3♂2♀ (KS), Japan • Mount Teraji-yama, Tobikoshi-shindô, on the border between Gifu Prefecture, Hida-shi, Kamioka-cho, Utsubo and Toyama Prefecture, Toyama-shi, Arimine, 4-X-2008, Hiroshi Nishida leg.

Figures 6–9. 

Right lateral (a), ventral (b), dorsal (c), left lateral (d), and posterodorsal (e) views of the endophallus of Nebria spp. 6. N. niohozana from Mount Teraji-yama; 7. N. furcata from Kuraiyama-tôge Pass; 8. N. dichotoma holotype from Mount Takanosu-yama; 9. N. dichotoma from near Hiraya-tôge Pass. Abbreviations: da, dorsoapical lobe; db, dorsobasal lobe; dm, dorsomedian lobe; gp, gonopore protrusion; la, left lateroapical lobe; lb, left laterobasal lobe; ra, right lateroapical lobe; rb, right laterobasal lobe; vb, ventrobasal swelling. Asterisk indicates that the gonopore protrusion or lobes are not fully everted.

In the Japanese Alps, this species had been recorded only from Mount Shirouma-dake, near the northern end of the mountain range (Sasakawa 2020). The present collection record indicates that N. niohozana is more widely distributed in the northern part of the Japanese Alps than previously recognized. No distinct differences in the endophallus shape were found between the Shirouma-dake specimen and the present specimen.

1♂1♀ (KS), Japan • Gifu Prefecture, Gero-shi, Hagiwaracho, Yamanokuchi, Kuraiyama-tôge Pass, alt. 1100 m, 2-VII-2011, Kazue Ito leg.; 5♀ (KS), Japan • Gifu Prefecture, Gero-shi, Hagiwaracho, Yamanokuchi, Mount Kurai-yama, near the upper reaches of Araragi-ko Lake (Kuguno Disaster Prevention Dam), alt. 1100 m, 18-VI-2009, Hiroshi Nishida leg.

This is the easternmost collection record of this species. The collection site is by definition not included in the Japanese Alps, but it is located in a mountain range adjacent to the northern part of the Japanese Alps (Fig. 1). The endophallus of the examined specimen was more similar to that of specimens from the distant type locality than to specimens from the nearest locality, Hida-shi (e.g., shapes of the dorsobasal lobe and the basal protrusion of the dorsoapical lobe), although the comparison was based on a small number of specimens from each locality (one or two males).

1♂1♀ (KS), Japan • Aichi Prefecture, Toyota-shi, Otagicho, Dando-dani Valley, 27-V-1989, Hiroshi & Masami Nishida leg.; 2♂ (KS), Japan • Nagano Prefecture, Hiraya-mura, near Hiraya-tôge Pass, 14-VI-2002, Hiroshi & Takeyuki Nishida leg.

Dando-dani Valley is located on the southern slope of Mount Takanosu-yama and can be regarded as virtually identical to the type locality. The record from Hiraya-tôge Pass is the northernmost collection record for this species, the first record outside the type locality, and the first record from Nagano Prefecture. The endophallus of the Hiraya-tôge Pass specimen differed slightly from that of specimens from the type locality in the shape of the backward sub-lobe at the apical 2/5 of the dorsoapical lobe (slightly bifid in the Hiraya-tôge Pass specimen versus simply rounded in the specimens from the type locality) and in the sizes of sub-lobes of the dorsomedian lobe and the base of the dorsoapical lobe. In the original description of N. dichotoma, the morphology of the gonopore protrusion could not be examined because of insufficient inversion of this structure. Therefore, the present result provides the first description of the gonopore protrusion of this species. Its gonopore protrusion was simple cylindrical, relatively long, and strongly bent at the base toward the base of the endophallus.

Holotype : ♂ (KS), Japan • Shizuoka Prefecture, Hamamatsu-shi, Mount Ryûtô-san, alt. 1200 m, 3-X-2006 [no collector data]. Paratypes: 1♀ (KS), same data as the holotype; 1♂ (KS), Japan • Shizuoka Prefecture, Shizuoka-shi, Aoi-ku, Mount Sasa-yama, alt. 1500 m, 29-X-2005, Masato Mori leg.; 2♂1♀ (KS), Japan • Shizuoka Prefecture, Tenryu-shi, Sakumacho, Mount Idoguchi-yama, the upper reaches of Aizuki-gawa River, 18-V-1996, Hiroshi Nishida leg.

Figures 10–12. 

Right lateral (a), ventral (b), dorsal (c), left lateral (d), and posterodorsal (e) views of the endophallus of Nebria dracocephala sp. nov. from Mount Ryûtô-san (10), Mount Idoguchi-yama (11), and Mount Sasa-yama (12).

Similar to the locally adjacent species N. dichotoma, but distinguished by the shape of the endophallus, particularly the left lateroapical lobe composed of two sub-lobes and the dorsoapical lobe with apex not bifurcated (bifurcated in N. dichotoma).

Body length: ♂, 9.95–10.73 mm, 10.26 ± 0.34 mm (n = 4); ♀, 10.89–11.13 mm, 11.01 ± 0.17 mm (n = 2). Sternum 7 with two setae on each ventrolateral side in both male and female. Other external structures, coloration, and chaetotaxy as in other related species that had been previously regarded as N. reflexa (Sasakawa 2020). Dorsobasal lobe absent. Dorsomedian lobe undeveloped, with a minute, slender protrusion or a weak, wide swelling. Dorsoapical lobe with a simple-shaped (i.e., not bifurcated) protrusion at the basal part, the size of which varies among localities, ranging from smaller than that of the dorsobasal lobe to almost the same size as the laterobasal lobes; the dorsomedian area with a large protrusion, the apex of which is simply rounded or bifid, varying among localities; the apical portion bent ventrally, with the dorsal bend discontinuous in lateral view, more or less swollen. Laterobasal lobes large; the left lobe more rectangular in shape than the right in ventral view; at the distal end of both right and left lobes, the endophallus-base side corner more protruding than the opposite side; ventrobasal surface largely swollen on both right and left lobes, distinctly visible in lateral view. Ventrobasal swelling absent. Right lateroapical lobe markedly wide in dorsal view, with the endophallus-base and -apex side corners protruding, resulting in a T-shape in dorsal view; the endophallus-base side corner more slender and protruding than the opposite side. Left lateroapical lobe composed of two sub-lobes, one on the left dorsolateral side and the other on the left lateral side of the endophallus; the sub-lobe on the dorsolateral side T-shaped, with the endophallus-apex side corner more robust and protruding than the opposite side; the sub-lobe on the lateral side large, with a simply rounded or slightly bifid apex, varying among localities. Gonopore protrusion bent at the base, directed toward the endophallus base; the observable part simply curved cylindrical, with no additional structures such as protrusions. Relative sizes of some lobes and protrusions are as follows: dorsomedian lobe < endophallus-apex side corner of left dorsolateral sub-lobe of left lateroapical lobe ≈ endophallus-base side corner of right lateroapical lobe < swelling of ventrobasal surface of laterobasal lobes < left lateral sub-lobe of left lateroapical lobe ≤ laterobasal lobe (including swelling of ventrobasal surface) ≤ protrusion on dorsomedian area of dorsoapical lobe.

There was marked variation in endophallus morphology, both qualitatively (shape) and quantitatively (size). However, all specimens shared basic structures, such as the left preapical lobe composed of two sub-lobes and the dorsoapical lobe with protrusions on the dorsobasal, dorsomedian, and dorsoapical sides, none of which are possessed by other closely related species (Sasakawa 2020, 2023b; Sasakawa and Itô 2021). Given their morphological complexity, these shared characters are probably synapomorphies uniting the three populations rather than symplesiomorphies. Therefore, the three populations were treated as a single species. Morita and Hirai (2010) recorded “N. reflexa” from the Abe-tôge Pass, 9 km northwest of Mount Sasa-yama, and this specimen is also most likely N. dracocephala.

The specific name is a combination of the Latin draco (dragon) and the Latin cephalus, -a, -um (head), derived from the type locality Ryûtô-san, which means “dragon’s head mountain” in Japanese.

Holotype : ♂ (KS), Japan • Tokushima Prefecture, Tsurugi-cho, Ichiu, near Tsurugisan Ski Area, alt. 1000 m, 14-VII-2006, Masato Mori leg. Paratypes: 1♂1♀ (KS), Japan • Tokushima Prefecture, Higashimiyoshi-cho, Nishisho, near Sajiki-tôge Pass, alt. 800 m, 10-VII-1999, Masato Mori leg.

Figures 13–15. 

Right lateral (a), ventral (b), dorsal (c), left lateral (d), and posterodorsal (e) views of the endophallus of Nebria spp.; 13. N. elephanta sp. nov. holotype from near Tsurugisan Ski Area; 14. N. hikosana from Ryûmon-kyô Gorge; 15. N. hikosana from Mount Seburi-san.

Similar to the locally adjacent species N. chugokuensis Sasakawa, 2020 and N. uenoi Nakane, 1963, but distinguished from N. chugokuensis by a smaller body size (in N. chugokuensis, 9.97–10.32 mm for males and 10.69–11.75 mm for females; Sasakawa 2020) and the absence of a concavity on the ventral surface of the aedeagal apex (present in N. chugokuensis), and from N. uenoi by the shape of the endophallus, in particular, the developed gonopore protrusion (undeveloped in N. uenoi) and the absence of lateroapical lobes on both sides (present in N. uenoi).

Body length: ♂, 9.70–9.81 mm, 9.75 ± 0.08 mm (n = 2); ♀, 10.54 mm (n = 1). Sternum 7 with two setae on each ventrolateral side in the male, and three in the female. Other external structures, coloration, and chaetotaxy as in other related species that had been previously regarded as N. reflexa (Sasakawa 2020). Dorsobasal lobe large, semi-spherical. Dorsomedian lobe absent. Dorsoapical lobe scalene trapezium-shaped in dorsal view, with the left and right sides concave; the left side more concave than the right; the left apical corner more protruding than the right. Laterobasal lobes with the ventrobasal surface largely swollen in a semi-spherical shape; the other part less swollen but wide, reaching the base of the dorsobasal lobe. Ventrobasal swelling large, semi-ellipsoid. Lateroapical lobes absent on both right and left sides. Gonopore protrusion markedly large and long, with total length along the median line from the base of the protrusion to the gonopore (apical end of the protrusion) longer than that from the base of the endophallus (ostium of the aedeagus) to the base of the protrusion; basal half thick, oriented in the same direction as the median line of the endophallus; apical half slender, strongly bent right laterally. Relative sizes of some lobes and protrusions are as follows: ventrobasal swelling ≤ swelling of ventrobasal surface of laterobasal lobes ≈ dorsobasal lobe < dorsoapical lobe < basal half of gonopore protrusion.

The description of setae on female sternum 7 as three pairs was based on only one specimen. This character state must be confirmed based on additional specimens, because in Carabidae, the number of setae on the body surface, including sternum 7, often varies due to additional setae, etc. The present collection sites are located on the northern slope of Mount Tsurugi-san in the eastern part of Shikoku. Considering the relatively wide distribution of related species and the continuity of the forest environment, which seems to be suitable for this group, N. elephanta is assumed to be widely distributed around Mount Tsurugi-san. In Shikoku, there is also a record of “N. reflexa reflexa” from Mount Takanawa-san in the western part, 110 km from Mount Tsurugi-san (Yoshitomi et al. 2012). The identity of the Takanawa-san specimens remains to be clarified in a future study.

The specific name is derived from the Latin elephantus, -a, -um (elephant) and refers to the elongated gonopore protrusion of this species, which resembles the trunk of an elephant.

• Holotype ♂ (NARO), “Col.A.HABU / Mt. Hiko / Fukuoka P. / V. 23, 1950 // Holotype / Nebria / hikosana / HABU // ♂”

3♂ (KMNH), Japan • Fukuoka Prefecture, Soeda-machi, Hikosan, Mount Hiko-san, Yasuo Takakura leg. (1♂ [no further locality information], 8-VI-1982; 2♂, Ryûmon-kyô Gorge, 8-V-1989); 4♂5♀ (KS), Japan • Saga Prefecture, Kanzaki-shi, Mount Seburi-san, alt. 950 m, 15-V-2001, Masato Mori leg.; 1♀ (KMNH), Japan • Fukuoka Prefecture, Fukuoka-shi~Saga Prefecture, Kanzaki-shi, Mount Seburi-san, 7-V-1977, Munemichi Fukamachi leg.

Similar to the locally adjacent species N. chugokuensis, but distinguished by one pair of setae on the ventral side of the sterna 7 in the male (two pairs in N. chugokuensis) and the absence of a concavity on the ventral surface of the aedeagal apex (present in N. chugokuensis).

Body length: ♂, 9.14–9.92 mm, 9.64 ± 0.27 mm (n = 8); ♀, 10.27–10.89 mm, 10.56 ± 0.24 mm (n = 6). Sternum 7 with one seta on each ventrolateral side in the male, and two in the female. Other external structures, coloration, and chaetotaxy as in other related species that had been previously regarded as N. reflexa (Sasakawa 2020). Dorsobasal lobe small and slightly swollen. Dorsomedian lobe semi-prolate-spheroid, weakly bent toward the endophallus base or not bent and simply directed dorsally. Dorsoapical lobe elongated, apically narrowed gradually, weakly and uniformly bent ventrally; apex widely rounded; dorsobasal surface slightly swollen in some specimens. Laterobasal lobes small, nearly spherical; in ventral view, the left and right laterobasal lobes nearly attached to each other, concealing the aedeagus apex. Ventrobasal swelling small, semi-spheroid, attached to the fused basal portion of the left and right laterobasal lobes in ventral view. Right lateroapical lobe moderate in size, bifurcated in a V-shape in dorsal view; subapical lobes both conical and about the same size. Left lateroapical lobe conical, except for the surface on the endophallus-base side in dorsal/ventral view, which is slightly swollen. Gonopore protrusion large, directed ventrally, with an additional protrusion on the left side at the middle; additional protrusion straight, directed left laterally, gradually narrowed apically, with a length more than half that of the main body of the gonopore protrusion. Relative sizes of some lobes and protrusions are as follows: subapical lobes of the right lateroapical lobe < laterobasal lobes ≈ ventrobasal swelling < additional protrusion of the gonopore protrusion ≤ dorsomedian lobe < main body of the gonopore protrusion < dorsoapical lobe.

There is disagreement as to whether the taxon hikosana should be treated as a subspecies of N. reflexa or as a distinct species, but neither treatment has been based on concrete evidence. Here, hikosana is regarded as a distinct species rather than a subspecies of another species, based on marked differences in endophallus morphology compared with other related species, particularly N. chugokuensis, the closest distributed species to N. hikosana (Sasakawa 2020). In Kyushu, there is also a record of this species from Mount Shaka-dake, located 30 km south of Mount Hiko-san and belonging to a different mountain range (Imasaka et al. 2019), but its identity requires confirmation in future studies.