akasaka nanae

Transcription

akasaka nanae

TMU Bulletin of Natural History, No. 3: 55-99. December 20, 1997.
Subspeciation and distribution pattern of
Carabus albrechti Morawitz in Japan
(Coleoptera, Carabidae)
by
Yasuoki Takarni' and Ryasuke 1shikan[a2
1
Department of Natural History, Graduate School of Science, Tokyo Metropolitan University,
Minami-ohsawa 1-1, Hachioji-shi, Tokyo 192-0364, Japan
takami-yasuoki@c.metro-u.ac.jp
'(20-op Olympia 320, Jingcmae 6-35-3, Shibuya-ku, Tokyo 150-0001, Japan
Abstract: The subspecies of Curabus (O/~owiuptc~ru.s)
u l b r ~ l i t iwhich
,
is distributed in northeastern Honshu east of the Itoigawa-Shizuoka Tectonic Line and Hokkaido in Japan, are revised
taxonomically. Twelve subspecies are recognized, three of which, C. u. yurr~u~~chLi,
C. (1. ltagui
and C . a. ts~(k~~bunus
are described as new. The phylogenetic relationships among the subspecies are reconstructed by cladistic analysis based on the adult morphological characters, and the
process of their differentiation is infemed.
Key words: Coleopteri, Cxabidae, Curubu.~(Ohonzo~>t~~ru.s)
ulbrc:c/ttl,subspeciation, distribution pattern, new subspecies, Japan, cladistics, hybrid zone, phylogeny, reproductive isolation.
Contents
Introduction .......................................................................................................................
Recognition of subspecies..................................................................................................
Description ...... ........................................................... .............. .......... .......... ......................
Key to subspecies of Curabt4,salbrec5htL....... ........,. .... ..... . .... ............ .......... .... ............ ........
Phylogei~y.......... ...... . ........ ........ .......... ...... . .. .... .........., ........ . ..., ..... ......, ..... ..,..... .....,....' , ....
Method............................................................,.,........ .,...... ......... .. .......... ............. .......
Evaluation of diagnostic characters..............................................................................
Result of cladistic analysis............................................................................................
Discussion ...................................................... ....... .......... ................. .......... ........ ... ..... . .......
Subspeciation in the Chcbu District ..............................................................................
Subspeciatiorl in the tt,ji-[her11part of Kant6 and the southern part of T6hoku
~
,
,
,
,
,
..........................................................................................................................................
Subspeciation in the northern part of the T6hoku District and Hokkaido......................
Establishme~~t
of reproductive isolation........................................................................
Conclusion............................... ....................... .......... ............... ........... ...... ......... .......... ......
Acknowledgements ..................................................................................... ........................
References...,........................,,........,...,............................ ...................................................
Appendix 1 : Character n~atricesused in the cladistic analysis............................................
Appendix 2: Index map of nort11-eastern Japan ..................................................................
~
Introduction
Carabta alhrechii Morawit~is a small to medium-sized species of the subgenus
Ohortzopterzi.~,
and is widely distributed in northeastern Japan east of the Itoigawa-Shizuoka
Tectonic Line of Honshu. This is the only Ohomoptcrz~.s-speciesoccurring a< far north as
Hokkaido. Although sporadic in distribution, it is usually one of conllnon species where it
occurs. It shows considerable geographical variation and therefore has been subdivided
into a series of subspecies. These subspecies were not strictly defined ~~~orphologically,
however, and the range of each subspecies was not sufficiently outlined until recently
(Ishikawa and Takami, 19961, mainly because tile morphological differentiation among
thein is not always conspicuous compared with other species of the subgenus 0hornopteru.s.
Carubu.s albrechti [=CarubuLs(Ohomopterzis) albre(~lztialbrcjchti] was originally described by Morawitz (1 862) based on the specimens from Hakodate (Hokkaido). Breuning
(1932-1937) placed this species in the subgenus Ap(>tornopterzi.sof the genus Carabzi.s in his
Monographic der Gattung Casabus L. Emden (1932) described Carahus (Olzonzoptet-us)
Albrechti Freyi from Sado Is. Nakane (1952a, b & 1953) treated C. lc+t~i~sia~~u.s
and C.
yamato as subspecies of Apotofizopteru.~albrechti. Later, he described esakiatzu,~as a subspecies of A. japonicus (=Carabzi.s (0hornopterzi.s) cilbrechti c>.sakicuzits)(Nakane, 1960).
Later, he regarded A. albrechti as a subspecies of A. japo~zi~~us,
in which C. le~isianzd.~,
C.
ya~natoand C. dczism were also included together as subspecies of that species (Nakane,
1962). Isllikawa ( I 966) described Apoto~~topteru.~
albrechti okur?zuruias a subspecies of A.
albrechti, as it clearly differs from A. j. e.sakiatzzk.s. Subsequently, he revised the species in
question under the name of A . ulbrechti, and placed C. daisen as a subspecies of that species
(Ishikawa, 1969).
Ishikawa (1973) included all the Japanese species wllicl~were assigned to Apotot?zo17teru.s by Brewing and Nakane, respectively, in the subgen~~s
0honzopterucsof the genus
Caral?u,s. Thus, A. albrechti was combined again with Curubu,~ Later, he described
Curabzi.~(Ohomo/)terzi.s) albr~>chti
tohokzkerzsis as an intermediate geographical form between C. (1. albrechti and C. a . esakiunu.s (Ishikawa, 1984). In 1985, lle illustrated in color
most subspecies of C. uZbrechti with brief descriptions, but the range ofthe species and their
geographic variation were not clearly defined. Recently, Ishikawa and Takami (1996) described four subspecies of this species: C. a . hidakanzts, C. a . itoi, C. a . echigo and C. a.
awashinzae. The former two subspecies were discriminated from C . a . a1l)rechti in
Hokkaido. The latter two were discritni~latedfrom C. a. orhigo in Honshu and C. a .
c~washi~zae
in Awashima Is. respectively, which were not clearly discriminated so far from
C. a. csakianus, C. a. okurnurai and C. a. j'reyi.
In the present study, criteria are applied to geographic populatio~~s
of Carabu.~albrechti
to allow distinguishing of subspecies. Some 5430 specimens from 37 I localities were examined. Phylogeny of the subspecies as thus recognized is reconstructed and the process of
subspecific differentiation is discussed.
Recognition of subspecies
Subspecies are morphologically distinguishable geographic races of' species that are not
isolated reproductively from others (Ishikawa, 1992). Such geographic races, if not allopatrically distributed, may be connected with others by intermediate populations at the lransi-
Fig. 1. Distribution ofthe subspecies of Curuhus ulbrc>chtiMorawitz based on the specinlens examined. I . C.
a. albrechti Morawitz; 2, C . u. hidukutz~~s
Ishikawa & Tdkami: 3, C. u . itoi Ishikawa & Takami; 4, C , u.
tohokuc?n.sis Ishikawa; 5, C . u. yurna~ichiisubsp. nov.; 6. C. (1. huSyui subsp. mv.: 7 , C. u , (~~~u.s/zirnuc~
Ishikawa & Takami; 8, C . a. t.suk~ibunu~s
subsp. nov.; 9, C. u.jreji Emden: 10, C, u, osukiunu.s (Nakane);
1 I , C. a. echigo Ishikawa & Takaini: 12, C. u . okikmlirui (Ishikawa). Shaded areas show transitional or
hybrid zones between subspecies. Meshed area, see Discussion.
58
Y . Tk~rniand R. lshikdwa
tion zones which were formed by secondary meeting of and hybridization between them.
The subspecies as thus recognized possibly comprises intra-subspecific populations which
may be the true units of speciation though not or barely distinguished from others morphologically. Current phylogenetic studies adopt subspecies (or species and higher taxa) as
Operational Taxonomic Units (OTUs), which are the fundamental units in phylogenetic
analysis and should naturally be monophyletic. However, morphologically recognized subspecies are not always monophyletic in terms of cladistics, because the monophyly of a
group is evidenced as a result of phylogenetic analysis and is not obvious ci priori.
In this study, subspecies are defined as the sn~allestevolutionary unit which is distinguishable morphologically. The subspecies comprises intra-subspecific populations which
share any morphological character(s) within the range of the subspecies. The morphological characters shared by each population, whether derived (apomorphic) or underived
(plesiomorphic). are coded as subspecific characters in the matrix for cladistic analysis.
Therefore, the monophyly of a subspecies is assumed by its own morphological homogeneity.
Description
Carabus (Ohomopterus) albrechti Morawitz, 1862
All the subspecies share the following characters, unless otherwise stated.
Upper surface of body dark copper. Palpi, tibiae and tarsi more or less rufous. Antennae
of male longer than in female and exceeding middle of elytra, underside of 5'7th segments
with distinct hairless depressions which are as long as their length. Head impunctate from
front to neck. Inner margin of male protibia distinctly and angulately convex at middle.
Lateral margins of pronotum cordately sinuous; postero-lateral corners rounded but distinctly produced; more than two marginal setae on either side. Elytra slender and weakly
convex; oval in female, and more oblong in male; primary foveae sniall; striae weakly
notched.
Male genitajia with apical portion of aedeagus narrowed, broader in outer view, and
gradually tapered from base to rounded tip, with deep groove on its outer margin which is
outlined by carinae; digitate part of copulatory piece narrow, subparallel at basal half, narrowed to blunt tip, weakly constricted at base.
Female genitalia with outer plate of vaginal apophysis broad and triangular; inner plate
of vaginal apophysis shallow cup-shaped, with shallow median groove; anterior r i n ~narrow.
Carabus (Ohomopterus) albrechti albrechti Morawitz
Carab~4,sA/brechti Morawitz, 1862. pp. 237-238; type-locality: Hdsodate
Carahus Albrechti: Morawitz, 1863, pp. 10-1 I .
Carabu,~(A/)otomopteru,s)ulbrechti (part.): Brewing, 1932, pp. 235-236.
Apotomopt<<ru.~
ulbre(+zfi (purt.): Nakane, 1952, pp. 50-5 I.
A[~otomol~ter~ds
ulbrechfi (part.): Nakane, 1953, p. 96 (48).
Apofomoplerus jupor~~cu.~
ulbr~chti(part.): Nakane. 1962, p. 39.
Apotomopteru.~ulbrer!~ti ulbrechfi hurt.): lshikawa, 1969. pp. 520-521, figs. I & 21.
Curubus (Qtzomopfer~d~s)
ulbrechti ulbrechti Q~uri.):lshikawa, 1985, p. 21.
Curubus (O/tomopteru,s)ulbrechti ajbrochti: Ishikawa & Takami, 1996, pp. 39-4 I, figs. 1-3.
Curubus (0hornopteru.s) ulbrec/iti ulbrctc/zti (part'): linuri & Mizusawa, 1996. pp. 20 & 107
fig. 44 (I); designation of lectotype.
This subspecies of the Oshi~naPeninsula in Hokkaido and Okushiri Is. was not distinguished from C. a. hidakatzuLsand C. a. itoi of Hokkaido until recently (Ishikawa &
Takami,1996). C. a. albrechti differs from them in the shape of the apical part of aedeagus
and of pronoturn, the characters which it shares with several subspecies of Honshu.
Length: ?, 2 1.5-25.5 trim; 2 , 20.5-23.5 rnm.
Upper surface of body copper or dark copper, rarely green. Legs n~ostlywholly black, or
tibiae and rarely tarsi partly somewhat rufous. Male antennae short, not reaching middle of
elytra; underside of 5-7th segments with or without vaguely outlined hairless depressions,
about 112-213 as long as each segment. Head weakly punctate on vertex, with coarse rugae
from front to neck. Inner margin of male protibia weakly convex but not angulate (Fig. 14).
Pronoturn small; lateral margins weakly cordately sinuous; disc convex; 3-5 (rarely 7 or 8)
marginal setae; postero-lateral corners angulately produced (Fig. 10). Elytra oblong, short
and convex; striae at most very weakly notched.
Male genitalia: Apical portion of aedeagus long and thin, finger-shaped, thinnest at
middle in lateral view; its outer margin weakly grooved or flat, o~~tlined
by rudimentary
ridges; copulatory piece broad and short (Fig. 2).
Female genitalia: Outer plate of vaginal apophysis triangular, 213 as broad as long: inner
plate with anterior rim broad, disk not grooved medially (Fig. 3).
Variation: In the specimens from high elevations (Mt. Daisengendake), the upper surface
of the body is uniformly bright copper, but in those from the lower altitudes (including
Okushiri Is.), the copper color is darker. Specimens from Okushiri Is. are generally larger.
The relative width of the pronoturn shows a tendency to become greater towards the north in
the south-easternmost part of Hokkaido (Oshima Peninsula) (see Discussion and Fig. 77).
Range: Most of the Oshima Peninsula, Hokkaido (including Okushiri Is.).
Specimens examined: 45 ( 2 4 9 T, 21 2 2) from the following localities: [Oshitna Province]: Hakodateshi: 1 q , Akagawa, vii 27 I968 (Y. Yasuda); 3 3 2 , ditto, vii 29 I968 (Y. Yasuda); 1 ? ,ditto, viii 16 I968 (Y.
Yasudaj; 19,ditto, viii 18 1968 (Y. Yasudaj; 1 9 , 2 2 2,ditto, viii 22 1969 (Y' Ota & S. Sato). Nanae-ch6:
1 $,Ohnuma-k6en, vii 28 I966 (K, Suga); 1 9 ,
ditto, viii 2 I985 (K. Kubota). Fukushima-ch6: 4 9 9 , 4 2 2 ,
Mt. Daisengendake, vii 30 1966 (K. Suga); 1 2 , ditto, vii 26 1969 (Y.Yasuda); 3 $ -?, 1 2 , ditto, 600 m,
(Kaminokuni-ch6), vii 23-25 1985 (K. Kubota). Y&umo-ch6: 12, Unseki-t6ge, 440 in, vii 31 I985 (K.
Kubota). [Shiribeshi Province1: Shimatnaki-mura: 4 9 ? , 2 2 2,Honme, 100 in, vii 28 I985 (K. Kubota); 3
99,5 2 2, ditto, vii 31 1985 (K. Kubopi); I 2 , Tomari, I00 in, vii 31-viii 1 I985 (K. Kubotaj. [liiyama
Provincel: Kumaishi-ch6: I $, Miichi-onsen, 440 m. vii 25 1985 (K. Kubota). [Okushiri Provincel: Okushirich6: 1 9 , Is. Okushiri, ix I6 1964 (W. Munakata); 2-9 -? , 12,ditto, viii 6 I969 (S. Sato & K. Itsuji).
60
Y. Takami and R. Ishikawa
The specimens are preserved in the R. Ishikawa Collection deposited at Department of Natural History,
Tokyo Metropolitan University.
Carabus (Ohomopterus) albrechti hidakanus Ishikawa & Takami
Apotomopterus albrerhti (pan.):Nakane, 1952, pp. 50-51, figs. 32-Sapporo & 33-Tomakomai.
Apotomopterus alhrerhti (part.): Nakane, 1953, p. 96 (48 in Japanese text), fig. 14: D-a, a'.
Apotomopterusjaponicus iilhrechti {iiart.): Nakane, 1962, p. 39, fig. 370-4, pi. IV: 61.
Apotomopterus ulbrechtl albrechti (part.): Ishikawa, 1969, pp. 520-521.
Carab~is(Ohomopterus) alhrechti albrechti (part.): Ishikawa, 1985, p. 21, pi. 4: fig. la.
Carabus (Olwmopterus) ulhrechti hidakanus Ishikawa & Takami, 1996, pp. 4 1-43, figs. 4-6,914; type-locality: Biratori, Hokkaido.
Carabus (Ohorriopterus) alhrechti albrechii {part.): Imura & Mizusawa, 1996, pp. 20 & 107,
fig. 44 (2).
This subspecies was not distinguished from C. a. albrechti until recently (Ishikawa and
Takami, 1996), though it shares aedeagal and pronotal characters with C. a. tohokuensis.
Distinguished by uniformly long thin apical portion of aedeagus.
Length: ?,23-26 mm; !,20.5-23 mm.
Upper surface of body polychromatic, dark copper, greenish copper, green or black.
Legs wholly black, but tibiae rarely somewhat rufous. Male antennae short, not reaching
middle of elytra; hairless area of underside of 5-7th segments concave but rudimentary,
about 113-1/2 as long as each segment. Head impunctate or at most weakly punctate from
front to neck. Inner margin of male protibia weakly convex but not angulate (Fig. 15).
Pronotum broad; disk less convex, with 3-5 marginal setae; postero-lateral corners weakly,
but broadly produced (Fig. 11). Elytra oblong and strongly convex; striae at most very
weakly notched.
Male genitalia: Apical portion of aedeagus very long, uniformly thin in lateral view; its
outer margin not grooved, barely outlined by rudimentary ridge; copulatory piece long (Fig.
4).
Female genitalia: Outer plate of vaginal apophysis long and narrow; anterior rim of inner
plate broad but narrower than C. a. albrechti; disc without median groove (Fig. 5).
Range: Hokkaido, Pacific coast from the environs of Sapporo east to the southern part of
Hidaka Province.
Specimens examined: 344 (178% ?, 1 6 6 2 2 ) from the frollowing localities: [Hidaka Province):
Biratori-ch6: 4 0 9 ?, 43 2 2, Biratori, 60 rn, viii 8-12 1976 (R. & F. Ishikawa) (paratypes from type locality). Erimo-ch6: 4 ? ?, 5 2 2,Oiwake-toge, vii 24 1969 (R. Ishikawa) (paratypes); 1 2 , Karniutahetsu, vii 28
1961 (T. Okumura) (paratype); 1 2, ditto, viii 13 196-5 (Y. Syu) (paratype): 8-? ?, 1 3 \ Horoman, Sarnanich6, viii 21-25 1980 (R. & F. Ishikawa) (paratypes). Samani-ch6: 1 2 , Mt. Apoidake, vii 29 1964 (T.
Okurnura) (paratype). Urakawa-ch6: 2 9 ?, 1 J\ Nishicha, vii 25 1964 (T. Okumura) (paratypes); 1 2 ,
Ikandai, 150 m, viii 17-19 1976 (R. & F. Ishikawa) (paratype). Mitsuishi-ch6: 3 -?, 1 0 2 i?, Toyooka, vi 21
1992 (K. Miyashita) (paratypes). Niikappu-ch6: 8 <? <?, 25 2 2 , TaiyO, 180 m, vii 10-13 1976 (R. & F.
Ishikawa) (paratypes); 21 <? ? , 9 $ j\ Wakazono, 100 m, viii 12-18 1978 (R. & F. Ishikawa) (paratypes); 19
Subspeciation and distribution pattern o f Curahus ufhrechti
Figs. 2-9. Carabus albrechti albrechti Morawitz ( 2 , 3 ) , C . a. hidakanus Ishikawa & Takami (4, S),and C. a.
itoc Ishikawa & Takami (6-9). [Copulatory piece and apical portion of aedeagus ( 2 , 4 , 6 and 8), and outer
and inner plates of vaginal apophysis (3, 5, 7 and 9 ) ) . 2 & 3, Akagawa, Hakodate-shi; 4 & 5, Biratori,
Biratori-ch6; 6 & 7, Rushin, Urahoro-ch6; 8 & 9, Mt. Rausudake, Rausu-ch6.
62
-?- ?, 6 2 2 , Biu, 200 m, viii 13-18 1978 (R. & F. Ishikawa) (paratypes); 5 9-?, 2 d" 2, ditto, viii 1 1969 (R.
Ishikawa) (paratypes). Monbetsu-ch6: 1 9 ,Miwa, viii 1 1969 (R. Ishikawa) (paratype). [Iburi Province):
Hobetsu-ch6: 1.59 9 , 11 2 j\ Niwa, 100 m, viii 8-12 1976 (R. & F. Ishikawa) (paratypes). Tomakomai-shi:
49
4 2 2 , Morappu, vii 17 1959 (R. Ishikawa) (paratypes): 3 9 ?, 1 0 2 J', ditto, vii 29 1959 (R. & F.
Ishikawa) (paratypes); 4 3 9 ?, 3 2 2 c?\ ditto, vii 6 1960 (R. & F. Ishikawa) (paratypes). [Ishikari Province):
Chitose-shi:! 2 , Shukuhai, v 30 1989 (T. Sugawara) (paratype). Sapporo-shi: 1 9 , Mt. Moiwayama, vii 17
1959 (K, Morimoto) (paratype); 1 9 , 2 2 2 , ditto, vi 14 1986 (K. Haga) (paratypes).
v,
Carabus (Ohomopterus)albrechti itoi Ishikawa & Takami
Apoiomopterus albrechti (part.): Nakane, 1953, p. 96 (48).
Apotomopterus japonicus albrechti [part.): Nakane, 1962, p. 39.
Apotomopterus albrechti albrechti (part.): Ishikawa, 1969. pp. 520-521.
Carabi~s(Ohomopterus} ulhrechti ulhrechti @art.): Ishikawa, 1985, p. 21.
Carabus (Ohomopterus) albrechti itoi Ishikawa & Takami, 1996, pp. 43-44, figs. 7-8, 15- 18;
type-locality: Rushin, Hokkaido.
The subspecies is distinguished from closely related C. a. hidakcttzus by the shorter and
thicker apical portion of the aedeagus. However, it possibly includes more than one discrete
geographical races. There are several small populations distributed sporadically in the east-
Figs. 10-17. Carab14~valbrechti albrechti Morawitz (10, 14), C . a. hidakunus Ishikawa & Takami
(1 1, IS), and C . a . itoi Ishikawa & Takarni (1 2, 13, 16 and 17). IPronoturn (10-1 3), and male
protibia (1 4-1 7)]. 10 & 14, Akagawa, Hakodate-shi; 1 1 & 1 5, Biratori, Biratori-ch6; 1 2 & 16,
Rushin, Urahoro-ch6; 1 3 & 17, Mt. Rausudake, Rausu-ct16.
Subspeciation and distribution pattern of Carahus albrcchti
63
ern part of Hokkaido which are slightly different from one another (see Discussion and
Ishikawa and Takami, 1996).
Length: -?-,
23-25 mm; j\ 22-23.5 mm.
Upper surface dark copper. Legs black, sometimes slightly rufous. Antennae short,
slightly exceeding basal 1/3 of elytra even in male; hairless area of male antennae vague,
less distinct than in C. a. hidakantis. Inner margin of male protibia weakly convex and not
angulate (Figs. 16 & 17). Head almost impunctate on front. Pronotum broad, with 2-4
(rarely 5 in male) marginal setae; lateral margins well convex anteriorly, postero-lateral
corners less produced than in C. a. hidahnus (Figs. 12 & 13). Elytra oblong and strongly
convex; striae at most very weakly notched.
Male genitalia: Apical portion of aedeagus long and thick; its outer margin not grooved,
barely outlined by rudimentary ridge; lateral margins of copulatory piece raised near base
(Figs. 6 & 8).
Female genitalia: Vaginal apophysis with outer plate broader than in C . a. hidukanus;
inner plate as in C. a. hidakanus (Figs. 7 & 9).
Variation: Specimens from the Shiretoko Peninsula differ from those from the type locality as follows: Head with coarser rugae from front to neck. Pronotum with transverse
rugae by median line. Apical part of aedeagus more cylindrical, feebly bent beyond middle;
lateral margins of copulatory piece more distinctly raised (Fig. 8). Outer plate of vaginal
apophysis shorter; inner plate with shallow median groove (Fig. 9).
Ranges: Hokkaido: the Shiranuka Hills and the Shiretoko Peninsula.
Specimens examined: 14 (7 Â¥? Â¥?-,2 o f ) from the following localities: [Tokachi Province]: Urahoro-ch6:
6 9 ?-, 5 2 2, Rushin, vi 19 1994 (K. It61 (paratypes from type locality). INemuro Province]: Rausu-ch6: 1
9,Mt. Rausudake, viii 2 1964 (T. Okumura) (paratype); 1 2 , ditto, viii 23 1963 (T. Okumura) (paratype); 1
2 , ditto, viii 3 1964 (T. Okumura) (paratype).
Carabus (Ohomopterus) albrechti tohokuensis Ishikawa
Carabus (Ohomopterus) albrechti tohokuensis (part.) Ishikawa, 1984, pp. 3-5: type-locality:
Matsuzono, Iwate Prefecture.
Carabus (Ohomopterus) albrechti tohokuensis (part.):Ishikawa, 1985, p. 21, PI. 4; I b.
Carabus (Ohomopte.rus)albrechti tuhok~iensis:Imura & Mizusawa, 1996, pp. 20 & 107, fig. 44
(3).
This subspecies is widely distributed on the Pacific coast of the T6hoku District where it
shows noticeable geographic variation. Especially the size of body varies considerably by
the locality, but the apical part of aedeagus is rather uniform within the range.
Length: ?, 19-24.5 mm; 2 , 18.5-23 mm.
Upper surface of body copper, black or green individuals unknown. Legs nearly wholly
black. Male antennae very short, barely reaching beyond 1/3 of elytra; hairless depressions
64
on underside of 5-7th segments rudimentary, weakly concave and about 1/3-1/2 as long as
each segment. Head impunctate, but partly with shallow rugae. Inner margin of male
protibia very weakly convex (Figs. 28 & 29). Pronotum looking round in outline; lateral
margins roundly arched and less sinuous; disc well convex, with 3-5 marginal setae;
postero-lateral corners weakly but broadly produced and rounded (Figs. 24 & 25). Elytra
short and strongly convex above; lateral margins rounded; striae at most very weakly
notched.
Male genitalia: Apical portion of aedeagus cylindrical, its outer margin not grooved;
copulatory piece long (Figs. 18 & 20).
Female genitalia: Outer plate of vaginal apophysis with lateral margins arched, less convergent posteriorly; inner plate with anterior rim broad, median groove absent (Figs. 19 &
2 1).
Ranges: The Pacific coast of the T6hoku District, the greater part of Iwate and restricted
areas of Aomori, Akita and Miyagi prefectures. In the environs of Hirosaki-shi, Aomori
Prefecture, it intergrades with C. a. yamauchii (Fig. 26).
Specimens examined: 285 (158 ? ?, 125 of of) from the following localities: [Aomori Prefecture]: Mutsushi: 11 9, 14 2 d\ Nagasaki, 100 m, Mt. Osorezan, viii 27-28 1995 (Y. Takami & T. Ishida); 1 ?, 1 of, near
Ipponsugi, Mt. Osorezan, viii 27-28 1995 (Y. Takami & T . Ishida). Ohata-machi: 4 ? ?, Akataki, ix 27 1987
(S. Yamauchi). Higashidfiri-mura: 1 0 9 9-,8 2 2 , Odanosawa, vii 1 1987 (S. Yamauchi); 5? ? , HiyamizutOge, x 7 1987 (S. Yamauchi). Gonohe-shi: 2 ? 9,Shitogishi, viii 25-26 1983 (R. & F. Ishikawa) (paratypes).
Hachinohe-shi: 1 2, no further data, vi 5 1983 (J. Kojima) (paratype). Nagawa-ch6: 1 9 , Mt. Nakuidake, 200
m, Mizusawa, viii 24-26 1983 (R. & F. Ishikawa) (paratype). Towada-shi: 5? -?, 1 2 m , Mise, vii 23 1990
(S. Yamauchi); 2 ? ? , 2 2 2 , ditto, vii 29 1990 (S. Yamauchi); 9 9<? , 10 2 2 , Sashikubo, vii 20 1990 (S.
Yamauchi); 5 '?^-, 7 <? 2, ditto, vii 23 1990 (S. Yamauchi). Shingo-mura: 14 ? ?, 13 2 J", Shikata, vii 29
1990 (S. Yamauchi); 1 ?, Gongennotaki, vii 23 1990 (S. Yamauchi); 1 -?. ditto, vii 29 1990 (S. Yamauchi); 1
?, Mt. Nanamori, vii 20 1990 (S. Yamauchi). Towadako-machi: 1 ?, Tsuta, 470 m, Mt. Hakkodasan, viii 1416 1985 (J. Kojima): Aomori-shi: 1 2 , Jogakura, vi 26 1994 (R. Hatano). [Iwate Prefecture]: Morioka-shi: 55
? 7,3 6 2 d\ Matsuzono, 200 m, vi 28-30 1980-ix 4 1981 (Y. Maeta) (paratypes from type locality); 12,
Mt. Futatsumoriyama, iv 6 1963 (T. Abe) (paratype). Iwaizumi-cho: 2 ? ? ,Shigawatarido Cave. Akka, vii 27
1967 (S. Ukno) (paratypes); 1 ?, Ryusenkutsu Cave, Sawamagari, vii 3 1954 (S. Ueno) (paratype); 12,
Osawa, north of Miyako-shi, iv 7 1963 (T. Abe) (paratype). Niisato-mura: 4 9 9 ,Genbeidaira, 800 tn, ix 1521 1981 (K. 0kubo) (paratypes). Miyako-shi: 4 9 9 , Matsuyama, 10 ni, ix 12-15 1981 (R. & F. Ishikawa)
(paratypes). Kawai-mura: 1 J\ Kuzubesawa, 550 m, vi 14 1987 (K. Haga). Yamada-shi: 19,Shimotanabe,
70 m, Toyomane, iv 3 1980 (R. & F. Ishikawa) (paratype). T6wa-ch6: 2 of S\ Yokomine. by Taseko Lake, viii
23-24 1983 (R. & F. Ishikawa). Kitakami-shi: 3 9 ?, 2 3 2 , Sawamagari. 150m, Yokokawame, ix 8-1 1 1984
(R. & F. Ishikawa) (paratypes); 1 ?, 1 of, Iwasawa, 180 m,ix 7-10 1984 (R. & F. Ishikawa). Yuda-machi: 1 9,
Torasawa, 270 m, Shitamae, ix 8- 10 1984 (R. & F. Ishikawa). Ofunato-shi: 5 ? -9,
2 C? of, NakaitayO, 40 m,
Hikoroichi, iv 4 1980 (R. & F. Ishikawa) (paratypes). [Akita Prefecture]: Tashiro-machi: 1 2 , Obuchi, vii 8
1990 (S. Yarnauchi); 1 9 , Sotokawara, 60 m, viii 28-30 1982 (R. & F. Ishikawa) (paratype). IMiyagi Prefecture]: Kesennuma-shi: 1 ?, 2 of 2, Kanetori, 120 m, iv 2 1976 (R. & F. Ishikawa).
Other specimens examined: [Population transitional with C . a. yamauchii]: 6? ?, 6 2 j\ Tennosawa,
Hirosaki-shi, Aomori Prefecture, vii 21 1995 (R. Hatano); 1 J\ ditto, vi 15 1992 (R. Hatano).
Subspeciation and distribution pattern of Cur~zbii~'i
ulbrechti
Carabus (Ohomopterus)albrechti yamauchii subsp. nov.
Curub~is(Ohomopterus)albrechti tohokumsis (part.) Ishikawa, 1984, pp. 3-5.
Carahtis (Ohomopterus) ulhrechti tohokuensis (part.): Ishikawa, 1985, p. 21, PI. 4; I b
This new subspecies is distinguished from C. a.tohokuensis by narrower pronoturn with
more angulate postero-lateral corners, slender elytra and the shape of aedeagus.
Length: 9, 20.5-23 tnm; 2,20-21.5 mm.
Upper surface of body copper or dark copper. Legs mostly wholly black. Male antennae
barely reaching beyond middle of elytra; underside of 6-7th segments with hairless depres-
Figs. 18-23. Carobus ulbrechtl tohokuensis Ishikawa (18-21), and C . a. yutnuuchii subsp. nov. (22, 23),.
[Copulatory piece and apical portion of aedeagus (1 8, 20 and 22), and outer and inner plates of vaginal
apophysis (19,21 and 23)l. 18 & 19, Matsuzono, Morioka-shi; 20 & 21, Nagasaka, Mt. Osorezan, Mutsushi; 22 & 23, M t . Shikarigatake, Ajigasawa-rnachi,
66
sion, about 1/2-2/3 as long as each segment, but vague on 5th segment. Head with vertex
weakly punctate or impunctate. Inner margin of male protibia weakly convex (Fig. 30).
Pronotum narrow; lateral margins weakly cordately sinuous; disc with 3-5 marginal setae;
postero-lateral corners distinctly and angulately produced (Fig. 27). Elytra slender and less
convex; striae weakly notched.
Male genitalia: Apical portion of aedeagus weakly grooved or flat, outlined by rudimentary ridge on its outer margin; copulatory piece long (Fig. 22).
Female genitalia: Outer plate of vaginal apophysis weakly narrowed posteriorly with
lateral margins arched; inner plate with shallow median groove, anterior rim broad (Fig.
23).
Range: The Shirakami Mountains on the border between Aomori and Akita prefectures
near the Japan Sea.
Etymology: Named after Mr. Satoshi Yamauchi, who collected the type series.
Holotype: 2,Mi. Shikarigadake, Ajigasawa-machi, Aomori Prefecture, vii 27 1985 (S. Yamauchi) deposited at Tokyo Metropolitan University.
Paratypes from type locality: 6? ?, 1 2 2 2 , same data as holotype.
Other paratypes: [Aomori Prefecture]: 39 9 , 2 2 J\ Ofuna-rind& Fukaura-machi, i v 16 1996 (R.
Hatano); 2 2 2, Kurosaki, Iwasaki-mura, v 17 1996 (R. Hatano): 2 ? ?, dkawa, Nishirneya-mura, viii 21
1984 (S. Yamauchi) (paratypes of C. a. toh(~ki~ensis).
IAkita Prefecture]: 3 9 9 , Yamaya, Noshiro-shi, vii 8
1990 (S. Yamauchi): 1 ?-, Taraky6, Fujisato-machi, vii 8 1990 ( S . Yamauchi); 1 -?, dkawamegawa, Tashiromachi, vii 8 1990 (S. Yamauchi).
Figs. 24-30. Carahus albrechti tohokuensis Sshikawa (24, 25, 28 and 29). C. a . t o h k u e n s i s transitional to C. a. yurnaurhii (26), and C. a. yamauchii subsp. nov. (27,301. [Pronotum (24-27), and
male protibia (28-30)J. 24 & 28, Matsuzono, Morioka-shi; 25 & 29, Nagasaka, Mt. Osorezan,
Mutsu-shi; 26, TennOsawa, Hirosaki-shi; 27 & 30, Mt. Shikarigatake, Ajigasawa-machi.
Subspeciation and distribution pattern of C a t - u h s ulbrvchti
67
Other specimens examined: [Population transitional with C. a . tohokitmsis\: see the description of C. a .
tolwkuert.~i.~.
Carabus (Ohomopterus) albrechti hagai subsp. nov.
Carabus (Ohomoptvrus) albrrchti tohokucn.si.s (part.) Ishikawa, 1984, pp. 3-5.
Curabus (OJzomopterus) ulbrechti tohokuensis (jmrt.): Ishikawa, 1985, p. 2 1, PI. 4; 1 b,
This new subspecies is distinguished from C. a. tohok~~etzsis
by the slender body, the
feature of male antennae, strongly notched elytral striae and the shape of the aedeagus.
Length: ? , 19.5-24 mm; j\ 19.5-22.5 mm.
Upper surface of body dark copper. Legs wholly black or tibiae rufous. Male antennae
barely reaching beyond middle of elytra; underside of 5-7th segments with hairless depressions, about 112-213 as long as each segment. Head weakly punctate on vertex. Inner
margin of male protibia roundly convex (Figs. 43 & 44). Lateral margins of pronotum
cordately sinuous, with 3-5 marginal setae; postero-lateral corners distinctly produced and
rounded (Figs. 39 & 40). Elytra slender, with shoulder weakly produced and weakly convex above; striae strongly notched.
Male genitalia: Apical portion of aedeagus with narrow flat part on its outer margin
outlined by rudimentary ridges; copulatory piece long (Figs. 3 1 & 33).
Female genitalia: Outer plate of vaginal apophysis triangular; inner plate with shallow
median groove, anterior rim broad (Figs. 32 & 34).
Range: The central part of the T6hoku District, from the Aganogawa River in Niigata
Prefecture and the northern part of Fukushima Prefecture north to the middle of Akita and
Miyagi prefectures, where it presumably intergrades with C. a.tohokuensis.
Etymology: Named after Mr. Kaoru Haga, who collected the type series.
Holotype: 2 , Shinmachi, Yuzawa-shi, Akita Prefecture, viii 10-13 1986 (K. Haga) deposited at Tokyo
Metropolitan University.
Paratypes from type locality: 17 9 '?,5 2 c?\ same data as holotype.
Other paratypes: 1 2, Sakai, Itado, Higashiyuri-machi, Akita Prefecture, ix 9-10 1984 (R. & F. Ishikawa)
(paratype of C. a . tohokuensis); 1 ? , 1 d\ Sunakoseki, 400 m, Nishikawa-machi. Yamagata Prefecture, vii 24
1981 (K. Haga); 2 9 9,1 2, Mizutani, Shibata-shi, Niigata Prefecture, vi 18 1967 (H. Koike). Miyagi Prefecture: 3 9 ? , 3 J" J", Matsushima-ch6, xi 15 1974 (R. & F. Ishikawa) (paratypes of C. a . tol7okiivnsis); 1 ? , 1
cf, Taihakusan, Sendai-shi, iv 9 1963 (T. Abe) (paratypes o f C . a. tohokuensis); 6-? 9 ,4 2 2 , Mt.
Takamoriyama, 80 m, Rifu-ch6 - Sendai-shi, iv 1 1 1975 (R. & V. Ishikawa); 4 9 ? , 4 rf" 2, Gongennomori,
Sendai-shi, iv 26 1993 (M. Ujiie & Y. Shimamura); 2 3 3,ditto, xi 6 1994 (M. Ujiie); 3 9,1 2 , Misumi,
500 m, near Togatta, Shiroishi-shi, xi 19 1966 (R. & F. Ishikawa) (paratypes of C. a . toJzokueri.sis); 2 9 -?, 2 cf
,'J Edano, Kakuda-shi, iv 12 1963 (T. Abe) (paratypes of C. a. tohokuensis}; 4 9 y,3 2 2,0uchi-tnura,
Marumori-machi, xii 21 1963 (T. Abe) (paratypes of C. a . to/zc)k~(ensis).
Other specimens examined: [Populations transitional with C. a . t,sukuhani~.s]:Fukushiina Prefecture: lo'?
9 , 10 2 2,Tachi, Haramachi-shi, ix 24-25 1995 (M. Ujiie & Y. Shimamura); 7 7--? , 3 2 2, Minamiyunoki,
Kashima-machi, ix 24-25 1995 (M. Ujiie & Y. Shimamura); 2 2 J\ Yufune, litate-mum, ix 24-25 1995 (M.
Figs. 31-38. Carahus albrechti hag& subsp. nov. (31-34), C. u. aw~ashimaeIshikawa & Takami
(35, 361, and C. a.freyi Emden (37,38). [Copulatory piece and apical portion of aedeagus (31,
33, 35 and 37), and outer and inner plates of vaginal apophysis (32, 34, 3 6 and 38)j. 3 1 & 32,
Shinmachi, Yuzawa-shi; 3 3 & 34, Gongen-no-mori, Sendai-shi; 35 & 36, Is. Awashima; 37,
Shinbogawa, Kanai-machi, Is. Sado; 38, Katagami, Niibo-mura, Is. Sado.
Subspeciation and distribution pattern of Curabus albrcrhti
Figs. 39-46. Carubus ulbrechti hugai suhsp. nov. (39,40,43 and 44), C. a. uw~ushimueIshikawa & Takami
(41, 45), and C. a. f r q i Emden (42, 46). [Pronotum (39-42), and male protibia (43-46)l. 39 & 43,
Shinmachi, Yuzawa-shi; 40 & 44, Gongen-no-mori, Sendai-shi; 41 & 45, Is. Awashima; 42, AikawaAono-tOge, Is. Sado; 46, Katagami, Niibo-mum, Is. Sado.
Ujiie & Y. Shimamura); 1 S\ Okura, litate-mura, ix 24-25 1995 (M. Ujiie & Y. Shirnamura).
The specimens are preserved in the R . Ishikawa Collection deposited at Department of Natural History,
Carabus (Ohomopterus) albrechti awashimae Ishikawa & Takami
Apotomopterus jupotz~cusfreyi (part.): Nakane, 1962, pp. 37-39.
Apotomopter~~s
alhrechti (part.):Ishikawa, 1969, pp. 520-521.
Carabus (Ohomopterus)ulbrechti uwushimae Ishikawa & Takami, 1996, pp. 46-47, figs. 25,
26 & 30; type-locality: Awashima Is., Niigaia Prefecture.
This subspecies endemic to Awashima Is., off the coast of Niigata Prefecture, is distinguished superficially from C. a. freyi of Sado Is. by strongly notched elytral striae and
uniformly dark brownish or reddish copper on the upper surface of the body.
Length: 9 , 22.5-24.5 mm; g\ 20.5-22.5 mm.
Upper surface of body dark brownish or reddish copper; greenish or black individuals
unknown. Legs and palpi wholly black. Head weakly punctate on vertex. Male antennae
short, barely reaching middle of elytra; underside of 5-7th segments with vaguely outlined
hairless areas, about 1/3-112 as long as each segment. Inner margin of male protibia roundly
convex (Fig. 45). Pronotum with postero-lateral corners strongly produced but rounded;
disc with 3-4 marginal setae; punctation of disc sparser medially (Fig. 41). Elytra less
70
convex above with shoulders strongly produced, striae strongly notched.
Male genitalia: Apical part of aedeagus thicker, with flat area on its outer margin; copulatory piece with rounded apex, constricted at base (Fig. 35).
Female genitalia: Outer plate of vaginal apophysis weakly narrowed posteriorly; inner
plate with anterior rim broad, with shallow median groove (Fig. 36).
Range: Awashima Is., Niigata Prefecture.
Specimens examined: 37 (17 2 9 , 2 0 2 2)from the following localities: [Niigata Prefecture]: Awashima
Is.: 13 f 9.1 6 2 3\Is. Awashinia, x 25-28 1963 (R. Ishikawa) (paratypes from type locality); 4 '? ?, 4
Kamaya, vii 21-23 1991 (T. Suda) (paratypes from type locality).
m,
Carabus (Ohomopterus) albrechti freyi Emden
Carabuts (0hotnopter~i.s)Alhret'hti Freyi Ernden, 1932, p. 62: type-locality: "Japan: Insel Sado"
Curahus (Apotorwpterus) alhrerhti {part.): Breuning, 1932, p. 235-236.
A/wiomopt~~rus
ulhwchti (purt.): Breuning, 1934, p. 31.
Apo~ftmpteru.s,jupo~iii~~~.s
j'reyi: Nakane, 1962, p. 37-39, Fig. 3%-w.
A~~tomoptorusju~~on'u'usf'reyi
(part): Nakane, 1963. p. 1 1- 12.
Ai~tmnoptrrusa,!hrechtifreyi: Ishikawa, 1969, p. 520-521, figs. 2 & 22..
Caruhu.~(Ohomopterus) albrerhti f r q i : Ishikawa, 1985, p. 21, pi. 4: Ie.
Ciiruhiis (0homopteru.s) ulbrecktifrey': Imura & Mizusawa, 1996, pp. 20 & 107, fig. 44 (4).
This subspecies which is endemic to Sado Is. is distinct superficially in having elongate,
parallel-sided elytra with smoother striae. The upper surface of the body is polychromatic
though dark.
Length: ?- , 20.5-25 mm; 2 , 19.5-24 mm.
Upper surface of body polychromatic, dark copper or greenish copper. Legs and palpi
wholly black. Head nearly impunctate. Male antennae barely reaching beyond middle of
elytra; underside of 5-7th segments concave, with hairless areas about 113-112 as long as
each segment. Inner margin of male protibia angulately convex (Fig. 46). Pronotum with
posteio-lateral corners strongly produced and rounded; disc with 3-4 marginal setae; punctures fused with one another to form wrinkles (Fig. 42). Elytra slender, weakly convex
above: lateral margins less arched and subparallel in male; striae weakly notched.
Male genitalia: Apical portion of aedeagus with flat part on st3 outer margin; copulatory
piece long, feebly constricted at the base, (Fig. 37).
Female genitalia: Outer plate of vaginal apophysis broad and triangular; inner plate with
anterior rim broad, smooth at bottom without median groove (Fig. 38).
Range: Sado Is., Niigata Prefecture.
Specimens examined: 360 (237 -?- 9, 123 2 2 )from the following localities: [Niigata Prefecture]: Sado
Is.: Aihawa-machi: 3-? 9-,
3 d7' 2, Aikawa-Aono-t6ge. iv 21 1960 (R.& F. Ishikawa). Niibo-mum: 3 ? I?, 1
2,Katagami, 3 m, v 12-15 1987 (R.& F. Ishikawa). Kanni-machi: 1849 ?, 80 o" d", Shinbogawa, 110 m, v
Subspeciation and distribution pattern ofCuruhu'i u l b m hti
71
13-15 1987 (R. & F. Ishikawa). Akadomari-niura: 3 ?$-, 1 d\ Kainikawaino, 360 in, iv 13 1996 (Y. Takami).
Mano-machi: 3 0 9 -?-, 28 3' 2, Yoshioka, 50 m, v 12-14 1987 (R. & F. Ishikawa): 7 ? ?, 5 2 2, Shizudaira,
200 ni, iv 13 1996 (M. Ujiie, Y. Shiniainura& Y. Takami): 1 ? ,ditto, 240 in (Y. Takami). Hatano-machi: I ? ,
I 0".Maruyama, 220 in, iv 14 1996 (Y. Takami); 1 3\Koi-iianoue, 320 m, iv 14 1996 (Y. Takaini); 2 ? -9,
I 2,
Bogaura-rinc16, 260 m, iv 14 1996 (Y. Takaini): 3 ? ? , 2 3 2, Hase, 160 m, iv 14 1996 (Y. Takami).
Carabus (Ohomopterus)albrechti esakianus (Nakane)
Apotomopteriis ulbrechti vsakii (nee Csiki): Nakane, 1952a, p. 13; range: "Kanto" (in Japanese).
Apotompterus alhrechti race esakb (nec Csiki): Nakane, 1952b, p. 50, Fig. "Takaosan"; range:
"Kant6" (in Japanese).
Apotomopterus albrechti esakii (nec Csiki): Nakane, 1953, p. 96, Fig. 15; range: "confined in
Kwanto District including Tokyo".
Apotomopterus japonicus esukiunus Nakane, 1960, pp. A99, A 100, A 106; range: "Kwanto
Plain". New name for "Subspecies" "in the Kwanto Plain".
Apotomopterus japonicus esakianus: Nakane, 196 1 , p. 1. ["nom. nov." for "Curahus (A .)
japonicus ab. esakii Breuning (nec Csiki).... 1932'' and "A. a1t)rvchti esukii Nakane (nec
Csiki)", 1953.1
Apotomopterusjaponicus esukiunus {part.): Nakane, 1962, p.37, Figs. P;. IV, 59 and 60; range:
"Yamanashi Prefecture and the areas around the Kanto Plain" (in Japanese).
Apotonzopteru.sjupo~z~cus
esukianiis: Nakane, 1963, pp. 1 1 - 12, PI. 6,2h, 21; range: "Kanto District" (in Japanese).
Apotomopterus albrechli e,sakianu.s {iMrf.):Ishikawa, 1969, pp. 520-521, Fig. 23.
Curabr~.~
(0homopterii.s) albrechti esakianus (part.): Ishikawa, 1985, p. 21, PI. 4, 1c & Id. (in
Japanese).
Carabu.~{Ohomopterii,~)ulbrechti esukiunu.~(part.): Imura & Mizusawa, 1996, pp. 107. (in
Japanese).
Nakane (1952a) discriminated the populations of Carahus albrechti with "red legs" of
'Kanto Heiyal'(=Plain)as "alhrechti esukii Csiki" from those with wholly black legs of the
adjacent regions. Following this (Nakane, 1952b). he called the same populations from the
'Kanto Heichi" (=Plain) the "race esakil Csiki" and illustrated the aedeagi and copulatory
pieces of the specimens from "Masuton~i"(Yamanashi Prefecture) and "Takaosan" (Tokyo), and the male protibki of the former. In his third paper (Nakane, 1953) treating this
subspecies, he defined Apolornopterus ulbrechti muhii and illustrated the aedeagi and copulatory pieces of the specimens from "Ibaragi, Hitachi", "Masutomi, Kai" and of "Tanzawa,
Sagami". However, the locality "Masutomi" is located in Yamanashi Prefecture in the
Chfibu District, and the populations of C. albrechti of the area including this region are
distinct from those of the Kant6 District as they were named later A. albrechti okiimwai
(Ishikawa, 1966) (=C. albrechti okumurui (Ishikawa)), and the figures of "Tanzawa" are
Breuning.
not of C . albrechti. but of C . lewisia~~u.~
Since the name esakii Csiki (replacement name for lewisi Gkhin, 1885) was given to the
72
species which Nahane (1953) described as ^hizuokat?t~.sis,Nakane ( 1961) proposed a new
name esakiuniis for "C. (
4
juponii'us ab. m k i i Breuning (nec Csiki)....1932" and for "A.
alhrechti csukil Nakane (nee Csiki)", 1953. However, since no available name had been
given to this subspecies, his proposal of new name was in fact the proposal of a new subspecies. Nakane ( 1 960) used the name esakiunus one year before this proposal for this subspecies of "Kanto Plain".
In the revision of the Japanese Carabinae, Nakane (1962) redescribed A . ulbrechti
esakiams found in Yamanashi Prefecture and the areas around the "Kanto Plain" and presented the figures of genitalia which are the same as shown in his previous paper (Nakane,
1953), but illustrated in the color plate two male specimens from "Takao, Tokyo" and
"'I~ainagawa,Kanagawa" as A. ulbrec hti esukiatz~~s.
The type-locality of A. japorzicus esakianus was never designated definitely by the author of this subspecies, and the range of the subspecies was outlined vaguely "Kanto"
(19523 and 195%). "Kwanto District" ( 1953), "Kwanto Plain" ( 1 960), "Yamanashi Prefecture and the areas around the Kanto Plain" (1962) and finally "Kanto District" (1963).
We propose here to designate as lectotype of A . ,japouicii^ esakiut~ii.\Nakane, a male
from "Takaosan" of which copulatory piece was illustrated for the first time by the author
(Nakane, 1952b), since this is the earliest concrete indication of this subspecies from the
given range, "Kanto Heiya" (ICZN, Article 74 (4). The type-locality "Takao" (=Mt.
Takaosan, Hacliioji-shi, Tokyo) as thus designated is only one of the two localities referred
to but situated in the Kanto Plain (Nakane. 1952b).
Length: -?,20.5-23.5 mm; 2, 19.5-21.5 mm.
Upper surface of body polychromatic, bright or dark copper, greenish copper, green or
black. Legs with tibiae and tarsi more or less rufous or wholly black. Male antennae exceeding middle of elytra; underside of 5-7th segments distinctly concave, with hairless areas a-i long as each segment. Head almost impiinctate from front to neck. Inner margin of
male protibia angulately convex (Figs. 59 & 60). Pronotum appearing round in outline,
postero-lateral corners strongly produced and rounded, with 3-6 marginal setae (Figs. 55 &
56). Elytra slender, lateral margins less arched at anterior half; striae weakly notched.
Male genitalia: Apical portion of aedeagus with finger-shaped tip, its outer margin
weakly grooved; copulatory piece triangular, but shorter (Figs. 47 & 49).
Female genitalia: Outer plate of vaginal apophysis triangular; inner plate with shallow
median groove; anterior rim broad (Figs. 48 & 50).
Distribution of color variation: The ratio of greenish individuals is higher towards the
northwestern part of Kant6 District than elsewhere. The specimens from the environs of
Karuizawa-machi, Nagano Prefecture, are predon~inantlylustrous green.
Range: The hilly regions around the Kant6 Plain of Gunma and Saitan~aprefectures and
part of Tochigi, Nagano, Yamanashi, Kanagawa, Tokyo and Chiba prefectures. At the
northern marginal areas of its range, middle part of Tochigi Prefecture, it intergrades with
C . u. t.sukitharz~i.\. In Ueda-shi, Nagano Prefecture, it is parapatric with C. a. okz~rt~z~rai
at the
Kamigawa River. In the systems of the Tamagawa and Sagamigawa rivers including the
Tanzawa Mountains, it i s parapatric with C. 1. lewisianus. In Chiba Prefecture, its range is
Subspeciation and distribution pattern of Carahns ulhwchti
73
restricted to the BQsQPeninsula at the environs of Mt. Kano~an,where it is surrounded by
C. lewisianus awakazusanus parapatrically (Shida, 1984).
Specimens examined: 1765 (1 152? 9 ,613 3' 3') from the following localities: [Tokyo Metropolis] :
Hachioji-shi: 59- -?, 4 m , Mt. Takaosan, x 27 1958 (R. Ishikawa): 4 ? ?, ditto, xi 6 1961 (R. & F.
Ishikawa); 21 ? -? , 13 <? 2,ditto, xi 14 1963 (R. Ishikawa): 4Â -9,
3 3' 2,ditto, x 29-30 1964 (R. Ishikawa);
ditto, xii 3 1961 (H.Maruokaj; 5 ? 9,
1 J\ ditto, v 31 1963 (S. Kondo); 5 ? ? , 3 2 2,ditto,
1 3 9 ?, 8
ditto, xi 6 1961: 9-? ?-, 9 2 2 , Oclarumi,
viii 5 1964 (S. Kondoj; 2 9 %, ditto, viii 1995 (T. Sait6); 4 9 9-,
ditto, iv 6 1964 (R. Ishikawa); 6? ?, 3 2 2,ogata, xi 28 1959 (R. Ishikawa); 2-? ?, 4 2 2,Tachimachi, viii
8 I984 (A. Makimoto); 2'??, 4 2 2 , Kawamachi, 210 m. iii 5 1994 (Y. Takarnij: 3 'i' ?, 3 3' 2 , ditto, iii 13
1994 (Y. Takami & M. Momiyamaj; 35 ? ?, 20 2" 2, ditto, v 28-30 1994 (Y. Takami); 2 ? ?, 7 c? 2". ditto,
iii 28 1995 (Y. Takami). Itsukaichi-shi: 2-? ?, 1 2, Aokidaira, 250 m, vi 13-20 1984 (Y. Inouej: 23 9--?-, 9^
2, Nitago, 240 In, vi 13-20 1984 (Y. Inoue). 0me-shi: 2 2 3'. nr Umegaya-tOge, 240-320 m, ix 27-28 1985
(T. Shiina). Okutama-eh6: 60? ? , 3 8 2 2 , Mt. Kawanoriyama, 1 150-1350 rn, vi 9-30 1984 (T. Shimornura);
8 ? ? , 8 2 2 , ditto, 736 m, v 8-15 1985 (A. Makimotoj; 1 j\ ditto, 777 rn,v 8-15 1985 (A. Makimoto): 1 9 ,
di~to,435 m, v 15 1985 (A, Makimoto); 32 ? ?, 13 2 2, Bonornine-Nagaotnaru, 900-950 rn, vii 3- 10 1984
(T. Shimomura); 1 9 , Semizawa, 280 m, dtabagawa, vii 2-3 1984 (R. Ishikawa); 1 2 , Konakachaya. 350 m,
Otabagawa, vii 2-3 1984 (R. Ishikawa): 1 1 -? ?. 1 2 , Yokosuzu-one, 900 m,Mitsudokke, vi 10- 1 1 1984 (T.
Shimomura); 31 ? ?, 1 9 2 2,ditto, vi ll-vii 1 1984 (T. Shimomuraj; 109 ?. 2 2 2 , ditto, 1000 m, vi 10-1 1
1984 (T. Shimomura); 5 ? ? , 4 2 2 , ditto, vi 1 1-vii 1 1984 (T. Shimomura); 3 ? -? , ditto, 1300 m, vi 11-vii
I 1984 (T, Shimomura); I ? , ditto, 1380 m, vi 10-11 1984 (T. Shimomura); 3?- ? , I 2,ditto, 1380 m, vi I Ivii 1 1984 (T. Shimomura); 10? ?, 3 2 2 , Mitsudokke-Mt. Sobatsubuyama, 1450-1473 m. vi 10-1 1 1984
(T. Shimomuraj; 31 ? -?, I3 <^^, ditto, vi 1 1-vii I 1984 (T. Shimomura); 4-9 ? . 1 J\ Mitsudokke-Mt.
Toridaniyama, 1506-1605 m, v 22-29 1985 (A. Makimoto); 7Â ?, 4 2 c7'. ditto, v 29-vi 7 1985 (A.
Makimoto): 3?- 9,3 2 c?\ Ogawadani, 900 m. v 19 1984 (T. Shimomura); 23 ? 9,
\lS 2. Kotozawa, 270
m, Mt, Mitakesan, viii 22-24 1985 (R. Ishikawa); 17 -? ? , 5 if j\ Mt. 0takesan. 1000- 1200 m, vi 7 1984 (T.
Shimomura); 67 9 9,
13 2 2 , Mt. Mitakesan-Mt, ~takesan,1000-1250 ni, vi 7-19 1984 (T. Shimomura); 15
?- -?-, 7 2 S\ ditto, v 9-23 1985 (T. Shimomura); 4-? ? , 1 2. Mt. Gozenyama, 1250-1400 m, v 10-24 1985
(T. Shimomura); 20-?- ?, 1 6 2 2, ditto, vi 3-4 1985 (T. Shimomura): 63 ? 9 , 8 3 3\Atami, 600-650 m, vi
10-25 1984 (A. Makimoto); 2 9 -?, Atami-Mt. Kuratoyama, 930-950 in, vi 25 1984 (A. Makimotoj; 1 0 9 ?-,
5 2 d\ Kayanoki-one, 1250-1350 m, Mt. Takanosuyama, vi 10-17 1984 (A. Makimoto); 2 5 9 -? , 1 6 2 2 ,
ditto, vi 17-25 1984 (A. Makimoto); 2 % ?, 1 J", ditto, 1550- 1600 in,vi 10-17 1984 (A. Makimoto); 2 9 ?-,
ditto, 1550-1600 m, vi 17-25 1984 (A. Makimoto); 3 ? ?, 2 2 S\ ditto. 1600-1700 m, vi 10-17 1984 (A.
Makimoto); 19, 5 2 2, ditto, 1620-1730 m, vi 17-25 1984 (A. Makimoto); 4 ? '?-,Mt. MutsuishiyamaMizuneyama, 1260- 1450 m. v 11-25 1985 (T. Shimomura); 1 ? , Mt. Kumotoriyama, 1700- 1870 m, vi 1 1-14
1985 (T. Shimomura); 1 9 , 1 2 , nr Okutama-station, 400 m, x 2-8 1985 (T. Shiina); 1 ? , Murosawa-tunnel,
600 m, nr Atami, x 2-8 1985 (T. Shiina); 2 2 2, Umezawa, 300 In, Kawai, ix 27-28 1985 (T. Shiina).
Hinohara-mura: 2 9 ? , I 2 , Asama-one Tozanguchi, 520 m, Minamiakigawa, vi 13-20 1984 (Y. Inoue); 1 9,
1 2, Kamikawanori, 420 m, Minamiakigawa, vi 13-20 1984 (Y. Inoue); 2 ? -? , 1 if, Asama-tOge-michi, 460
m, Minamiakigawa, vi 13-20 1984 (Y. Inoue); 2 9 ?, Kashiwagino, 320 tn, Minamiakigawa, vi 13-20 1984
( Y . Inoue); 2% $, d n o , 340 In, Nang6, Minamiakigawa, vi 13-20 1984 (Y. Inoue); & ?-, 12,
Minamiakigawa, 480 m, v 19-27 1985 (T. Shimon~ura);1 -?, Tokisaka, 480 m, Kitaakigawa. vi 13-20 1984
(Y. lnoue); 8 ? ? , 2 m , Motog6,320 m, Motojuku, vi 13-20 1984 (Y. Inoue); 6 ? -? , 1 2, Sasa-one, 1092
in, v 19-27 1985 (T. Shimomura); 1 9, 1 2,ditto, 1052 m, v 19-27 1985 (T. Shimomura): 4 ? ? , 3 2 2,ditto,
S\ditto, 1042 nl, v 19-27 1985 (T. Shimornura): 6 -? -?-,
1080 m, v 19-27 1985 (T. Shimomura); 15 ? ? ,G
9 3 2 , ditto, 920 ni, v 19-27 1985 (T. Shimomuraj; 1-1'. ditto, 850 m, v 19-27 1985 (T. Shimomura).
[Yamanashi Prefecture 1: Tabayama-mura: 2 -? ?- , 2 2 if, Sanjonoyu, 1490 m, x 3-4 1985 (T. Shimomura).
m,
74
[Kanagawa Prefecture]: Tsukui-machi: 9 ? -?,2 (7 d\ Mt. Ogurayama, ii 22 1959 (R. lshikawa); 9 ? ? , 5 (7
(A Shida-tOge, xi 24 1961 (R. Ishikawa); 2 2 9 9 , 13 2 j\ditto, 280-300 m, iv 25-27 1995 (Y, Takami); 5 5 9
?, 29 2 (7, ditto, vi 20-22 1996 (Y. Takami). Sagamiko-machi: 4 % ? , 1 2 , Mt. SekirOsan, 450 m. ii 23 1994
(Y. Takami). Shiroyama-machi: 1 9 , Matsukaze, 210 m, , ii 28 1994 (Y. Takami). Isehara-shi: 1 J", Oiwake,
&ma, x 1 1958 (R. Ishikawa). [Chiba Prefecturel: Futtu-shi: 15? ?, 5 2 (7, Mt. Kan6za11, iii 4 1959 (R. &
F. Ishikawa); 1 (7. ditto, ii 1 1961 (R. Ishikawa); 3-? ?, 5 2 2, near Mother-Bokuj6, 260-270 m, Mt.
KanOzan, iv 22-24 1996 (Y. Takami). Kimitsu-shi: 2 9 ?, 2 2 O\ Nishino, ii 1 1961 (R. Ishikawa). [Saitama
Prefecture]: Minano-machi: 9 ? , 8 J" J", Hirakusa, 390 m,Misawa iii 11 1980 (R. Ishikawa). Ohtaki-mura:
3 9 $-, Wasabizawa, 1050 m, near Karisaka-t6ge, vi 6-7 1986 (K. Nemoto). Kodama-machi: 1 ? , no further
data. iv 29 1985 (A. Shimim). [Gunma Prefecture): Manba-machi: 2-? ? , 6 J'J', Shiozawa-t6ge, 1070-1 120
m, Mt. Mikaboyama, v 23-x 10 1985 (K. Nemoto); 11 ? -?, 5 3' 3',ditto, ix 11-13 1985 (R. & F. Ishikawa); 3
9 9,1 2,ditto, 970 m. ix 11-13 1985 (R. & F. Ishikawa); 1 ?, 1 j\ Sendagaya, 1300 m, Mt. Akagunayama,
i 11-13 1985 (R. & F. lshikawa); 2 2 2 , west of Mt. Akagunayama, 1390-1490 m, ix 11-12 1985 (R. & F.
Ishikawa); 1 -?- , Kurinokidaira, 830 m, Mt. Akagunayama, ix 1 1-13 1985 (R. & F. Ishikawa). My6gi-machi: 5
9 ? , 8 2 rf\ Mt. Mycgisan, xii 2 1962 (R. Ishikawa). Naganohara-niachi:2? ? , Kitakaruizawa, vii 25-27
1992 (S. Yasugi); 2 ? $, 1 2 , Furumori, 720 m, ix 18-19 1995 (Y. Takami); 2 - 9 ?, 3^' J", Kaise, viii 31-ix
1 1993 (R. & F. lshikawa). Tsumagoi-mura: 5 5 ? , 11 3' cf, Uenoyama, 930 m, Mihara, vii 17-18 1996 (R. &
F. Ishikawa). Kuni-mura: 2 9 ?, 3 2 2, env. Namasu, 830 m, Kozame, vii 18-19 1996 (R. & F. Ishikawa).
NakanojO-rnachi: 2 4 9 ?. 13 2 J\ Makiba, 590 rn, Kamisawatari, ix 12-14 1995 (R. & F. Ishikawa).
, m, vi 26-28 1996 (R. & F. Ishikawa): 42 ? ?, 14 J' 2 , ditto, vii. 2Minakami-machi: 1 9, 2 cf S\ ~ a n a540
3, 1997 (R. & F. Ishikawa); 2 -? -f-,2 2 2, Yubiso, 570 m, vi 26-28 1996 (R. & F. Ishikawa); I ?, YubisokOen, 610 m, vi 26-28 1996 (R. & F. Ishikawa); I ? , 1 cf, Machigasawa, 840-870 rn,vii. 1-2, 1997 (R. & F.
Ishikawa). Akagi-mura: 3 % 9,Okikado, 490 m, Sakae, vi 26-27 1996 (R. & F. Ishikawa); 2 0 9 I?, 3 2 2 2,
Mochikashiwagi, 450 m,vi 26-27 1996 (R. & F. Ishikawa). Kiryu-shi: 1 4 9 -?-, Fukiage, Kawauchi-ch6, vii
17-19, 1996 (M. Ujiie); 4 9 9 , 2 J" J", Serishi, vi 6 1960 (R. Ishikawa); 1 ? , Umehara, iii 12 1980 (T. Asami).
[Nagano Prefecture]: Saku-shi: 1 9 , 4 2 J", Aidate, 780-790 in, Uchiyamaky6, ix 12-13 1990 (R. & F.
Ishikawa); 1 0 9 I?, 7 2 2 , Uchiyama-t6ge-Mt. Arafuneyama, 1060-1 150 ni, ix 12-13 1990 (R. & F.
Ishikawa). Karuizawa-machi: 6 9 9,Hotchi, 930 m,ix 17-19 1995 (Y. Takami); 1 ? , 2 J' (7, near Kokuminshukusha, 935 m, x 17-19 1995 (Y. Takami); 3 9 ?, 2 2 j\ Usui-toge, 1190 m, (on borders betw. Matsuidamachi, Gunma Prefecture), ix 18-19 1995 (Y. Takami); 1 2 , Minenochaya, 1400 111. Mt. Asamayama, xi 13
1959 (E. lshikawa); 1 ?, ditto, 1410 m, ix 18-19 1995 (Y. Takami); 2 ? -?,2 2 2, Sengataki-nishiku, viii 25
3 J" J", ditto, 1050 m, vii 11-12 1996 (Y. Takami). T6bu-machi: 19,Narahara, viii
1991 (N. Fujino); 8 ^ ,
2 1976(K. Nemoto); 1 9, ditto, 1300 m, vii 10-12 1996 (Y. Takami). ITochigi Prefecture]: Ashikaga-shi: 1 ?,
Fujisaka-toge, 190 m, Ashikaga-shi, iii 12 1980 (T. Asami); 16 9 ?, 5 2 j\ onoma, Tsukiya-ch6, vii 17-19
1996 (M. Ujiie). Nishikata-mura: 27? ?, 18 2 2 , Magami, 230 m, v 23-25 1995 (Y. Takami); 10 ?- ? , 8 2
2, ditto, 200 m, v 23-25 1995 (Y. Takami). Tsuga-machi: 92 ? ?, 51 2 3,Nogami, 170 m, v 23-25 1995
(Y .Takami).
~ ~ . ~ ] : Prefecture: 8 9-?, 9
Other specimens examined: [Populations transitional with C. a. t s ~ t k i ~ h a n Tochigi
2 J\ Nagai, Yaita-shi, vi 8-9 1994 (R. & F. Ishikawa); 2-? ?, Sanuki, 260 m, Shioya-machi, v 26-27 1993
(R. & F. Ishikawa); I ? , Tobanoyu, 430 m, Shioya-machi, v 19-20 1993 (R. & F. Ishikawa): 1<?.
Karniterashima, Shioya-machi, ii 20 1993 (M, Ujiie); 7 ? ? , 4 (7 2, Kanpaku, 200 m,Kamikawachi-machi, v
26-27 1993 (R. & F. Ishikawa); 18? 9, 11 2 rf\ Idozawa, 250 m, Shinoi, liyama, Utsunomiya-shi, v 26-27
1993 (R. & F. Ishikawa). [Supposed hybrid population between C . a . esukianus and C. a. okumuraij: S ? 9,
6 2 2, Shiminnomori, 850 m, Osairi, Ueda-shi, Nagano Prefecture, ix 25-27 1996 (Y. Takami).
Subspeciation and distribution pattern of Carcihii\ dht-i>chti
Carabus (Ohornopterus) albrechti tsukubanus subsp. nov
Apotornoprus alhrerhti i<sakii (part):Nakane, 1953, p. 96 (48), Figs. 13: D-d, 5.
Apotomopterus jafwnicus t~sukiunus(part): Nakane, 1962, pp. 37-38, Figs. 37: a, b.
Carabus (Ohmoptcrux)ulbrv.t.'hti esukiuniis (part. ): Imura & Mizusawa, 1996, pp. 20 & 107,
fig. 44 (5).
This subspecies is similar to C. a . esakianus and C. a. hagai, but is different from them in
more slender body, thinner apical part of aedeagus and male antennae without hairless depressions.
Length: ? , 20.5-24 mm; 2 , 19.5-22 mm.
Upper surface of body dark copper or greenish black, less lustrous. Legs wholly black,
but in rare case tibiae partly somewhat rufous. Male antennae barely reaching beyond
middle of elytra; underside of 5-7th segments without hairless areas. Head almost
iinpunctale from front to neck. Inner margin of male protibia roundly convex (Figs. 6 1 &
62). Pronoturn with 4-6 marginal setae; disc more convex than in C. a. ~~si/kiat;ns:
posterolateral corners well produced but rounded (Figs. 57 & 58). Elytra slender and less convex,
shoulders less convex than in C . a. esakianus: striae weakly notched.
Male genitalia: Apical portion of aedeagus similar to that of C a. esakianus, but thinner
than in that subspecies, finger shaped, its outer margin slightly, narrowly grooved or flat;
copulatory piece triangular, but shorter (Figs. 5 1 & 53).
shallow median groove, anterior rim broad (Figs. 52 & 54).
Distribution of color variation: In the northern part of the range, greenish black individuals were not found.
Range: Mt. Tsukubasan, the Abukuma and Yamizo Mountains, the northern part of the
Kant6 District, Tochigi and Ibaraki prefectures, and most part of Fukushima Prefecture,
between the ranges of C. a. esakianus and C. a. hagai.
Etymology: Named after the type locality.
Holotype: if, Mt. Tsukubasan, Ibaraki Prefecture, xi 22 1959 (R. Ishikawa) deposited at Tokyo Metropolitan University.
Paratypes from type locality: 37 -? ^?-,
34 rf1 S'.same data as holotype: 3 9 ?, 3 <!' 2, v 12 199 1 (T. Toba).
Other paratypes: [Ibaraki Prefecture): 5 9 2 , 3 2 2,Yubukuro-t6ge. Yasato-machi, v 9 1994 (A. Toba); 3
? 9 , 4 (? c f , Gozenyama-mura, xi 16 1959 (R. & F. Ishikawa); 2-? -? , Mogaki, 40 m, Noda, Gozenyamamura, v 17- 18 1993 (R. & F. Ishikawa); 2 9 T, Mt. Yamizosan, 1000 m, Daigo-machi, iv 14- 15 1992 (R. & F.
Ishikawa); 5 ?- ?, Mt. Hanazonoyama, 800 m, vii 18-19 1987 (K. Nemoto): 2? ? . 6 d'2, Azuhata,
Hanakawa-ch6, Kitaibaraki-shi, vii 7-9 1996 (M. Ujiie). [Tochigi Prefecture!: 2 2 d\ Hodokubo, 120 m,
Kuroda, Motegi-machi, v 17- 18 1993 (R. & F. Ishikawa): 17 ? y , 9 3' 3\ Hirose, 120 m. BatO-machi, v 13-15
1992 (R. & F. Ishikawa): 1 ?, 1 c?\ ditto, 200 in, v 13- 15 1992 (R. & F. Ishikawa); 5 ? -?,2 2 d\ Kurumegi,
230 m, Koisago, Kurobane-machi, v 13-15 1992 (R. & F. Ishikawa); 1 9 , Takatori, 380 m. Kurobane-machi,
v 13- 15 1992 (R. & F. Ishikawa); 19? 9 ,1 6d7' 2 , Akasaka, Yaita-shi, vi 7-8 1994 (R. & F. Ishikawa): 109? , 6 2 2 , Terayama-dam, Yaita-shi, vi 8-9 1994 (R. & F. Ishikawa); 3 1 ? -7, 2 0 2 cf, Nagaihigashi, 195 m,
Naka, Yaita-shi, v 19-20 1993 (R. & F. Ishikawa); I?, 12, Kurodawara, Nasu-machi, iv 2 1961 (R.
Figs. 47-54. Carahus aLbrechti esakianus (Nakane) (47-SO), and C . a. tsukuhams subsp. nov. (51 54). [Copulatory piece and apical portion of aedeagus (47, 49, 5 1 and 53), and outer and inner
plates of vaginal apophysis (48, 50, 52 and 54jJ. 47 & 48, kawamachi, Hachioji-shi; 49 & 50,
Nogami, Tsuga-machi; 51 & 52, Mt. Tsukubasan; 53, Moriko, Nasu-machi; 54, Fujishio, Nasumachi.
Subspeciation and distribution pattern of C a r r t b ~ ~albrechti
s
77
Ishikawa); 1 ? , 3 J'2 , Nasu-machi, vii 28-viii 1 1966 (R. & F. Ishikawa): 2 9 ? , 5 3' 2, ditto, 630-660 m, vi
16-17 1984 (R. Ishikawa); 2 9 ?, 1 2 , Hachiman, 1000 m, Nasu-machi, viii 18-20 1985 (K. Nemoto); 1 ?,
near Yahata-onsen, 1020 m, Nasu-machi, v 23-25 1995 (Y. Takami); 1 ?, near Asahi-onsen, 1160-1180 m,
Nasu-machi, v 23-25 1995 (Y. Takami); 18 9 9 , 2 3 2 3\Moriko, 530 m, Takakuotsu, Nasu-machi, v 23-25
1995 (Y. Takamij; 8 ? ?, 3 J'J', H6j6,425 m,Takakuhei, Nasu-machi, v 24-25 1995 (Y. Takami); 22? ?,
6 2 2, Fujishio, 400 m, Takakuhei, Nasu-machi, v 24-25 1995 (Y. Takami); 1 -? , Tono, 375 nl, Teragohei,
Nasu-machi, v 24-25 1995 (Y. Takami); 1 3 , Kiriliiri, NikkO-shi, x i 1 1 1958 (R. Ishikawa); 3? ?,
Kirifurinotaki,750 m, NikkO-shi, xi 9 1995 (M. Ujiie & K. Kubota & Y. Takami); 4 '? ? , 6 J" 2 , Shimizubara,
750 m, Fujiwara-machi, ix 1-4 1983 (R. & F. Ishikawa); 4 ? ?, 2=,
SannO-loge, 920 m, FukushimaTochigi Prefectures, ix 1-4 1983 (R. & F. Ishikawa). [Fukushinia Prefecture): 37 9-? , 2 8 2 J\ Shoja, 890 m,
Tateiwa-mura, ix 1-4 1983 (R. & F. Ishikawa); 7 9 -?, 1 1 2 j\ Uwano, 660 m, Tajima-machi, v 25-26 1987
1 3,Sakai,
(R. Ishikawa): 1 ?, TMU lodge, 660-670 m, Tajima-machi, viii 21-23 1995 (Y. Takamij; 2 9 9,
500 m, NangO-mura, ix 4-5 1983 (R. & F. Ishikawa); 7$ ?, 6 c? 2,ditto, 540 m,i x 13-14 1995 (Y. Takami);
1 ?, ShOwanomori, 640 m, Nakamizawa, ShOwa-mura, ix 14-15 1995 (Y. Takami); 3 9 9-,
Tarafu, 570 m,
Kaneyama-machi, ix 3-5 1983 (R. & F. Ishikawa); 1 2 , ditto, ix 13-14 1995 (Y. Takami); 4 9 9 , 5 2 2 ,
Shirasaka, 390 m, Shirakawa-shi, v 24-25 1995 (Y. Takami); 1 ? , 1 3'.Hirohata. Kanayama, OmotegO-mum,
xi 16 1982 (R. & F. Ishikawa); 1 9 , Eda, Iwaki-shi, v 31 1987 (K. Haga); 7 ? ?, 3^^,
Akane-rindO,
Yokokawa, Haramachi-shi, v 29 1988; 5 9 ?, 1 j\ Dake, 640-1000 m, Mt. Adatarayama, iv 27-29 1963 (R.
Ishikawa) (paratypes of C. a. tohokuensis).
Other specimens examined: [Populations transitional with C . a. hagai]:see the description of C. a. hugai.
[Populations transitional with C. a. esakianus}: see the description of C. a. esakianus. [Supposed hybrid populations between C. a. tsukubanus and C . a. echigoj: Fukushima Prefecture: 10 ? ?,7 2 2, Ohara, 660 m, Inamura (the east of the Inagawa River), ix 1 1-1 2 1995 (Y. Takami); 7 9 ? , 5 (TV, Miyazawa, 610-620 m, Inamura (the west of the Inagawa River), ix 11-12 1995 (Y. Takami); 19,1 J\ Uchikawa, 590 m, Ina-mura (the
west of the Inagawa River), ix 12-13 1995 (Y. Takami).
Figs. 55-62. Carabus dbrechti e.sakiartuLs(Nakane) (55,56,59 and 60), and C. a. tsukuhanus subsp. nov. (57,
58,61 and 62). [Pronotum (55-58). and male protibia (59-62)J.55 & 59, kawamachi, HachiOji-shi; 56 &
60, Magami, Nishikata-mura; 57 & 61, Mt. Tsukubasan; 58, Fujishio, Nasu-machi; 62, Moriko, Nasumachi.
78
Curabus (Ohornopterus)albrechti echigo Ishikawa & Takami
Apotonwpterns ulbrechti (put.):Nakane, 1953, p. 96 (48). fig. 14: D-4.
Apotomopt(~riisjaponicusfreyi(part.): Nakane, 1962, pp. 37-39, fig. 37w.
A,wtomopterus japonicus freyi (part.): Nakane, 1963, pp. 1 1- 12.
Apotompterus alhrechtifn'yi {part.): Ishikawa, 1969, pp. 520-52 1.
Carabus (Ohomopterus) ulbrechti e s u k ~ a ~ (part.):
~ u s Ishikawa, 1985, p. 21.
Curabus (Ohomopteru.~)alhrechti echigo lshikawa & Takami, 1996, pp. 44-46, figs. 19-24,
27-29; type-locality: Mugurazawa, Yunotani-mura, Niigata Prefecture.
Length: 9 ,20-23 mm; j\ 19.5-2 1.5 mrn.
Upper surface of body bright or dark copper, greenish copper or black. Legs and palpi
wholly black. Male antennae exceeding middle of elytra; underside of 5-7th segments with
shallow hairless depressions, about 112-2/3 as long as each segment. Inner margin of male
protibia angulately produced (Fig. 72). Pronotum with 3-4 (rarely 5 ) marginal setae; lateral
margins distinctly emarginate posteriorly behind middle, with postero-lateral corners
strongly produced backwards but rounded (Fig. 69). Elytra narrow, broadest behind
middle; weakly convex above; shoulders weakly convex, lateral margins nearly straight
anteriorly especially in male; striae very weakly notched.
Male genitalia: Apical portion of aedeagus thick and tapered from basal ostium area in
outer view, with distinct groove on its outer margin; copulatory piece with pointed apex
(Fig. 63).
Female genitalia: Outer plate of vaginal apophysis triangular but narrower; inner plate
with broader outer rim, with shallow median groove (Fig. 64).
Variation: The specimens transitional to C. a. ok~imuraihave narrower copulatory piece
(Fig. 65), a broader outer plate and a narrower anterior rim of the inner plate of vaginal
apophysis (Fig. 66).
Range: The southern part of Niigata Prefecture so far north to Aganogawa River, and its
adjacent regions down south to Hakuba-mura, the Saikawa River and Iiyama-shi in Nagano
Prefecture, where it intergrades with C. a. okumurai.
Specimens examined: 466 (311 ? 9,1 5 5 2 2 ) from the following localities: IFukushima Prefecture]:
Kaneyama-machi: 69 9,2 2 2, Kamiigusa, 400 m, ix 3-5 1983 (R. & F. Ishikawa) (paratypes). Tadamimachi: 1 -? , Mt. Asakusa-dake, 750 m, vii 18 1979 (K. Nemoto) (paratype); 1 2 , ditto, 19 vii 1979 (K.
Nemoto) (paratype); 2 9 9,2 m , ditto, 20 vii 1979 (K. Nemoto) (paratypes). [Niigata Prefecture):
Yunotani-mura: 8 9 ? , 12,Mugurazawa, 190 m. v 31-vi 1 1993 (R. & F. Ishikawa) (paratypes from type
locality): Sanjyo-shi: 1 ?-,Mt. Tsukiokayama, v 22 1967 (H. Koike) (paratype). Teradoinari-machi: 3-? ?-. 1
2 , Watabe, 10 m, vi 8-9 1993 (R. & F. Ishikawa) (paratypes). Nagaoka-shi: I 9 ,I 3\ Mottate-tdge, 400-420
m, x 7-9 1992 (R. & F. Ishikawa) (paratypes). Kashiwazaki-shi: 25 9 . 6 2 3\WNW foot of Mt. Yoneyania,
250 in, viii 17- 18 1991 (T. Suda) (paratypes); 4 ? f , 1 J\ Kamikata, 10 m, vi 9-1 0 1993 (R. & F. Ishikawa)
(paratypes). Horinouchi-machi: 2 2 9 9,
2 2 2 , Nagaya, 150 m, v 3 1-vi 1 1993 (R. & F. Ishikawa). Irihirose-
Subspeciation and distribution pattern of C u r u b u s ulbrw/zti
79
mura: 1 9, 1 2 , Kuromatagawa River-Ashizawa-cleai, vi 1-2 1996 (K. Haga). Tsunan-machi: 2 ? ?, 2 2 2 ,
Tanaka, 260 m, Ueda, vi 1-2 1993 (R. & F. Ishikawa) (paratypes). Matsudai-machi: 12,GimyO, viii 30-31
I996 (M. Ujiie). JOetsu-shi: 7 ? ?, 4
Nishitono, 40 m, vi 13-14 1990 (R. & F. Ishikawa) (paratypes); 1
, Aratezawa, Nagahama, viii 30-31 1996 (M. Ujiie); 5 2 9 9 ,35 2 J\ Gotenyama-machi, 20 m, v 11-12
1997 (T. Narumi & R. & F. Ishikawa); 6 ? 9 , 1 j\ Rinsenji, 25 In, Nakamonmae, v 1 1- 12 1997 (T. Narumi &
R. & F. Ishikawa). Maki-mura: 5 9 ? , Furusato-mura, 200 m, Ikefune, 1 1 - 12 v 1997 (T. Narumi & R. & F.
Ishikawa). Nou-shi: 10 9 9 , 4 2 2,Iwadaira, 60-90 m, Komi, vi 13-14 1990 (R. & F. Ishikawa) (paratypes).
Ohmi-rnachi: 1 9 , 1 3 , Rind& 10-50 m, Suzawa, vi 13-15 1989 (R. & F. Ishikawa) (paratypes): 9 9 ? , 5 2
\ env. Hatsudensho, 10 m, Suzawa, vi 14-15 1989 (R. & F. Ishikawa) (paratypes). Itoigawa-shi: 6 '? ?,
Hakamaiwa, 90 m, v 28-29 1991 (R. & F. Ishikawa) (paratypes); 23 ?- ?, 6 (7 2, NakajOho, 350 m, v 28-29
1991 (R. & F. Ishikawa) (paratypes); 7 ? 9 , 1 3\ Kuzuba-tOge, 450 m, vi 12-13 1990 (R. & F. Ishikawa)
(paratypes). My6kCi Mts.: MyOkO-mura: 3 9 9,1 J\ Tsubam6, 1000-1080 in, vi 12-15 1986 (R. & F.
Isliikawa) (paratypes); 2 9 ? , 2 2 2 , near Sekimi-tOge, 970 m, vi 13-14 1986 (R. & F. Ishikawa) (paratypes);
9 9,1 2 r f V , route to Seki (from Akakura), 845 m, vi 24-25 1988 (R. & F. Ishikawa) (paratypes).
MyOkOkOgen-machi: I ? , Akakura, 820 m, vi 13-14 1986 (R. & F. Ishikawa) (paratype); k?, Mt.
Sasagamine, viii 17 1988 (H. Otobe). [Toyama Prefecturel: Asahi-rnachi: 4 9 ?, 6 d" 2, Ogawa-onsen, 280290 m, ix 17-21 1996 (Y. Takami). [Nagano Prefecturel: liyama-shi: 2? ?, 3 2 2 , Sasazawa NE by
Nozawaonsen, 480 m, , ix 6-7 1990 (R. Ishikawa) (paratypes); 2 9 ?, Hakusanjinjya, Kuwanagawa, ix 7 1993
(M. Ujiie) (paratypes); 11 ? ? , 12 2 3 , Habiroyama, 22-24 viii 1995 (M. Ujiie) (paratypes). Nozawaonsenmura: 5 9 9 , 3 2 2,Mushiu, viii 22-24 1995 (M. Ujiie) (paratypes). Togakushi-mura: 1 d\ Taneike, 11401 8 0 m, vi 23-24 1988 (R. & F. Ishikawa) (paratype); 2 9 (?. 1 2, Koshimizugahara, 1254 m, vi 23-24 1988
(R. & F. Ishikawa) (paratypes). Otari-mura: 21 -? 9, 11 2 2, Otarionsen-Kamaike, 970-1070 m, vi 14-15
Kuruma, 450 m, vi 14-15 1990 (R. & F. Ishikawa)
1990 (R. & F. Ishikawa) (paratypes); 10 <? 9, 2
(paratypes); 3 ? ? , ditto, 400-430 m, vi 7-8 1995 (Y. Takami) (paratypes); 12 '? ? , 63" d\ Sotozawa, 500550 m. vi 19-20 1990 (R. & F. Ishikawa) (paratypes); 3 ? ?, Ishihara, 550 m, v 27-29 199 1 (R. & F. Ishikawa)
(paratypes); 2 9 q , I 2 , Dorosaki, 660 m, Chikuni, vi 12-13 1990 (R. & F. Ishikawa) (paratypes): 8 ? <?,10
,
Tsuchikura, 680 m, v 27-30 1991 (R. & F. Ishikawa) (paratypes); 8 ? ?, 5 2 j\ ditto, 760 m, v 27-30
1991 (R. & F. Ishikawa) (paratypes).
Other specimens examined: [Populations transitional with C. a. okurnurw\: Nagano Prefecture: 17 <? ? , 8
3 2, Kayou by Rt.l48,650 m, Hakuba-mura, vi 12-13 1990 (R. & F. Ishikawa); 1 9, 2 2 J\ Sanosaka, 850
rn, Omachi-shi, vii 1-2 1986 (R. & F. Ishikawa); 2 9 9,
3 <? 2 , Kurosawa-rind& 1 100 m, Omachi-shi, vi 2223 1988 (R. & F. Ishikawa); 1 4 9 9 , 3 2 2,Hashizume, 660-690 m, Naniai, Nagano-shi, vii 2-3 1996 (Y.
Takami); 2 2 9 -9, 1 2 2 j\ Shinonoiutabi, 670-680 m, Nagano-shi, vii 1-3 1996 (Y. Takami); 3 ? ? , 4 d" d",
-Maguse, 750 m,Kijimadaira-mura, ix 24-25 1996 (Y. Takami), [Supposed hybrid populations between C . a .
echigo and C. a . t.sukubanus\: see the description of C . a. tsukuhanus. [Supposed relict populations of C. a .
okimzurai\: 1 9,1 2 , Mukaiyama, 1000 m, Yuzawa-machi, NiigataPrefecture, vi 3 1995 (K. Haga) (paratypes
of C. a . echigo); 1 9 , 2 2 2 , Koakazawa, 770 m, Sakae-mura, Nagano Prefecture, vi 2-3 1993 (R. & F.
Ishikawa) (paratypes of C. a . echigo); 1 1 9 <?,
4 2 2,ditto; 5 -? ?, 3 3' 2 , Wayama, 1000 m, Sakae-mura,
Nagano Prefecture, vi 2-3 1993 (R. & F. Ishikawa) (paratypes of C. a. echigo).
Tokyo Metropol itan University.
mi
m,
Carabus (Ohomopterus) albrechti okumurai (Ishikawa)
Apotomopteriis albrechti (part.):Nakane, 1952, p. 50, figs. 32; Masutomi, 33; Masutomi.
Apotomopterus ulbrechti esakii (part): Nakane, 1953, p. 96 (48). Figs. 13; D-e, 6.
Apotomopterus ulbrec'hti okumuru; Ishikawa, 1966, pp. 45 1- 453, Figs. 1-5; type-locality: Mt.
80
Y. Takami and R . Ishikawa
2 mni
Figs. 63-73. Curab~t.~
albrechii ech@ Ishikawa & Takami (63,64,69 and 72), C. a. echini transitional to C.
a. okikmurai (65, 66 and 70) and C. a. okumurai (Ishikawa) (67, 68, 71 and 73). [Copulatory piece and
apical portion of aedeagus (63, 65 and 67), outer and inner plates of vaginal apophysis (64, 66 and 68),
pronoturn (69-71), and male protibia (72, 73)l. 63, 64,69 & 72, Mugurazawa, Yunotani-mura: 65, 66 &
70, Kayou, Hakuba-mura; 67 & 68, Kobuchizawa-cho;71, Inukiri-the, Enzan-shi; 73, Yanagisawa-t@e,
Enzan-shi.
Subspeciation and distribution pattern of Curahu.~
alhrechii
Daibosatsu, Yamanashi Prefecture.
Apotomopterus albrechti okumuru'r:Ishikawa, 1969, pp. 520- 521, Figs. 4, 24.
Carahus (Ohomopterus)alhrechtc okumurui: Ishikawa, 1985, p. 21, PI. 4; I f.
Carahus (Ohumopterus) albrechti ok~4murai:Imura & Mizusawa, 1996, pp. 20 & 107, fig. 44
(6).
This subspecies is easily distinguished from other subspecies by the round tips of the
elytra of which the outer margins are evenly arched.
Length: ? ,20-23 mm; 2 , 19.5-21.5 mm.
Upper surface of body polychromatic, bright or dark copper, greenish or bluish copper or
black, usually lustrous. Palpi tibiae and tarsi usually wholly or partly rufous. Male antennae exceeding middle of elytra; underside of 57th segments with hairless depressions as
long as each segment. Inner margin of male protibia strongly, angulately convex (Fig. 73).
Pronoturn with 3-6 marginal setae; postero-lateral corners distinctly produced but rounded
(Fig. 7 1). Elytra strongly convex above; lateral margins evenly arched; apices rounded and
separated from each other; striae weakly notched.
Male genitalia: Apical portion of aedeagus broad, gradually tapered in dorsal view, with
distinct groove on its outer margin outlined with carinae; copulatory piece narrow, constricted at base, with rounded apex (Fig. 67).
shallow median groove, anterior rim narrow (Fig. 68).
Variation: Specimens from the right (east) bank of the Chikumagawa River, Nagano
Prefecture, are larger and more glossy.
Range: The greater part of Nagano and Yarnanashi prefectures north to the environs of
Hakuba-mura, the Saikawa River and liyama-shi in Nagano Prefecture where i t intergrades
with C. a. echigo. In Ueda-shi, Nagano Prefecture, it is parapatric with C . a. esukiarzus at
the Kamigawa River. In the eastern portion of Yamanashi Prefecture, it is parapatric with
C. I. lewisianus.
Specimens examined: 1258 (7579 ? , 501 of 2)from the following localities: !Yamanashi Prefecture]:
Enzan-shi/Kosuge-mura: 2 2 9 9, 2 0 2 2, Mt. Daibosaturei-Ishimaru-tbge, 2056-1957 m, vi 11-vii 1 1984
( Y . Fukazawa); 3 m , Marukawa-toge, 1660-1800 m, vi 11-vii 1 1984 ( Y . Fukazawa). Enzan-shi: I ? ,
Yanugisawa-toge, vi 9 197 1 (R. & F. Ishikawa); 1 ?, 2 2 S\ Saiki-rind6, 1530-1550 In, Yanagisawa-tbge, vi
10-1 1 1994 (Y. Takami); 9 9 l?, 6 2 2,Ninose, 1160 m, v 21-29 1985 (S. Uchida); 9 9 ?, 11 c? 2, ditto,
1160 m, v 29-vi 11 1985 (S. Uchida); 6 ? ? , 3 2 2 , ditto, 1200 m, viii 23-24 1985 (R. Ishikawa); 1 ?, 1 2 , nr
Miharasi, Sannose, vi 24-vii 11 1985 (A. Makimoto & Y. Inoue); 1 ?, Sawarakubo-~kubosawa,1010 m, v
21-29 1984 (S. Uchida); 3 2 2,Bunkiten-Gobindaira, 1600 m, vi 24-vii 1 1 1984 (A. Makimoto & Y. Inoue);
2 ? , 5 of of, Sh6ran-toge-Gobindaira-bunkiten, 1450 m, vi 24-vii 1 1 1984 (A. Makimoto & Y. Inoue); 1 ? ,
Sanj6shinbashi (R bk., Sensuidani-side), 830 m, iv 30-v I3 1985 (S. Uchida); 1 9 , 1 3\ditto, v 12-29 1985 (8.
Uchida); 3 ? ? , 2 2 S\ ditto, v 29-vi 1 1 1985 (S. Uchida): 1 ?, ditto, viii 22-24 1985 (R. Ishikawa); 6 9 ?, 1
, Inukiri-tbge, Ichinose-takahashi, vi 10-1 1 1994 (Y. Takami). 0tsuki-shi, 3 9 ?, Tatsuno, 340 m,
Yanagawa, iv 27-28 1992 (R. & F. Ishikawa); 1 9-? ?, 5 2 3\Magi, 560 m, v 22-23 1992 (R. & F. Ishikawa);
36 9 9, 17 2 2, R bank of Asarigawa River, 430 m, Okuyama, vi 2-3 1992 (R. & F. Ishikawa); 14? ? , 9 c?
2. Okunosawa, 700 m, v 22-23 1992 (R. & F. Ishikawa); 7 ? ?, 8 3" 3\Takayama, 400 rn, Hanasaki, v 29-
82
1 J\ Asakawabashi, 530 m, Nanaho-ch6. v 29-30 1992 (R. & F.
30 1992 (R. & F. Ishikawa); 2 9 9-,
Iwatono, 400 m. Nigioka-ch6, vi 2-3 1992 (R. & F. Ishikawa): 1 ? ,Okuhata, 4 10 m,
Ishikawa): 7 ? , 2
Magi, v 8-9 1995 (Y. Takami); 1 9 , 1 j\ Oiwake, 670 m, Kuronoda, v 8-9 1995 (Y. Takarni); 1 <?\ Oiwake,
690 m, Kuronoda, v 8-9 1995 (R. & F. Ishikawa). Ohtsuki-shi/Kawaguchiko-machi:I 'f , Seihachi-tbge, Mt.
Mitsutogeyama, iv 26 1965 (T. Okumura); 1 2 ditto, x 1 1 1965 (T. Okumura). Kawaguchiko-rnachi: 27? ?,
8 3" 2. Nichigersujinja, 900 m, Wade. ish hi, vii 10-1 1 1992 (R. & F. Ishikawa). Fujiyoshida-%hi:33 ?- ?, 21
f 2, Hachimanjinja, 800 m, Asahi, vi 2-3 1992 (R. & F. Ishikawa). Kasugai-ch6: 2 ? ?, 2 m , Mt.
Kabu~oyama,iv 12-13 1962 (T. Okumura). KOfu-shi: I ?, Taraga-loge, xi 4 1975 (R. Ishikawa). Ashigawamura: 2 ? ? , Kurodake-tozanguchi, 1400 m, vii 7-8 1993 (R. & F. Ishikawa); 2 ? ?. 3 d71 J\ Donbei-t6ge,
1440 in, vii 7-8 1993 (U. & F. Ishikawa). Kushigata-machi: 1 -9,Mi. Kushigatayania, viii 22 1968 (Niki).
Ashiyasu-mura: 1 9 , Ashiyasu-onsen, 750 m, vi 7-8 1990 (R. & F. Ishikawa). Nirasaki-shi:: 2 9 9,
Harayama-jinja, 425 m, Aoki, Seitetsu-machi, vi 1-3 1994 (Y. Takami); 1
? , 6'7- 2 , Aoki, 450 m,
Scitetsu-machi, vi 7-8 1990 (R. & F. Ishikawa); 1 2 , Aoki, 430 m, Seitetsu-niachi, vi 1-3 1994 (Y. Takami);
2 ? 'f , Tozawa-rind6, Shimomarui, Maruno, v 30-3 1 1990 (R. & F. Ishikawa). Akeno-mura: 17 ? ? , 1 3"east of Nagai, 680 m, v 30-31 1990 (R. & F. Ishikawa). Hakushii-machi: 10-? ? , 7 2 2 , Mt. Komagatake,
810 m, N of Chikuu, vi 6-7 1990 (U. & F. Ishikawa); 149- ?, 2 2 3". ditto, 730 m, vi 13-14 1990 (R. & F.
shikawa); 3 3 ? <?,
29 j" d\ ditto, vi 1-2 1994 (R. & F. Ishikawa); 2-? ? , 4 2 2,ditto, 740 m, vi 1-2 1994 (Y.
Takami); 1 1 ?- 9 ,1 S\ ditto, vii 2-4 1994 (Y. Takami); 3 <? 9 ,1 2 , Shiozawa-k6sen. 770 m, vi 27-28 1991
(R. & F. Ishikawa); 8 9 9, 3 (7' j\ Yamaguchi, 700 m, vi 27-28 1991 (R. & F. Ishikawa). Kobuchizawa-chO:
41 $ 2.2 2 2 2, Shinohara, 1020 m. vi 27-28 1991 (R. & F. Ishikawa); 9 ? ?, I? 2 , Kannondaira, 15701730 111, viii 29-30 199 1 (R. & F. Ishikawa); $ 0 9 9 , 39 23\ Kobuchizawa I. C. 990 In, vi. 24-25, 1995 (R.
& F. Ishikawa); 41 'f' ? , 2 2 rf" 2, ditto, 990 m, vi 24-26 1995 (Y. Takanii); 1 1 ? 9 , 6 2 d\ditto, 980 m. vi 2526 1995 (,Y. Takami); 229- 9, 4 6 3 d\ ShOkO, 830 m, vi 25-26 1995 (Y. Takami & R. & F. Ishikawa).
[Nagano Prefecture 1: Fujimi-machi: 16 ? 9,7 (7' 3\ Sanrigahara, 1200- 1300 in, Mi. Yatsugatake, vi 17- 18
1987 (A. Makimoto); 1 ?, 1 J\ Fujimi-kogen, viii 21 1987 (F. Hayashi); 17 9 ?, 1 6 2 Matsume, 1035 m,
vii 10-12 1991 (U. & F. Lshikawa); 19 9 -?, 15 2 j\odaira, 988 m, vii 10-1 1 1991 (R. & F. Ishikawa); 1 2,
FujimikOen, 960 m, vi 2-3 1995 (R. & F. Ishikawa); 6'f- ?, 15 c7' 2 , ditto, vi. 25-26. 1995 (R. & F. Ishikawa);
4 ? s) , 4 j\NyGgasa-ko, 910 m, ~ s a w avi, 2-3 1994 (R. & F. Ishikawa); 8 ?~ ? , 6 c? 3,NyGgasako Lake,
vi 2-3 1994 (Y. Takami); 9 9 -?, 11 2 2, Mt. Nytigasayama, 1 1 10 m, vi 28-30 1993 (R. & F. Ishikawa). Haramum: 262 9, 4 J' 2, Tatsusawaseki, 1250 m, vi 26-27 1991 (U. & F. Ishikawa). Chino-shi: 5 9 ? , 4 2 2 ,
Kanazawa-shimoni, 850 m, vii 11-12 1991 (R. & F. Ishikawa); 4 Â¡2 , 1 rf\ 6sawa, 920 m, Kanazawa, vi 2-3
1994 (Y.Takami); 3 9 ?, 1 J\ Suwataisha-kamisha-mae~niya, 780 n1, Koinachiya, vi 1-2 1994 (R. & F.
Ishikw a); 1 9 ,ditto, (Y. Takami); 2 ? ?, 1 2 , ditto, vii 2-3 1994 (Y. Takami): 9 ? 9 , 10 2 3, Ankokuji,
870 ni, vii 10-1 1 1991 (R. & F. Ishikawa); 2 2 2 , ditto, vi 1-2 1991 (R. & F. Ishikawa): 4 9 ?, 1 2,ditto, vi
1-2 1994 (Y. Takami); 1 ? , 3 3 j\ ditto, vii 2-3 1994 (Y. Takami); 2 2 2 , S of Ankokuji. 860 m, vi 1-2 1994
(R. & F. Ishikawa). Suwa-shi: 2 2 d\ Nozokiishiguchi. 1050 m, vii 9-10 1991 (U. & F. Ishikawa). Okaya-shi:
Higashi 3-chOme, 800 m, Kawagishi, vi 15-16 1992 (R. & F. Ishikawa). Minowa-machi: I?-, 1
32 ?,1
J\ Mikkarnachi, 750 m, v 10-12 1985 (R. & F. Ishikawa). Ina-shi: 6-?- 9 . 3 J' 3..
RokudO-tsutsumi, 750 m, v
12- 13 1985 (R. & F. Ishikawa); I 9 , Hatogaike, 970 m, Yokoyama, vi 19-20 1996 (R. & F. Ishikawa). Hasemum: 9 5; -?, 11 J' 2 , Karei-Kogen, 1170 m, vi. 23-24, 1994 (R. & F. Ishikawa), Narakawa-mura/Kiso-mura:
1 9 . 2 2 2,Torii-(age, iv 21 1963 (2.Nai-use) (paratypes). Kiso-mura: I ? , 2 2 J\ Matsubara, ix 17-18 1996
(M. Ljiie); 1 9 ,1 rf", Yabuhara, 950 m, v 21-22 1997 (M. Ujiie). Shiojiri-shi: 2 9 9 ,1 2, Seba, xii 4 1965 (T.
Okiimura). Azumi-mura: 3 9? , 3 J' J\ Sawando, 1000 m, vi 8-1 0 1988 (R. & F. Ishikawa); 1 3\ Yugawado,
1050 m,vi 8-10 1988 (R. & F. Ishikawa). Azusagawa-mura: 3 9 -?, 3 rf" 2, Oirizawa, 780-840 m, Taya,
Leno, vi 6-8 1995 (Y. Takami). Misato-mura: 1 'f', 1 2. Kitaogura. 760 ITI, xi 11 1976 (R. & F. Ishikawa); I 9,
ditto, vii 10-1 1 1983 (R. & F. Ishikawa); 8
8 (7' d\ J6shinji. 720 rn, Ogura, vi 6-8 1995 (Y. Takami); 1 2,
Higabhi-tOge, 900-920 m, xi 11 1976 (R. & F. Ishikawa). Hotaka-tnachi: 3 9? , 2 2 2, Nakabusa, 930 m, v
30-vi 2 1985 (R. & F. Ishikawa). Shiga-mura: I ? , Aida, no date (S. Asawa). Ueda-shi: 4 ? 9,3
m,
\a
w,
m,
Subspeciation and distribution pattern of Curahus alhn'diti
83
TOshOji, 600 m, Urano, ix 19-20 1996 (Y. Takami). Kamiyaniada-machi: 14 ? ?, 8 S" J\ KyOwa, 580 rn, vii
1-3 1996 (Y. Takami). Sakaki-machi: 8 ? ?. 5 2 j\ Nazawagawa, 540 rn, vii 1-3 1996 (Y. Takami). Suzakashi: 8 9 9, 8 2 31, Maeyama-tsutsuji-keen, 450 m. Hatch& vii 2-3 1996 (Y. Takami).
Other specimens examined: [ A supposed hybrid population between C. a. okumurui and C . a. e s a k i u m ~ s ~ :
1 d\ ChhO-kita, 570-580 m, ix 20-21 1996 (Y. Takami); 1 3'.ditto, ix 25-27 1996 (Y. Takami). [Populations
transitional to C. a. echlgo]:see the description of C. a. vchigo.
Key to the subspecies of Carabus albrechti
1 (2) Apices of elytra rounded and not meeting at suture ..................okumurai (Ishikawa)
2 (1) Apices of elytra pointed and meeting at suture.
3 (4) Apical part of aedeagus with distinct groove on its outer margin ..............................
echigo Ishikawa & Takami
...........................................................................................
4 (17) Apical part of aedeagus without groove but with flat part on its outer margin;
postero-lateral corners of pronotum well produced.
5 (6) Elytra short and broad with shoulders well convex.....................aihrechti Morawitz
6 (5) Elytra oblong with lateral margins less arched and nearly straight at anterior half.
7 ( 8 ) Antennae without hairless depressions ..................................tsukubunus subsp. nov.
8 (7) Antennae with hairless depressions.
9 (12) Elytra with strongly notched striae.
10 (1 1) Copulatory piece with rounded apex; constricted near base .....................................
..............................................................................
uw(~~shinuic
Ishikawa & Takami
1 1 (10) Apex of copulatory piece not rounded .........................................
hugui subsp. nov.
12 (9) Elytra with weakly or not notched striae.
13 (14) Pronotum narrower; with angulate postero-lateral corners .......................................
yamauchu. . subsp. nov.
...................................................................................................
14 (13) Pronotum wider; with rounded postero-lateral corners.
15 (16) Elytra less convex above; lateral margins subparallel .........................freyi Emden
16 (15) Elytra more convex above; lateral margins nearly straight at anterior half but not
subparallel.. .......................................................................................
.esukianus (Nakane)
17 (3) Apical portion of aedeagus unmodified and without flat part on its outer margin;
postero-lateral corners of pronotum weakly or not produced.
18 (19) Elytra oval with lateral margins evenly arched ......................tohokuensis Ishikawa
19 (1 8) Elytra short and broad with shoulders well convex.
20 (21) Apical portion of aedeagus longer and uniformly thin .............................................
.................................................................................
hidukaizz4.s Ishikawa & Takami
21 (20) Apical portion of aedeagus shorter, thicker and cylindrical .....................................
.................................................................................................
itoi Ishi kawa & Takami
Y. Takami and R . Ishikawa
Phylogeny
Method
In order to reconstruct the phylogenetic relationships among the subspecies of C.
alhrechti, analyses were made by cladistic method on the basis of 15 morphological characters of the adult which were used for distinguishing the subspecies.
First, simple parsimony analysis (Wagner parsimony) was performed, in which reversals
and convergences were counted equally. Globally parsimonious trees within the alhrechti
species group were sought in order to investigate the monophyly of C . alhrechti (=ingroup)
(Maddison el al., 1984). C. daisen okianus was used as the outgroup. Second, the CaminSokal parsimony analysis was performed with character polarities determined, in which the
monophyly of C. albrechti is assumed. The polarity of each character was determined by
outgroup comparison (Maddison et al., loc. cit.). In several characters whose polarities
were difficult to determine by outgroup comparison, their polarities were determined by the
so-called commonality principle. The characters are discussed in the 'Evaluation of diagnostic characters' and are shown in the Appendix 1. The outgroup in the latter analysis was
arranged automatically as the hypothetical ancestor (i.e., all the character states of the
outgroup were coded as "0" ). The results of two alternative analyses, in which character
polarities were determined or not, were compared.
The following species or subspecies constituting the albrechti species group were used
for outgroup comparison: C. lewisianus lewisianus, C. 1. awakazusanus, C. kimurai and C.
yamato. C. daisen okiunus was added to the outgroup because the molecular analysis based
on the mitochondria1 DNA sequences suggests the close relationships between this subspecies of C. daisen and the above-mentioned three species (Su et al., 1996).
All the coded characters were recoded binarily (see Appendix l), and then, the data matrix was analyzed by PHYLIP 3% (Felsenstein, 1986-1993). The analysis was under the
PENNY program using the branch-and-bound search. In the first analysis, however, there
were too many OTUs for the PENNY program to search. Therefore, the analysis was performed by the Mix program using heuristic search. A strict consensus tree was obtained by
computing under the CONSENS program. Character reconstructions were established using the DELTRAN optimization.
Evaluation of diagnostic characters
Morphological features are selected, and are evaluated to code (i.e. (1. 0) etc.) for cladistic analyses.
Size of body. Body is small to medium-sized (19.5-26 mm in length). Smaller body size
is assumed to be plesiomorphic, simply because related species C. lewisianus, C. kimurai
and C. yamato are all small-sized (17-23 mm in length). This character is not coded, because it varies within each subspecies, and the polarity can not be determined definitely.
Color of upper surface of body. Upper surface of body is predominantly copper and
Subspeciation and distribution pattern of Curah14.salhrechti
85
lustrous, but polychromatic, e.g. dark or bright copper, greenish copper, green, greenish
black or black, though the ratio of different color vary by population.
Head.
1) Sculpture: In the species belonging to the subtribe Carabina, head is distinctly punctale and wrinkled from front to neck (1. 0). However in the present species, punctation of
head is apparently reduced: not, or at most partly, punctate from front to neck (1. 1).
2) Antenna:
a) Length: Male antennae are longer than female antennae and exceed the middle of
elytra in most species of the subgenus Ohomoplerus (2. 0). In several subspecies of C.
alhrechti, they are short and barely reach the middle of the elytra (2. 1); in C . a. albrechti, C.
a. tohokuensis, C. a. hidakatzu.~and C . a. itoi, they are shorter and do not reach the middle of
the elytra (2. 2). The states (2. 1) and (2. 2 ) are assumed to be apomorphic.
b) Hairless depression of male antenna1 segments: In the male antennae, there are
hairless depressions on the underside of the 5-7th segments. This state is assumed to be
plesiomorphic because many species of the subtribe Carabina have them (3.0). In several
subspecies of C. alhrechti, however, they are narrow (3. 1) or rudimentary or absent (3.2).
Pronotum.
1 ) Outline:
a) Lateral margin: The lateral margins of the pronotutn are more or less cordately
sinuous, that is, they are arcuately convex anteriorly and concavely convergent posteriorly.
The female pronoturn is broader and its lateral margins are more roundly convex anteriorly
than in the male.
b) Postero-lateral corner: The postero-lateral corners are distinctly produced, more or
less lobate or angulate, but rounded in most subspecies and this state is assumed to be
plesiomorphic (4. 0 and 5. 0). In C. a. albrechti and C. a. yamauchii, they are produced
angulately (4. I), and in C. a. tohokuensis, C. a. hidakanus and C. a. itoi , they are weakly or
not produced (5. 1). States (4. 1) and (5. 1) are apomorphic. This character is coded as
characters 4 and 5 because it is a bifurcate character.
2) Punctation of disk: The disk is distinctly punctate in the present species. In several
subspecies, punctures are fused with one another or condensed to form transverse rugae by
the longitudinal median line.
3) Marginal setae: There are long setae at the lateral margins of the pronotum. The
bisetose condition, i.e. one seta at middle and another at the postero-lateral corner, is predominant, and therefore, presumed to be the plesiomorphic state of the subtribe Carabina (6.
0). The pronotum of the albrechti species-group except C . ulhrechti is trisetose, that is,
there are two anterior setae and one posterior (6. 1). In C. ulhrec hfi, pronotal setae are
multiple. This condition is secondarily derived from the trisetose state by multiplication of
the anterior setae (6. 2).
Elytra.
1 ) Shape:
a) Outline: The shape of the elytra is oval, more oblong in male. The lateral margins are
less convex at the anterior half especially in the male of most subspecies of C . alhrechti and
86
its related species (7. 0). In C. a. tohokuensis and C . a. okunzurai, the lateral margins are
evenly convexly arched and not nearly straight on the anterior half. This state is aponiorphic (7. 1). In the subspecies of Hokkaido, namely, C. a. alhrechti, C. a. hidakanus and
C. a. itoi, the elytra are broader and shorter, with shoulders strongly produced. This condition is most apomorphic (7. 2).
b) Apices: The elytra of the subspecies except C. a. okumurai have pointed apices
which meet at the suture (8.0). In C. a. okurnitrai , the apices of the elytra are rounded and
apart from each other ( 8 . 1).
c) Convexity: The elytra are more or less convex above in all the subspecies. The
C.
weakly convex state is assumed to be plesiomorphic, because the elytra of C. lcw~i,sicznus,
kimurai and C. yamcito, are all weakly convex. The strongly convex state is assumed to be
apomorphic. This condition is commonly shared by the "secondary species" or "secondary
subspecies" (sensu Ishikawa, 1979) of the subgenus Ohomopterus.
2 ) Sculpture:
a) Intervals: Elytral intervals of C. albrechti are all triploid homodynamic as in the
other species of the subgenus Ohomopterus. Primary foveae are small, rarely rudimentary,
or partly or almost wholly lost in C . a. e s a k i a w .
b) Notches of striae: In C. a. awashimae and C. a. bpi, the striae are strongly notched
(9. 1). This state is assumed to be apomorphic. In other subspecies, they are weakly
notched or smooth (9.0).
Appendages (legs and palpi).
1 ) Color: The appendages are wholly black, or the palpi, tibiae and tarsi are wholly or
partly more or less rufous. Appendages with brownish or reddish parts are characteristic of
the primary species or subspecies (Ishikawa, 1979) of the subgenus Ohomopterus. This
state is assumed to be plesiomorphic. In C. albrechti, however, no subspecies is known to
be composed only of individuals with rufous appendages. Therefore, this character is not
used for the cladistic analysis of the subspecies.
2) Male protibia: The inner margin of male protibia is more or less convex or angulate
medially. The strongly convex and angulate state is plesiomorphic (10. 0). The states not
angulate though distinctly convex (10. 1) and weakly convex (10. 2) are apomorphic.
Male genitalia.
1 ) Apical portion of aedeagus: The aedeagus is narrowed distally from the membranous
ostium area to form a digitate apical portion which bears characteristic features for each
subspecies.
a) Longitudinal groove of outer margin: The presence of a distinct groove on the outer
margin which is outlined on either side by a carina is plesiomorphic (1 1.0). The shallow or
flat, not concave (1 1. I ) , or unmodified ( 1 1. 2) condition of the groove are apomorphic.
b) Shape in outer view: In C. a. ok14murai and sister-species, the apical portion of
aedeagus is broad and flattened at base and hence gradually tapered to apex (12.0). In other
subspecies, however, it is weakly ( 1 2. 1) or strongly (12. 2) narrowed beyond the membranous ostium area. These states are assumed to be apomorphic.
2) Copulatory piece: The copulatory piece is composed of the basal plate inbedded in
Subspeciation and distribution pattern of Curuhus ulbrci hti
87
the membranous wall of the endophallus and the digitate part arising vertically from the
distal part of the basal plate and bent back nearly parallel with it. The digitate part is narrow,
flat and tapered to apical point with features characteristic of subspecies.
a) Apex: The apex of copulatory piece is blunt in most subspecies. In C. a. okumurai
and C . a. awashimae, it is rounded, but in C. a. e c h i p , it is pointed.
b) Basal area of the digitate portion: The digitate portion of the copulatory piece of C,
a. okurnz~raiand C. a. awa.~himueis constricted near the base. In C. a. itoi, the lateral
margins of copulatory piece are raised near the base.
Female genitalia.
1) Vaginal apophysis:
a) Outer plate: In the related species, the outer plate of vaginal apophysis is short and
broad. Its lateral margins are subparallel in sister-species (13. 0); in C. albrechti, it is
subtriangular, strongly narrowed backwards at its posterior half (1 3. 1). In several other
subspecies, it is more strongly and evenly narrowed, with the lateral margins weakly arched
(13. 2).
b) Inner plate:
b-i) Anterior rim: The outer margin of the shallow, cup-shaped inner plate is not raised
at the anterior half in most species of the subgenus Ohomopterus (14.0). In C . albrechti and
its allied species, the anterior margin of inner plate is raised to form a narrow (14. 1) or
broad (14. 2) rim.
b-ii) Median groove; In most species of the subgenus Ohomopterus, the disc of inner
plate has a median longitudinal groove which is outlined laterally by distinct ridges (1 5.0).
The median groove is barely outlined(15. 1) or absent (15. 2) in the apomorphic state.
Result of cladistic analysis
The monophyly of C. albrechti is supported by simple parsimony analysis, but some
polytomies are included in the group under study. In the Camin-Sokal parsmony analysis in
which character polarities were determined, the result agreed logically with that of simple
parsimony analysis, and exhibited greater resolution. Therefore, the result obtained with
determined character polarities was adopted as the phylogenetic hypothesis among the subspecies of C. albrechti.
Three topologies of the most parsimonious trees were obtained by cladistic analysis.
The strict consensus tree i s qhown in Fig. 74 with the DELTRAN reconstructions, in which
only the subspecies of C. albrechti are shown and other species are included as OUTGROUPS. The consistency index (CI) of the tree is 0.70.
The monophyly of C. ulbrechti (= the component 1) is supported by the degeneration of
head punctures (character 1; the word 'character' is omitted in the following discussion),
multiplied marginal setae of pronoturn (61, narrowed outer plate of the vaginal apophysis
( 3 ) and the presence of a shallow median groove of the inner plate of the vaginal apophysis
1 5).
In component 1, C. a. okumurai is defined by two characters of elytra, viz., evenly
88
arched lateral margins (7) and the rounded apices (8). Character 7 is shared as a homoplasy
with the component 8. Character 8 is a unique autapomorphy of this subspecies. Component 2 is defined by the much narrowed apical portion of the aedeagus (12) and the broad
anterior rim of the inner plate of the vaginal apophysis (14).
In component 2, C. a echigo is defined by the narrowed hairless depressions of the male
antennae (2). Character 2 is a homoplasy shared with component 4. Component 3 is defined by the presence of a shallow longitudinal groove on the outer margin of the apical
portion of the aedeagus (1 1) and much narrowed apical part of aedeagus (12).
In component 3, C. a. esakianus is not supported by any autapomorphy. Component 4 is
defined by two characters of the antennae, viz., the narrowed hairless depressions of the
male antennae (2) and the shortened length (3). Character 2 is a shared homoplasy with C.
a. e m o in component 2.
I n component 4, C . a. freyi is defined by the degeneration of the median groove of the
inriel plate of vaginal apophysis (15). Character 15 is a shared homoplasy with component
8. Component 5 is defined by the weakly convex inner margin of the male protibia (10).
In component 5, the polytomy consisting of C. a. tsukiihatzus establishes component 6
and 7. C. a. tsukubanus is defined by the degenerated hairless depressions of the male
antennae (2). Character 2 is a shared homoplasy with component 9. C . a. awashimae and
C . (i hagai representing component 6 are supported by the presence of strong notches of the
elytral striae (9). Component 7 is defined by the weakly convex inner margin of the male
protibia (1 0).
In component 6, C. a. awashimae is defined by the narrowed outer plate of the vaginal
apophysis (13) but C. a. hagai has no autapomorphy. Character 13 is a shared homoplasy
vv ith C\ a yamauchii and component 9.
In component 7, C. a. yamauchii is defined by the angulate postero-lateral corners of the
pronotum (4) and the narrowed outer plate of the vaginal apophysis (13). Character 4 is a
hoinoplasy shared with C . a. albrechti, and character 13 is a shared homoplasy with C. a.
a-va^iimue and with component 9. Component 8 is defined by the shortened antennae (31,
the evenly arched lateral margins of the elytra (7) and the degeneration of the median
groove of the inner plate of vaginal apophysis (15). Character 7 and 15 form shared homoplasies with C. a. okumurai and C . a. freyi respectively
In component 8, C . a. alhrechti is defined by the angulately produced postero-lateral
corners of the pronotum (4) and the broader elytra (7). Characters 4 and 7 are shared homoplasies with C . a. yamauchii and component 10, respectivelv. Component 9 is supported
by the degeneration of the hairless depressions of the male antennae (2). the less produced
posicro-lateral corners of the pronotum (S), the unmodified tip of the aedeagus ( 1 1 ) and the
nanowed outer plate of the vaginal apophysis ( 1 3). Character 2 is a shared homoplasy with
C. a. tsztk~i/~anus,
and character 13 with C. a. awashimae and C. a. yanzaz~chii.
in component 9, C. a. tohokuensis has no autapomorphy. Component 10 is defined by
broad elytra (7). Character 7 is a shared homoplasy with C. a. alhrechti.
In component 10. C. a. hidaktzus and C . a. itoi are not defined by coded characters of
thest own. However, they are distinguished by characteristic shape of the apical portion of
Subspeciation and distribution pattern of Carahns alhrerhti
Fig. 74. Strict consensus tree of the most parsimonious trees obtained by cladistic analysis with DELTRAN
reconstructions. Solid bars refer to synapomorphies; simple bars refer to homoplasies; characters are
shown by small numbers on each bar. The components are shown by large numbers under bars.
aedeagus.
Discussion
Subspeciation in the Chuhu District
The result of cladistic analyses suggests that C. ulbrechti was first established in the
Chiibu and Kant6 districts in central Honshu, where the common ancestor of component 1
gave rise to C. a. okumurai and component 2. C. a. echigo and C . u. m k i u n u s were differentiated allopatrically from the ancestors of components 2 and 3, respectively.
C. a. okumurai and C. a. echigo are parapatric in the Chfibu District (Fig. 75). On the
borders of these subspecies, there are morphological dines in the shape of the aedeagus and
apices of the elytra. The zone, in which the morphological dine of the aedeagus occurs, is
about 30-50 km in width, whereas the transitional zone of the elytral character is wider than
that of the aedeagus (Fig. 76). On the other hand, the ranges of C. u. o k u m i and of C. a.
aakiunus appear to be divided by the Kamigawa River in Ueda-$hi, Nagano Prefecture
(Fig. 75). This is different from the case of C. a. okumurai and C. a. echigo. A clinal
transition is not found in this area, but there is slight morphological variation presumably
owing to hybridization between the populations from either side of the Kamigawa River.
Moreover, the population density of these subspecies is very low in this area. The hybrid
zone is presumed to be maintained by the balance between dispersal from both populations
and selection against hybrids (Barton and Hewitt, 1985). The dispersal abilities of several
carabid beetles were investigated (Kubota, 1996), and they have the abilities in the same
90
degree within species. Therefore, the different widths of the hybrid zone are due to the
different rates of selection against hybrids. This means that the subspecies which have
narrower hybrid zones at their borders are more strongly isolated reproductively than those
having wider hybrid zones. Therefore, the degree of reproductive isolation between C. a.
okumurai and C. a. esakianus is presumed to be higher than that between C. a. ~ ~ k ; i m ; ~ r a i
and C. a. echigo. The result of cladistic analysis shows that the period after the branching
event between C. a. o h u r a i and C. a. e c h i p is equal to that between C. a. okurnurai and
C . a. esakianus. It shows that the level of reproductive isolation is not always correlated
with the period after the branching event.
The isolated populations with rounded apices of elytra are found in the southernmost part
of the range of C. a. e c h i p , the valleys of the Nakatsugawa and Kiyotsugawa rivers on the
borders between Niigata and Nagano prefectures (Fig. 1 and 75; meshed area). They are
compared with C. a. okumurai because of having its unique character. They are separated
from the range of C. a. okurnurui by mountains, but are contiguous to the range of C . a.
echigo along the valleys forming a cline in the shape of the apices of the elytra. This distribution pattern suggests that C. a. okumurai once had a more northerly distribution than it
has at present and invaded these valleys from lower land. After the invasion of C. a.
okumurai, C . a. echigo spread its range southward, enclosing the populations of C. a.
okumurai. Then, relict populations of C . a. okumurai was remained in the valleys. A fluctuation of the borders in such a way may have established broad and scattered transitional
areas as they occur at present.
Subspeciation in the northern part @anti? and the southern part of Tohoku
The ancestor of component 5 established in the T6hoku District and was split there,
while its sister group entered Sado Island to become C. a. freyi.
In northern Kant6 and the T6hoku District, C. a. tsukubanus and the ancestor of components 6 and 7 were differentiated allopatrically. Then, C. a. tsukubunus met and intergraded
with C. a. esakianus at the middle of Tochigi Prefecture, where the morphological dine
between them is apparent particularly in the size of the hairless depression of the male
antennae which increases toward the south in the transitional zone. Similarly, C. a.
tsukubanus intergrades with C. a. hagai at the northern part of its range where a similar
dine is recognized. In the west, C. a. tsukubanu.~is parapatric with C. a. echigo at the
Inagawa River. The morphological variation of the aedeagal characters, considered as the
result of hybridization, is apparent only in the populations from both sides of the Inagawa
River. Therefore, the degree of reproductive isolation between C. a. tsukubanus and C. a.
echigo is higher than that among C. a. tsukubanus, C. a. esakianus and C. a. hagai. It is
partly correlated with the length of the period after subspeciation, namely, the branching
event.
In the central part of the T6hoku District, the ancestor of component 6 invaded Awashima Island off the northern part of Niigata Prefecture, where it was isolated and differentiated to C. a. awashimae. On the other hand, C . a. h q a i has no unique apomorphy, and
may be the ancestor of component 6 or a close relative. Therefore, C. a. awashimae is a
Subspeciation and distribution pattern of Carubu.s alhrechti
Fig. 75. Distribution pattern of Carabus ulbrechti okumurai, C. a. ewe and C. a. esakianu.~in the
ChGbu District. Closely related species distributed parapatrically, C. le~vis/anus,
C. kimurai and
C. yamuto, are also shown. Shaded areas refer to hybrid zones between subspecies inferred from
genital morphology. Meshed area, see Discussion.
Fig. 76. Clinal change ofelytral character between Carahus ulbrechti oh~tmuruiand C. a . a hzgo in
the Chiibu District. Black areas refer to the frequencies of individuals which have the rounded
apices of elytra. White areas refer to the frequencies of individuals with normally pointed apices.
Gray areas refer to the frequencies of intermediate form between them.
Subspeciation and distribution pattern of Curubus ulbrerhti
93
direct derivative of C. a. hagai. Their close relationship is also supported by the distribution
pattern of these two subspecies.
Subspeciation in the northern part of the Tbhoku District and Hokkaido
The ancestor of component 7 occupied the northern half of the T6hoku District, and
invaded Hokkaido several times from there. It is considered that the invasions to Hokkaido
and C. a. itoi (= the ancestor
by C. a. albrechti and the common ancestor of C. a. hidakc~~zz~.~
of component 10) had been made independently because they are not monophyletic (Fig.
74). C. a. albrechti was derived from the ancestor of component 8 and invaded the Oshima
Peninsula of Hokkaido. On the other hand, the ancestor of component 10 was derived from
ancestor 9, which is most possibly C. a. tohokuensis having no autapomorphy of its own,
and somehow invaded Hidaka Province and the eastern half of Hokkaido. After the invasion, the population became established in the eastern half of Hokkaido but was reduced in
size to become sporadic in distribution. In particular, C. a. itoi is known only from two
localities very far from each other. This suggests that C. a. itoi become isolated within the
range once established. Since the specimens from the two localities are slightly different,
this subspecies seems to comprise several diversified, isolated populations. The phylogenetic relationships of component 10 should be revised in the future based on additional
Pronotal width /length k S. D.
.---/
\,
,is^,
Honshu
Fig. 77. Clinal change of male pronotal shape in the Oshima Peninsula. Ajigasawa-machi is located
in the range of Carahus albrechti yamuuc hii, and other three are in the range of C . a. albrechti.
Averages are presented by open rectangles with S. D.
94
material from more localities, because these two populations of C . a. itoi are not monophyletic (Ishikawa & Takami, 1996).
C. a. yamauchii and C. a. albrechli have in common the small, narrow pronotum with
strongly angulate postero-lateral corners (character 4. 1). The result of cladistic analysis
shows that these characters developed independently (Fig. 74) or may be a symplesiomorphy. However, the secondary contact and hybridization between them possibly
has caused their parallel specialization. The Tsugaru Peninsula, the northernmost part of
Honshu, was connected more than once with the Oshima Peninsula and C. a. albrechti
shows aclinal change in the pronotal character in this region (Fig. 77). This hypothesis may
include a circular argument because the cladistic analysis was made including the pronotal
character. However, the topology of the tree did not change in the alternative analyses in
which the pronotal character of C. a. albrechti was deleted or its state was changed. Therefore, even though C. a. albrechti had the different state of the pronotal character in the past,
the phylogenetic relationships among the subspecies of C. albrechti was fixed. Under these
considerations, the hybridization hypothesis concerning the parallel evolution of the
pronotal character seems to be very plausible.
Establishment of reproductive isolation
The phylogenetic relationships of the subspecies obtained by cladistic analysis and the
relationships between subspecies are shown in Fig. 78. There are only two combinations of
subspecies, viz., C. a. okumurai with C. a. esakianus, and C . a. echigo with C . a.
tsukubanus, which have narrow hybrid zones between them. They are inferred to have
become more strongly isolated reproductively than the others. Other combinations of subspecies have wider transitional zones between them, so that they are infered to be less iso-
albrechti
hidakanus
itoi
tohokuensis
yamauchii
awashimae
1
hagai
tsukubanus
frey i
esakianus
echigo
ukumurai
-
1.
1
Fig. 78. Relationship between phylogeny and hybridization. Bold brackets refer to the combinations more
reproductively isolated; simple brackets refer to the combinations less reproductively isolated. Solid rectangle is a putative event concerning the establishment of reproductive isolation.
Subspeciation and distribution pattern of Curubus ulbrcchii
95
lated reproductively. Therefore, the occurrence of some event, which was concerned with
the establishment of reproductive isolation, is predicted in the ancestor of component 3
(Fig. 78). Consequently, the hybrid fertility or viability is reduced between the subspecies
derived before and after this event. Such a punctuated evolution of reproductive isolation is
predicted theoretically as fixation of favorable mutations, which is not the by-product of
selective change in additively inherited polygenic traits (Charlesworth et al., 1987), or may
be due to cytoplasmic incompatibility by the infection of symbiotic microorganisms in cytoplasm (Thompson, 1987). For the verification of these hypotheses, experimental hybridization between different combinations of subspecies not occurring in nature should be attempted.
Conclusion
The differentiation within C. albrechti to the subspecies level has completed by northward movement after repeated dispersals from central Honshu where this species was first
established. It is assumed that this process was complete before the divergence of the other
Ohomopterus species, since C. albrechti is the only species found as far north as Hokkaido
where it diverged to establish three allopatric subspecies. To show the process of the differentiation within C. albrechti more precisely, the investigation of molecular phylogenies on
the basis of maternally and zygotically inherited gene sequences would be needed together
with discussion in relation to the geological history of the Japanese Archipelago.
In additon, this species will offer valuable material for studies of hybridization and speciation, because it shows various levels of hybridization in areas or zones where the ranges
of subspecies are contiguous.
Acknowledgements
We express our gratitude to Professor Dr. T. Yamasaki, Department of Natural History, Tokyo Metropolitan University for his help for our study and support to the publication of this paper. Takami, the senior
author, is indebted to him for his guidance in the course of the present study. We particularly thank Dr. Marius
Wasbauer (Brookings, U. S. A.) for reading the entire text in its original form, and Dr. K. Kubota (University
of Tokyo), Dr. T. Sota (Shinshu University) and Mr. M. Ujiie (Kashiwa) for their cooperation in the present
study in various ways. Mr. 0. Tominaga (Nara) provided the senior author with valuable information on the
distribution of certain subspecies. We also thank Dr. T. Torii (Tokyo Metropolitan University) for valuable
discussions concerning cladistic analysis. Our thanks should also be extended to the following persons for the
gift of the specimens examined which were indispensable to the present study. Most of them were donated to
Ishikawa, the junior author for a long period: Mr. T. Abe (Tokyo), Dr. T. Asami (Tachikawa Junior College),
Mr. S. Asawa (Matsumoto), Mr. N. Fujino, Mr. Y. Fukazawa, Mr. K. Haga (Yuzawa), Mr. R. Hatano
(Hirosaki), Dr. F. Hayashi (Tokyo Metropolitan University), Mr. Y. Inoue (Kasugai), Mr. T. Ishida (Uji), Mr.
E. Ishikawa (Tokyo), Mr. K. It6 (Memuro), Mr. K. Itsuji, Mr. H. Koike (Shibata), Mr. J. Kojima, Mr. S.
Kondo, Dr. K. Kubota (University of Tokyo), Dr. Y. Maeta (Shimane University), Mr. A. Makimoto, Mr. H.
Maruoka, Mr. K. Miyashita (Ebetsu), Ms. M. Momiyama (Machida), Dr. K . Morimoto (Kyushu University),
Dr. M. Munakata (Hakodate), Mr. T. Narumi (Joetsu), Mr. Z. Naruse, Dr. K. Nemoto (Tokyo University), Mr.
Niki, Mr. K. Okubo, Mr. T. Okumura (Yokohama), Mr. Y. Ota, Mr. H. Otobe, Mr. T. Sait6 (Tokyo Metropoli-
96
tan University), Mr. S. Sat& Mr. T. Shiina, Mr. Y. Shimamura (Kamakura), Dr. A. Shimizu (Keio High
School). Mr. T. Suda (Sakura), Mr. K. Suga (Tokyo), Dr. T. Sugawara (Matsumoto), Mr. Y. Syu (Yokohama),
Mr. A. Toba (Tokyo), Dr. S. Uchida (Biwako Museum), Dr. S. Ueno (National Sciense Museum, Tokyo), Mr.
M. Ujiie (Kashiwa), Mr. S. Yamauchi (Aomori Prefectural Museum), Dr. Y. Yasuda (Sendai), and Dr. S.
Yasugi (Tokyo Metropolitan University).
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1962. Carabidae 1. Insecta Japonica, 2 (3). 2+98 pp., 6 pis., Hokuryukan, Tokyo. (In Japanese, with
English summary.)
1963. The Family Carabidae, In T. Nakane, K. Ohbayashi, S. Nomura & Y . Kurosawa (Eds.),
Iconogruphia Insectorum Japonicorum, Colore nuturali editu, Vol. 2 (Coleoptera): 3-54, Pis. 3-27.
Hokuryukan, Tokyo. (In Japanese.)
uwukuz~~.sunus
in the central
Shida, T. 1984. Distribution of Carahus ulbrechti esakianus and C . 1~~w~i.siurtu.s
Chiba, (34):
part of the B6s6 Peninsula, Chiba Prefecture. Bull. Biol. Soc'. Chibu (Chiha-seihutsi~.s/t~),
22-29. (In Japanese.)
Su, Z-H., Tominaga, O., Ohama, T., Kajiwara, E., Ishikawa, R., Okada, T. S., Nakamura, K. and Osawa, S.
1996. Parallel evolution in radiation of Ohomopterus ground beetles inferred from mitochondria1
ND5 gene sequences. J. Md.Evol., 43: 662-671.
Thompson, J. N. 1987. Simbiont-induced speciation. Biol. .I. L h n . Soc.. 32: 385-393.
Errata
Page
56
56
60
60
60
63
63
70
70
78
78
83
83
84
84
87
90
95
95
96
Line
For
17
17
9 from bottom
8 from bottom
8 from bottom
17 from bottom
17 from bottom
8
8
6 from bottom
6 from bottom
12
14
15
11 from bottom
15 from bottom
13
6
20
11
Read
(= Carabus
esakianus)
frollowing localities:
[=Carabus
esakianus]
following localities:
4388
44<^8
paratypes from
holotype and paratypes from
50^
6c?d'
paratypes from
1688
1 7 88
paratypes from
2 m
holoty pe and paratypes from
Apical portion
Apical portion
in Appendix 1.
by the MIX program
parsimony
C.a. okumurai
Charlesworth et al.
In addition
18
paratypes from
Apical part
Apical part
in the Appendix 1.
by the Mix program
parsmony
C. a. okumurai
Charlesworth et al.
In additon
Breuning, S. 1934.
1934.
Appendix 1
Character matrices used in the cladistic analysis. a, Original matrix described in 'Evaluation of diagnostic
characters'; b, Matrix analyzed by PHYLIP, which is recoded binarily.
OTU
a.
albrechti
hidahnus
itoi
tohokuensis
yamauchii
awashimae
hagai
tsukubanus
freyi
esakianus
echlgo
okumurai
C. I. lewisianus
C. /. awakazusanus
C, kimurul
C. yamato
C. daisen okiunus
b.
albrechti
hidahnus
itoi
tohokuensis
yamauchii
awashimae
hagai
tsukubanus
freyi
esakianus
echigo
okumurai
C. 1. lewisianus
C. kimurai
C. yamato
C. daisen okiunus
1
I
2
3
4
5
6
Characters
7 8 9
10 11 12 13 14
IS
Subspeciation and distribution pattern of Curubif-,ulhrechti
Appendix 2
Index map of north-eastern Japan, in which Carabus albrechti is distributed. 1 , Aomori Prefecture; 2,
Iwate Pref.; 3, Akita Pref.; 4, Miyagi Pref.; 5, Yamagata Pref.; 6, Fukushima Pref.: 7, Ibaraki Pref.; 8,
Tochigi Pref.; 9, Gunma Pref.; 10, Saitama Pref.; 11, Chiba Pref.; 12, Tokyo Metropolis; 13, Kanagawa
Pref.; 14, Niigata Pref.; 15, Nagano Pref.: 16, Yamanashi Pref.
Okushiri Is.
T6hoku District

akasaka nanae

Transcription

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