Calchas gruberi Fet, Soleglad et Kovařík, 2009

Fet, Victor, Soleglad, Michael E. & Kovařík, František, 2009, Etudes on iurids, II. Revision of genus Calchas Birula, 1899, with the description of two new species (Scorpiones: Iuridae), Euscorpius 82 (82), pp. 1-72 : 57-68

Calchas gruberi Fet, Soleglad et Kovařík , sp. nov.

( Figs. 3 , 5–7 , 9–11 , 14 , 17–21 , 25–26 , 32–38 , 59–75 ; Tables 2–3)

REFERENCES:

Calchas nordmanni : Kinzelbach, 1980: 169–174 (in part), figs. 1–2, 5 (map locality 16); Kinzelbach, 1982: 58 (in part); Kinzelbach, 1985: Map IV (in part); Crucitti, 1999: 87 (in part); Kovařík, 1999: 40, 42 (in part); Fet & Braunwalder, 2000: 18 (in part); Sissom & Fet, 2000: 418–419 (in part); Crucitti & Cicuzza, 2001: fig. 7 (in part); Stathi & Mylonas, 2001: 290, 293; Soleglad & Fet, 2003 a: 5, fig. 2; Soleglad & Fet, 2003 b: 7 (in part), figs. 19, 23, 52, 96; Fet et al., 2004: 24, figs. 1–4, 41–42 (Megisti listed in error; should be Anamur, NMW 0841); Fet et al., 2006a: 269–271, figs. 6–7; Fet & Soleglad, 2008: 256 (in part); Kaltsas et al., 2008: 227 (in part); Soleglad et al., 2009: 2 (in part), fig. 1.

Paraiurus nordmanni : Vachon & Kinzelbach, 1987: 99, 10 2, fig. 6 (in part); Sissom, 1988: 272.

Holotype: ♀ ( NMW 0847 ), TURKEY, Mersin Province: Mamure Kalesi, Anamur , under stones, 15 May 1969 (F. Ressl) . Paratypes (6 ♂, 10 ♀, 4 juv.): see list below.

Note: the type locality, Mamure Kalesi, is a famous waterfront Roman fortress, further strengthened by the Ottomans. It is located on Cape Anamur, the southernmost point of the Anatolian Peninsula.

Diagnosis. Small-sized scorpion with medium chelae, 26–36 mm in length, pectinal tooth counts 8–9 (8) male and 7–8 (7) female. Coloration variable, from yellow-orange with little patterns to dark brown and variegated patterns. Telson with bulbous vesicle, with long, wide curved aculeus; subaculear setal pair (SSP) located at vesicle/aculeus juncture. Seven and six inner denticles (ID) and eight and seven median denticle (MD) groups on the movable and fixed fingers, respectively. Fixed finger of chela longer than palm, trichobothrium it located on distal third of fixed finger. Metasomal segment V two times longer than wide.

Distribution. TURKEY: south (Antalya and Mersin Provinces). GREECE: Samos Island, Megisti Island. (see maps in Figs. 38 , 59 ).

Etymology. The species name is a patronym honoring our colleague Dr. Jürgen Gruber, the prominent opilionologist and a veteran curator of Arachnida in 3.Zoologische Abteilung, Naturhistorisches Museum Wien, Austria.

FEMALE. Description based on holotype female from Mamure Kalesi , Anamur, Turkey. Measurements of this holotype plus four paratypes are presented in Table 3. See Figure 60 for a dorsal view of the female holotype .

COLORATION. Basic color of carapace, tergites, metasoma, telson, pedipalps, and legs yellow-orange; carinae of pedipalps and metasoma, and cheliceral dentition a light red; chelal finger dentition and telson aculeus a dark reddish-brown; leg condyles red; sternites and pectines pale yellow. No variegated patterns present.

CARAPACE ( Fig. 61 ). Anterior edge with a small median indentation, equipped with four small irregularly placed setae (others may be broken off); interocular area somewhat rough with scattered granulation, though smooth around the immediate area of the median eyes; posterior lateral aspects covered with medium to large granules. Mediolateral ocular carinae present, extending to the lateral eyes; lateral eyes number two, the posterior eye a little larger. Median eyes and tubercle somewhat small, positioned considerably anterior of middle with the following length and width formulas: 118|420 and 54|338.

MESOSOMA ( Fig. 62 ). Tergites I–III lightly granulated, IV–VI with heavier granulation, primarily on posterior half; tergite VII covered with coarse granules with two pairs of granulate carinae. Sternites III–VI smooth and lustrous, VII covered with small granules; two pair of weak, vestigial carinae present on segment VII. Stigmata ( Fig. 62 ) are short sub-oval in shape, angled 45° in an anterointernal direction.

METASOMA ( Figs. 66–67 ). Segments I–IV: dorsal and dorsolateral carinae crenulate; dorsal (I–IV) and dorsolateral (I–III) carinae terminate with spine, smaller on dorsolateral; lateral carinae crenulate on I and obsolete on II–IV; ventrolateral and ventromedian carinae crenulate. Dorsolateral carinae of segment IV terminate at articulation condyle. Segment V: dorsolateral carinae crenulate; lateral carinae irregularly granulated for two-thirds of posterior aspect; ventrolateral and single ventromedian carinae crenulate; ventromedian carina not bifurcated, terminating in straight line ( Fig. 67 ). Intercarinal areas essentially smooth on I–III, scattered with granulation ventrally on IV–V. Metasoma essentially void of setation.

TELSON ( Figs. 66–67, 68 , paratype male from Anamur). Bulbous vesicle with long widely curved aculeus. Vesicle surface covered ventrally with medium sized granules, heavier basally; scattered setae located on ventral surface of vesicle; subaculear setal pair (SSP) located at vesicle/aculeus juncture.

PECTINES ( Fig. 64 ). Well-developed segments exhibiting length|width formula 400|160 (length taken at anterior lamellae|width at widest point including teeth). Sclerite construction complex, three anterior lamellae and 3/3 middle lamellae; fulcra of medium development. Teeth number 7/7. Sensory areas developed along most of tooth inner length on all teeth, including basal tooth. Delicate thin, white setae found on anterior lamellae and distal pectinal tooth. Basal piece large, with deep indentation along anterior edge, length|width formula 80|100.

PREPECTINAL PLATE ( Fig. 64 ). Conspicuous lustrous plate, approximately as wide as a gential operculum sclerite; length/width ratio 230/550.

GENITAL OPERCULUM ( Fig. 64 ). Sclerites much wider than long, fused medially. Posterior medial area contracts anteriorly. Genital papillae absent (see discussion on male below).

STERNUM ( Fig. 64 ). Type 2, posterior emargination present, well-defined convex lateral lobes, apex visible but not conspicuous; slightly wider than long, length| width formula 115|135; sclerite tapers anteriorly, posterior-width|anterior-width formula 135|93.

CHELICERAE ( Fig. 69 , paratype male from Anamur). Movable finger dorsal edge with one large subdistal (sd) denticle; ventral edge with three small crenulations (va), two pigmented, on the distal half, and one large pigmented accessory denticle at finger base; ventral edge with heavy setal brush covering well-developed serrula with over 20 contiguous tines, terminating just before distal tip. Ventral distal denticle (vd) considerably 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. 65 , 71 ). Moderately chelate, heavily carinate species, no scalloping on chelal fingers, thus not exhibiting sexual dimorphism in this structure. Femur: Dorsointernal carina serrate, dorsoexternal and ventrointerior carinae crenulate, ventroexternal rounded. Dorsal, ventral, and external surfaces rough, internal surface granulated. Patella: Dorsointernal, dorsoexternal, and ventrointernal carinae crenulate, ventroexternal carina granulate, exteromedian carina irregularly crenulate. Dorsal and ventral surfaces rough; external surface with exteromedian carina; internal surface smooth except for weakly developed doubled DPS and VPS. Chelal carinae: Complies with the “8- carinae configuration”. Digital (D1) carina strong, smooth to granulate; dorsosecondary (D3) present on basal half only, covered with large granules; dorsomarginal (D4) strong, round, continuous, with large granules; dorsointernal (D5) weak, sparsely granulated; ventroexternal (V1) strong and granulated, terminating at external condyle of movable finger; ventrointernal (V3) medium development, continuous to internal condyle, covered with small granules; external (E) strong, continuous, and granulated; internal (I) weak, rounded, not continuous, with small granules. Chelal finger dentition ( Figs. 65 ): median denticle (MD) row groups oblique and slightly imbricating, numbering 7 and 8; 6/6 and 7/7 internal denticles (ID) and 6/6 and 7/7 outer denticles (OD) on fixed and movable fingers, respectively. No accessory denticles present. Number of MD denticles on movable finger is 76. Trichobothrial patterns ( Fig. 71 ): Type C, orthobothriotaxic. See discussion of trichobothria differences between Calchas species which highlights the pattern of this female type.

LEGS ( Figs. 62 ). Both pedal spurs present on all legs; tibial spurs present on legs III and IV. Tarsus covered heavily with large socketed setae on ventral surface.

HEMISPERMATOPHORE. Unknown, not found in available males.

Male Paratype ( Figs. 68–70 , from Anamur). Adult males are approximately the same size as the female, 26.60–27.40 [3] for males, compared to 28.75–31.90 [2] for females. The genders do not exhibit any significant morphometric differences: for example, metasomal segments (L/W), where gender differences are commonly found, mean-value differences ranged only

0.8–2.4 %. Only the chelal palm showed mean-value differences exceeding ten percent; chela length / palm width 14.9 % and chela length / palm length 12 % (in both cases, based on the female with the longer chela). Pectinal tooth counts in the male exceed the female by one tooth, male 8–9 (8.44) [18], female 7–8 (7.29) [24] (see histograms in Fig. 36 ). Note that one of the males with nine pectinal teeth is from the Greek island of Samos, off the western coast of Turkey, somewhat removed from the primary range of C. gruberi (see maps in Figs. 38 , 59 ). The genital operculum of the male is dramatically different from that in the female ( Figs. 64, 70 ). The sclerites, subtriangular in shape, are as long as or longer than wide in the male whereas in the female, the sclerites are short and wide, more than twice as wide as long. Whereas the sclerites are fused medially in the female, they are separated their entire length in the male, exposing significantly developed genital papillae. The prepectinal plate, so conspicuous in the female, is absent in males ( Figs. 64, 70 ). Figures 72–73 show dorsal and ventral views of two dark colored C. gruberi female specimens: one from Antalya (dry, Fig. 72 ) another from Akseki (live specimen, Fig. 73 ) as well as the locality area where the latter specimen was collected; and Figures 74–75 show dorsal and ventral views of light colored female and male specimens from Kemer, Turkey.

Type material. TURKEY. Holotype: 1 ♀ ( NMW 0847), Mersin Province: Mamure Kalesi, Anamur, 36.078°N, 32.8 34°E, under stones, 15 May 1969, F. Ressl leg. Paratypes (6 ♂, 10 ♀, 4 juv.): Mersin Province: 2 ♀ ( NMW 0847), Mamure Kalesi, Anamur, under stones, 15 May 1969, F. Ressl leg.; 1 ♂, 1 ♂ juv. (partially dismembered and used for SEM) ( NMW 0841), 15 km by road W of Anamur, 36.078°N, 32.817°E, 18 May 1969, G. Pretzmann leg. Antalya Province: 1 ♂, 1 ♀ ( NMW 0848), Belkıs (Aspendos), 36.939°N, 31.172°E, near ruins, under stones, 16 May 1965, F. Ressl leg.; 1 ♂, 1 ♀ ( NMW 0838/ VF; specimens partially dismembered and used for SEM), mountains N of Antalya, 36.913°N, 30.69°E, 19 May 1969, J. Koller & F. Ressl leg. (together with Iurus ); 1 juv. (damaged) ( MNHN RS 7024), Olympos, 36.403°N, 30.474°E, 30 July 1974; 1 ♀, near Antalya, 1994 ( FKCP); 2 ♀, 1 juv. ( FKCP), 12 km S of Akseki, 11–12 May 2006, F. Kovařík leg.; 3 ♂, 2 ♀, 1 juv. ( FKCP), Kemer, May 2009; 1 ♀ ( NMM 0250), 15 km NE of Kumluca, 36.546°N, 30.283°E, 12 August 1972, R. Kinzelbach leg. ( Kinzelbach, 1982: 58; the same specimen was reported as “ 20 km N of Kumluca, 12 August 1972 ” by Kinzelbach, 1980).

Note: see Pretzmann (1972) for a detailed itinerary and map of the NMW expedition to Turkey in 1969.

Other material examined: GREECE, 1 ♀ juv. (NMHC 81.1.7.9/ VF; specimen dismembered and partially used for DNA studies), Megisti (=Kastelorizo) Island, 36.149°N, 29.594°E, I. Stathi leg. ; 1 ♂ ( FMNH), Samos Island , 37.757°N, 26.977°E, Mt. Spiliani, 2 km N Pithagorion, S slope, 23 April 1979, A. Riedel leg.

Note: we do not include Greek island specimens, identified here as C. gruberi , in the type series of C. gruberi ; a further detailed study of Samos and Megisti populations is warranted.

Other specimens/localities (material not examined): TURKEY, Antalya Province: 1 specimen ( NMW 0845 ), 4 6 km by road N of Antalya, 865 m, mountain pass, 21

May 1969, G. Pretzmann leg. GREECE, Megisti (=Kastelorizo) Island (Stathi & Mylonas, 2001: “big population”).

Biogeography

Kaltsas et al. (2008: 238) mentioned that “the biogeography of the monotypic genus Calchas still remains a mystery to scientists.” We are glad to offer here a window to this mystery.

Figures 38 , 39 , 42 , and 59 present the current ranges of three species described and discussed in the present paper, based on all known records. The observed disjunction follows modern ecological barriers. The ecologically diverse territory of the Anatolian Peninsula (historical Asia Minor) is divided into a number of biogeographic provinces or regions. Three disjunct species of Calchas tend to fall under three separate regions as described for the Anatolian scorpiofauna by Crucitti & Cicuzza (2001).

Already the range of Calchas nordmanni depicted by Kinzelbach (1985) and Vachon & Kinzelbach (1987: 99, fig. 6) reflected three widely disjunct populations; additional records expanded boundaries of their ranges, but the disjunction remains. The geographic range of Calchas nordmanni Birula, 1899 (sensu stricto) is limited to Çoruh River valley (Black Sea watershed) in northeastern Turkey, i.e. mainly in the very east of Black Sea Region; it does not cross the mountain ranges stretching between Erzurum and Kars. The geographic range of Calchas birulai sp. nov. covers a large area in southeastern Turkey, across Gaziantep, Adıyaman, Malatya, Sanlıurfa, Diyarbakır, Mardin, and Siirt Provinces, and reaches to northern Iraq. This range covers the entire East Anatolian Region as defined by Crucitti & Cicuzza (2001). C. nordmanni and C. birulai are clearly separated by the high mountain ranges of Güneydoğu Toroslar, Hakkari, Munzur, and Bingöl. The geographic range of Calchas gruberi sp. nov. at this moment is limited to a small portion of southern Turkey along the Mediterranean coast, from Kumluca to Anamur, within the Mediterranean Region. On the east, it is well separated by the Taurus Mountains from the range of C. birulai . The populations from Greek islands (Samos and Megisti) are currently also assigned to C. gruberi but should be a subject of a more detailed study, with a possible clarification of Samos locality.

The disjunct distribution of Calchas , therefore, appears to be a textbook illustration of allopatric speciation, with three species that seem to occupy different ecological regions. Although it is hard to speculate about the timing of this disjunction, it does not have to be very recent (i.e. Pleistocene). Modern evolution of scorpiofauna in the Aegean-Anatolian area, at species level and below, was traced in the recent DNA-based studies of Greek researchers (Parmakelis et al., 2006a, 2006b). It is believed to have been influenced by tectonic fragmentation for both a widespread Mesobuthus gibbosus ( Buthidae ) and a more localized Iurus dufoureius ( Iuridae ). It is generally accepted that the Aegean-Anatolian area was represented by a single landmass (Agäis) in the Upper to Middle Miocene (23–12 Mya), with the subsequent fragmentation due to tectonic events of 12–5 Mya (Stathi & Mylonas, 2001).

Hrbek et al. (2004) in their detailed DNA-based work on historical biogeography of Central Anatolian fish, conclude that “geologically complex areas of the Near East contain many phylogenetically deeply divergent lineages, some showing reproductive isolation, that are otherwise morphologically difficult to differentiate. We suggest this is most likely due to fragmentation of previously contiguous areas into a parapatric series of ecologically equivalent regions.” The case of Calchas seems to be more straightforward: there are three distinct morphospecies in three ecologically different regions.

Figures

Figure 3: Carapace and close-up of median and left lateral eyes in Calchas gruberi, sp. nov., female, Antalya, Turkey. Lateral eyes and mediolateral ocular carina indicated.

Figure 5: Prepectinal plate in Calchas. Top. Close-up of prepectinal plate of Calchas birulai, sp. nov., female, Nemrut Dağı, Turkey. Bottom. Detached prepectinal plate of C. gruberi, sp. nov., female, Antalya, Turkey, and close-up showing surface of the plate.

Figure 6: Pecten of Calchas gruberi, sp. nov., female, Antalya, Turkey, showing close-up of peg sensilla.

Figure 7: Fourth left lung stigma in Calchas gruberi, sp. nov., female, Antalya, Turkey.

Figure 9: Cheliceral movable finger, ventral view, showing well developed serrula. Top. Calchas birulai, sp. nov., female, Nemrut Dağı, Turkey. Bottom. C. gruberi, sp. nov., juvenile male, Antalya, Turkey.

Figure 10: Close-up of trichobothrium areola and bristle base, Calchas gruberi, sp. nov. Top and Bottom Left. Adult female, Antalya, Turkey, showing patellar trichobothrium with bristle partially raised from areola. Bottom Right. Adult female, Antalya, Turkey, showing normal setting of bristle.

Figure 11: Pedipalp right patella showing Dorsal (DPS) and Ventral (VPS) Patellar Spurs. Calchas gruberi, sp. nov. Top. Dorsal view (left), juvenile male, Antalya, Turkey, showing doubled DPS and accompanying setae. Internal view (right, dorsal edge indicated by trichobothrium i), female, Antalya, Turkey, showing doubled DPS and VPS plus accompanying setal areolae. Bottom. Close-up of internal base of patella showing spurs and setal areolae (s).

Figure 14: Constellation array in Calchas gruberi, sp. nov., showing 15 sensilla. Left. Juvenile male, Antalya, Turkey. Right. Adult male, Antalya, Turkey.

Figure 17: Leg tarsus spination in Calchas gruberi, sp. nov., ventral surface, juvenile male, Antalya, Turkey.

Figure 18: Leg tarsus setation in Calchas gruberi, sp. nov., ventral surface. Left. Adult male, Antalya, Turkey. Right. Adult female, Antalya, Turkey, pedal spurs indicated.

Figure 19: Distal aspect of leg basitarsus in Calchas gruberi, sp. nov., Antalya, Turkey. Left. Adult male, showing “rosette” structure with seven denticles. Right-Bottom. Juvenile male, “rosette” with five denticles. Right-Top. Slit sensilla (indicated by ss) and “rosette”.

Figure 20: Telson, lateral view, of Calchas species. Top. C. birulai, sp. nov., female, Nemrut Dağı, Turkey. Bottom. C. gruberi, sp. nov., female, Antalya, Turkey.

Figure 21: Close-up of the telson of Calchas showing two different placements of the subaculear setal pair (SSP) with respect to the vesicle/aculeus juncture. Top. Calchas birulai, sp. nov., female, Nemrut Dağı, Turkey, SSP placed on aculeus base, distal of the juncture. Bottom. C. gruberi, sp. nov., female, Anamur, Turkey, SSP placed at juncture. Ves/Acu = vesicle/aculeus juncture, VDSP = ventral distal setal pair, DDSP = dorsal distal setal pair.

Figure 25: Chelal movable finger of Calchas species showing dentition. Note differences in the number of inner denticles (ID) and median denticle (MD) groups between the species: C. nordmanni, female, Tortum, Turkey, and C. birulai, sp. nov., male holotype, Mardin, Turkey, with six ID and seven MD groups as compared to C. gruberi, sp. nov., female holotype, Mamure Kalesi, Anamur, Turkey, with seven ID and eight MD groups.

Figures 26–31: Subaculear setal pair (SSP) in genus Calchas, lateral view unless otherwise noted (note, only the SSP, VDSP, and DDSP setal pairs are shown). Conspicuous in these illustrations is the aculear placement of the SSP in species C. nordmanni and C. birulai. In C. gruberi, the SSP is located at the vesicle/aculeus juncture, common to most scorpions. 26. C. gruberi, sp. nov., female, Mamure Kalesi, Anamur, Turkey. 27. C. birulai, sp. nov., male holotype, Mardin, Turkey. 28. C. nordmanni, female, Tortum, Turkey. 29. C. nordmanni, male, Turkey. 30. C. nordmanni, female, Tortum, Turkey, viewed from the aculeus. 31. C. nordmanni, male, Tortum, Turkey, ventral view. VDSP = ventral distal setal pair, DDSP = dorsal distal setal pair.

Figure 32: Relative position of chelal trichobothrium dst in Calchas with respect to trichobothria dt and db. Trichobothrium dst mean position indicated by open circle based on 30 samples (other fixed finger trichobothria not shown). Method of measurement: dt|dst = distance between dt and dst, dst|db = distance between dst and db. Histogram of morphometric ratio shows mean (vertical bar), minimum, maximum, and standard error (black rectangle) ranges. We see two significant dst positional differences in these species: In C. birulai, trichobothrium dst is approximately at midpoint between dt and db whereas in C. nordmanni and C. gruberi, dst is considerably distal of midpoint, most exaggerated in C. gruberi.

Figure 33: Relative position of chelal trichobothrium it in Calchas based on its location with respect to the fixed finger length. Trichobothrium it mean position indicated by open circle based on 30 samples. Method of measurement: ff_LENG = fixed finger length, it_POS = it position. Histogram of morphometric ratio shows mean (vertical bar), minimum, maximum, and standard error (black rectangle) ranges. We see two significant it positional differences in these species: In C. birulai and C. nordmanni, trichobothrium it is approximately at finger midpoint or proximal, most exaggerated in C. birulai, whereas in C. gruberi trichobothrium it is considerably distal of finger midpoint.

Figure 34: External surface of the chela highlighting trichobothria positional differences in Calchas species. Four specific areas (A–D) are delineated showing major trichobothrial location differences between species C. gruberi as compared to C. birulai and C. nordmanni. A: in C. gruberi trichobothrium eb is placed considerably more distally on finger than in the other two species. B: in C. gruberi the pattern formed by trichobothria Et3–Et5–db essentially forms a straight line with db more distally placed, situated parallel to the finger juncture; in addition, trichobothrium Et4 is located considerably dorsally of Et3. C & D: in C. gruberi trichobothrium Dt and Db are positioned considerably more distally than in the other two species.

Figure 35: Sternopectinal area in Calchas species showing the genital operculum, prepectinal plate (in female), and genital papillae (in male). Note, in the male, the large long subtriangular shaped genital operculum with separated sclerites exposing significant genital papillae, as compared to the female, we see a narrow operculum with fused sclerites. Top-Left. C. nordmanni, female, Tortum, Turkey. Top-Right. C. birulai, sp. nov., female, Mardin, Turkey. Bottom-Left. C. gruberi, sp. nov., female, 12 km S. Akseki, Turkey. Bottom-Right. C. nordmanni, male, Turkey.

Figure 36: Distribution of pectinal tooth counts in genus Calchas based on 82 specimens (32 males and 50 females). Number of teeth on bottom, number of samples on top. Data is based on material examined and Birula (1900, 1905, 1911, and 1912).

Figure 37: Key morphometric ratio differences in species of Calchas based on 28 sets of measurements. Top. Metasomal segment V proportions (length/width) show that metasomal segment V is considerably more slender in C. nordmanni than in the other two species, exhibiting considerable standard error as well as absolute range separation. Differences in mean values between C. nordmanni and the other two species, male|female, is as follows: C. birulai = 24.2 %| 27.1 %, C. gruberi = 27.4 %|23.8 %. Bottom. Telson proportions (telson length / vesicle length) show that species C. nordmanni and C. birulai have a longer vesicle than that seen in C. gruberi. The mean value differences between C. gruberi and the other two species (combined) is 13.8 % male and 16.2 % female. Also of interest in both sets of ratios, we see that the differences between the genders across species are negligible.

Figure 38: Distribution of all reported instances of genus Calchas. Open icons depict localities examined in this study; closed icons depict remainder of published localities; icons with enclosed dot indicate type locality. Note C. birulai localities from northern Iraq and possibly Syria (? = not confirmed).

Figure 39: Large-scale range of Calchas nordmanni, white circles. See general Calchas map in Fig. 38.

Figure 42: Large-scale range of confirmed localities of Calchas birulai, sp. nov. (open circles). See general Calchas map in Fig. 38.

Figure 59: Large-scale range of Calchas gruberi, sp. nov. (open circles). See general Calchas map in Fig. 38.

Figure 60: Calchas gruberi, sp. nov., female holotype (NMW), dorsal view (32 mm), Mamure Kalesi, Anamur, Turkey.

Figures 61–70: Calchas gruberi, sp. nov. 61–67. Female holotype (NMW), Mamure Kalesi, Anamur, Turkey. 61. Carapace showing close-up of lateral eyes. 62. Leg IV (left, ventral view), showing the tibial spur. 63. Stigma III. 64. Sternum, genital operculum, prepectinal plate, and pectines. 65. Chelal fixed and movable finger dentition. 66. Telson and metasomal segments IV–V, lateral view. 67. Metasomal segment V and telson, ventral view. 68–70. Male paratype (NMW), Anamur, Turkey. 68. Telson, ventral view. 69. Chelicerae, vental and dorsal views. 70. Sternum, genital operculum, and pectines (note lack of prepectinal plate conspicuous in female).

Figure 71: Trichobothrial pattern of Calchas gruberi, sp. nov., female holotype (NMW), Mamure Kalesi, Anamur, Turkey.

Figure 72: Calchas gruberi, sp. nov., dorsal and ventral views. Adult female paratype (FKCP), Antalya, Turkey.

Figure 73: Adult female paratype (FKCP) of Calchas gruberi, sp. nov., (top) and collection locality, 12 km S. Akseki, Turkey (bottom).

Figure 74: Calchas gruberi, sp. nov., dorsal and ventral views. Adult paratype female (FKCP) (34 mm), Kemer, Turkey.

Figure 75: Calchas gruberi, sp. nov., dorsal and ventral views. Adult paratype male (FKCP) (36 mm), Kemer, Turkey.