Glossoscolex (Praedrilus) lutocolus, Bartz, Marie Luise Carolina, James, Samuel Wooster, Pasini, Amarildo & Brown, George Gardner, 2012

Bartz, Marie Luise Carolina, James, Samuel Wooster, Pasini, Amarildo & Brown, George Gardner, 2012, New earthworm species of Glossoscolex Leuckart, 1835 and Fimoscolex Michaelsen, 1900 (Clitellata: Glossoscolecidae) from Northern Paraná, Brazil, Zootaxa 3458, pp. 59-85 : 61-65

Glossoscolex (Praedrilus) lutocolus n. sp. Bartz & James

( Fig. 1 a,b, Table 1)

Holotype. COFM BRPR 0050 adult, spring seepage area outskirts of Ibiaci district, Primeiro de Maio, Paraná, Brazil; 22º56.43’S, 51º01.24’W, 354 masl, 5 March 2005, G.G. Brown and S.W. James colls.

Paratypes. A - COFM BRPR 0008 one adult, same collecting data as holotype; B - COFM BRPR 0003 one adult, swamp, Reserva Particular do Patrimônio Natural Estadual Mata São Pedro, Lupionópolis, Paraná, Brazil; 22º41.79’S, 51º40.82’W, 314 masl, 29 April 2004, G.G. Brown and S.W. James colls.

Other material. COFM BRPR 0334 one juvenile, same location as holotype; COFM BRPR 0010 one subadult and MZUSP 1400 one adult, swamp, Reserva Particular do Patrimônio Natural Estadual Mata São Pedro, Lupionópolis, Paraná, Brazil; 22º41.79’S, 51º40.82’W, 314 masl, 29 April 2004, G.G. Brown and S.W. James colls; COFM BRPR 0005 two adults and MZUSP 1401 two adults, in front of river, wet area, Parque Estadual Ibicatu, Centenário do Sul, Paraná, Brazil; 22º46.97’S, 51º29.28’W, 331 masl, 29 April 2004, G.G. Brown and S.W. James colls.; COFM BRPR 0006 four adults, swamp, Colorado Farm, Paiquerê district, Londrina, Paraná, Brazil; 23º29.90’S, 51º02.42’W, 552 masl, 30 April 2004, G.G. Brown and S.W. James colls.; COFM BRPR 0007 one adult and one preclitellate, wet area, previously under rice cultivation, Sítio Igrejinha, Paiquerê, Londrina, Paraná, Brazil; 23º29.90’S, 51º02.42’W, 552 masl, 30 April 2004, G.G. Brown and S.W. James colls.; COFM BRPR 0051 two adults and MZUSP 1402 two adults, wetland next swamp, Jaguapitã, Paraná, Brazil, 23º45.22’S, 51º37.43’W, 485 masl, 26 November 2003, N.P. Benito and Y. Rios colls.; COFM BRPR 0072 three adults and MZUSP 1403 one adult and one preclitellate, wetland next to swamp, Jaguapitã, Paraná, Brazil, 23º45.22’S, 51º37.43’W, 485 masl, 26 November 2003, N.P. Benito and Y. Rios colls.

Etymology. The species is named for its preference of habitat, the soft alluvial mud of swamps and marshes.

Description. Dimensions: Holotype 164 mm by 6.1 mm at x, 5.0 mm at clitellum, 5.9 mm at xl, 219 segments; paratypes A - 185 mm by 7.2 mm at x, 7.8 mm at clitellum, 7.5 mm at xl, 196 segments and B - 119 mm by 6.4 mm at x, 7.1 mm at clitellum, 6.2 mm at xl, 147 segments. Body cylindrical. Setae closely paired throughout; genital setae absent; setal formula AA:AB:BC:CD = 18:2:10:1.5 at xl, DD> 1/2 circumference throughout. Setae aa commence on iv–viii, cd on iv–viii, usually not on same segment. Prostomium prolobous, pre- and postsetal secondary annulations present xi–xiv. Unpigmented, clitellum slightly yellowish or reddish. Ovipores postsetal, just above b in xiv; male pores 1.3–1.5 mm apart on xvi within paired ovate to angular porophores; broad midventral raised areas in BB of xv, larger paired raised genital markings on xvi. Clitellum saddle to just above b, xvi, xvii–xxv ( Fig. 1 a). Nephropores just above b.

Septa 5/6 thin, 6/7–10/11 equally thick and muscular, 11/12 and remaining septa membranous, septa 12/13/14 with white sac-like glandular investment anterior face of 12/13, posterior face of 13/14; 13/14 often with posterior bulges containing iridescent white material; posterior face of 12/13, anterior face of 13/14 coated with fine villous white material. Septa 12/13/14 united by circumesophageal membrane isolating villous interior from other septal contents of xiii, which are medial to membrane. Alimentary canal with large gizzard in vi; esophagus with high lamellae, large blood sinuses vii–ix, valvular in xiv; intestinal origin xv; typhlosole origin xv, end cxc–ccxc, strongly zig-zag folded xv–xix, xxi, zig-zag with ventral edge bent over to form pockets from xx to region of xxx, after gradually straightening to become simple thick lamina. Calciferous glands paired xii, composite-tubular type, lenticular (slightly elongate in Ibicatu material), sessile on dorsal esophageal wall; blood vessels to gland include large branch of dorsal vessel to approximate center of each gland, two coalescing vessels from ventral gland margin to extra-esophageal vessel. Gland opening to esophagus near dorsum, large opening with lip along its ventral margin. Holonephric, vesiculate; ducts to body wall near level of b.

Vascular system with ventral trunk, single dorsal trunk, lateral vessels in vii–ix, latero-esophageal hearts in x–xi. Extra-esophageal vessel visible near pharyngeal glands, pass along ventral-lateral face of gizzard and esophagus, ending in calciferous glands; supraesophageal vessel in x–xi.

Ovaries, ovarian funnels free in xiii adjacent to seminal vesicle tube passing through xiii; spermathecae absent. Male sexual system metandric, testes and funnels in single midventral subesophageal sac in xi; small pouches of septum 11/12 contain iridescent male funnels; medial to these pouches under hearts of xi, pass narrow tubes to seminal vesicles; seminal vesicles expand from narrow tubes in xiv, penetrate septa and range posteriorly along intestine to xxx–lxxx but mostly about xl–lv; seminal vesicles simple elongate sacs with parallel blood vessels on median side of longitudinal axis of vesicle; vasa deferentia long, looped from xi, form dense zig-zag on body wall en route to ventro-lateral face of large long oval muscular copulatory bulbs, join bulbs at level of xiv–xv; bulbs extend over xiv–xviii but occupy septally defined space of xvi. Copulatory bulbs with thin muscular outer layer, dense, delicate corrugated glandular inner surface with small lumen leading to male pore; no transverse muscle bands crossing over bulbs; bulbs with anterior, posterior apices attached by short stout muscles to body wall.

Remarks. Glossoscolex (Praedrilus) lutocolus belongs in the sub-genus G. (Praedrilus) Righi (1971). Righi mentioned this subgenus in Righi (1984), in relation to the description of G. matogrossensis (it has male pores in xv–xvi), but because of an indistinct and variable anterior clitellum boundary, Righi considered this species between subgenera Glossoscolex s.s. and Praedrilus. In Righi (1995) he stated that the subgenus Praedrilus could not be maintained because he thought that the clitellum of his G. t u p i i specimens was not fully developed, and that the position of the male pores (xvii) would then place G. t u p i i in the Glossoscolex (Glossoscolex) truncatus group ( Rosa, 1895), although he did not mention G. matogrossensis . Between 1971 and 1995 Righi found no further Glossoscolex species with the clitellum beginning behind the male pores, which was the basis for the Praedrilus subgenus designation. However we now have more species with preclitellar male pores, so the subgenus may yet be valid. The differences between G. (P.) lutocolus and G. (P.) tupii are as follows, with the characteristics of the latter in parentheses: length 150–230 mm (260–330 mm), number of segments 290–360 (450–620), setae beginning between iv and vii (setae beginning in segment xxx), setal ratios 17:1:9:1 (4:1.5: 1.6:1), DD> 1/2 circumference (DD<1/2 circumference), septum 11/12 present (septum 11/12 lacking), hearts of xi free (hearts of xi enclosed in testes sacs), testes sacs ventral, united (testes sacs circumesophageal), no muscle band over copulatory bulbs (two muscle bands, one large, over copulatory bulbs), septa bounding segment xiii with many small sacs on segment xii and xiv sides, sometimes with iridescent contents in sacs, thick white villous coating on xiii sides (no such development in septa 12/13/14). In G. (P.) tupii and some other members of the genus, septum 11/12 has been modified to create a testes sac encompassing the entire contents of segment xi. Although the number of specimens of G. (P.) tupii is not really sufficient to allow a good comparison—the original description is based on 3 clitellate specimens only—it appears that generally the seminal vesicles of the new species are longer. In cases where the segmental extent is comparable to G. (P.) tupii it is because numerous loops prevent extension further back.

Glossoscolex (P.) lutocolus corresponds to Glossoscolex n. sp. 21 and Glossoscolex n. sp. 10, as cited in Brown and James (2007a), Brown et al. (2004, 2008), James and Brown (2006, 2008), Fragoso and Brown (2007) and Sautter et al. (2006, 2007).

All the specimen lots represented in the material had members preserved in alcohol for DNA extraction, although none of the individuals examined morphologically has been sampled for DNA. The alcohol specimens collected from six different sites formed a distinct monophyletic cluster in a neighbor-joining analysis of the COI barcode region, with genetic distances below 6% within the cluster ( Fig. 2 ).

TABLE 1. Comparison of characters of the new species (in bold) of Glossoscolex (Praedrilus) , Glossoscolex (Glossocolex) and Fimoscolex , the type species of each genus and subgenus (marked with asterisk) and the species compared in the remarks. All species of the G. . (G.) truncatus group are included. Segments in Arabic numerals.

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Figures

FIGURE 1. New species of Glossoscolex. A, B. Glossoscolex (Praedrilus) lutocolus n. sp. C, D. Glossoscolex (Praedrilus) uliginosus n. sp. E, F. Glossoscolex (Praedrilus) itaguajensis n. sp. FM = female pores; MP = male pores; A, C, E lateral view, B, D, F ventral view.

FIGURE 2. Cluster of the neighbor-joining analysis of the COI barcode region from Glossoscolex (P.) lutocolus, G. (P.) uliginosus and G. (G.) palus. Phylogenetic relationship at the genus level ar not adequately represented.