Scorpiops kovariki, Tang & Ouyang & Liu & Šťáhlavský, 2024

Scorpiops kovariki sp. n.

( Figures 39–72 View Figures 39–40 View Figures 41–44 View Figures 45–50 View Figures 51–61 View Figures 62–71 View Figures 72–73 ; Tables 1–2) http://zoobank.org/urn:lsid:zoobank.org:act:AC035EC1-

C699-4C7E-B78D-90D93E591779

TYPE LOCALITY AND TYPE DEPOSITORY. CHINA, Tibet Autonomous Region, Nyingchi (or Linzhi, Nyingtri) prefecture-level City, Zayü (or Chayu, Zayul) County , 28°33'27.2''N 98°17'36.5''E (28.5575607°N 98.29347779°E), 2492 m a. s. l.; VT GoogleMaps .

TYPE MATERIAL. CHINA, Tibet Autonomous Region, Nyingchi City , Zayü County, 28°33'27.2''N 98°17'36.5''E, 2492 m a. s. l., 15 July 2023, 1♀ (holotype), leg. Tongtong. GoogleMaps

ETYMOLOGY. The specific epithet is a patronym in honor of Czech scorpiologist František Kovařík, a major contributor to the taxonomy of Scorpiopidae , who has also been supporting the first author in her past publications. The nexus between the concrete species and and its nomenclature was established based on its evident resemblance to the congener described by F. Kovařík, S. jendeki . Chinese equivalent: ※氏ĀH (roughly as “Kovařík’s resemblant scorpion” in English; see Tang (2022a) for the rules of designation).

DIAGNOSIS (♀). Total length ca. 33 mm for female. Base color uniformly brownish red to blackish brown; chela with fingers and carinae blackish and manus reddish, telson brownish red, genital plate light brown, pectines light yellow. Cuticular surface (somatic) matte and sparsely but coarsely granular; pedipalp chela manus lustrous. Carapace with anteromedian notch semicircular; ocular tubercle distinct within ocular islet, elliptical in profile; ocular subislets weakly granular and planar; interocular sulcus absent; superciliary carinae smooth. Cheliceral fixed finger with basal and median denticles forming into a bicuspid trunk. PTC 4 in female; fulcra absent; pectines with two marginal and one middle lamellae present (P3); sternites III– VI lustrous, VII granular with four distinct carinae. Pedipalp patella with weak prolateral apophysis; 16–17 external and 5–6 ventral trichobothria. Chela with 4 V series trichobothria; Eb 3 located in proximal half of manus between Dt and Db (type D); Ch-L/W ca. 2.4 in female. Dentate margin of movable fingers not undulate in female, with 9–10 OD, 49–51 MD, 6–7 IAD and 6 ID. Telotarsi of leg I– VI with ca. 6–12 stout median ventral spinules nearly in a single row. Metasoma I – V with 10-8-8-8-7 carinae. Telson moderately short and moderately bulbous, densely covered with fine granules; T-L/D ca. 3.05 in female; annular ring developed .

DESCRIPTION (♀).Overall habitus ( Figs. 39–40 View Figures 39–40 ), prosoma, mesosoma, metasoma and telson, chelicera and legs were photographed under white light. Additional UV fluorescence imaging was applied for prosoma and mesosoma.

Prosoma and mesosoma ( Figs. 41–50 View Figures 41–44 View Figures 45–50 , 62–63 View Figures 62–71 ). Prosoma: Carapace with 3 pairs of lateral ocelli (two larger, one smaller); median ocular tubercle distinct within ocular islet, elliptical in profile; subislets weakly granular, approximately planar; interocular sulcus absent; superciliary carinae smooth. Dorsomedian part of the carapace (ca. 1/2 of total area) planar, with anterolateral boundaries demarcated by coarse granules; lateral surfaces slanting downwards. Entire carapace moderately covered with fine to median-sized granules, weakly lustrous at non-granular regions; larger and somewhat interfused granules concentrated anteriorly at edges flanking anteromedian sulcus, and above and posterior to lateral ocelli; distinct carinae absent. Anterior margin of carapace with a shallow, semicircular median notch leading to a shallow, smooth anteromedian sulcus; circumocular and posteromedian sulci smooth and matte; lateral surfaces with a pair of indistinct central lateral sulci and prominent posterior lateral sulci, both smooth and matte; posterior marginal sulcus deep. Chelicerae with dorsal surface smooth and ventral surface densely setose on 2/3 area; all denticles of cheliceral fingers relatively blunt but might had been caused by abrasion; basal and median denticles on fixed finger conjoined on a common trunk. Mesosoma: All tergites (except for pretergites which are moderately lustrous) covered by a layer of matte, greyish mask, contrasting with the smooth granules that reflect the base color; all tergites covered with moderate to large, somewhat flattened granules, becoming larger closer to posterior margins, with one distinct median carina (less prominent on I–II while concealed by random granules on VII). Sternites III–IV highly lustrous, with sparse macrosetae and fluorescent microsetae as well as two axisymmetric, curved furrows; sternite VII strongly granulated with four granular carinae; respiratory spiracles oval. Genital operculum divided into two halves, without genital papillae at the base. Pectines with two marginal and one middle lamellae present, distal furrow on the right pectine not fully delineated; pectine teeth number 4/4; area of peg sensilla concentrated along the external edge of each pectinal tooth, minute in size; fulcra absent; fluorescent microsetae present.

Metasoma and telson ( Figs. 64–67 View Figures 62–71 ). Metasoma sparsely hirsute and weakly granulated; carinae formed by larger granules. Metasoma I with 10 carinae, II–IV with 8 carinae, and V with 7 carinae; median lateral carinae of metasoma II– IV decomposed into random granules and do not form distinct carinae; all carinae serrated, especially the ventrolateral and ventromedian carinae on metasoma V. Anal arch armed with sharp granules. Telson lustrous, very sparsely hirsute, moderately short and bulbous; all surfaces nearly smooth; prominent annular ring developed. Vesicle with lateral surface furnished by one longitudinal sulci close to dorsal surface, and ventral surface with two parallel longitudinal sulci; lateral sulci deeper than ventral sulci. Aculeus smooth, short, weakly curved (tip broken).

Pedipalps ( Figs. 51–61 View Figures 51–61 ). Pedipalps sparsely hirsute, intercarinal surfaces scattered with generally sparse granules and patches of lattice microstructures. Patella with 17/16 external (4/3 et, 4 est, 2 em, 2 esb, 5 eb; one esb closer to em series) and 5/6 ventral trichobothria. Chela with 4 V series trichobothria; trichobothrium Eb 3 located in proximal half of manus between trichobothria Dt and Db. Femur with 6 highly granular carinae and a coarsely granulated dorsal surface; retromedian and retroventral carinae incomplete, respectively deconstructed into discrete granules proximally and distally; granules larger on promedian, retrodorsal and proventral carinae. Patella with 5 lustrous, granular or costate carinae and weakly granulated intercarinal surfaces; prodorsal carina granular; proventral carina costate-granular; retrodorsal, retromedian and retroventral carinae costate; granules do form a distinct reticulate configuration on ventral surface; prolateral surface with a triangular apophysis not armed by prominent spiniform granules. Manus sparsely adorned with small to moderate intercarinal granules, which form a reticulate configuration with large hollows on dorsal surface; dorsal secondary carina indicated, gradually diffusing to dispersed granules; subdigital and ventrointernal carinae obsolete; dorsal internal and dorsal marginal carinae evidently separated, with dorsal internal carina indicated by large but flat granules; dorsal marginal, digital, external secondary, ventroexternal, ventromedian and interomedian carinae strongly costate and lustrous. Dentate margin of movable fingers not undulate (proximal lobe absent), adorned by 10/9 OD, 51/49 MD, 6/7 IAD (same size as OD and ID) and 6/6 ID; IAD series creates an incomplete second row along MD series, paired with OD distally, which divide the entire MD series into several subrows, each ended in a slightly enlarged MD.

Legs ( Figs. 68–71 View Figures 62–71 ). Tibia and tarsomeres of legs with several macrosetae not arranged into bristle combs. Basitarsi of legs I–II with two rows of dense, short spinules, becoming vestigial on legs III and IV; a pair of pedal spurs present on the distal margin of all basitarsi (one lost due to injury on IV). Telotarsi of legs I–IV with a row of short, stout ventromedian spinules (ca. 6–12 in number). Ungues moderately short, stout and curved. Carinae on femur and patella distinct, granular to costate-granular.

Measurements. See Table 1.

AFFINITIES. S. kovariki sp. n. can be confidently differentiated from all its Tibetan congeners, even those dark colored species with stout chelae, including S. atomatus , S. ingens , S. langxian Qi et al., 2005 , S. lourencoi , S. margerisonae and S. tibetanus , by a completely different pectine morphology (short and light yellow, comprising only four teeth; Figs. 46, 49 View Figures 45–50 ), alongside a plethora of other pronounced distinctions. Intriguingly, this species showcases a remarkable resemblance with S. jendeki from Yunnan. S. jendeki is the most widespread Scorpiops species in Yunnan with its northmost occurrence in Gongshan County, Nujiang Lisu Autonomous Prefecture (Tang, 2023: table 2; adjacent with the type locality of S. kovariki sp. n.); recent investigation also revealed its occurrence in Kachang, Yingjiang County (pers. comm., Tongtong). Both species are small in size, with short, robust pedipalps bearing movable fingers devoid of proximal lobes, as well as relatively short pectines and sparsely granulated tergites. Detailed photographs of S. jendeki can be referred from Tang (2023: figs. 97–100, 137, 146, 155, 164, 173–174, 196–199). Cursory examination on the holotype female detected a pair of proportionally small median ocelli and lustrous pedipalp manus and telson in contrast to S. jendeki , which led us to suspect its identity. To eliminate the possibility that the glossy appearance of the chela and telson observed in the holotype of S. kovariki sp. n. was not unique to this species, additional specimens of S. jendeki previously collected from Yunnan were reexamined (partially listed in Tang, 2023: 2). It was ascertained that, at least on the basis of the 21 specimens examined, S. jendeki is not characterized by a smoothness on chelae. Comparative photos of those specimens, with the holotype of S. kovariki sp. n., are included in the supplementary PDF file.

Following a scrutiny upon the holotype of S. kovariki sp. n., several morphological distinctions from S. jendeki , confined to the comparison between adult females, are proposed: (1) carapacial anteromedian notch proportionally shallow and semicircular (vs. typically deep and dome-shaped in S. jendeki ; cf. Fig. 48 View Figures 45–50 vs. Tang, 2023: fig. 146); (2) ocular islet proportionally longer with respect to the median ocelli but smaller with respect to the carapace, with an elliptical ocular tubercle (vs. shorter and larger with a rounded ocular tubercle in S. jendeki ; cf. Fig. 48 View Figures 45–50 vs. Tang, 2023: fig. 146); (3) relatively small median ocelli with respect to the carapace (cf. Fig. 48 View Figures 45–50 vs. Tang, 2023: fig. 146); (4) carapace somewhat more flattened and less triangular (cf. Fig. 48 View Figures 45–50 vs. Tang, 2023: fig. 146); (5) pedipalp chela manus lustrous (vs. matte in S. jendeki ; cf. Figs. 72–73 View Figures 72–73 ); (6) dorsal internal carinae of chela manus with fewer and weaker granules (vs. typically prominent and strongly spiniform in S. jendeki ; cf. Fig. 51 View Figures 51–61 vs. Tang, 2023: fig. 99); (7) region along the dorsal marginal on dorsal surface of manus not adorned by darkened granules (vs. densely adorned by darkened granules in S. jendeki ; cf. Fig. 51 View Figures 51–61 vs. Tang, 2023: fig. 99); (8) dorsal marginal, external secondary and ventromedian carinae of chela manus highly costate (vs. costate-granular to granular in S. jendeki ; cf. Figs. 51–52 View Figures 51–61 vs. Tang, 2023: figs. 99–100); (9) retrodorsal carinae of pedipalp patella costate (vs. granular to costate-granular in S. jendeki ; cf. Fig. 54 View Figures 51–61 vs. Tang, 2023: fig. 174); (10) mesosoma with a distinct median carina and denser and coarser granules exhibiting stronger between-granule size contrast (vs. median carina less discernable, with granules being sparse and ambiguous; cf. Fig 42 View Figures 41–44 vs. Tang, 2023: fig. 164, and Figs. 72– 73 View Figures 72–73 ); (11) telson slightly more lustrous (cf. Figs. 72–73 View Figures 72–73 ); (12) pectines with less dense fluorescent microsetae (cf. Fig. 49 View Figures 45–50 vs. Tang, 2023: fig. 199).

With reference to Kovařík et al. (2020: 129, table 9), the only reliable quantitative disparity between the two species appears to lie in the count of IAD (6–7 in S. kovariki sp. n. vs. 10 in S. jendeki ). 18 specimens of S. jendeki were counted for their finger denticles, with two of which represented by a single finger respectively (number of fingers n = 34, including the one examined in Kovařík et al. (2020)). While their illustration indeed displayed an evident count of 10 IAD (Kovařík et al., 2020: 18, fig. 115), a noticeable level of intricacy was observed in the materials examined herein. Specifically, additional denticles may be present at the proximo-internal side flanking the main MD series. They represent a category of complex denticles that had been considered as a part of the MD series in Tang (2022b, 2023). There may be no explicit hiatus between those denticles and the preceding definitive IAD as shown in Kovařík et al. (2020). Unbiasedly incorporating all these denticles into the IAD series would result in a considerable degree of variability in the count of IAD for S. jendeki (ca. 9–18, mean ± standard deviation = 13.35±1.98). On the other hand, a biased count rendered a total range of ca. 7–10 (mean ± standard deviation = 8.44±0.66), with the determination based on the subjectively perceived interdental distance and whether the pertinent denticle is paired with an OD on the opposing side. Given the potential ambiguity in distinguishing between IAD and MD, their sum (mean ± standard deviation = 65.74±2.73) was therefore calculated for comparison with that of S. kovariki sp. n. The examined 34 movable fingers of S. jendeki yielded a biased MD (defined as sum minus biased IAD) of 57.29±2.74 (mean ± standard deviation; range 50–62). As an additional reference, the IAD and MD counts of S. kovariki sp. n. was further compared the biased IAD and MD counts of S. jendeki . A simple linear regression test was performed to detect the possible correlation between left and right finger denticle counts of the 16 examined S. jendeki specimens: (1) sum comparison, R 2 = 0.3655 (Pearson’s r = 0.6046), p = 0.01311; (2) biased IAD comparison, R 2 = 0.04368 (Pearson’s r = -0.209), p = 0.4373; (3) biased MD comparison, R 2 = 0.4607 (Pearson’s r = 0.6788), p = 0.00384. Since one of the correlations was weak, we first treated these 32 fingers as independent and included the two solitary fingers (hence n = 34 in total). Those data were compared against the data (two fingers) of S. kovariki sp. n., which yielded a statistically significant difference (two-sample Mann-Whitney U test): (1) sum comparison, p = 0.02018; (2) biased IAD comparison, p = 0.01165; (3) biased MD comparison, p = 0.02398. Alternatively, we compared the means between the two species (for S. jendeki , the data of two solitary fingers represented their own mean, hence n = 18). Shapiro-Wilk test was first taken to confirm normality in the count of S. jendeki : (1) sum data confirmed normal: W = 0.9833, p = 0.9774; (2) biased IAD data confirmed non-normal (symmetrical but leptokurtic): W = 0.8687, p = 0.01696; (3) biased MD data confirmed normal: W = 0.9812, p = 0.9613. Consequently, a one-sample t-test was used for the sum and biased MD comparison, which yielded p -values of 1.088e- 14 and 3.197e- 10 respectively. An outlier detection for the dataset comprising the mean IAD of all specimens of both species (dataset n = 19) was conducted based on Tukey’s fences (k =1.5). Data 6.5 (mean IAD of S. kovariki sp. n.) and 10 (of one S. jendeki , which was a solitary value, not a mean) were identified as outliers. When the two solitary counts of two S. jendeki were discarded (dataset n = 17), the single outlier was S. kovariki sp. n. If the data of S. jendeki (i.e., 16 mean and 2 solitary) were compared with the two raw data of S. kovariki sp. n. by two-sample Mann-Whitney U test, a significant p -value of 0.02007 was obtained (discarding the two solitary values yielded a lower p -value, 0.0192). In all cases, the two counts were found significantly different between the two species. However, these results must be taken with caution as the sample size for S. kovariki sp. n. is exceedingly small, and our analysis was based on the assumption that the data of that species represent its population mean.

Regrettably, our molecular analysis revealed a low 16S genetic distance between the two species (0.02, compared to the within-species variation of S. longimanus , 0–0.05, as well as the between-species distance of other species; Table 3). However, a low genetic distance is not always a negation of heterospecificity, and phenotypes can exhibit high rates of speciation even when genetic divergence is relatively low between species (Tarvin et al., 2017). Low genetic divergence often favors the assumption that the observed phenotypic difference may be due to intraspecific variation. Empirically, intraspecific phenotypic variation in the genus Scorpiops is typically limited to a few diagnostic characters (e.g., total length, trichobothria count, PTC, and ratiometrics of pedipalp chela, finger lobe and telson). Another variable meristic is the pedipalp finger denticle count, where our new species putatively represents an outlier from the variation range of S. jendeki . While quantitative characters display obvious variations in this genus, several qualitative characters remain very stable and conserved where their phenotypic plasticity is unprecedent in Scorpiops (i.e., no within-species polymorphism), particularly the carination (costate or granular). Those characters were leveraged in the previous species described by the first author, which is now further validated via molecular data ( Table 3). Our new species exhibited similar distinctions from its closest congener. As a result, we reckon that the holotype female does not represent a new record of S. jendeki in Xizang, but rather a distinct yet closely related species. The most salient characters that distinguish S. kovariki sp. n. from S. jendeki are the lustrous pedipalp chela and costate carination in the former (see supplementary PDF for more photos).

BEHAVIORAL NOTES. The holotype of S. kovariki sp. n. had given birth to a brood of scorplings, currently reared by the second author. The first author’s observation unveiled a temperament characterized by considerable sensitivity, reminiscent of that exhibited by the holotype S. matthewi sp. n. However, the holotype S. kovariki sp. n. displayed a more bellicose demeanor, often resorting to attack the perceived predatory stimulus using its pedipalps and telson. In contrast to adult females of S. jendeki , where individuals typically adopted a motionless stance, S. kovariki sp. n. tended to exhibit a preference for a fight-or-flight response, with cataleptic behavior being rarely observed. Such ethological discrepancy might be species-dependent.

DISTRIBUTION. Known only from the type locality, but might extend to Yunnan Province.