Alpiscorpius germanus (C. L. Koch, 1837 ) Kovařík & Štundlová & Fet & Šťáhlavský, 2019

Kovařík, František, Štundlová, Jana, Fet, Victor & Šťáhlavský, František, 2019, Seven new Alpine species of the genus Alpiscorpius Gantenbein et al., 1999, stat. n. (Scorpiones: Euscorpiidae), Euscorpius 287, pp. 1-29 : 10-12

Alpiscorpius germanus (C. L. Koch, 1837) , comb. n.

( Figures 7–12, 71, Tables 1, 4, 5) http://zoobank.org/urn:lsid:zoobank.org:act:C0E34258-

E851-48CC-9692-128CDE8226E5

Scorpius germanus C. L. Koch, 1837: 110–112 , pl. CVIII, figs. 250–252.

Euscorpius germanus “forma tipica”: Capra, 1939: 203.

Euscorpius germanus germanus: Di Caporiacco, 1950: 211 (in part); Bonacina, 1980: 58; Fet et al., 2004: 55 (in part).

Euscorpius germanus marcuzzii: Valle et al., 1971: 96 (in part); Bonacina, 1980: 58 (in part); Fet et al., 2004: 55 (in part).

Euscorpius germanus “T-Form”: Scherabon, 1987: 87.

Euscorpius (Alpiscorpius) germanus: Gantenbein et al., 1999: 55 ; Gantenbein et al., 2000: 858 (in part); Fet & Braunwalder, 2005: 28.

Euscorpius germanus: Komposch & Scherabon, 1999: 621 ; Komposch et al., 2001: 267; Soleglad & Sissom, 2001: 48; Kofler, 2002: 137; Fet et al., 2004: 55 (in part); Komposch, 2004: 444; Vignoli & Salomone, 2008: 205 (in part); Komposch, 2009: 359; Fet, 2010: 6 (in part); Tropea et al., 2015: 3; Fet et al., 2016: 6.

Euscorpius (Euscorpius) germanus: Fet & Sissom, 2000: 366–368 (in part; complete references list until 1998).

Euscorpius (Euscorpius) germanus germanus: Fet & Sissom, 2000: 366–368 (in part).

Euscorpius (Euscorpius) germanus marcuzzii: Fet & Sissom, 2000: 368 (in part).

Euscorpius marcuzzii: Mancini, 2018: 17 .

Euscorpius (Alpiscorpius) germanus Karyotypic race I (Ege 46a): ŠtundlovÁ et al., 2019: 156.

= Euscorpius germanus marcuzzii Valle et al., 1971: 96 , syn. n. http: //zoobank. org/ urn : lsid: zoobank. org: act: 86E73B4F-7774-4795-970D-95E27DBD9D50

TYPE LOCALITY AND TYPE DEPOSITORY. Italy, South Tyrol: BriXen (=Bressanone); neotype ♂ designated by Gantenbein et al. (2000: 858), NMBS, No. BG-109-07 .

MATERIAL EXAMINED ( NMPC): 29♂ 12♀. Austria, Carinthia: Nassfeld Pass, 46.560°N 13.278°E, 2♂ (Nos. S 045, S046); Walchenbach, 47.273°N 10.782°E, 3♂ (Nos. S 425, S468, S527) 1♂. Italy, Friuli-Venetia Giulia: Gerchia, 46.247°N 12.883°E, 3♂ (Nos. S 083, S086, S087); Lestans, 46.165°N 12.880°E, 2♂ (Nos. S 082, S088); South Tyrol : Montechiaro, 46.632°N 10.564°E, 3♂ (Nos. S 185, S263, S274); Schluderns, 46.669°N 10.593°E, 2♂ (Nos. S 254, S264) 1♂; Trentino: Ometto, 45.753°N 11.117°E, 1♂ (No. S 283) 1♀; Passo Rolle, 46.293°N 11.771°E, 1♂ 6♀ (Nos. S 1002, S1003, S1004, S1059, S1060, S1061, S1077); Quellenhof, 46.747°N 11.216°E, 4♂ (Nos. S 444, S432, S437, S445) 2♂ 4♀; Veneto: Peaio, 46.414°N 12.277°E, 1♂ (No. S 847); Soffranco, 46.280°N 12.244°E, 3♂ 2♀ (Nos. S 184, S1006, S1008, S1010, S1011) .

DIAGNOSIS. We limit Alpiscorpius germanus (C. L. Koch, 1837) to the “Karyotypic race I (Ege 46a)” defined by ŠtundlovÁ et al. (2019), which has 2n = 46 chromosomes. The karyotype is composed of 36 metacentric and 10 submetacentric chromosomes (fig. S1D in ŠtundlovÁ et al., 2019). The rDNA clusters are situated in the subterminal region of the long arms of the metacentric chromosome pair 11 (fig. S1D in ŠtundlovÁ et al., 2019). Number of pectinal teeth (Dp) in male usually 8 (ca. 60% of eXamined specimens). Number of patellar ventral trichobothria (Pv) 5. Reduction of patellar eXternal trichobothria from 4 to 3 common in series et (>10%) and very common in series eb a (up to 25%); see also Notes.

VARIABILITY. We scored standard phenotypic markers for 26 eXamined specimens (24♂, 2♀), with the following variation observed:

Dp in males (n=24): 8/3 (1), 6/6 (1), 7/7 (3), 7/8 (1), 8/8 (11), 8/9 (1), 9/8 (3), 9/9 (3); in total (not counting one aberrant Dp =3) 6 in 4.26% (2); 7 in 14.89% (7), 8 in 59.57% (28), and 9 in 21.28% (10); mean = 7.98, SD = 0.74.

Dp in females (n=2): 6/6 (1), 8/8 (1).

Pv (n=26): 5/3 (1), 5/5 (25); in total (not counting one aberrant Pv =3) 5 in 100% (51); mean = 5.00, SD =0.0.

et (n=26): 3/3 (2), 3/4 (1), 4/3 (2), 4/4 (20), 4/5 (1); in total, 3 in 10.87% (13.46%), 4 in 84.62% (18), and 5 in 1.92% (1); mean = 3.89, SD = 0.38.

eb a (n=26): 3/3 (3), 3/4 (4), 4/3 (3), 4/4 (16); in total, 3 in 25% (13) and 4 in 75% (39); mean = 3.75, SD = 0.43.

DISTRIBUTION. Austria, Italy (east of the Adige River: Friuli- Venetia Giulia, South Tyrol, Trentino, Veneto), Switzerland (Grisons: Val Müstair) ( Fig. 71). A. germanus is the northernmost species of the genus Alpiscorpius .

NOTES. Populations of A. germanus from Italy were studied by Bonacina (1980) and Gantenbein et al. (2000). Gantenbein et al. (2000) published 16S mtDNA sequences for 11 populations of “ Euscorpius (Alpiscorpius) germanus ” ( Switzerland: Grisons: Santa Maria, now Val Müstair); Italy: South Tyrol: Schluderns, Völs; Trentino: Vetriolo, Mezzocanale, Veneto: Borca di Cadore; Austria: Starkenbach, Tarrenz, Kranzach, Dellach, Oberdrauburg), all of which closely match DNA sequences of Ege 46a (= A. germanus ) obtained independently by ŠtundlovÁ et al. (2019). Since A. germanus is restricted in our study, the Italian populations studied by Bonacina (1980) should be reassessed, and their phenotypic markers scored from the material deposited in MCSNB.

Graham et al. (2012: 42) published a CoxI mtDNA sequence for a population of “ Euscorpius (Alpiscorpius) germanus ” from Egna (Neumarkt) ( Italy, Trentino), collected by F. ŠťÁhlavský, which closely matches those of Ege 46a (= A. germanus ) obtained independently by ŠtundlovÁ et al. (2019).

Note that A. germanus (C. L. Koch, 1837) s. str. is not found in Slovenia; instead, populations from Slovenia formerly listed under “ Euscorpius germanus ” are described below as two new species, A. kappa sp. n. ( Slovenia) and A. lambda sp. n. ( Slovenia, Italy).

Scherabon (1984, 1987) scored the standard phenotypic markers for a large series (over 200 specimens) of this species in Austria (as “T-Form” of Euscorpius germanus ), with the following results (recalculated here; et and eb a data from Scherabon, 1984):

Dp in males (n=74): 7/7 (2), 7/8 (4), 8/7 (8), 8/8 (45), 8/9 (3), 9/8 (5), 9/9 (6), 10/10 (1); in total, 7 in 10.81% (16), 8 in 74.32% (110), 9 in 13.51% (20), and 10 in 1.35%(2); mean = 8.08, SD = 0.65.

Dp in females (n=158): 4/6 (1), 6/4 (1), 5/5 (9), 5/6 (11), 6/5 (7), 6/6 (91), 6/7 (14), 7/6 (5), 7/7 (18), 8/7 (1); in total, 4 in 0.63% (2), 5 in 11.39% (36), 6 in 69.94% (221), 7 in 17.72% (56), and 8 in 0.32% (1); mean = 6.06, SD = 0.57.

Pv (n=238): 3/5 (1), 4/4 (2), 4/5 (7), 5/4 (6), 5/5 (211), 5/6 (3), 6/5 (7), 6/6 (1); in total, 3 in 0.21% (1), 4 in 3.57% (17), 5 in 93.70% (446), and 6 in 2.52% (12); mean = 4.99, SD = 0.26.

et (n=238): 3/4 (13), 4/3 (5), 4/4 (213) 4/5 (3), 5/4 (4); in total, 3 in 7.56% (18), 4 in 89.50% (213), and 5 in in 2.94% (7); mean = 3.98, SD = 0.23.

eb a (n=238): 0/4 (1), 2/3 (1), 4/2 (1), 3/3 (5), 3/4 (15), 4/3 (8), 4/4 (206), 5/4 (1); in total (not counting one aberrant), 2 in 0.42% (2), 3 in 7.37% (34), 4 in 92.0% (437), and 5 in 0.21% (1); mean = 3.91, SD = 0.34.

The reduction from 4 to 3 in both et group (7.56%) and eb a group (7.37%) in the marginal Austrian populations appears to be less than in eXamined Italian populations.

ŠtundlovÁ et al. (2019) have not studied the karyotypes of the Swiss population from Grisons (Santa Maria, now Val Müstair; see Braunwalder, 2001; Fet & Braunwalder, 2005) but the 16S mtDNA data ( Gantenbein et al., 2000) confirms that they belong to A. germanus (C. L. Koch, 1837) sensu stricto, as defined in this study. Fet & Braunwalder (2005) scored the standard phenotypic markers for 46 specimens of the Val Müstair population, with the following results:

Dp in males (n=12): 8.32, SD = 0.53.

Dp in females (n=25): mean = 6.46, SD = 0.63.

Pv (n=44): mean=4.92, SD = 0.27.

et (n=44): 3 in 6.82% (6) and 4 in 93.18% (82); mean = 3.94, SD = 0.93.

A great stability of Pv =5 within this species in all studied populations is notable compared to Pv = 6 in A. alpha and a bimodal distribution of this trait (5 to 6) in A. beta . The reduction from 4 to 3 in et group (6.82%) in the marginal Swiss population is slightly less than in eXamined Italian populations.

A special discussion is needed of the enigmatic taXon Euscorpius germanus marcuzzii Valle et al., 1971 . It was first recorded by Marcuzzi & Fabris (1957) from the Dolomites (northeastern Italy) as a population with 20 trichobothria (as opposed to common 21) on the eXternal aspect of the pedipalp patella. Valle et al. (1971: 95-96) very briefly (in just one line) described and named a new subspecies from the “refugial massifs of limestone Venetian Pre-Alps ( Italy) and northern Slovenia ”, without designating type specimens or eXact locality (a large series eXists in MCSNB that could be considered as syntypes). The sole morphological character originally distinguishing this taXon from E. g. germanus was the eXistence of 3 trichobothria instead of 4 in the accessory basal group (eb a = 3) on the eXternal aspect of pedipalp.

Recently, Mancini (2018) elevated E. g. marcuzzii to species level (without any justification) and designated the male lectotype from Bassano del Grappa (Vicenza, Veneto, Italy, 45.7667°N 11.7333°E, 1959, leg. R. Bianchi, MCSNB 333; paralectotypes 1♂ 2♀, same label, MCSNB 328, 330, 332), in the Venetian Pre-Alps (Ital. ‘Prealpi Venete’). Mancini also listed three paralectotypes from the same locality, thus limiting the type series to the type locality. Bassano del Grappa, which lies east of Adige River, is close to the range of A. germanus populations analyzed by ŠtundlovÁ et al. (2019, fig. 1), e.g. Ometto (45.753°N 11.117°E). Therefore, Euscorpius germanus marcuzzii Valle et al., 1971 , is a synonym of Alpiscorpius germanus (C. L. Koch, 1837) , syn. n., sensu stricto.

Bonacina (1980) mentioned a number of populations from northeastern Italy (Goriziano area) and Slovenia, some as “pure” E. g. marcuzzii , and some “miXed with E. g. germanus ” (as well as “hybrid”, i.e. asymmetric eb a =3/4 or 4/ 3 in the same individual!). Gantenbein et al. (2000) treated E. g. marcuzzii as a dubious taXon. Their DNA analysis of E. germanus from Italy in fact included asymmetrical specimens (eb a = 3/4 or 4/3) which were found to be rare at Bezzecca (only one find) but were much more common at Auronzo di Cadore, Belluno, Schluderns, Voltago, Vetriolo, and Völs. None of the populations mentioned corresponded to a monophyletic, clearly divergent clade.

At the same time, the Slovenian specimens of “ E. germanus ” studied by Gantenbein et al. (2000) (Črniče) did form a separate DNA clade; this population had both eb a = 3 and eb a = 4 specimens. The 16S rRNA DNA data for this population matches that for our A. lambda sp. n.; see below for discussion of its variability.

Moreover, the same reduction has been detected in the “ mingrelicus compleX” of this genus. While it is rare in the Anatolian-Caucasian part of the range ( Tropea et al., 2015), our data on newly described species of the “ gamma group” in Slovenia (see below) indicate that it is common at least in some species ( A. omikron sp. n.); see below.

Thus, eb a = 3 reduction represents a local phenotypic variation in some (but not all) species of Alpiscorpius , and is more or less prevalent in certain geographic areas. It could provide a useful marker for microevolutionary studies at population level; the eXistence of fluctuating asymmetry suggests incomplete dominance/penetrance of this phenotype (see Discussion)