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Corresponding author: Kôji Sasakawa ( ksasa@chiba-u.jp ) Academic editor: Ana Sofia P. S. Reboleira
© 2022 Kôji Sasakawa, Hirotarô Itô.
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Citation:
Sasakawa K, Itô H (2022) The identities of two species in the Pterostichus macrogenys species group of subterranean carabid beetles (Coleoptera, Carabidae) revealed by external morphometric analysis and comparative genital morphology. Subterranean Biology 43: 61-71. https://doi.org/10.3897/subtbiol.43.80969
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The Pterostichus macrogenys species group is an endemic subterranean Japanese carabid lineage that shows marked regional differentiation, but unresolved taxonomic issues remain, even at the species level. Based on morphological examinations of the genital structures of newly collected specimens and an external morphometric analysis of all the species concerned, P. falcispinus Sasakawa, 2005 syn. nov. is synonymized with P. asahinus Habu & Baba, 1960, and P. awashimaensis sp. nov. is described from Awashima Island, a small island off the coast of Honshu. Based on external morphometrics, P. awashimaensis was determined to be most similar to P. yahikosanus Sasakawa, 2009, but its external and genital features differ distinctly from the latter species and it is thought to be more ancestral.
Discriminant analysis, ground beetle, Japan, new species, new synonym, Nialoe
The macrogenys species group of Pterostichus subgenus Nialoe Tanaka, 1958 (s. lat., i.e., Nialoe sensu
Distribution of the Pterostichus macrogenys species group in northern Chubu, northern Kanto, and southern Tohoku Districts, Honshu, based on specimens with unambiguous identities, modified from
This study sought to resolve two taxonomic issues in the macrogenys species group: the relationship between P. asahinus Habu & Baba, 1960 and P. falcispinus Sasakawa, 2005, and the identity of the population on Awashima Island, a small island off the coast of Honshu. The fact that the holotype of P. asahinus and the available specimen from the Awashima population are both females, which provide less taxonomic information than males, has hampered resolution of these taxonomic issues. To overcome this, we performed external morphometric analysis, a useful taxonomic method that can also be applied to females (e.g.,
Specimens newly examined here were collected using subterranean baited traps, which were also used in our previous studies (
The discriminant analysis was used to investigate morphometrically whether the identity of the P. asahinus holotype and our newly defined P. asahinus is supported and to which species the Awashima specimen is most similar. The following specimens were examined together with the Awashima specimen and P. asahinus holotype: P. adatarasanus 3♀, P. asahinus (redefined here to include P. falcispinus) 17♀, P. chokaisanus 2♀, P. eboshiyamanus 3♀, P. iwakiensis 7♀, P. monolineatus 1♀, P. ohsawacavus 1♀, P. takadateyamanus 25♀, and P. yahikosanus 6♀. These samples include all species to which the Awashima specimen and P. asahinus holotype may belong and for which female specimens are available. Scaled photos obtained previously (
The male endophallus was everted and fully inflated by injecting toothpaste from the base of the aedeagus (
Pterostichus (Paralianoe) macrogenys asahinus:
Pterostichus (Paralianoe) asahinus:
Pterostichus macrogenys:
Pterostichus (Nialoe) asahinus:
Pterostichus (Nialoe) falcispinus:
1♂, Miomote, alt. 276 m, on the right bank of Miomotegawa River, Murakami-shi, Niigata Prefecture, Japan (38.273211°N, 139.779922°E), 12.vi.–17.vii.2021, Hirotarô Itô leg., in the collection of HI.
Pterostichus asahinus male from Miomote, on the right bank of the Miomotegawa River A habitus dorsal view B–E Endophallus left lateral (B) ventral (C) right lateral (D) and dorsal (E) views F–H right paramere left lateral (F) apical (G) and dorsal (H) views. go, gonopore; lal, left apical lobe; lpl, left preapical lobe; rpl, right preapical lobe.
Pterostichus (Paralianoe) macrogenys macrogenys:
Pterostichus macrogenys:
Holotype : ♀, Mt. Koshibayama, alt. 235 m, Awashima Island, Awashimaura-mura, Niigata Prefecture, Japan, 24.v.–14.vi.2015, Hirotarô Itô leg., deposited in the Laboratory of Zoology, Department of Science Education, Faculty of Education, Chiba University, Chiba, Japan.
In the discriminant function analysis, the Awashima specimen was classified as P. yahikosanus. On the scatterplots of the first two canonical variates, the specimen was located outside of, but close to, the area of P. yahikosanus (Fig.
Pterostichus awashimaensis sp. nov. holotype female (A, C, E) and P. yahikosanus female from Mt. Yahikosan (B, D, F) A, B habitus dorsal view C, D pronotum dorsal view E, F vagina anterior view. am, apophysis of the median oviduct; as, apophysis of the seminal canal; m, median oviduct; p, pigmentation on the innermost part of the vagina; sp, spermatheca.
BLm 13.79 mm; BLl 12.66 mm; BLc 12.13 mm. Head, pronotum, and elytra dark brown; appendages reddish brown. Dorsal surface almost smooth except for laterobasal impressions of the pronotum, the anterior half of which bear several transverse wrinkles and the posterior half of which are weakly punctate.
Head large, widest at tempora, which are distinctly swollen; width at the widest point larger than pronotal posterior margin width; length from clypeal apex to neck base longer than pronotum length along the median line. Left mandible larger than the right and curved at the apical 1/4; length between mandible apex and posterolateral end on dorsal side slightly shorter than 2.5 times the anterior width of the clypeus. Eyes weakly convex, with the anterior-posterior length longer than 1/2 length of antennal segment 1. Antennal segment 2 with two setae.
Pronotum cordate, notably flat, widest at apical 1/5. Lateral margins arcuate on apical 2/3, slightly sinuate on basal 1/3; two marginal setae on each lateral side, anterior setae near widest pronotal point, and posterior setae near hind angles. Anterior margin emarginated, with curvature approximately the same as that of apical 2/3 of lateral margins; anterior angles widely rounded and barely produced. Hind angles obtuse. Posterior margin only slightly emarginated at median area, weakly but distinctly arcuate posteriorly behind the laterobasal impressions. Median line impressed in the middle, not reaching either the anterior or posterior margins; laterobasal impressions single, shallow.
Elytra sides almost parallel, less convex; shoulder distinct, but not denticulate; apices rounded; scutellar stria present, connected to stria 1 on the right side but not on the left; 1 setigerous puncture on stria 1 at the level of the posterior end of scutellum; two setigerous punctures on interval 3, anterior one slightly before the middle and posterior one on apical 1/5, both adjoining stria 2. Hind wings completely atrophied. First fore tarsomere no adhesive hairs on ventral side.
Female genital structures identical to those of other consubgeners; apophyses of seminal canal and median oviduct fully sclerotized; pigmentation on the innermost part of the vagina present, but less than half the size of apophyses of seminal canal and median oviduct; other parts of vagina lack conspicuous pigmentation.
We thank Y. Mitsuduka (Yamagata) for allowing us to examine specimens in his collection. This study was partly supported by the Japan Society for the Promotion of Science to KS (no. 17K15171).
Information on specimens examined and the raw data for discriminant analysis.
ID † | Species | Collection site § | PL | PAW | PW | PPW |
---|---|---|---|---|---|---|
mttk071 | adatarasanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.82 | 3.55 | 3.93 | 3.08 |
mttk073 | adatarasanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.85 | 3.64 | 4.16 | 2.97 |
mttk077 | adatarasanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.66 | 3.40 | 3.91 | 2.93 |
cHIT_0004 | asahinus | Ishikiri Cave, Tainai-shi, NI | 2.83 | 3.98 | 4.48 | 3.44 |
mttk028 | asahinus | Kurosawagawa forest road, Mamurogawa-machi, YG | 2.43 | 3.42 | 3.94 | 3.07 |
mttk079 | asahinus | Togawa, Tozawa-mura, YG | 2.45 | 3.42 | 3.87 | 2.92 |
mttk026 | asahinus | Ôtori, Tsuruoka-shi, YG | 2.60 | 3.88 | 4.35 | 3.23 |
mttk040 | asahinus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.9 | 4.20 | 4.72 | 3.67 |
mttk070 | asahinus | Mt. Takadateyama, Tsuruoka-shi, YG | 3.08 | 4.25 | 4.83 | 3.87 |
mttk072 | asahinus | Mt. Takadateyama, Tsuruoka-shi, YG | 3.35 | 4.56 | 5.20 | 4.00 |
mttk033 | asahinus | Nukumidaira, Oguni-machi, YG | 3.14 | 4.66 | 5.08 | 3.98 |
mttk055 | asahinus | Nukumidaira, Oguni-machi, YG | 3.08 | 4.73 | 5.13 | 3.96 |
mttk056 | asahinus | Nukumidaira, Oguni-machi, YG | 2.98 | 4.37 | 4.88 | 3.71 |
mttk057 | asahinus | Nukumidaira, Oguni-machi, YG | 3.10 | 4.46 | 4.92 | 3.90 |
mttk059 | asahinus | Nukumidaira, Oguni-machi, YG | 2.97 | 4.35 | 4.77 | 3.76 |
mttk060 | asahinus | Nukumidaira, Oguni-machi, YG | 2.82 | 4.16 | 4.65 | 3.56 |
mttk061 | asahinus | Nukumidaira, Oguni-machi, YG | 3.18 | 4.39 | 5.19 | 3.74 |
mttk062 | asahinus | Nukumidaira, Oguni-machi, YG | 3.23 | 5.06 | 5.43 | 4.04 |
mttk063 | asahinus | Nukumidaira, Oguni-machi, YG | 3.08 | 4.37 | 5.00 | 3.73 |
mttk064 | asahinus | Nukumidaira, Oguni-machi, YG | 3.05 | 4.67 | 5.04 | 3.85 |
mttk030 | chokaisanus | Sakunami, Sendai-shi, MG | 2.54 | 3.07 | 3.61 | 2.46 |
mttk080 | chokaisanus | Mt. Chokaisan, Yuza-machi, YG | 2.25 | 2.86 | 3.34 | 2.40 |
mttk052 | eboshiyamanus | Yunosawa, Iide-machi, YG | 2.47 | 3.02 | 3.43 | 2.47 |
mttk053 | eboshiyamanus | Yunosawa, Iide-machi, YG | 2.38 | 3.09 | 3.47 | 2.61 |
mttk054 | eboshiyamanus | Yunosawa, Iide-machi, YG | 2.63 | 3.47 | 3.81 | 2.83 |
cHIT_0002 | iwakiensis | Miyahisa, Tainai-shi, NI | 2.53 | 3.37 | 3.82 | 2.84 |
cHIT_0005 | iwakiensis | Ishikiri Cave, Tainai-shi, NI | 2.38 | 3.35 | 3.87 | 2.93 |
iwakiensisPT ‡ | iwakiensis | Iritôno, Iwaki-shi, FS | 2.95 | 3.60 | 4.29 | 3.07 |
mttk034 | iwakiensis | Nukumidaira, Oguni-machi, YG | 2.43 | 3.04 | 3.65 | 2.73 |
mttk058 | iwakiensis | Nukumidaira, Oguni-machi, YG | 2.31 | 3.06 | 3.53 | 2.57 |
mttk065 | iwakiensis | Nukumidaira, Oguni-machi, YG | 2.60 | 3.36 | 3.90 | 2.75 |
mttk066 | iwakiensis | Nukumidaira, Oguni-machi, YG | 2.61 | 3.32 | 3.75 | 2.74 |
mttk037 | monolineatus | Aobaminami, Marumori-machi, MG | 2.77 | 3.55 | 3.94 | 2.96 |
cHIT_0053 | ohsawacavus | Ohsawa Cave, Gosen-shi, NI | 2.79 | 3.98 | 4.22 | 3.26 |
cHIT_0017 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.37 | 3.06 | 3.53 | 2.64 |
cHIT_0018 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.52 | 3.39 | 3.84 | 2.79 |
cHIT_0019 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.60 | 3.57 | 4.01 | 2.88 |
cHIT_0020 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.59 | 3.57 | 3.99 | 2.98 |
cHIT_0021 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.56 | 3.57 | 3.96 | 2.93 |
cHIT_0022 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.60 | 3.48 | 3.86 | 2.86 |
cHIT_0023 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.34 | 3.09 | 3.51 | 2.69 |
cHIT_0024 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.80 | 3.93 | 4.19 | 3.02 |
cHIT_0025 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.63 | 3.53 | 4.07 | 3.05 |
cHIT_0026 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.31 | 3.20 | 3.64 | 2.69 |
cHIT_0027 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.60 | 3.57 | 4.01 | 2.92 |
cHIT_0028 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.50 | 3.44 | 3.87 | 2.87 |
cHIT_0029 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.46 | 3.42 | 3.87 | 2.83 |
cHIT_0030 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.40 | 3.24 | 3.65 | 2.78 |
cHIT_0031 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.61 | 3.50 | 3.90 | 2.82 |
cHIT_0032 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.52 | 3.70 | 4.01 | 2.94 |
cHIT_0034 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.72 | 3.73 | 4.10 | 2.90 |
cHIT_0035 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.46 | 3.44 | 3.83 | 2.77 |
cHIT_0036 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.51 | 3.32 | 3.81 | 2.84 |
cHIT_0037 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.25 | 3.07 | 3.52 | 2.67 |
cHIT_0039 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.46 | 3.31 | 3.67 | 2.80 |
mttk041 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.67 | 3.59 | 3.99 | 2.88 |
mttk074 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.59 | 3.76 | 4.14 | 2.89 |
mttk075 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.50 | 3.60 | 3.99 | 2.82 |
mttk076 | takadateyamanus | Mt. Takadateyama, Tsuruoka-shi, YG | 2.55 | 3.59 | 3.95 | 2.90 |
tateishiyamanusPTf ‡ | tateishiyamanus | Kuratani-sawa, Nishiaizu-machi, FS | 2.50 | 3.07 | 3.51 | 2.55 |
cHIT_0044 | yahikosanus | Mt. Yahikosan, Nagaoka-shi, NI | 2.56 | 2.98 | 3.52 | 2.42 |
cHIT_0045 | yahikosanus | Mt. Yahikosan, Nagaoka-shi, NI | 2.59 | 3.18 | 3.65 | 2.67 |
cHIT_0046 | yahikosanus | Mt. Yahikosan, Nagaoka-shi, NI | 2.55 | 3.15 | 3.62 | 2.53 |
cHIT_0048 | yahikosanus | Mt. Yahikosan, Nagaoka-shi, NI | 2.50 | 3.10 | 3.56 | 2.40 |
cHIT_0049 | yahikosanus | Mt. Yahikosan, Nagaoka-shi, NI | 2.68 | 3.29 | 3.77 | 2.74 |
cHIT_0050 | yahikosanus | Mt. Yahikosan, Nagaoka-shi, NI | 2.62 | 3.29 | 3.73 | 2.66 |
asahinusHT ‡ | asahinus | Mt. Dôrokujinpô, Murakami-shi NI | 2.51 | 3.51 | 4.14 | 3.10 |
awashima_F01 ‡ | awashimaensis | Awashima Island, Awashimaura-mura, NI | 2.50 | 2.92 | 3.38 | 2.48 |