Nemastoma bidentatum sparsum Gruber & Martens, 1968

Novak, Tone, Novak, Ljuba Slana, Kozel, Peter, Schaider, Miriam Gudrun, Komposch, Christian, Lipovšek, Saška, Podlesnik, Jan, Paušič, Igor & Raspotnig, Günther, 2021, Hidden diversity within the Nemastoma bidentatum Roewer, 1914 complex (Opiliones: Nemastomatidae) Part I: Morphological evidence, European Journal of Taxonomy 777, pp. 1-67 : 11-16

Nemastoma bidentatum sparsum Gruber & Martens, 1968

Figs 2–3 , 4B , 5B , 6B , 7C , 8C , 9B , 10B , 11A, C , 12B , 13B ; Tables 3–4

Nemastoma bidentatum sparsum Gruber & Martens, 1968: 146–147 , figs 1–2, 7−11 (original description) [partim; the type series also includes N. bidentatum gruberi ssp. nov.].

Nemastoma bidentatum – Roewer 1914: 141 [partim: type series also includes N. bidentatum bidentatum and N. triste ]. — Šilhavý 1956: 131, figs 243–246. — Loksa 1962: 267, figs 3a–f. — Gruber & Martens 1968: 146–147, figs 1–2, 7−11. — Staręga 1976: 313–315, figs 25–27. — Martens 1978: 111–112, figs 143−145, 147, 150; 2006: 182–184, figs 20, 21c–d, 22a–h. — Novak & Gruber 2000: 286. — Novak et al. 2002: 136 [partim: Radenci]; 2006: 269. — Komposch & Gruber 2004: 485.

— Novak 2004b: 245 [partim: Breznica, Slavonski brod, Spačva near Županja]; 2005a: 315 [partim: Bosanski Brod]. — Schönhofer & Holle 2007: 25, figs 2a–i. — Schönhofer 2013: 36.

Diagnosis (according to Gruber & Martens 1968, slightly modified)

Male Ch basal article with nearly straight dorsal margin in front of dorsal indentation, and short, roundly oblong Ch-Apo, with small apical pinnacle. Pa-Fe moderately club-shaped (Pa-Fe min:max width ~1:2.5), with 2 distal, pointed spines, spine 2 the largest. Pa-Pt Apo conical. Glans equilaterally triangular. Rec sem with 14 balloon-like vesicles.

Material examined

CROATIA – XM05 • 1 ♀; Jurovčak ; 3 Apr. 1983; T. Novak and L. Slana Novak leg. (LSN 7/1983, TN rev. 2008); PMSL .

SLOVENIA – WM06 • 1 ♂; Cvitrško sedlo, Otiški vrh ; 28 May 2008; T. Novak leg. (160/2011); PMSL. – WM42 • 1 ♂, 2 ♀♀, 1 juv.; Mestinje; 4 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 43/1983); PMSL • 1 ♂, 1 ♀; ibid .; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1138/2002); PMSL • 1 ♀; Šmarje pri Jelšah; 6 Apr. 1986 (1169/1997, rev. 2015); PMSL • 1 ♀; Zgornje Poljčane; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1160/2002); PMSL. – WM45 • 1 ♂; Ledine ; 25 Jul. 1987; F. Janžekovič leg. (7/1999); PMSL • 1 ♂; Maribor; 24 Aug. 1986; Malavašić and Koren leg. (265/2002); PMSL. – WM47 • 1 ♀; Kresnica ; 20 Apr. 2008; L. Slana Novak and T. Novak leg. (162/2011); PMSL. – WM52 • 1 ♂; Maršečka vas ; 11 Oct. 1997; L. Slana Novak and T. Novak leg. (1893/1998); PMSL. – WM53 • 2 ♂♂, 1 ♀; Pečke ; 24 Jun. 2007; T. Novak leg. (45/2007); PMSL • 1 ♂, 2 ♀♀; Šega, Makole; 9 Apr. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 70/1983); PMSL. – WM56 • 1 ♀; Kaniža; 20 May 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 80/1983); PMSL • 1 ♀; Mt. Piramida , Maribor, 1−30 Nov. 1978; (909/1997, rev. 2015); PMSL • 1 ♀; Spodnje Dobrenje ; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (165/2011); PMSL • Ranca , Pesnica. – WM57 • 1 ♂; Ceršak ; 20 Apr. 2008; L. Slana Novak and T. Novak leg. (113/2011); PMSL • 1 ♂; Kozjak pri Ceršaku ; 20 Apr. 2008; L. Slana Novak and T. Novak leg. (116/2011); PMSL. – WM66 • 2 ♀♀; Črnetina ; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (163/2011); PMSL • 1 ♂; Kremberg; 4 Jun. 1984; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 343/1985); PMSL • 2 ♂♂; ibid .; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (114/2011); PMSL • 1 ♀; Spodnji Žerjavci ; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (115/2011); PMSL. – WM67 • 1 ♂; Sv. Ana ; 3 Jul. 2002; T. Novak leg. (799/2002); PMSL • 1 ♀; Nasova ; 3 Jun. 1984; L. Slana Novak and T. Novak leg. (314/2002, rev. 2015); PMSL • 1 ♂; Stogovci; 13 Apr. 1984; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 323/1985); PMSL • 1 ♂, 1 ♀; Zgornja Velka; 20 Mar. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 8/1983); PMSL. – WM73 • 1 ♀; Repišče ; 24 Jun. 2007; T. Novak leg. (31/2007, rev. 2015); PMSL. – WM74 • 1 ♀; Podvinci ; 26 Oct. 1994; M. Varžič leg. (21/1999); PMSL • 1 ♂; Žamenci ; 22 Oct. 1994; M. Varžič leg. (18/1999); PMSL • 1 ♂; ibid .; (20/1999); PMSL • 1 ♂; ibid .; 23 Oct. 1994; M. Varžič leg. (19/1999, rev. 2015); PMSL. – WM75 • 1 ♂; Trnovce ; 2 Jun. 2002; L. Slana Novak and T. Novak leg. (760/2002); PMSL • 7 ♂♂, 4 ♀♀; Velika Slavšina ; 13 Oct. 2005; L. Slana Novak and T. Novak leg. (158/2005, rev. 2018); PMSL • 5 ♂♂, 3 ♀♀; Vitomarci ; 30 Jun. 2002; T. Novak leg. (787/2002); PMSL • 1 ♀; ibid .; 3 Jul. 2002; T. Novak leg. (792/2002); PMSL • 1 ♂; ibid .; (797/2002); PMSL • 1 ♀; ibid .; 3 Jul. 2002; T. Novak leg. (794/2002, rev. 2008); PMSL • 1 ♀; ibid .; (796/2002); PMSL • 1 ♂; ibid .; (797/2002); PMSL • 1 ♂; ibid .; (800/2002); PMSL • 1 ♀; ibid .; 2 Oct. 2002; T. Novak leg. (1280/2002); PMSL • 1 ♂; ibid .; (1279/2002); PMSL. – WM77 • 1 ♀; Črešnjevci; 24 Apr. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 79/1983, TN rev. 2008); PMSL • 1 ♂, 1 ♀; Korovci ; 7 Apr. 1984; L. Slana Novak and T. Novak leg. (LSN 191/1985); PMSL • 1 ♂, 1 ♀; Lomanoše; 1 Oct. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 110/1983); PMSL. – WM78 • 1 ♂; Gerlinci ; 11 May 1986; T. Novak and L. Slana Novak leg. (LSN 139/1986); PMSL. – WM86 • 2 ♂♂, 6 ♀♀; Hrašenski vrh ; 9 Sep. 1983; T. Novak and L. Slana Novak leg. (LSN 115/1983); PMSL • 4 ♂♂, 3 ♀♀; Kapelski vrh; 17 Sep. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 112/1983); PMSL • 1 ♂; Murski vrh ; 19 May 1984; L. Slana Novak and T. Novak leg. (LSN 240/1984); PMSL • 1 ♀; Radenci; 25 Jul. 1983; M. Slana Novak and L. Slana Novak leg. (LSN 111/1983, TN rev. 2007); PMSL • 2 ♂♂, 2 ♀♀; ibid .; 20 Sep. 1983; L. Slana Novak leg. (LSN 119/1983); PMSL • 1 ♂; Rihtarovci ; 26 Feb. 1997; B. Drovenik leg. (94/2001); PMSL. – WM87 • 3 ♂♂, 3 ♀♀; Cankova; 15 Sep. 1983; T. Novak and L. Slana Novak leg. (LSN 114/1983); PMSL • 3 ♂♂, 3 ♀♀; Skakovci ; 27 Mar. 1983; T. Novak and L. Slana Novak leg. (LSN 4/1983); PMSL • 1 ♂; Sodišinci , Radenci; 19 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 68/1983); PMSL. – WM88 • 1 ♂; Gerlinci; 11 Aug. 1983; L. Slana Novak, M. Slana Novak and T. Novak leg. (1322/1998); PMSL. – WM89 • 2 ♂♂, 4 ♀♀, 1 juv.; Trdkova; 30 Jul. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 49/1983); PMSL. – WM94 • 1 ♀; Pavlovci ; 15 Apr. 2000; L. Slana Novak and T. Novak leg. (744/2002); PMSL. – WM95 • 1 ♀; Podgradje , Ljutomer; 27 May 2001; S. Brelih leg. (125/2001); PMSL. – WM96 • 1 juv.; Bratonci; 9 Aug. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 335/1985, TN rev. 2008); PMSL. – WM97 • 1 ♂; Andrejci ; 28 Jul. 1981; T. Novak leg. (1188/1981, JG det.); PMSL • 1 ♂; Moravci ; 16 Mar. 1984; T. Novak and L. Slana Novak leg. (LSN 243/1984); PMSL • 4 ♂♂; Murska Sobota; 24 Nov. 1982; T. Novak leg. (603/1982); PMSL • 3 ♀♀; Rakičan – Noršinci ; 23 Sep. 1984; L. Slana Novak leg. (322/1985); PMSL • 1 ♂, 2 ♀♀; ibid .; 13 Sep. 1981; T. Novak leg. (371/1982, JG det.); PMSL. – WM98 • 2 ♂♂, 3 ♀♀; Košarovci; 24 Jul. 1982; L. Slana Novak, T. Novak and M. Slana Novak leg. (529/1982, JG det.); PMSL • 1 ♀; ibid .; 1 Sep. 1984; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 318/1985); PMSL • 1 ♂, 3 ♀♀; Mačkovci; 31 Jul. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 67/1983); PMSL. – WM99 • 2 ♂♂; Lucova; 2 Jun. 1984; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 239/1984); PMSL. – XM05 • 1 ♀; Bobri ; 24 Aug. 1988; F. Velkovrh leg. (13/1995); PMSL • 2 ♂♂, 1 juv.; Dolnja Bistrica ; 17 Aug. 1996; I. Kovač leg. (SL 2/1997); PMSL • 2 ♀♀; ibid .; (SL 3/1997); PMSL. – XM06 • 2 ♂♂, 3 ♀♀; Banuta /Bánuta; 10 Aug. 1996; I. Kovač leg. (SL 5/1997); PMSL • 2 ♂♂, 1 ♀; ibid .; (SL 22/1997); PMSL • 2 ♂♂, 2 ♀♀; ibid .; (SL 21/1997); PMSL • 13 ♂♂, 15 ♀♀; ibid .; 13 Aug. 2012; T. Novak leg. (GR 3837−3849, 3792−3797, TN det.); PMSL • 4 ♂♂, 3 ♀♀; ibid .; (GR 3770−3776); PMSL • several specs; Polanski log, Mala Polana ; 17 Jul. 2000; T. Novak leg.; PMSL • 1 ♂, 1 ♀; Radmožanci / Radamos ; 17 Mar. 1984; T. Novak and L. Slana Novak leg. (LSN 109/1984); PMSL. – XM07 • 4 ♂♂, 3 ♀♀; Dobrovnik /Dobronak; 8 Aug. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 57/1983); PMSL • 1 ♀, 1 juv.; ibid .; 18 Aug. 1996; I. Kovač leg. (SL 1/1997); PMSL. – XM08 • 3 ♂♂, 1 ♀; Prosenjakovci / Pártosfalva; 8 Aug. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 50/1983); PMSL • 2 ♀♀; Središče / Szerdahely ; 12 Aug. 1996; I. Kovač leg. (SL 6/1997); PMSL • 4 ♂♂; ibid .; 24 Aug. 1996; I. Kovač leg. (SL 7/1997); PMSL • 2 ♀♀; ibid .; (SL 8/1997); PMSL • 1 ♂, 3 ♀♀; ibid .; (SL 20/1997); PMSL • 4 ♂♂; ibid .; (SL 23/1997); PMSL • 4 ♂♂; ibid .; (SL 61/1997); PMSL • 3 ♂♂; ibid .; (SL 108/1997); PMSL • 12 ♀♀; ibid .; (SL 105/1997); PMSL. – XM15 • 3 ♀♀; Benica / Benice ; 1Aug. 1996; I. Kovač leg. (SL 11/1997); PMSL • 3 ♀♀; ibid .; (SL 12/1997); PMSL • 2 ♂♂; ibid .; 19 Aug. 1996; I. Kovač leg. (SL 25/1997); PMSL • 2 ♀♀; ibid .; (SL 26/1997); PMSL • 5 ♀♀; ibid .; (SL 28/1997); PMSL • 3 ♂♂; ibid .; (SL 29/1997); PMSL • 2 ♂♂; ibid .; (SL 39/1997); PMSL • 2 ♀♀; ibid .; (SL 64/1997); PMSL • 1 ♂, 1 ♀; ibid .; (SL 65/1997); PMSL • 1 ♂, 3 ♀♀; Gaberje / Gyetyános ; 22 Aug. 1996; I. Kovač leg. (SL 9/1997); PMSL • 1 ♀; ibid .; (SL 10/1997); PMSL • 1 ♂; ibid .; (SL 15/1997); PMSL • 3 ♀♀; ibid .; (SL 16/1997); PMSL • 1 ♀; ibid .; 12 Oct. 1996; I. Kovač leg. (SL 30/1997); PMSL • 2 ♀♀; ibid .; (SL 55/1997); PMSL • 5 ♂♂; ibid .; 19 Oct. 1996 (SL 17/1997); PMSL • 4 ♂♂, 4 ♀♀; ibid .; (SL 24/1997); PMSL • 4 ♀♀; ibid .; (SL 103/1997); PMSL • 8 ♀♀; ibid .; (SL 104/1997); PMSL • 2 ♀♀; ibid .; (SL 106/1997); PMSL • 7 ♂♂; ibid .; (SL 107/1997); PMSL • 2 ♂♂; ibid .; (SL 109/1997); PMSL • 14 ♀♀; ibid .; 22 Aug. 1997; I. Kovač leg. (SL 27/1997); PMSL • 1 ♂, 1 juv.; ibid .; (SL 32/1997); PMSL • 1 ♂; ibid .; (SL 68/1997); PMSL • 4 ♂♂; ibid .; 24 Aug. 1997; I. Kovač leg. (SL 19/1997); PMSL • 2 ♀♀; ibid .; 28 Aug. 1997; I. Kovač leg. (SL 13/1997); PMSL • 2 ♂♂; ibid .; (SL 14/1997); PMSL • 1 ♂; ibid .; 12 Oct. 1997; I. Kovač leg. (SL 4/1997); PMSL • 4 ♂♂; ibid .; (SL 20/1997); PMSL • 1 ♀; ibid .; (SL 56/1997); PMSL • 1 ♂; ibid .; (SL 100/1997); PMSL • 4 ♀♀; ibid .; (SL 101/1997); PMSL. – XM16 • 2 ♂♂; Murska šuma ; 29 Aug. 1998; T. Novak leg. (1538/1998); PMSL • several specs; ibid .; 18 Jun. 2001; T. Novak leg.; PMSL • 1 ♂, 1 ♀; Dolgovaške Gorice / Hossúfaluhegy; 31 Mar. 1984; T. Novak, D. Kuhar and L. Slana Novak leg. (LSN 141/1984); PMSL • 4 ♂♂, 1 ♀; Lendavske gorice / Lendvahegy ; 17 Aug. 1996; I. Kovač leg. (SL 31/1997); PMSL. – XM25 • 1 ♂; Muriša, Murska šuma; B. Drovenik and A. Pirnat leg. (26/2005); PMSL .

Description (according to Gruber & Martens 1968, slightly modified)

Male (from Prosenjakovci)

BODY. Length 1.5−2.0. Males from Slovenia larger than those from the type locality. Length 1.96, width 1.44.

CHELICERAE. Ch basal article with nearly straight dorsal margin in front of dorsal indentation and small ventral hump, Ch-Apo short, roundly oblong, with convex anterior margin and small apical pinnacle ( Figs 4B , 6B , 9B ). Secretion field medially. Proximal article length 0.53, distal article length 0.59, max width 0.18, movable finger length 0.25.

PEDIPALPS. Pa-Tr with highly arched dorsal hump. Pa-Fe moderately club-shaped (Pa-Fe min:max width ~ 1:2.5), slightly ventrally bent, with 2 stout, pointed spines in distal Pa-Fe quarter; spine 2 the largest. Pa-Pt Apo triangular-conical. Pa-Ti simple, straight. Pa-Ta conically tapering ( Figs 5B , 6B ). Pa article lengths in Table 3.

PENIS. Pe length 1.6, Pe basis ¼ Pe length, glans equilaterally triangular ( Figs 7C , 8C ).

LEGS. Leg article lengths in Tables 3–4.

Female (from Prosenjakovci)

BODY. Body length 1.6−2.4. Females from Slovenia larger than those from the type locality. Body length 2.40, body width 1.97.

CHELICERAE. Ch basal article with low-arched dorsal margin in front of dorsal indentation, and straight ventral margin with triangular hump, Ch basal article length:width>2.6:1. Proximal article length 0.57, distal article length 0.63, max width 0.21, movable finger length 0.32 ( Fig. 13B ).

PEDIPALPS. Pa-Tr with moderately high, arched dorsal margin; Pa-Fe with distal bonce, Pa-Fe max:min width ~ 1.7:1; Pa-Ti elongated-oval (13B). Pa article lengths in Table 3.

OVIPOSITOR. Ovipositor length 1.0, Rec sem with 14 balloon-like vesicles ( Fig. 11A, C ).

LEGS. Leg article lengths in Tables 3–4.

Remarks

In Slovenia, N. b. sparsum lives in the northeast, at the margins of the Pannonian basin (Martens & Gruber 1968; Martens 1978; Kovač 1997). The type series deposited in Naturhistorisches Museum Wien ( Gruber & Martens 1968) also includes N. b. gruberi ssp. nov. distributed south of the Sava River in Slovenia and Croatia. All prior identifications of N. b. sparsum south of the Sava River ( Novak et al. 1995, 2002; Novak 2005b) refer to N. b. gruberi ssp. nov. Martens (1978: 112) reported a sparsum -like population living in the mountain pine association (Hyperico grisebachii-Pinetum mugo Horvat 1938 ) on Mt. Veliki Snežnik (the peak at 1796 m) at least up to the altitude of 1650 m a.s.l. These individuals may refer to N. b. gruberi ssp. nov., N. b. martensi ssp. nov. or N. b. sneznikensis ssp. nov. Individuals from Bosnia and Herzegovina (Mt. Plasa, Jajce) and Dalmatia, Croatia (Split) ( Gruber & Martens 1968) should be revised for identification. See remarks under N. b. gruberi ssp. nov. and N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov., N. b. martensi ssp. nov. and N. b. sneznikensis ssp. nov.

Distribution

Central and eastern Europe: Austria, Bulgaria, Bosnia and Herzegovina, Czech Republic, Denmark, Hungary, Croatia, Montenegro, Romania, Russia, Slovakia, Slovenia, Serbia and Turkey ( Gruber & Martens 1968; Staręga 1976; Martens 1978, 2006; Mučalica 1988; Klimeš 2000; Komposch 2002; Komposch & Gruber 2004; Schönhofer & Holle 2007). Vertical distribution: 120−1650 m a.s.l. ( Komposch 1999; Komposch & Gruber 2004), in Slovenia 155–644 m a.s.l. Type locality: Sonnenberg (47.88° N, 16.31° E, 480 m a.s.l.), Leitha Mts, Burgenland, Austria.

Ecology

Nemastoma b. sparsum is a lowland and hilly land subspecies inhabiting various types of acidophilous and neutrophilous forests of Fagus sylvatica , Carpinus betulus L. and Quercus petraea (Matt.) Liebl. , riverine forests, and thermophilous scrubs in north-eastern Slovenia, mostly on non-calcareous soils. Phenology: adults eurychronous.

Figures

Fig. 2. The complete distributional area of the Nemastoma bidentatum Roewer, 1914 complex, according to Gruber & Martens (1968), Martens (2006), Schönhofer & Holle (2007) and own data, with marked positions of the type localities of the taxa.

Fig. 3. Distributional areas of six subspecies (circles) and hybrids (triangles, squares) within the Nemastoma bidentatum Roewer, 1914 in Slovenia and adjacent regions. Note that many localities overlap on the map.

Fig. 4. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Ch basal articles (medial, lateral views).

Fig. 5. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Pa (medial view).

Fig. 6. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Synchroscopic images of the male Ch basal article and Pa-Fe (medial views, except G, right figure, which is in dorso-medial view).

Fig. 7. Nemastoma bidentatum Roewer, 1914 complex. Penes (lateral view). Note: The penis of N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 8. Nemastoma bidentatum Roewer, 1914 complex. Glans (dorsal and lateral views). Note: The penis of N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 9. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Variation of the Ch basal article (medial view) in species, subspecies and hybrids.

Fig. 10. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Variation of the Pa-Fe (A–B first image, C, D–E first image, F–G first image; H–J, L–M right, N–O: lateral view; the others in dorso-medial view to show the armament).

Fig. 11. Nemastoma bidentatum Roewer, 1914 complex, ♀♀; Rec sem (lateral view, right portion, except F left figure, which is in dorsal view). A. The frame encloses the ovipositor portion with Rec sem. Note: Female N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 12. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. From left to right: Ch basal article (medial, lateral views), Pa (medial view) and Pe tip (dorsal, lateral views). Arrows indicate the most identifying characters. Note: Pe of N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 13. Nemastoma bidentatum Roewer, 1914 complex, ♀♀. From left to right: Silhouettes of the Ch basal article, Pa (both medial view) and Rec sem. Arrows indicate the most identifying characters. Note: female N. pluridentatum (Hadži, 1973) stat. nov. is unknown.