This new species is most similar to A. acuminata and A. dominicana from Ecuador and Dominica, respectively, based on the general structure of tergum X (i.e., subtriangular in shape, with digitate apicomesal processes). It differs from A. acuminata by the shorter, thicker apicomesal processes on tergum X, and the shorter and rounder apicolateral processes on tergum X; in lateral view, the apicolateral processes in A. andina sp. n. are much shorter than the apicomesal processes, whereas in A. acuminata, both processes are roughly equal in length and the apicolateral processes are accuminate. Additionally, the inferior appendages in A. andina sp. n. are inflated mesally in lateral view, but not in A. acuminata. From A. dominicana, it differs by having a much narrower and shallower mesal cleft on tergum X between the apicomesal processes. Additionally, the posteromesal margin of the inferior appendages in ventral view is rounder and more pronounced in A. andina sp. n., whereas in A. dominicana, this margin is straight.

Adult male. Forewing length 9.8 ± 0.5 mm (n = 3). General color black, forewing membrane brown, covered in brown and white setae. Head with long, brown setae. Antennae with long, brown setae on scape and pedicel, flagellomeres with dark brown setae and ring of white setae basally. Maxillary palps brown, with long, brown hairs. Thorax black, with dark brown hairs. Forelegs brown, tarsomeres with white ring basally; mid legs dark brown with white setae; hind legs dark brown with white setae, interspersed with brown spines, increasing in thickness towards the tarsal segments. Tibial spur formula 0, 2, 2.

Segment IX annular, short, with anterior margin sinuous, posterior margin slightly produced mesally (Figure 2A). Preanal appendages shorter than tergum X, slender, digitate, setose (Figure 2A, B). Tergum X slightly notched apicomesally, basal portion of tergum X membranous, with two pairs of sclerotized apical processes; apicomesal processes digitate and directed upwards in lateral view (Figure 2A); apicolateral processes much shorter than apicomesal processes, thumb-like, directed laterad in dorsal and caudoventral views (Figure 2B, D); apices with short spicules (Figure 2A, B, D). Inferior appendages with basal portion of first segment broad, setose, highly pigmented, mesal margin rounded, apical portion digitate, straight, slightly inflated subapically in ventral view (Figure 2C), covered with stout, spine-like setae on its mesal surface; second article short, rectangular (Fig. 2A, C). Phallic apparatus simple, without any processes; phallobase tubular; phallotremal sclerites complex, consisting of pair of elongated sclerites basally (very faint in the specimens examined), hooked sclerite subapically, and U-shaped sclerite apically (rectangular in lateral view); endothecal membranes trilobed (Figure 2E).

Figure 2. 

Atanatolica andina sp. n., male genitalia. A Lateral B Segments IX and X, dorsal C Inferior appendage, ventral D Tergum X, caudoventral E Phallic apparatus, lateral (inset: phallic apparatus apex, ventral). Abbreviations: IX abdominal segment IX, X abdominal tergum X.

ECUADOR: Napo: Reserva Ecológica Cayambe-Coca waterfall, rd. to Oyacachi, 0.32621S, 78.1505W, 3690 m, 26.ii.2012, B Ríos-Touma, L Pita (UMSP) [UMSP000098741].

ECUADOR: Chimborazo: small roadside waterfall on Highway E-46 (via Riobamba - Macas), 2.17572S, 78.5047W, 3527 m, 2♂, 25.i.2015, R Holzenthal, B Ríos-Touma (MECN); Napo: unnamed trib. to Oyacachi R., ca. 5.2 mi W of Oyacachi, 0.229504S, 78.0059W, 2823 m, 1♂, 24.ii.2012, B Gill (NMNH), unnamed tributary to Papallacta River, Hwy. E-28, ca. 1 km SW Papallacta, 0.38589S, 78.1435W, 3246 m, 8♂, 6♀, 25.i.2012, B Gill (NMNH), stream, 2.73 Km W Papallacta, Hwy. E-28, 0.534639S, 78.2254W, 3982 m, 8♂, 25.i.2012, B Kondratieff, B Gill (NMNH).

Named after the Andean ranges where the specimens were collected.

Napo and Chimborazo Provinces (Ecuador) (Figure 8). The species occurs at the highest elevation ever recorded for the genus.

This species is related to A. aurea Holzenthal, 1988 from Colombia, and A. penai Holzenthal, 1988 from Bolivia, especially in the broad apicomesal processes of tergum X. The new species differs from A. aurea by the much shorter, acute, angulate apicomesal processes, and the rounded apicolateral processes on tergum X as well as the shape of the inferior appendages, especially the posteromesal margin of the first segment of this structure in ventral view. From A. penai, it differs by the shape of the apicomesal processes of tergum X, which in A. angulata sp. n. are strongly angulate. They also differ by the shape of the inferior appendages, which have a mesal bump on the inner surface in A. penai, but not in A. angulata sp. n.

Adult male. Forewing length 9 mm (n = 1). General color light brown, forewing membrane brown, covered in golden and brown setae. Head with long, white setae. Antennae with long, white setae on scape and pedicel, flagellomeres with dark brown setae and ring of white setae basally. Maxillary palps light brown, with long, white and short brown hairs. Thorax light brown with brown hairs. Forelegs with coxae, trochanter, and femur light brown with long, white setae; tibia and tarsomeres with white and brown setae; mid and hind legs light brown with white and brown setae, tibia and tarsomeres with two rows of dark spines ventrally. Tibial spur formula 0, 2, 2.

Segment IX annular, short, with anterior and posterior margins sinuous, setae on ventral and lateral surfaces not associated with warts (Figure 3A). Preanal appendages slightly shorter than tergum X, slender, digitate, setose (Figure 3A, 3B). Tergum X notched apicomesally, basal portion of tergum X membranous, with two pairs of sclerotized apical processes; apicomesal processes flattened, directed dorsad in lateral view (Figure 3A), strongly angulate, posterolateral apex acute in dorsal and caudoventral views (Figure 3B,) ; apicolateral processes capitate, directed dorsad in lateral view, slightly longer than the apicomesal processes, but equally as long in dorsal and caudoventral views (Figs 3A, B, D), digitate in dorsal and caudoventral views (Figure 3B, D). Inferior appendages with basal portion of first segment broad, setose, mesal margin forming an angle, apical portion elongate, digitate, slightly curved, and slightly inflated basally in ventral view (Figure 3C), covered with stout, spine-like setae on mesal surface; second article apparently fused with the first segment, represented by triangular semimembranous process (Figure 3A, C). Phallic apparatus simple, without any processes; phallobase short, tubular, curved downwards; phallotremal sclerite simple, spine-like, curved; endothecal membranes not everted (Figure 3E).

Figure 3. 

Atanatolica angulata sp. n., male genitalia. A Lateral B Segments IX and X, dorsal C Inferior appendage, ventral D Tergum X, caudoventral E Phallic apparatus, lateral.

ECUADOR: Napo: Río Jondachi, 30 km N Tena, 950 m, 10.ix.1990, OS Flint (NMNH) [USNMENT01295341].

The specific name angulata is a Latin adjective referring to the angulate apicomesal processes on tergum X.

Napo Province (Ecuador) (Figure 8).

This new species resembles A. homora Oláh, 2016 from Peru in that both possess paired, basodorsal membranous lobes on tergum X, but in A. homora these lobes bear peg-like setae and are narrower than in A. curvata sp. n. Also, the apicomesal processes of tergum X are long and capitate in A. homora (Oláh 2016, fig 26), but short and digitate in A. curvata sp. n. (Figure 4B, D).

Adult male. Forewing length 10.5 mm (n = 1). General color light brown, forewing membrane light brown, covered in golden setae throughout wing membrane and brown setae on costal margin. Head with long, yellow setae dorsally, brown setae ventrally. Antennae broken at second flagellomere, scape and pedicel with long, light brown setae. Maxillary palps light brown, with long, brown hairs. Thorax light brown with long, yellow setae. Forelegs light brown, tibia with brown setae, tarsomeres with white ring basally; mid and hind legs with yellow setae, and two rows of dark spines on tibia and tarsomeres. Tibial spur formula 0, 2, 2.

Segment IX annular, short, with anterior and posterior margins sinuous, with setae on ventral and lateral surfaces (ventral setae arising from a wart) (Fig. 4A). Preanal appendages shorter than tergum X, slender, digitate, setose (Figure 4A, 4B). Tergum X slightly notched apicomesally, basodorsal portion of tergum X membranous, produced into pair of membranous lobes; with two pairs of sclerotized apical processes; apicomesal processes digitate and directed dorsad in lateral view (Figure 4A); apicolateral processes shorter than apicomesal processes, apex subtriangular in dorsal view, directed laterad in dorsal and caudoventral views (Figure 4B, 4D). Inferior appendages with basal portion of first segment broad, setose, highly pigmented, mesal margin concave in ventral view (Figure 4C), apical portion digitate, curved mesad, inflated apically in ventral view, covered with stout, spine-like setae on its mesal surface; second article very small, short, triangular (Figures 4A, 4C). Phallic apparatus simple, without any processes; phallobase tubular; phallotremal sclerite simple, U-shaped in ventral view; endothecal membranes not everted (Figure 4D).

Figure 4. 

Atanatolica curvata sp. n., male genitalia. A Lateral B Segments IX and X, dorsal C Inferior appendage, ventral D Tergum X, caudoventral E Phallic apparatus, lateral (inset: phallotremal sclerite, ventral).

ECUADOR: Napo: 12 km W Baeza, 2380 m, 09.ix.1990, OS Flint (NMNH) [USNMENT01295342].

The specific name curvata is a Latin adjective that means curved and refers to the strongly curved inferior appendages in ventral view.

Napo Province (Ecuador) (Figure 8).

Morphology of the male genitalia of A. decouxi sp. n. is similar to A. cotopaxi Holzenthal, 1988 and A. muyupampa Holzenthal, 1988 from Ecuador and Bolivia, respectively. From A. cotopaxi, it differs by the slightly posteromesally produced segment IX, the longer preanal appendages reaching the apex of tergum X, and the shape and length of the apicolateral processes on tergum X, which are much shorter in A. cotopaxi. The putative larvae of A. decouxi sp. n. has spines on the anterior margin of the legs, similar to those found in A. cotopaxi, as illustrated by Holzenthal (1988), but the adults are light brown, whereas in A. cotopaxi, they are dark brown. Atanatolica muyupampa differs from A. decouxi sp. n. by the shorter and laterally directed apicomesal processes on tergum X, and the deeper mesal cleft on tergum X between the apicomesal processes; in A. decouxi sp. n. both of these processes are apically rugose. Additionally, the inferior appendage in A. decouxi sp. n. is straight in ventral view, but curved mesad in A. muyupampa.

Adult male. Forewing length 11 ± 0.5 mm (n = 3). General color light brown, forewing membrane light brown, covered in brown setae along the costal margin and yellow setae through the remainder of the forewing. Head with yellow and light brown hairs. Antennae with light brown hairs on the scape and pedicel, flagellomeres with dark brown setae and ring of white setae basally. Maxillary palps light brown, with long, brown hairs. Thorax brown with yellow and brown hairs. Fore and midlegs with coxae and trochanter with light brown hairs, remaining segments with dark brown setae and ring of yellow hairs basally. Hind legs with yellow hairs and interspersed brown spines, increasing in thickness towards the tarsal segments. Tibial spur formula 0, 2, 2.

Segment IX annular, short, with anterior margin sinuous, posterior margin slightly produced mesally (Figure 5A). Preanal appendages as long as tergum X, slender, digitate, setose (Figure 5A, B). Tergum X notched apicomesally, basal portion membranous, with two pairs of sclerotized apical processes; apicomesal processes digitate, slightly directed laterad in dorsal view (Figure 5B), apex rugose; apicolateral processes slightly shorter than apicomesal processes, thumb-like, directed laterad in dorsal and caudoventral views (Figure 5B, D), apex rugose. Inferior appendages with basal portion of first segment broad, setose, mesal margin rounded, apical portion digitate, almost straight in ventral view (Figure 5C), covered with stout, spine-like setae on its mesal surface; second article short, triangular, directed ventrad (Figure 5A, C). Phallic apparatus simple, without any processes; phallobase tubular; phallotremal sclerite complex, consisting of pair of elongated sclerites ventrally, hooked sclerite subapically, and U-shaped sclerite apically (subrectangular in lateral view); endothecal membranes trilobed (Figure 5E).

Figure 5. 

Atanatolica decouxi sp. n., male genitalia. A Lateral B Segments IX and X, dorsal C Inferior appendage, ventral D Tergum X, caudoventral E Phallic apparatus, lateral (inset: phallic apparatus apex, ventral).

Largest instars, assumed to be the 5^th, up to 13.8 mm in length (n = 89).

Head (Figure 6A) ovate, brown; eyes large; antennae very short; coronal suture very short, broad; ventral apotome a single sclerite, elongate rectangular, unpigmented; head with long, prominent primary setae in postgenal region and along anterior edge of frontoclypeal apotome; parietal region and frontoclypeus covered with short, closely appressed, clear setae; labrum quadrate, prominent, with row of many short to long primary setae along anterior third and many very short secondary setae along membranous apical edge; mandibles broadly triangular, without separate teeth, with smooth mesal scraping edge, patch of curved setae in mesal concavity (Figure 6A, B). Thorax (Figure 6A): pronotum slightly longer than wide, brown, except for unpigmented posterior edge; covered with two sclerites, with many long setae dorsally and laterally on anterior half, anterior edge with row of uniformly spaced short, spine-like setae. Mesonotal sclerites almost completely covering mesonotum, brown, with pair of small elongate-oval darkly pigmented anteromesal marks; with many long setae dorsally and laterally on anterior half, mesal setae forming W-shaped row. Metanotal sa1 (setal area 1) and sa2 sclerites completely fused, forming large single median plate with posterolateral corners extended and directed medially, with brown pigmentation mesally and small patches of pigmentation along posterior edge; covered with long setae; sa3 sclerites long, oval, brown, except for narrow unpigmented mesal edge, with long marginal setae. Meso- and metapleural sclerites large, brown; metasternum (Figure 6D) with pair of ventrolateral patches of ca. 20 long setae. Foretrochantin horn-shaped (Figure 7A). Legs elongate, cylindrical, robust, brown, setose; foreleg the shortest, hind leg the longest; short, spine-like setae present on anterior (mesal) surface of tibia and tarsus of foreleg (Figure 7A and inset) and tarsi of midleg and hind leg, hind tibia with incomplete suture at basal third (Figure 7B, C, and insets); tarsal claws short, thick. Abdomen: long and slender, abdominal gills not apparent; segment I with small, elongate-oval dorsal sclerite and dorsolateral setae, one long, one short (Figure 6A); lateral hump sclerite (Figure 6C) of segment I prominent, elongate, ventral portion heavily sclerotized and encompassing membranous, raised area covered with minute setae, dorsal portion very lightly sclerotized and extending almost to dorsum of segment; with anteromesal and ventrolateral rows of ca. 4–5 setae; abdominal fringe sinuous, very narrow, composed of minute spicules; with small lateral tubercles on segment VIII (identical as those illustrated by Henriques-Oliveira and Santos 2012: fig 2H); dorsal sclerite of segment IX semicircular in dorsal view (Figure 6E upper inset), with ca. 12 alternating short and long setae along posterior edge; anal prolegs (Figure 6E) each with narrow ventral plate in addition to small lateral sclerite and ventral sole plate, dorsolaterally with darkly pigmented secondary lateral sclerite; band of uniform small spines adjacent to anal opening; anal claw with robust primary hook and single dorsal accessory hook (Figure 6E lower inset).

Figure 6. 

Atanatolica decouxi sp. n., larva (tentative association). A Head, thorax, and abdominal; segment I, dorsal B Right mandible (enlarged), dorsal C Abdominal segment I, left lateral D Metasternum and abdominal segment I (partial), ventral E Abdominal segments IX and X, left lateral; upper inset: segment IX dorsal sclerite (enlarged), lower inset: anal claw (enlarged).

Figure 7. 

Atanatolica decouxi sp. n., larva (tentative association). A Right foreleg (inset: fore tibia and tarsus, anterior surface) B Right midleg (inset: tibia, anterior surface) C Right hind leg (inset: tibia, anterior surface) D Larval case, lateral (inset: posterior opening) E Pupal peduncle F Pupal anterior silken cap.

Elongate, narrow, gently curved and tapering, up to 20 mm long (Figure 7D); composed of small sand grains; posterior opening restricted to small opening by silken ring (Figure 7D inset); prior to pupation case fixed to substrate by short silken peduncle (Figure 7E), and anterior opening closed by silken cap with single opening (Figure 7F).

Larvae described here are tentatively assigned to A. decouxi sp. n. Unfortunately, no adult male metamorphotype pupae were collected to confirm the association. Larvae and adults were collected at the same site, but on different dates, and adults of only the single species were collected. In our previous collections of species in the genus and from museum material, it appears that species of Atanotolica do not co-occur at a site, lending support to this tentative association. The probable larva of A. decouxi is very similar to those described previously by Holzenthal (1988) and Henriques-Oliveira and Santos (2014). The larva of A. decouxi sp. n. described here is very similar to A. nordestina Henriques-Oliveira & Santos, 2014 in overall color and structure, but the morphology of the small spine-like setae on the anterior surfaces of the tibiae and tarsi may be distinctive; at least they are different from several species illustrated by Holzenthal (1988: figs 34–41) and most similar to those of A. cotopaxi. The lateral hump sclerite is also very similar to that described for A. cotopaxi by Holzenthal (1988: fig. 50) and A. nordestina (Henriques-Oliveira and Santos 2014: fig 2G). The case of A. cotopaxi is made of transparent silk with a few rock inclusions (Holzenthal 1988: fig 71), while those of A. decouxi sp. n. and A. nordestina are made entirely of rocks.

ECUADOR: Imbabura: Reserva Los Cedros, Río de la Plata, 0.32495N, 78.7808W, 1587 m, 15.iii.2012, B Ríos-Touma, G Bragado, T Policha (UMSP) [UMSP000158717].

ECUADOR: Imbabura: Reserva Los Cedros, Río de la Plata, 0.32495N, 78.7808W, 1587 m, 1♂, 1♀, 15.iii.2012, B Ríos-Touma, G Bragado, T Policha (UMSP), Reserva Los Cedros, tributary to Río Los Cedros, 0.30374N, 78.782W, 1312 m, 1♂, 2♀, 18-19.x.2011, R Holzenthal, B Ríos-Touma, A Encalada (MECN).

ECUADOR: Imbabura: Reserva Los Cedros, Río de la Plata, 0.32495N, 78.7808W, 1587 m, 84 larvae, 18.x.2011, R Holzenthal, B Ríos-Touma, A Encalada (MECN), 5 larvae (UMSP).

We dedicate this species to José DeCoux, an exceptional person who has been protecting Bosque Protector Los Cedros for more than three decades.

Imbabura Province (Ecuador) (Figure 8).

Larvae were found in high densities in the Río de la Plata on rocks adjacent to a large pool and in the riffle below the pool, forming groups of individuals (Figure 1B). All larvae were submerged. Larvae were observed feeding by scrapping periphytic algae growing on rocks. We were not able to differentiate separate stages of larvae collected, but the size distribution suggests continuous growth in this tropical region (Figure 9).

New distribution record

Atanatolica manabi Holzenthal, 1988:83 [Type locality: Ecuador, Manabi, Santo Domingo de los Colorados (79 km W); NMNH; ♂].

ECUADOR: Carchi: Río Hualchancito near Hacienda Primavera, 0.80279N, 78.21816W, 1200 m, 1♂, 1♀, 11.ix.2017, B Ríos-Touma (UMSP) (Figure 8).

The species was previously recorded from three males and several series of larvae collected from “Santo Domingo de los Colorados” and vicinity by workers from the Smithsonian Institution in the mid-1970s. The specimens from Carchi represent the only additonal records of the species since those collections. The male genitalia are identical to those illustrated for the holotype by Holzenthal (1988: fig. 21). In the Carchi specimens, the wings of the strikingly colored adults are in perfect condition. The forewings are brown with a large patch of golden hairs on the apical half extending as a band along the posterior (anal) margin to near its base (dorsal on specimen when wings folded at rest). The apical edge of the forewing bears a fringe of bright, white hairs. The legs and antennae are banded with white and brown hairs.

Figure 8. 

Distribution map of the new Atanatolica species described from Ecuador.

Figure 9. 

Body length (mm) (gray) and head width (mm) (black) of A. decouxi sp. n. larvae (n = 89) from Río de la Plata, Reserva Los Cedros, Imbabura Province, Ecuador.