The cradle of giants: insights into the origin of Scherotheca Bouché, 1972 (Lumbricidae, Crassiclitellata) with the descriptions of eight new species from Corsica, France Author Marchán, Daniel Fernández CEFE, Université de Montpellier, CNRS, EPHE, IRD, 1919, route de Mende, F- 34293 Montpellier cedex 5 (France) danifermch @ gmail. com danifermch@gmail.com Author Domínguez, Jorge Universidade de Vigo, Grupo de Ecoloxía Animal (GEA), E- 36310 Vigo (Spain) Author Hedde, Mickaël Eco & Sols, INRAE, IRD, CIRAD, SupAgro Montpellier, Campus SupAgro, Bâtiment 12, 2 place Viala, F- 34060 Montpellier cedex 2 (France) mickael. hedde @ inrae. fr hedde@inrae.fr Author Decaëns, Thibaud CEFE, Université de Montpellier, CNRS, EPHE, IRD, 1919, route de Mende, F- 34293 Montpellier cedex 5 (France) thibaud. decaens @ cefe. cnrs. fr decaens@cefe.cnrs.fr text Zoosystema 2023 2023-02-08 45 3 107 128 journal article 10.5252/zoosystema2023v45a3 bc090a4d-4bce-42e0-bf6b-fb4bd45ca811 1638-9387 7681295 urn:lsid:zoobank.org:pub:62C2AC78-741C-4675-8C3A-67E359EB369E Scherotheca altarocca Marchán, Decäens & Domínguez , n. sp. ( Fig. 3A ) urn:lsid:zoobank.org:act: 89EFE242-73AF-435E-86C8-67B17B93D253 TYPE MATERIAL. — Holotype . Corsica • Adult ; Corse du Sud , Zicavo , Castellu d’Ornucciu ; 41°52’22”N , 9°9’17”E ; 1422 m a.s.l. ; 3.IV.2021 ; T. Decaëns , D. Fernández Marchán leg.; BOLD Sample ID: DFM-0446 ; MNHN . Paratypes . Corsica • 3 adult specimens; Corse du Sud , Olivese (D26 road); 41°50’45”N , 9°4’55”E ; 1051 m a.s.l. ; 6.IV.2021 ; T . Decaëns, D. Fernández Marchán leg.; BOLD Sample ID: DFM-0512, DFM-0513, DFM-0514; MNHN • 3 adult specimens; Corse du Sud , Olivese (D69 road); 41°50’13”N , 9°5’45”E ; 1178 m a.s.l. ; 6.IV.2021 ; T . Decaëns, D. Fernández Marchán leg.; Eco&Sols; BOLD Sample ID: DFM-0266, DFM-0267, DFM-0268 ) • 2 adult specimens, 12 juvenile ; same data as for preceding; BOLD Sample ID: DFM-0269, DFM-0270, DFM-0271, DFM-0272, DFM-0274, DFM-0275, DFM-0276, DFM-0277, DFM-0278, DFM-0279, DFM-0280, DFM-0281, DFM-0282, DFM-0283; MNHN • 6 adult specimens; Corse du Sud , Zicavo , Castellu d’Ornucciu ; 41°52’22”N , 9°9’17”E ; 1422 m a.s.l. ; 3.IV.2021 ; T . Decaëns, D. Fernández Marchán leg.; BOLD Sample ID: DFM-0447, DFM-0448, DFM-0449, DFM-0450, DFM-0451, DFM-0452; MNHN • 3 adult specimens; Corse du Sud , Zicavo , Ponte di Valpine ; 41°52’33”N , 9°7’58”E ; 1248 m a.s.l. ; 03.IV.2021 ; T . Decaëns, D. Fernández Marchán leg.; BOLD Sample ID:, DFM-0443, DFM-0444, DFM-0445; MNHN . ETYMOLOGY. — The species name is derived from the natural region of Altarocca where the type specimens have been collected (name in apposition). TABLE 2. — Reproduction-related morphological characters of the newly described species of Corsican Scherotheca Bouché, 1972 and their closest relatives. Abbreviations: Sperm. pores , position of spermathecal pores; S.V., position of seminal vesicles; T.P. , position of tubercula pubertatis . Symbols: * , according to Pop (1947) : composite between the different populations included in the original description; ** , according to Bouché (1972) .

Species	Sperm. pores	Clitelum	T.P.	S.V.	Spermathecae
Sc. albomaculata Bouché, 1972	11/12-13/14 multiple	(26)27-37(38)	30-38	9,10,11,12	Multiple 11,12,13
Sc. altarocca Marchán ,	(11/12)12/13, 13/14 simple	(29)30-36(37)	(½ 32)33-36(37)	11,12	Simple (11),12,13
Decäens & Domínguez, n. sp.
Sc. boccaverghju Marchán ,	12/13, 13/14 multiple	27-36	31-34(½ 35)	9,10,11,12	Multiple 12,13
Decäens & Domínguez, n. sp.
Sc. brevisella Bouché, 1972	9/10-13/14 simple	½ 27-37	30-37	(10)11,12	Simple 9,10,11,12,13
Sc. capcorsana Marchán ,	12/13, 13/14 simple	(27)28-35(36)	½ 30-34	9,10,11,12	Simple 12,13
Decäens & Domínguez, n. sp.
Sc. corsicana* ( Pop, 1947 )	12/13,13/14 simple/	27-35,36	31-34	9,10,11,12	Simple/double/
double/triple	26-34	30,31-34,35 29-32	triple 12,13
Sc. corsicana **	12/13, 13/14 simple	(1/2 26)1/2 27-1/2	(1/2 30)31-1/2 34(34)	9,10,11,12	Simple 12,13
36(36)
Sc. corsicana popi Bouché, 1972	12/13, 13/14 simple	30-1/2 37	(33)34-1/2 37	9,10,11,12	Simple 13, 14
Sc. darioi Marchán, Decäens &	12/13, 13/14 simple	26-35	29-33	9,10,11,12	Simple 12,13
Domínguez, n. sp.
Sc. emiliae (Qiu & Bouché, 1998)	12/13, 13/14 simple	26-35	28-32(33)	11,12	Simple 12,13
n. comb.
Sc. litoralis Marchán, Decäens &	12/13, 13/14 simple	27-36	30-34(35)	9,10,11,12	Simple 12,13
Domínguez, n. sp.
Sc. marceli Marchán, Decäens &	12/13, 13/14 simple	26-35	(29)30-33	9,10,11,12	Simple 12,13
Domínguez, n. sp.
Sc. mausoleana Marchán ,	12/13, 13/14 simple	(26)27-36(37)	30-34(35)	9,10,11,12	Simple 12,13
Decäens & Domínguez, n. sp.
Sc. portcrosana Marchán &	12/13, 13/14 simple	26-35(½ 36)	30-33	9,10,11,12	Simple 12,13
Decäens, 2020
Sc. portonana Qiu & Bouché, 1998 12/13, 13/14 simple		(25)26-35	(28)29-32(33)	9,10,11,12	Simple 12,13
Sc. qiui Marchán, Decäens &	12/13, 13/14 simple	(26)27-35(36)	30-33(1/n 34)	*11,12	Simple 12,13
Domínguez, n. sp.
Sc. targionii (Baldasseroni, 1906)	11/12,12/13 simple/double	(26)27-40	27-40	11,12	Simple/double 11,12

DIAGNOSIS. — Scherotheca altarocca Marchán, Decäens & Domínguez , n. sp. can be distinguished from all of the other species of Scherotheca (except Sc. corsicana popi Qiu & Bouché, 1998 ) by the position of the clitellum in segments (29)30-36(37) and tubercula pubertatis in segments (½ 32)33-36(37) ( Tables 1 ; 2 ; Figs 2 ; 3A ). Although Sc. altarocca Marchán, Decäens & Domínguez , n. sp. resembles Scherotheca corsicana popi Qiu & Bouché, 1998 in characters such as the clitellum and tubercula pubertatis , it does not exactly fit the description made by Qiu & Bouché (1998a) , differing significantly in internal characters such as position of spermathecae and number of seminal vesicles (spermathecae in (11)12, 13 vs 13, 14 and 2 pairs of seminal vesicles in Sc. altarocca Marchán, Decäens & Domínguez , n. sp. vs 4 pairs in Sc. corsicana popi ). COI uncorrected average pairwise distances and topology of multilocus molecular phylogenetic trees supports the status of Sc. altarocca Marchán, Decäens & Domínguez , n. sp. as independent from other morphologically related species for which sequences are available. DISTRIBUTION AND ECOLOGY. — Scherotheca altarocca Marchán, Decäens & Domínguez , n. sp. has been found in the municipality of Olivese and Zicavo, in the natural region of Altarocca, Corse du Sud ( Fig. 4A ), in Fagus sylvatica L. forests and alpine grasslands at moderately high elevations ( 1051 to 1258 m ). DESCRIPTION External morphology Body pigmentation absent. White-beige in fixed specimens ( Fig. 3A ). Average length, 80 mm ( 66-95 mm , n = 12 adults ); body cylindrical in cross-section; average number of segments 149 ( 135-156 mm , n = 11 adults ; 135 segments in the holotype ). Average weight (fixed specimens), 1.26 g ( 0.81-1.78 g , n = 14 adults ). Prostomium epilobous, closed. Longitudinal furrows in segments 1 and 2. First dorsal pore at intersegmental furrow 10/11. Nephridial pores “en solfège” (irregularly distributed). Spermathecal pores at intersegmental furrows (11/12) 12/13 and 13/ 14 in setal line c . Male pores in segment 15, surrounded by a well-developed porophore. Female pores in segment 14. Clitellum saddle-shaped in segments (29)30-36(37). Tubercula pubertatis in segments (½ 32)33-36(37). Chaetae small and closely paired. Genital papillae in segments (10,11)12-16, (29,30)31,32,37,38(3 9,40,41,42). Internal anatomy Septa 5/6-10/11 thickened and muscular. Hearts in segments 6-11, oesophageal. Calciferous glands in segments 10-14, with diverticula in segment 10. Crop in segments 15-16, gizzard in segments 17-19. Typhlosole pinnate.Male sexual system holandric, testes and funnels (not enclosed in testes sacs, but with sperm present) located ventrally in segments 10 and 11. Two pairs of reniform seminal vesicles in segments 11 and 12; in one individual very inconspicuous, vestigial seminal vesicles present in segments 9 and 10. Ovaries and female funnels in segment 13, ovarian receptacles (ovisacs) in segment 14. Two or three pairs of small globular spermathecae in segments (11), 12 and 13 (intersegments 11/12, 12/13, 13/14). Nephridial bladders U-shaped, reclinate in segment 30. FIG. 2. — External and internal morphological differences between the newly described species of Corsican Scherotheca Bouché,1972 and their closest relatives. The three different shades represent absent, slight and moderate pigmentation. Open circles represent structures that only appear in some individuals. Structures shown in light and dark shades (genital papillae, clitellum and tubercula pubertatis ) represent the maximum and minimum extension of these characters, respectively. Phylogenetic relationships are the same as those shown in Figure 1. REMARKS This species displays some degree of morphological variability, with different combinations of extension of clitellum and tubercula pubertatis , position of genital papillae and spermathecae. These different morphotypes do not show any clear association with genetic lineages, and therefore they do not appear to correspond to differentiated taxa within the species described here. This species corresponds to Sc. corsicana L 6 in the checklist of Corsican earthworms of Marchán et al. (2022a) .