Andiodrilus nonuya sp. nov.

(Figs 1, 2 A,B, 3A–C, Table 1)

Material examined. Holotype: Clitellate specimen (CA-0013). Locality: Municipality of Belén de Los Andaquíes, rural community of Agua Dulce (01°20’34.2” N, 75°49’17.1” W), altitude: 280 m above sea level. Collection date: 25 June 2008. Collectors: A. Feijoo, L. Celis. Paratypes: 12 adults (CA-0014), two subadults, 44 juveniles. Locality: Municipality of Florencia, rural community of Alto Canelos (01°28’15.4”N, 75°32’49.7”W), altitude: 272 m above sea level (1 adult, 6 juveniles). Collection date: 20 June 2008. Locality: Municipality of Florencia, rural community of Balcanes (01°25’27.3” N, 75°30’23.0” W), altitude: 284 m above sea level (1 adult, 10 juveniles). Collection date: 18 June 2008. Locality: Municipality of Belén de Los Andaquíes, rural community of Agua Dulce (01°20’29.1” N, 75°49’09.5” W), altitude: 273 m above sea level (10 adults, 2 subadults, 28 juveniles). Collection date: 23 June 2008. Collectors: A. Feijoo, L. Celis.

Etymology. The new species is named in recognition of and homage to the Nonuyas indigenous people who live in the departments of Caquetá and Putumayo, Colombia. Noun in apposition.

Description. Endogeic species, with light brown-colored dorsum, milky white belly, and cylindrical body. The length of the conserved holotype is 93.5 mm, with 131 segments. Its diameter is 3.2 mm in the preclitellar region, 4.0 mm in the clitellum, and 3.0 mm in the mid-posterior region. Paratype: 82.5 mm long and 3.0 mm wide. Number of segments: 122. The prostomium is long, thin, and ends in a milky white lobe measuring 0.9 mm. The clitellum is saddle-shaped, located in segments 15–25 (=11 segments); tubercula pubertatis are between segments ¼ 19–¼ 24 (Fig. 1). Eight rows of closely paired setae are visible (a, b, c, d) as of segment 4. The distance between setae in the mid-body region (segments 40–50) is aa:ab:bc:cd:dd = 5.4:1.0:5.0:0.3:17. The common setae (Fig. 2 A) present a strong curvature with central marks. This thickness then decreases to finally form a sharply defined hook. The ventral setae ab from 18 to 23 are modified as genital setae (Fig. 2 B) about 1.4 mm long, 0.04 mm in diameter. Setae are slightly sigmoid with a strong curvature, continuing rectilinearly to end in a broadened lanceolate tip. Numerous semi-lunar grooves appear after 1/2, which then merge in the center forming bordered limits (Fig. 2 B left). The male pores are located in segment 21 and female pores in segment 14. The nephropores are aligned with setae c, d. Three pairs of spermathecal pores were observed on intersegments 6/7, 7/8, 8/9 aligned with seta c.

Internal anatomy. The anterior septa are strong and moderately thick between segments 6 and 10. The round muscular gizzard and thick musculature are found in the cavity of segment 6. Three pairs of extramural calciferous glands originate in a short, wide duct that opens dorsally on the back of the esophagus in segments 7–9; the glands are of lamellar structure ("Lamellentaschen") (Fig. 3 A).

The esophagus-intestine transition begins in segment 20 and the typhlosole begins in segment 24. This thick Sshaped lamella occupies 1/3 of the intestinal space. There are no intestinal caeca.

The dorsal blood vessel is moniliform in segments 15–21. There are three pairs of lateral commissures in 7–9 and three pairs of esophageal hearts in 11–13. There is one pair of holonephridia in each segment, located just above the line of setae c, d. Each nephridium of the postclitellar region consists of a single infundibulum and two foldable bands joined to the infundibulum (Fig. 3 B).

The voluminous and slightly lobed hypoesophageal testis sacs lie in segment 11, with one pair of seminal vesicles in 12. The deferent ducts are visible, located close to the ventral esophagus, opening in segment 14 into the intraparietal region and flowing into segment 21 to the male pores. The ovaries are in the ventral region of segment 13. Spermathecae (Fig. 3 C) are in segments 7, 8, 9. The sperm duct is short and the ampulla flat and round and has abundant sperm cells.

Remarks. Andiodrilus nonuya is closely related to A. icomi Righi, 1971, A. biolleyi Cognetti, 1904, A. guayabero Feijoo, 2008, A. kiri Righi, 1995, A. masigure Feijoo, 2008, A. pixao Righi, 1995, A. xaropi Righi, 1995, and A. cabalensis Righi, 1995, because of the extension of the clitellum and tubercula pubertatis. It is also very closely related to A. xaropi and A. cabalensis because of the presence of three pairs of hearts in 11–13. It differs, however, from the aforementioned species in the distance of the setae aa: ab: bc: cd: dd, the shape and ornamentation of the genital seta, the position and number of segments of the clitellum, the locations of tubercula pubertatis, the shape of spermathecae, and in the origin of the intestine (Table 1).

Species Author Length Segments Setal ratio Clitellum Tubercula Spermathecae Male pores Hearts

(mm) (no.) aa:ab:bc:cd:dd pubertatis

A. nonuya Feijoo 82.5–93.5 122–131 5.4:1.0:5.0:0.3:17 saddle, ¼ 19–¼ 24 Duct long, ampulla 21 11–13 & Celis 15–25 flattened