Euscorpius candiota Birula, 1903

Fet, Victor, Soleglad, Michael E., Parmakelis, Aristeidis, Kotsakiozi, Panayiota & Stathi, Iasmi, 2013, Three more species of Euscorpius confirmed for Greece (Scorpiones: Euscorpiidae), Euscorpius 165, pp. 1-27 : 11-16

Euscorpius candiota Birula, 1903

( Figures 21–32; Tables 1–2)

Euscorpius candiota Birula, 1903: 298–299 .

REFERENCES:

Euscorpius carpathicus: Penther, 1906: 61 ; Roewer, 1928: 452; Roewer, 1943: 236 (in part; Crete); Vachon, 1948: 63; Stathi & Mylonas, 2001: 289 (in part; Crete); Kovařík, 2002: 13 (in part; Crete).

Euscorpius candiota: Birula, 1917a: 129 , 198; Birula, 1917b: 168, 210; Werner, 1938: 173; Di Caporiacco, 1950: 182, 188; Kinzelbach, 1982: 63–64 (in part; Crete); Bonacina, 1983: 9; Fet, 1986: 6 (under question).

Euscorpius carpathicus candiota: Kinzelbach, 1975: 36– 37 , fig. 12, 16 (in part; Crete); Valle, 1975: 219; Lacroix, 1991: 19; Kritscher, 1993: 384–385 (in part; Crete); Fet & Braunwalder, 2000: 19; Fet & Sissom, 2000: 362; Gantenbein et al., 2001: 302; Fet et al., 2003a: 368; Fet & Soleglad, 2007: 419; Vignoli & Salomone, 2008: 196, fig. 4; Fet, 2010: 7; Tropea & Rossi, 2012: 27, 30; Tropea et al., 2012: 75.

Euscorpius carpathicus “Subgroup A 3”: Fet, 2000: 53 (in part; Crete).

Euscorpius “ carpathicus candiota ”: Fet & Soleglad, 2002: 3; Fet et al., 2002: 142; Fet, 2003: 272; Fet et al., 2003b: 151; Fet et al., 2003c: S250.

Euscorpius “ carpathicus ” candiota: Fet et al., 2004: 52 ; Kaltsas et al., 2008: 234.

Type material: Lectotype (designated here) ♀, Greece, Crete, Candia (now Heraklion), 24 October (8 November) 1898, leg. Dr. Bogolyubov ( ZISP 947); paralectotypes, 8 ♂ and 14 ♀, same label as lectotype ( ZISP 947). Birula (1903) listed 8 ♂ and 18 ♀. Fet (1986) in 1985 examined in ZISP the entire syntype series; of these only 5 ♀ were available on loan for current study. See Fig. 21 for labels accompanying the loaned type specimens ( ZISP 947).

Geographic range. Greece: Crete (Heraklion). See map in Fig. 1.

History of Study

Birula (1903, 1917a, 1917b) considered Euscorpius candiota an endemic insular species, which he compared to topotypical E. carpathicus from Banat in Romania (then “southern Hungary ”) as well as to E. tauricus (C. L. Koch, 1837) from Crimea. Interestingly, Birula (1903) did not even mention two mainland Greek Euscorpius he just described previously ( Birula, 1900), E. koschewnikowi and E. scaber , probably since E. candiota was very distinct from both in morphology, primarily in granulation and metasomal carination.

In a lumping trend that prevailed in following decades, Penther (1906: 61) synonymized E. candiota with E. carpathicus . Vachon (1948: 64–66) studied a series from Crete and noted high variation within the island; he also considered Birula’s species a synonym of E. carpathicus , specifically of a subspecies he called E. carpathicus mesotrichus Hadži, 1929 , an unavailable name (a junior homonym of E. italicus mesotrichus Hadži ; see Fet & Sissom, 2000).

Birula’s name E. candiota had a special position in Kinzelbach’s unsubstantiated hypothesis of hybridogenic speciation within subgenus Euscorpius . Kinzelbach (1975) maintained that E. candiota (or E. carpathicus candiota ) is an “intermediate”, hybrid form between E. carpathicus and “ E. mesotrichus ”. The hy- pothesis was erroneously based on a correct discovery by Kinzelbach (1975) that two species of Euscorpius s.str. were sympatric in Thessaly (e. g. see below under E. ossae , which is sympatric with E. sicanus on Mt. Ossa). There is, however, no support for hybridization, and no phylogenetic indication that E. candiota , or any other Euscorpius species , has a hybridogenic origin. Kinzelbach (1975) recognized E. carpathicus (with 8 or less ventral patellar trichobothria) and “ E. mesotrichus Hadži ” (with 10 or more ventral patellar trichobothria). We know today that each of these two taxa sensu Kinzelbach (1975) in fact contains many separate species, some still undescribed. To accommodate “inter- mediate populations” (which ranged from 8 to 11 ventral patellar trichobothria), Kinzelbach (1975) assumed that those are hybrids between his two species, and designated them as “subspecies E. carpathicus candiota Birula ”. Such taxonomic assignment broadened the range of a “mixed form” across Greece from Crete to Kerkyra.

Fet (1986) published pectinal and trichobothrial statistics of Birula’s types from ZISP and ZMMSU but was unsure of their taxonomic status. Kritscher (1993) was the first to collect a very large series on Crete ( NHMW), which he addressed as E. carpathicus candiota . Fet & Sissom (2000) limited E. c. candiota to Crete; they listed it as a subspecies of E. carpathicus but noted that its status was still uncertain.

Gantenbein et al. (2001), using 16S rDNA markers, first demonstrated that “western” populations of from France (E. “ carpathicus ” niciensis) and Italy (now E. concinnus ) were genetically distant from “eastern” E. carpathicus candiota Birula (Crete) . Same reasoning was used by Fet (2003) to illustrate an isolated position of the Crimean E. tauricus (C. L. Koch, 1837) . The species delimitation approaches (Parmakelis et al., in press) validated as a distinct evolutionary entity a population from the Dionysades (satellite islands located northeast of Crete) as belonging to “ E. candiota complex”. The exact taxonomy of Euscorpius on Crete proper remains unresolved. Additional 16S DNA marker data (unpublished) from within Crete show a potential diversification of this clade, which likely includes the northern population of E. candiota redescribed here from type locality (Heraklion).

Diagnosis. Medium sized (about 40 mm), brown in color species; pedipalps dark orange. Metasomal carinae on segments II–IV smooth to obsolete. Pedipalp patellar external trichobothria numbers: eb = 4, eba = 4, esb = 2, em = 4, est = 4 and et = 6 to 7; ventral aspect of patella 9 to 10. Pectinal tooth counts: females usually 7, males 8 to 9.

FEMALE. The following description is based primarily on the lectotype female from Candia (now Heraklion), Crete, Greece, with some structures described from a paralectotype female. Measurements of the lectotype female are presented in Table 2. See Fig. 22 for the dorsal and ventral views of the lectotype female. Lectotype female, an adult, has the right pedipalp detached, chela detached from femur and patella, 2 legs detached, and mesosoma slightly damaged.

COLORATION. Carapace and metasoma dark brown; tergites brown; pedipalp dark orange; telson yellow, aculeus red; legs yellow-orange; genital operculum, pectines, and basal piece yellow; and sternites brown. No patterns present.

CARAPACE ( Fig. 23). Anterior edge slightly convex; slight granulation on lateral edges below lateral eyes, otherwise, smooth and lustrous, lacking any indication of carinae. There are two lateral eyes. Median eyes and tubercle are small to medium in size, positioned slightly anterior of middle with the following length and width ratios: 0.422 (anterior edge to medium tubercle middle / carapace length) and 0.160 (width of median tubercle including eyes / width of carapace at that point).

MESOSOMA. Tergites I–VII smooth and lustrous; tergite VII lacking lateral and median carinal pairs. Sternites III–VII smooth and lustrous; VII lacking lateral and median carinae. Stigmata are very small, narrow elliptical.

METASOMA ( Fig. 24). Segment I wider than long in ratio 0.861. Segments I–IV: dorsal carinae are smooth with subtle granulation distally; dorsolateral carinae smooth to obsolete; lateral carinae obsolete; ventrolateral carinae obsolete on I–II, weak and smooth on III– IV; single ventromedian carina obsolete. Segment V: dorsolateral carinae rounded and rough; lateral carinae obsolete; ventrolateral and ventromedian carinae irregularly granulate. Anal arch lined with minute granules. Intercarinal areas generally smooth except for the distal ventral surface of segment V which is covered with scattered granules.

TELSON ( Fig. 27). Vesicle slightly swollen and elongated, with moderately curved aculeus. Vesicle essentially void of granules, very lustrous. Vesicular tabs smooth.

PECTINES ( Fig. 30, paralectotype female). Mediumdeveloped segments exhibiting length / width ratio 2.222 (length taken at anterior lamellae / width at widest point including teeth). Sclerite construction complex, three anterior lamellae and 5 middle lamellae; fulcra of medium development. Teeth number 7/7. Sensory areas developed along distal aspect on all teeth, including basal tooth. Basal piece large, length / width ratio 0.541.

GENITAL OPERCULUM ( Fig. 30, paralectotype female). Sclerites triangular, wider than long, connected for most of their length. Genital papillae absent.

STERNUM ( Fig. 30, paralectotype female). Type 2, posterior emargination present, moderately defined convex lateral lobes, apex visible but not conspicuous; wider than long, width to length ratio 0.80.

CHELICERAE ( Fig. 29, paralectotype female). Movable finger dorsal edge with two small subdistal (sd) denticles; ventral edge smooth; serrula not visible. Ventral distal denticle (vd) conspicuously longer than dorsal (dd). Fixed finger with four denticles, median (m) and basal (b) denticles conjoined on common trunk; no ventral accessory denticles present.

PEDIPALPS ( Figs. 25–26, paralectotype female Fig. 28). Well-developed chelae, moderately carinated, medium scalloping on chelal fingers. Femur: Dorsointernal and ventrointernal carinae serrated, dorsoexternal and ventroexternal crenulated. Dorsal surface scattered with small granules, ventral surface smooth except for a proximal cluster of small granules, internal surface with a line of 13+ granules, and external surface smooth. Patella: Dorsointernal carinae irregularly granulate, ventrointernal crenulate, dorsoexternal with lowprofile granules, ventroexternal granulated, and externomedian carina irregularly granulated. Dorsal and ventral surfaces essentially smooth; external surface rough; internal surface smooth with well-developed DPS and near obsolete VPS. Chelal carinae: Complies with the “10-carinae configuration”. Digital (D1) carina strong, smooth, slight traces of low-profile granulation; subdigital (D2) essentially obsolete, represented by 3 small granules; dorsosecondary (D3) obsolete, area quite flat; dorsomarginal (D4) smooth with large low-profile scattered granules; dorsointernal (D5) rounded to obsolete; ventroexternal (V1) strong, granulated, curving to external condyle of movable finger; ventromedian (V2) obsolete; ventrointernal (V3) strong, but very rounded and smooth; external (E) strong medially, rough. Chelal finger dentition (paralectotype female Fig. 28): Median denticle (MD) row groups in straight line; 6 ID s fixed finger and 7 on movable finger; 6 OD s on fixed finger and 7 on movable finger; 4 and 5 IAD s on fixed and movable fingers, respectively. Trichobothrial patterns ( Fig. 32): Type C, neobothriotaxic: chela ventral = 4/4; patellar eb = 4/4, eba =4/4, esb = 2/2; em = 4/4, est = 4/4, et = 6/6; patellar ventral = 9/10.

LEGS ( Fig. 31, paralectotype female). Both pedal spurs present on all legs, lacking spinelets; tibial spurs absent. Tarsus with delicate single row of nine spinules on ventral surface, terminating distally with one pair of spinules. Unguicular spine well-developed and pointed.

HEMISPERMATOPHORE. Hemispermatophore unknown for this species.

Variation. We examined in detail only a lectotype and four paralectotype females of E. candiota from Candia (now Heraklion), Crete. In addition, all 23 syntypes (8 ♂ 14 ♀, ZISP 947 ) were scored for trichobothria and pectinal teeth number by the first author years ago in ZISP ( Fet, 1986: 6). This putative species, or forms closely related to it, are widespread and common across Crete; however, the full assessment of variation (and redescription of males) within the island will be a subject of a separate study by our research group using molecular and morphological data (in progress) .

Within the syntype series ( Fet, 1986), variation was as follows. Pectinаl teeth number in males varied unequally between 8 (31.2 %) and 9 (68.8 %), with distribution 8 (5) and 9 (11) [n=16], mean 8.69, SD = 0.48. Pectinаl teeth number in females was usually 7 (86.7 %), with distribution 5 (2), 6 (1), 7 (26), and 8 (1) [n=30], mean 6.87, SD = 0.57. Number of ventral patellar trichobothria (Pv) varied almost equally between 9 (52.1 %) and 10 (45.8 %), with distribution 8 (1), 9 (25), and 10 (22) [for n=48 as reported by Fet, 1986: 6; should be n=46], mean 9.44, SD = 0.54. Number of external terminal patellar trichobothria (et) varied almost equally between 6 (43.5 %) and 7 (54.3 %), with distribution 5 (1), 6 (20), and 7 (25) [n=46], mean 6.52, SD = 0.55. Also, in two cases, number of external subterminal patellar trichobothria (est) was aberrant 3, otherwise (in 44 pedipalps, or 95.7 %) it was “standard” 4 [n = 46].