mark; tarsi light yellow–brown; palpal tarsal claw present. Leg macrosetae present on all segments, except metatarsi I-II and tarsi; trichobothria present on all segments except femur and patellae. Leg measurements: I 1.84 (0.49/0.23/0.41/0.37/0.34); II 1.72 (0.48/0.21/0.39/0.33/0.31); III 1.52 (0.45/0.19/0.31/0.29/0.28); IV 2.05 (0.58/0.21/0.48/0.41/0.37); leg formula: 4123.

Genitalia. Epigynal plate flat lightly sclerotized; with small, curved copulatory openings, set laterally (Figure 29B). Internal genitalia with short copulatory ducts; spermathecae elongated oval, on an elongated base; fertilization short internally directed (Figure 29C).

Distribution. Only known from the type locality in Cotopaxi Province.

Natural History. Specimens were collected between 1480 and 1485 m in a foothill evergreen mountain forest (BsPn01) of the Western Cordillera [9]. The species lives in sympatry with Amaloxenops minialista n. sp., Kasha papta n. sp. and Neohahnia pristirana n. sp.

Figure 29. Pristirana niederi new species, female holotype. (A) Habitus, dorsal and ventral view (arrow point to tracheal spiracle). (B) Epigynum, ventral view. (C) Internal genitalia, dorsal view. Scale bar = 0.5 mm.

Figure 29. Pristirana niederi new species, female holotype. (A) Habitus, dorsal and ventral view (arrow point to tracheal spiracle). (B) Epigynum, ventral view. (C) Internal genitalia, dorsal view. Scale bar = 0.5 mm.

3.1.16. Pristirana nowickii New Species

Figure 30, Figure 31, Figure 32 and Figure 33, Scheme 2.

Type Material

Holotype: ♀ from Ecuador, Cotopaxi province, San Francisco de la Pampas, OTONGA reserve (-00.41994°-70.00623°), 1997 m, 13–15 Nov. 2014, Berlese, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10895 (QCAZ). Paratypes: 2♂1♀ same data as holotype, ECFN 11259 (ZMH-A0020677); 2♀, OTONGA reserve (-00.41994°-70.00623°), 1997 m, 24–30 May 2014, hand collected, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10896 (QCAZ).

Etymology. The specific name is a patronym in honor of German botanist Dr. Christoph Nowicki for his contribution to the knowledge of the OTONGA reserve’s vascular epiphytes.

Diagnosis. Males and females most resemble P. barthlotti n. sp. but can be distinguished by their gray abdomen without chevrons (Figure 30A), while P. barthlotti n. sp. has four–five chevrons (Figure 34A). Furthermore, females are distinguished by spermathecae touching medially (Figure 31C) while in the latter, the spermathecae are separated by almost their diameter (Figure 35C); males are distinguished by the bent tip of retrolateral tibial apophysis (Figure 31B), tips of retrolateral tibial apophyses are curved in the latter (Figure 35B).

Description. Female (holotype). Total length: 1.14; carapace length: 0.59; carapace width: 0.42; abdomen length: 0.55.

Cephalothorax. Carapace pear-shaped, dark brown suffused with black pattern (Figure 30A); pars cephalica flat, pars thoracica inclined. Sternum dark brown suffused with black; truncated (Figure 32C); labium and endites light brown (Figure 30B), serrula with 34 teeth (Figure 32D). Clypeus: 0.06. Chelicerae light yellow; teeth not observed. Eight eyes surrounded by black pigmentation; AME 0.02, ALE, PLE, and PME 0.07.

Abdomen. Oval, dark gray, covered with long whitish setae (Figure 30A); ventral side dark gray; tracheal spiracle 5x further from the epigastric groove than the spinnerets (Figure 30B and Figure 32E, arrow); spinnerets aligned (Figure 33A); spinnerets whitish; PMS touching (Figure 30B); ALS and PLS basal segments of equal size; PLS apical segment shorter than basal (Figure 33A).

Legs. Femur and patellae whitish, tibiae to tarsi light brown (Figure 30A,B); macrosetae present on all segments, except metatarsi I-II and tarsi; trichobothria present on all segments except femur and patellae; tarsal organ rounded in a concavity (Figure 33B,D). Leg measurements: I 1.35 (0.38/0.16/0.28/0.26/0.27); II 1.30 (0.36/0.16/0.27/0.28/0.23); III 1.17 (0.32/0.15/0.26/0.23/0.21); IV 1.46 (0.39/0.18/0.32/0.32/0.25); leg formula: 4123; palpal tarsal claw present; three claws (Figure 33C).

Genitalia. Epigynal plate flat lightly sclerotized; with small, curved copulatory openings, set laterally (Figure 31C). Internal genitalia with short copulatory ducts; spermathecae oval on an elongated base; fertilization short, internally directed (Figure 31D).

Male (paratype): Total length: 1.15; carapace length: 0.60; carapace width: 0.42; abdomen length: 0.55.

Cephalothorax. As in female, slightly paler (Figure 30A). Sternum as in female, slightly paler (Figure 30B). Labium and endites as in female.

Clypeus: 0.06. Chelicerae light yellow; teeth not observed. Eight eyes surrounded by black pigmentation; AME 0.02, ALE, PLE, and PME 0.07.

Abdomen. As in female (Figure 30A).

Legs. As in female (Figure 30A,B). Leg measurements: I 1.35 (0.36/0.17/0.31/0.28/0.23); II 1.27 (0.36/0.17/0.26/0.25/0.23); III 1.13 (0.30/0.12/0.26/0.23/0.22); IV 1.41 (0.38/0.13/0.32/0.32/0.26); leg formula: 4123.

Genitalia. Palpal patella with pointed, sub-basal patellar apophysis; tibial apophysis curved, smooth with bent tip (Figure 31B); cymbium truncated, without retrolateral fold (Figure 31B); embolus elongated starting apically, encircling half the tegulum; median apophysis absent (Figure 31A); tegulum not protruding (Figure 31A,B).

Distribution. Only known from the type locality in Cotopaxi Province

Natural History. Specimens were collected in a low evergreen mountain forest (BsBn04) of the Western Cordillera [18] at 1997 m where it lives in sympatry with P. barthlotti n. sp., Neohahnia pristirana n. sp., and Neohahnia freibergi n. sp.

Figure 30. Pristirana nowickii new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 30. Pristirana nowickii new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 31. Pristirana nowickii new species. (A) Male paratype, palp ventral view. (B) Male paratype palp, retrolateral view. (C) Female holotype, epigynum ventral view. (D) Internal genitalia, dorsal view.

Figure 31. Pristirana nowickii new species. (A) Male paratype, palp ventral view. (B) Male paratype palp, retrolateral view. (C) Female holotype, epigynum ventral view. (D) Internal genitalia, dorsal view.

Figure 32. Pristirana nowickii new species, female (ECFN 10896) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Sternum, ventral view. (D) Endites and labium, ventral view. (E) Abdomen ventral view (arrow points to tracheal spiracle). (F) Epigynal plate, ventral view.

Figure 32. Pristirana nowickii new species, female (ECFN 10896) SEM. (A) Habitus, dorsal view. (B) Carapace, dorsal view. (C) Sternum, ventral view. (D) Endites and labium, ventral view. (E) Abdomen ventral view (arrow points to tracheal spiracle). (F) Epigynal plate, ventral view.

Figure 33. Pristirana nowickii new species, female (ECFN 10896) SEM. (A) Spinnerets, ventral view. (B) Tarsus I, dorsal view (arrow points to tarsal organ). (C) Claw I, dorsal view. (D) Tarsus II, apical view (arrow points to tarsal organ).

Figure 33. Pristirana nowickii new species, female (ECFN 10896) SEM. (A) Spinnerets, ventral view. (B) Tarsus I, dorsal view (arrow points to tarsal organ). (C) Claw I, dorsal view. (D) Tarsus II, apical view (arrow points to tarsal organ).

3.1.17. Pristirana barthlotti New Species

Figure 34 and Figure 35, Scheme 2.

Type Material

Holotype: ♀ from Ecuador, Cotopaxi province, San Francisco de la Pampas, OTONGA reserve (-00.41994°-7900623°), 1997 m, 12 Nov. 2014, beating trees, N. Dupérré, E. E. Tapia, C. A. Tapia, ECFN 10901 (QCAZ). Paratypes: same data as holotype, 1♂1♀, ECFN 11260 (QCAZ); 1♂1♀, ECFN 11261 (ZMH-A0020678); 2♂3♀, ECFN 11262 (USNM).

Etymology. The specific name is a patronym in honor of the German botanist Dr. Wilhelm Barthlott for his contribution to the knowledge of the OTONGA reserve’s epiphyte diversity.

Diagnosis. Males and females most resemble P. niederi n. sp., and P. nowickii n. sp., but are differentiated by their dark abdomen with chevrons (Figure 34A), while abdomens lack chevrons in P. nowickii n. sp. (Figure 30A), and abdomens are light brown in P. niederi n. sp. (Figure 29A). Furthermore, females are distinguished from both species by their rounded spermathecae, separated by almost their width (Figure 35D), touching in P. nowickii n. sp. (Figure 31D), and elongated in P. niederi n. sp. (Figure 29C). Males are further distinguished from P. nowickii n. sp. by their retrolateral tibial apophysis tip curved (Figure 35B), while bent in the latter (Figure 31B).

Description. Female (holotype). Total length: 1.74; carapace length: 0.66; carapace width: 0.51; abdomen length: 1.08.

Cephalothorax. Carapace pear-shaped, brown suffused with black pattern (Figure 34A); pars cephalica flat rounded, pars thoracica incline. Sternum brown strongly suffused with black, truncated apically (Figure 34A); labium and endites light brown (Figure 34A); serrula present. Clypeus: 0.06. Chelicerae light brown; promargin three teeth, retromargin two teeth. Eight eyes surrounded by black pigmentation; AME 0.02, ALE 0.07, PLE 008, PME 0.08.

Abdomen. Oval, dark gray with pattern of four–five chevrons (Figure 34A); ventrally dark gray; tracheal spiracle recurved, 13x further from the epigastric groove than the spinnerets (Figure 34B, arrow); spinnerets aligned, uniformly whitish; PMS touching; ALS basal segment equal to PLS basal segment; PLS both segment of almost equal size (Figure 34B).

Legs. Whitish with two black marks ventrally on femur and tibiae, on black mark on metatarsi (Figure 34A,B). Legs macrosetae present on all segments, except metatarsi and tarsi; trichobothria present on all segments except femur and patellae. Leg measurements: I 2.40 (0.62/0.23/0.59/0.56/0.40); II 2.20 (0.59/0.19/0.54/0.49/0.39); III 2.11 (0.60/0.20/0.49/0.41/0.42); IV 2.61 (0.39/0.18/0.32/0.32/0.25); leg formula: 4123; palpal tarsal claw present.

Male (paratype). Total length: 1.74; carapace length: 0.71; carapace width: 0.52; abdomen length: 0.7.

Cephalothorax. As in female (Figure 34A). Sternum slightly darker than female (Figure 34B); chelicerae, endites and labium as in female. Clypeus: 0.07. Eight eyes surrounded by black pigmentation; AME 0.02, ALE 0.07, PLE 008, PME 0.08.

Abdomen. As in female (Figure 34A).

Legs. As in female. Leg measurements: I 2.40 (0.58/0.19/0.57/0.60/0.46); II 2.20 (0.63/0.21/0.51/0.44/0.41); III 1.97 (0.53/0.16/0.48/0.45/0.35); IV 2.52 (0.72/0.21/0.66/0.52/0.41); leg formula: 4123.

Genitalia. Palpal patella with pointed, sub-basal patellar apophysis; tibial apophysis curved, smooth with curved tip (Figure 35B); cymbium truncated, without retrolateral fold (Figure 35B); embolus elongated starting basally, circling 1× the tegulum; median apophysis absent (Figure 35A); tegulum not protruding (Figure 35A,B)

Distribution. Only known from the type locality in Cotopaxi Province.

Natural History. Specimens were collected in a low evergreen mountain forest (BsBn04) of the Western Cordillera [18] at 1997 m where it lives in sympatry with P. nowickii n. sp., Neohahnia pristirana n. sp., and Neohahnia freibergi n. sp.

Figure 34. Pristirana barthlotti new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 34. Pristirana barthlotti new species. (A) Female holotype and male paratype, habitus dorsal view. (B) Female holotype and male paratype, habitus ventral view (arrows point to tracheal spiracle). Scale bars: 0.5 mm.

Figure 35. Pristirana barthlotti new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

Figure 35. Pristirana barthlotti new species. (A) Male paratype, palp ventral view. (B) Male paratype, palp retrolateral view. (C) Female holotype, epigynum ventral view. (D) Female, internal genitalia dorsal view.

3.2. Key to South American Hahniidae genera

1. Males and females with rastellum ([6]; plate 50, fig. 5)…………………………………………	Cybaeolus
Males and females without rastellum………………………………………………………………..	2
2. Males and females with abdominal ventral patch of thick, short setae ([9]; fig. 1G) …………………………………………………………………………………………………………	Austrohahnia
- Males and females without with abdominal ventral patch of thick, short setae………………….	3
3. Males and females with white plumose setae on dorsum of abdomen (Figure 4A)……..	Kasha n. gen.
- Males and females without white plumose setae on dorsum of abdomen………………………	4
4. Males and females with spinnerets not aligned in row (Figure 24B and Figure 28A)…………………….	5
- Males and females with spinnerets aligned in row (Figure 24B)…………………………………..	(males) 6; (females) 12
5. Females with dorsal side of abdomen covered with white spatulate setae ([6]; p. 306, pl. 50, fig. 2)…………………………………………………………………………………………………….	Lizarba
- Males and females without dorsal side of abdomen covered with white spatulate setae (Figure 24A and Figure 28A) ……………………………………………………………………………………………	Paramito n. gen.
Males
6. Male palp without retrolateral tibial apophysis ([7]; fig. 9)…………………………………….	Harmiella
- Male palp with retrolateral tibial apophysis (Figure 11B and Figure 31B)…………………………………	7
7. Male palp with retrolateral cymbial furrow (Figure 20B)………………………………………	8
- Male palp without retrolateral cymbial furrow (Figure 35B)……………………………………..	9
8. Male patellar apophysis simple or bifid (Figure 15D and Figure 20B)…………………………………..	Neohahnia
- Male patellar apophysis trifid ([23]; fig. 54A,B)…………………………………………	Hahnia heterophthalma
9. Palpal cymbium rounded, with short RTA and short patellar apophysis ([24]; fig. 9E)………………………………………………………………………………………………………..	Intihuatana
Palpal cymbium and apophyses not as such……………………….……………..….…….………	10
10. Palp without patellar apophysis ([5]; fig. 15)….….….….…….…………………………………	Amaloxenops
- Palp with patellar apophysis (Figure 5B and Figure 11B)…………………………………………………….	11
11. Palpal cymbium truncated, RTA curved (Figure 31B and Figure 35B)………………………………….	Pristirana n. gen.
- Palpal cymbium pointed, RTA elongated ([25]; fig. 28)….…………………………………………	Hahnia tatei
Females
12. Epigynum with medial slit-like copulatory opening ([24]; fig. 8D) ……………………………	Hahnia michaleseni
- Epigynum without medial slit-like opening ………………………….…………….........................	13
13. Internal genitalia with spermathecae on elongated bases (Figure 29C and Figure 31D)…………………	Pristirana n. gen.
- Internal genitalia with primary spermathecae without elongated bases (Figure 5D and Figure 11D)……..	14
14. Internal genitalia with secondary spermathecae present (Figure 11D and Figure 22D)…………………	Neohahnia
- Internal genitalia without secondary spermathecae………...………...………….............................	15
15. Epigynum with copulatory openings well separated (4× from each other), spermathecae rounded ([23]; fig. 76C)……………...……….........................................................................................	Intihuatana
- Epigynum with copulatory openings joined or close together (2× from each other), spermathecae elongated ([23]; fig. 89B and 90A; Figure 1B)………………………………………………………	Amaloxenops

4. Discussion

4.1. Generic Description and Diagnosis

Throughout South America, Hahniidae are understudied; only one contemporary genus has been revised, Austrohahnia, by Rubio et al. [9]. Consequently, most genera and species’ generic descriptions and diagnoses are incomplete, often lacking illustrations. For instance, the genus Neohahnia was established in 1917 [3] by Mello-Leitão without any illustrations, and the type species N. sylviae, as well as N. palmicola Mello-Leitão, 1917, were destroyed (WSC 2023). Neohahnia ernsti (Simon, 1898) was described based on specimens (male and female) from St-Vincent by Simon [20], also without any illustrations. Later on, Petrunkevitch ([26]; p. 78; fig. 67, 68) described a female from Puerto Rico stating that he was not sure that the female was of the same species in the absence of males. Unfortunately, the identification cannot be confirmed since Petrunkevitch did not provide illustration of the epigynum. Lehtinen ([11]: p. 251) transferred the species, and redescribed the genus based on specimens of “Hahnia ernsti ♂♀syntypes? from Paris-London in part; and Paris & Hamburg from all parts of the Neotropical world”, and he illustrated the male palp and female epigynum ([11]; fig. 366, 370). Unfortunately, we did not find any specimens in Hamburg, and therefore cannot compare them with Lehtinen’s [11] description and illustrations. To redefine the genus and clearly establish the identity of the type species, new specimens from the type locality should be collected. In the meantime, we have provided a diagnosis of the genus based on the characters provided by Mello-Leitão [3], Lehtinen [11], and Heimer and Müller [8], and our observations.

Another problematic genus in South America is the genus Hahnia. This diverse group (102 valid species) is widely distributed (North America to Central America, Asia, and Africa), with four species found in South America. Once more, two species were described without illustrations, Hahnia heterophthalma Simon, 1905 [27] and Hahnia simoni Mello-Leitão 1919 [28]. One species, Hahnia michaelseni, is misplaced in the genus. The female epigynum with an elongated vertical slit-like opening is unique ([24]; fig. 8d). It should be noted that the female and male illustrated by Vellard [29] are misidentified. The female genitalia, the eyes, and the spinnerets do not match the type specimen ([29]; figs. 18–25). According to Vellard himself, the type could not be examined, no illustrations were provided by Simon, and he was unsure but preferred to assign his specimens to Simon’s species ([29]; p. 137). Additionally, Schiapelli and Gerschman’s illustration of a female epigynum ([30]; fig. 35) is unconvincing and should be considered as a possible misidentification.

In addition, the placement of Hahnia tatei (Gertsch, 1934) is uncertain. The species was first described in Bigois, based on males and females, but the female epigynum was not illustrated and the specimens are badly mutilated ([25]; p. 14). Lehtinen [11] synonymized Bigois with Alistra and transferred the species to a new genus Unzickeria, alongside Hahnia okefinokensis Chamberlin & Ivie, 1934 [25]. Opell and Betty [13] synonymized Unzickeria under Hahnia ([13]; p. 420), and as a result, the Gertsch species was also transferred to Hahnia. There are some differences between Hahnia tatei and other members of Hahnia, such as the tracheal spiracle position (5× further from epigastric grove) and spinnerets ([25]; p. 14); however, the type specimen and females have to be examined to establish the generic position of the species.

To help with the Hahniidae generic identifications herein, we provide a key to all South American genera. The key could not have been completed if not for Catley [23], who provided illustrations, diagnoses, and detailed descriptions of various type material (e.g., Intihuatana antarctica (Simon, 1902); Hahnia heterophthalma Simon, 1905 as well as other important species. Furthermore, Catley ([23]; p. 227) mentions that the male of Intihuatana antarctica (Simon, 1902) [31] has a median apophysis that was missed both by Lehtinen [11] and Dupérré and Harms [24]. The paralectotype was revised once more, and Catley’s observation is correct. The median apophysis is present in the species. Hahnia simoni Mello-Leitão, 1919 was not included in the key because we were unable to examine its type, and its original description lacks sufficient information to distinguish it from its congeners.

4.2. Morphology

The family Hahniidae is usually morphologically characterized by the spinneret aligned in one row (Figure 4B and Figure 8A). Most members of the family possess this unique characteristic, but some genera present a different spinneret conformation (e.g., Cicurina, Chorizomma, Lizarba, and Cybaeolus). Most genera occurring in Ecuador have their spinneret aligned in one row (Figure 4B and Figure 8A), except for the new genus Paramito n. gen. (Figure 24B). This new genus also differs from its Ecuadorian congener by the presence of a median apophysis, a ventral lamella on the cymbium (Figure 25A), and the male-modified chelicerae (Figure 26H). Based on its morphology, the new genus does not belong to the subfamily Hahniinae Bertkau, 1878, nor to the subfamily Cybaeolinae Lehtinen, 1967. There is a possibility that the genus belongs to the subfamily Cicurinae Pickard-Cambridge, 1893 [32]. However, the placement of the genus in any subfamily cannot be determined until more specimens are examined.

Interestingly Hahniidae, Hahniinae are often described with few or completely lacking spines [33], whereas all the species found in Ecuador have leg spines (macrosetae). Males and females from the genus Kasha n. gen. are well-armed, with two elongated macrosetae on all leg patellae (Figure 6H). Other members of the group, such as Austrohahnia, are characterized by a patch of setae on the ventral surface of their abdomen, while Kasha n. gen. is characterized by the presence of plumose setae on the abdomen (Figure 6F), a characteristic unique to the genus. Hahniids are also characterized by variability in their eye number and size. Within the same genus, eyes vary from six to eight (e.g., Neohahnia, Amaloxenops), and in most cases, the AME are small (Figure 2B) to minute (Figure 6C) compared to the other eyes. In Ecuador, only three species have eight eyes of equal size (N. pristirana n. sp., N. chalupas n. sp., and N. freibergi n. sp.) albeit N. freibergi n. sp. eyes are smaller than its congener (0.03 vs. 0.08).

4.3. Distribution and Habitat

Before the discovery of Hahniids in Ecuador, most species known to occur in South America were found in the Southern Region (Argentina, Chile). The discovery and description of 13 new species from Northern South America and the first record of the family in Ecuador shed new light on the diversity and distribution of this small family. In Ecuador, species occur in three ecoregions (the coast, Andes, and Amazonian) but most species are found in the Andes (Scheme 2). This could be due to sampling bias, but this trend was also observed in other spider genera such as Linothele and Mysmenopsis [34,35]. Three species occur in the Andean Paramó, specimens were collected in grassland habitats up to 4018 m, but most species were collected in evergreen forests between 884 m and 2225 m. Species were collected in sympatry, up to four species at the same locality, but our data suggest that the species coexist due to microhabitat niche separation. Some species are arboreal; Neohania catleyi n. sp., Neohahnia pristirana n. sp. (collected from epiphytes), and Pristirana barthlotti n. sp. (collected by beating trees), while other species are litter-dwelling species (Neohahnia piemontana n. sp., N. paramo n. sp., N chalupas n. sp., Paramito papallacta n. sp., and P. oyacachi n. sp.). Evergreen forests are known to be rich ecosystems, humid to hyper-humid [36] with high diversity and complexity of mosses and epiphytes [37] creating a diversity of microhabitats for small spiders, allowing sympatry.

5. Conclusions

This is the first step in studying the small family Hahniidae in Ecuador, providing a glimpse into its diversity. Although Ecuador is a small country, it has 65 distinct ecosystems. Herein, we have described Hahniidae species from five different ecosystems. Nevertheless, the 13 new species described herein have almost doubled our understanding of the South American Hahniidae fauna.