Three new “caecate” earthworm species from Sulawesi, Indonesia (Oligochaeta, Megascolecidae)

Abstract Three new earthworm species are described from Sulawesi, Indonesia. Two belong to the genus Pithemera Sims & Easton, 1972, namely P.suwastikai Fahri, Amaliah & Atmowidi, sp. n. and P.tadulako Fahri, Amaliah & Atmowidi, sp. n. The new species, P.suwastikaisp. n. is distinguished by a medium size (135–165 mm long, 4.5–6.5 mm diameter), four pairs of spermathecal pores in 5/6/7/8/9, 7–12 setae between male pores, no genital markings, holandry, and simple intestinal caeca. Pithemeratadulakosp. n. is recognized by a large size (217–340 mm long, 13–15 mm diameter), two pairs of spermathecal pores in 7/8/9, no setae between male pores, no genital markings, holandry, and simple intestinal caeca. Another new species, Metaphirerusydii Fahri, Amaliah & Nguyen, sp. n., is diagnosed by its large size (250–280 mm long,12–16 mm diameter), two pairs of spermathecal pores in 7/8/9, no setae between male porophores, presence of genital markings in the male region, holandry, and complex intestinal caeca. Additionally, an identification key to “caecate” species is provided to the Sulawesi’s fauna.

The earthworms were killed in formalin 2%, transferred to 4% formalin for fixation for approximately 24 hours, and then transferred to fresh 4% formalin for longterm preservation and morphological studies. Specimens were dissected from the dor-

Pithemera suwastikai
Etymology. The species is named after Dr I Nengah Suwastika, head of the Biology Department of Tadulako University for his kind support of our study on Sulawesi earthworms.
The openings of male pores small, on the setal ring of xviii, paired and discharging directly onto surface of xviii (Fig. 2E); ventral distance between male pores approx. 0.2x body circumference. Genital markings absent in the male region.
Holandric. Testes sacs paired in x-xi, small, separated, ventral, yellowish. Seminal vesicles in xi-xii. Ovaries paired on xiii. Prostate gland racemose paired within xvi-xx or xvii-xx. Prostatic ducts long and U-shaped (Fig. 2D). No accessory glands. Habitat and ecology. Specimens were collected in yards at Lomba village, Banggai district, Central Sulawesi, Indonesia at 5 m a.s.l. The species was found in sandy soils at depths of 0-10 cm, around the roots of banana trees planted 1-2 m away from houses. No other specimens were collected despite continued collecting effort. The specimens were collected at an elevation of 50 meters above sea level. When collected, earthworms were moving actively on the ground.
Remarks. This new species belongs to the Pithemera pacifica group characterized by having the first spermathecal pores in 5/6 and four thecal segments (Sims and Easton 1972). This group currently consists of three species, P. pacifica (Beddard, 1899) from New Guinea and Samoa, P. dahli (Ude, 1905) from New Guinea, and P. donvictorianoi (Aspe & James, 2015) from the Philippines. Although Pithemera suwastikai sp. n. is classified into Pithemera pacifica group because of the presence of four thecal segments (5/6-8/9), the new species is totally differentiated from P. pacifica (Beddard, 1899) in having larger size (length 135-165 mm vs. up to 56 mm), holandric (vs. metandric), a single female pore (vs. a pair of female pores (see James et al. 2004)), clitellum in xiv-xvi without setae (vs. in xiv-xv/xvi with setae), absence of genital markings (vs. presence of genital markings).
Etymology. The species is named after the well-known Tadulako University in central Sulawesi. 'Tadulako' means a 'leader' in indigenous culture of central Sulawesi.
Description. External characters. Body generally cylindrical; large size, length 217-340 mm, diameter 13-14 mm at x and 15 mm at xx, segments 120-123. Darkish blue on dorsum, purplish brown around clitellum and yellowish white on ventrum, especially for living specimens (Fig. 3A). After fixation, color grayish blue on dorsum and yellowish white on ventrum.
Male pores small, forming vertical lines, paired and discharging directly onto surface of xviii (Fig. 3E); copulatory pouches absent; ventral distance between pores approx. 0.02x body circumference. Genital markings absent in the male region.
Habitat and ecology. Specimens were collected in Puncak Jiti, Toro village, South Kulawi sub-district, Sigi district at 1,370 m a.s.l. and near Kalimpa'a Lake, Lore Lindu National Park at 1,600 m a.s.l. This species was found on the surface of soils after rains that contains a lot of leaf litter.
Remarks. The new species is totally different from species groups divided by Sims and Easton (1972) in first spermathecal pores in 7/8 and two thecal segments. The new species is clearly different from all other Pithemera species in its very large size (length 217-340 mm, diameter 13-14 mm at x and 15 mm at xx, segments 120-123), and color of living specimens (dark blue on dorsum, purplish brown around clitellum, and yellowish white on ventrum, color after fixation grayish blue on dorsum and yellowish white on ventrum).
Remarks. This genus currently contains approximately 198 species widely distributed in the Oriental regions from Japan southwards through the Indo-Australasian archipelago to the rain forests of Australasia eastwards through Oceania (http://taxo. drilobase.org/index.php?title=Metaphire).  Description. External characters. Body generally cylindrical. Large size, length 250-280 mm, diameter 12-16 mm at x and 13-15 mm at xx, segments 117-119. Purplish blue on dorsum through the lateral region, clitellum purplish brown on dorsum through the lateral region and paler on ventrum, especially for living specimens ( Fig. 4A-C). After fixation, color grayish blue on dorsum and yellowish white on ventrum.
Male pores deeply located inside copulatory pouches in the setal ring of xviii. The openings of copulatory pouches horizontally elongated, with swollen edges; ventral distance between male porophores 0.02x body circumference (Fig. 4G). Three pairs of small, rounded genital markings including two in front of the setal rings of xvii and xx, and one medially located in the setal ring of xix (Fig. 4G). Ventral distance between genital markings in xvii and xx about 0.07x body circumference, and less than 0.01x body circumference in xix.
Spermathecae paired in viii and ix. Spermathecal ampulla large, oval shaped; ducts stout and short, 1/5 as long as ampulla. Diverticulum cylindrical, much shorter than ampulla, attached to ducts; distal part enlarged to a rounded seminal chamber (Fig.  4E). No nephridia on spermathecal ducts. No accessory glands in the spermathecal region.
Holandric. Testes sacs paired in x-xi, large, separated, ventral, yellowish. The copulatory pouches hidden inside body wall. Seminal vesicles in xi-xii. Ovaries paired on xiii, oviduct and ovisac developing in xiii. Prostate glands racemose, paired in xvii-xix (Fig. 4F). Prostatic duct U-shaped, invisible from dorsal view since being hidden under prostate glands. No accessory glands.
Habitat and ecology. Specimens were collected in Mt. Torompupu (Quarlessi Mountain), natural forest at 1,120 m a.s.l. and 1,360 m a.s.l. This species was found on soil surface in piles of leaf litter after rains. According to local hunters in Mt. Torompupu, the species can be mostly found after rains and usually observed on old trails in the forest. We did not find this species at an altitude of less than 1,000 m a.s.l.; presumably, the species may be distributed in an altitude of more than 1,000 m a.s.l.