Accep ed by J. Klimaszewski: 5 Ma . 2013; published: 24 Ap . 2013
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ZOOTAXA
ISSN 1175-5326 (p in edi ion)
ISSN 1175-5334 (online edi ion)
Copy igh © 2013 Magnolia P ess
Zoo axa 3641 (3): 201–222
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Re ision o he Subgenus Cop ocha a Mulsan & Rey o he Genus Aleocha a
G a enho s om Japan (Coleop e a: S aphylinidae: Aleocha inae)
SHÛHEI YAMAMOTO1, 2 & MUNETOSHI MARUYAMA2
1En omological Labo a o y, G adua e School o Bio esou ce and Bioen i onmen al Sciences, Kyushu Uni e si y, Hakozaki 6-10-1,
Fukuoka, 812-8581 Japan.
E-mail: s.yamamo [email protected]
2The Kyushu Uni e si y Museum, Hakozaki 6-10-1, Fukuoka, 812-8581 Japan
Abs ac
A axonomic e ision o he subgenus Cop ocha a Mulsan & Rey, 1874 o he genus Aleocha a G a enho s , 1802 in
Japan is p esen ed. The ollowing h ee species a e ecognized: Aleocha a (C.) e na Say, 1833, A. (C.) bino a a K aa z,
1856 and A. (C.) squali ho ax Sha p, 1888, o which A. bino a a is new o Japan. All p e ious eco ds o “A. (C.) bipus-
ula a (Linnaeus, 1760)” should be ega ded as misiden i ica ions o ei he A. e na o A. bino a a. New eco ds a e added
o , A. squali ho ax, a li o al species. All species a e edesc ibed, igu ed, keyed and mapped.
Key wo ds: Aleocha ini, biodi e si y, iden i ica ion key, edesc ip ion, o e bee le, axonomy
In oduc ion
The o e bee le genus Aleocha a G a enho s , 1802 is a la ge g oup in Aleocha inae, comp ising mo e han 450
species in 19 subgene a (Pa k & Ahn, 2010; Yamamo o & Ma uyama, 2012). Aleocha a is dis ibu ed
h oughou he wo ld, excep o An a c ica (Klimaszewski, 1984; Maus e al., 2001). Aleocha a is one o he
axonomically di icul g oups in Coleop e a. The genus includes many la ge adul s and commonly ound
species o Aleocha inae; hus, p ecise and simple iden i ica ion me hods a e equi ed. Close simila i ies in
ex e nal and in e nal s uc u es, and cosmopoli an dis ibu ions o se e al species make species iden i ica ion
di icul .
La ae o Aleocha a a e known o be pa asi oids o cyclo haphous Dip e a, and some species, especially
membe s o he subgenus Cop ocha a Mulsan & Rey, 1874, a e expec ed o be biological con ol agen s o
no o ious pes lies (e.g., Klimaszewski & Jansen, 1993, 1994; Maus e al., 1998, 2001; Fou ne e al., 2000). Thus,
mos Aleocha a species ypically occu in ly-in es ed habi a s such as animal d oppings, decaying plan ma e ial,
and ca ion (Klimaszweski, 1984; Yamamo o & Ma uyama, 2009, 2012).
Aleocha a species in Asia a e a om adequa ely desc ibed, and axonomic knowledge o Japanese Aleocha a
is s ill incomple e; only 25 species ha e been eco ded (Be nhaue & Schee pel z, 1926; Sme ana, 2004;
Yamamo o & Ma uyama, 2009, 2012).
The species o Cop ocha a ha e been p oblema ic e en in Eu ope and No h Ame ica, and o da e, no one has
conduc ed a axonomic e iew o he Japanese species o he subgenus. The pu pose o his s udy, he hi d se ies o
con ibu ions o he Japanese Aleocha a auna, is o cla i y he axonomic iden i y o Japanese Cop ocha a species
and o in es iga e geog aphical a ia ion wi hin each species. All h ee species a e edesc ibed wi h diagnosis and
igu es. Diagnos ic keys and dis ibu ion maps a e also p o ided. Figu es showing collec ion si es o wo species
a e included.
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Ma e ials and me hods
Deposi ion o ma e ials
App oxima ely 700 d y specimens o adul indi iduals we e examined. Mos o he ma e ial is deposi ed in he
ollowing public ins i u ions and p i a e collec ions.
Ins i u ions:
BMNH The Na u al His o y Museum, London, UK (R. Boo h).
CBM The Na u al His o y Museum and Ins i u e, Chiba, Japan (A. Sai ô).
KUM The Kyushu Uni e si y Museum, Fukuoka, Japan (S.-I. Naomi & S. Nomu a Ro e Bee le Collec ion,
and M. Ma uyama Collec ion) (M. Ma uyama).
SCM The Sagamiha a Ci y Museum, Kanagawa, Japan (H. Mo iya).
P i a e collec ions:
cHaga Kao u Haga (Tôkyô, Japan).
cHay Yasuhiko Hayashi (Hyôgo, Japan).
cI o Ta eo I ô (Kyô o, Japan).
cOno Hi oki Ono (Chiba, Japan).
cShi Yasu oshi Shiba a (Tôkyô, Japan).
cWa Takashi Wa anabe (Kanagawa, Japan).
cWa Y Yasuaki Wa anabe (Tôkyô, Japan).
cYam Shûhei Yamamo o (Fukuoka, Japan).
Measu emen s and e minology
The echnical p ocedu es, e minology, and o he me hods adop ed he e we e gi en in de ail in Ma uyama (2006)
and Yamamo o and Ma uyama (2012). Wo ds o geni al pa s we e adop ed mainly om Welch (1997) and Maus
(1998).
The specimen da a used in his s udy la gely did no ollow he o iginal spellings and place names w i en on
specimen labels. We co ec ed o mo e p ecise and de ailed in o ma ion, excep o some ype ma e ials. The
o iginal spellings on he label a ached o he specimen we e adop ed in such cases (a e Yamamo o & Ma uyama,
2012).
Abb e ia ions o mo phology and ma e ials
Geni al s uc u es a e abb e ia ed as ollows: ai, apical in agina ion o spe ma heca; bs, basal swelling o median
lobe; sa, apical po ion o spe ma hecal s em; sb, basal po ion o spe ma hecal s em; sga, a achmen o
spe ma hecal duc ; sh, head o spe ma heca; sm, memb anous po ion o spe ma hecal duc ; and sn, neck o
spe ma heca (a e Welch, 1997: Figs. 1, 4; see Figs. 18, 21, 29 in he p esen pape ).
Condi ions o specimen labels a e abb e ia ed as ollows: HW, hand w i en; PC, pape ca d; PRL, pu ple
ound label pinned by a cu a o ; RRL, ed ound label pinned by a cu a o .
Ely al colo a ia ions in Aleocha a squali ho ax a e abb e ia ed as ollows: GT, g ay ype; YT, yellow ype.
Glossa y o Japanese local names
Japanese place names gene ally ollow he same spelling used in Japan excep o some English nouns. Wo ds
nea ly co espond as ollows: -mu a ( illage); -chô, -machi ( own); -ku (wa d/dis ic ); -shi (ci y); -gun (coun y); -
o, - u, -ken (p e ec u e); -dake, - ake, -san, -zan, -yama (moun ain); -misaki, -zaki (cape); -gawa ( i e , s eam); -
kaigan, -hama (seasho e beach); -shima (island); -sho ô (islands, a chipelago).
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FIGURES 1–4. Japanese species o Aleocha a (Cop ocha a) in habi us. 1. Aleocha a (Cop ocha a) e na; 2. A. (C.) bino a a;
3–4. A. (C.) squali ho ax: g ay ype (Fig. 3) and yellow ype (Fig. 4).
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FIGURES 5–13. Japanese species o Aleocha a (Cop ocha a). 5–7. Aleocha a (Cop ocha a) e na; 8–10. A. (C.) bino a a;
11–13. A. (C.) squali ho ax. (5, 8, 11: ely a; 6, 9, 12: in e coxal p ocess o me a en i e; 7, 10, 13: meso en i e and
me a en i e).
Sys ema ics
Genus Aleocha a G a enho s , 1802
See Klimaszewski (1984), Sme ana (2004), and Gouix and Klimaszewski (2007) o synonymic in o ma ion and
e e ences. A de ailed desc ip ion is p o ided by Klimaszewski (1984) and a sho desc ip ion is p o ided by
Yamamo o and Ma uyama (2012).
Subgenus Cop ocha a Mulsan & Rey, 1874
Cop ocha a Mulsan & Rey, 1874: 146; Klimaszewski, 1984: 14; Lohse, 1989: 236; Klimaszewski & Jansen, 1994: 148;
Welch, 1997: 3; Maus, 1998: 83; Maus & Ashe, 1998 (online); Klimaszewski e al., 2000: 237; Sme ana, 2004: 355; Gouix
& Klimaszewski, 2007: 23.
Ba yodma Thomson, 1858: 31.
Eucha ina Casey, 1906: 165. [homonym].
Funda Blackwelde , 1952: 166. [ eplacemen name].
Meco hopalus Solie , 1849: 347.
Skenocha a Be nhaue & Schee pel z, 1926: 795.
See Klimaszewski (1984) and Maus (1998) o u he e e ences.
Type species: Aleocha a bilinea a Gyllenhal, 1810.
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Diagnosis. The subgenus Cop ocha a can be dis inguished om he o he subgene a o Aleocha a by he
combina ion o ollowing cha ac e s a es (see de ails in Klimaszewski, 1984; Klimaszewski & Jansen, 1994;
Maus, 1998): an ennae hick wi h segmen s V–X clea ly ans e se; wo longi udinal, pa allel o subpa allel ows
o mo e o less imp essed punc u es on midline o p ono um; ely a in some species wi h o ange o yellow spo ;
meso en i e wi h comple e ca ina; spe ma heca usually mul iply coiled pos e io ly, a ying om 1 o mo e han
100 coils; median lobe o aedeagus wi h lagellum and dis inc i ely a anged scle i es (Figs. 18–19), lacking
subapico- en al p ojec ions (see Yamamo o & Ma uyama, 2012: Fig. 18).
Rema ks. The subgenus Cop ocha a includes 37 species wo ldwide and 18 species om he Palea c ic egion
(Maus, 1998, 2000; Sme ana, 2004). This subgenus is widely dis ibu ed in all zoogeog aphical egions
(Klimaszewski, 1984). The axonomy o he subgenus Cop ocha a is di icul due o ex e nal simila i y,
conside able a ia ion wi hin species, and inco ec in e p e a ions o he con igu a ion o male geni alia (Maus,
1998). Fo example, species iden i ica ions in England we e con used un il he la e 1980s (Welch, 1990, 1997).
Despi e hese axonomic di icul ies, species belonging o he subgenus ha e a ac ed special a en ions as
biological con ol agen s o pes lies. Mos s udies on he subgenus ha e used A. (C.) bilinea a Gyllenhal, 1810,
A. (C.) bipus ula a (Linnaeus, 1760), and A. (C.) e na Say, 1833. One ad an age o using he subgenus
Cop ocha a o biological con ol is ha hey a e na i e species wi hin a wide dis ibu ion ange, which educes
ecological impac on ecosys ems.
Phylogene ic ela ionships among species o he subgenus Cop ocha a a e ela i ely well known. Maus and
Ashe (1998), based on mo phological cha ac e s, and Maus e al. (2001), based on molecula da a, p oduced
phylogene ic ees o he subgenus. Howe e , Maus and Ashe (1998) no ed di icul ies in he examina ion o
phylogene ic ela ionships wi hin he subgenus, such as he high equencies o pa allelism and homoplasy in he
g oup, and mos o he apomo phic cha ac e s a es ha e e ol ed independen ly in di e en lineages wi hin he
subgenus.
Th ee species o he subgenus Cop ocha a ha e been eco ded in Japan, A. bipus ula a, A. squali ho ax Sha p,
1888, and A. e na, he la e eliably only om Gunma and Kanagawa P e ec u es (Maus, 1998; Pa k e al., 2011).
“Aleocha a bipus ula a”, on he o he hand, has been widely eco ded om Japan. The li o al species, A.
squali ho ax was o iginally desc ibed om Hokkaidô and Honshû (Sha p, 1888).
Key o species om he subgenus Cop ocha a in Japan
1. Body (Figs. 3–4) g ay, hick, spindle shaped; do sal su ace co e ed wi h g anula mic os uc u es and o ebody no glossy a
all. In e coxal p ocess o me a en i e (Figs. 12–13) na ow and long, poin ed apically. Ely a (Fig. 11) wi h pos e io ma gins
deeply no ched la e ally; su ace ough, densely co e ed wi h sho bu obus se ae. Te gi e VIII (Figs. 34–35) wi h nume ous
la ge o al p ojec ions on su ace. Collec ed om decaying seaweed on seasho e beach. [Male]: median lobe o aedeagus as in
Figs. 38–39; scle i e Z la ge, wi hou a achmen a apex; lagellum longe han median lobe o aedeagus. [Female]: p oximal
po ion o spe ma heca (Fig. 41) wi h ex emely nume ous coils. . . . . . . . Aleocha a (Cop ocha a) squali ho ax Sha p, 1888
- Body black o blackish b own, slende , pa allel o subpa allel sided; do sal su ace no co e ed wi h g anula mic os uc u es,
and clea ly glossy. In e coxal p ocess o me a en i e wide and ounded apically. Ely a wi h pos e io ma gins ounded, usu-
ally wi h yellowish po ion. Te gi e VIII wi hou la ge o al spines on su ace, wi h se a e o unca e pos e io ma gin. Spe -
ma heca p oximally wi h 20 coils a mos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Do sal su ace smoo h and appa en ly glossy (Fig. 1). P ono um wi h shallow and inconspicuous punc u es and wi h hin se ae.
Two ows o punc u es along midline ine. Ely a (Fig. 5) wi h a pai o small eddish o yellowish spo s nea pos e io ma gin
and su u e, bea ing wi h nume ous hin and inconspicuous se ae. Common species in Japan. [Male]: s e ni e VIII (Fig. 16)
simple and weakly poin ed. Median lobe o aedeagus (Figs. 18–19): apex o apical lobe o median lobe blun in la e al iew;
lagellum long, sligh ly longe han median lobe o aedeagus; scle i e Z la ge, wi h long and p ominen a achmen , wa ing
owa d apex. [Female]: coiled po ion o spe ma heca (Fig. 21) no g ea ly ex ended la e ally.. . . . . . . A. (C.) e na Say, 1833
- Body mo e slende . Ely a en i ely yellow excep o an e io ma gins nea p ono um (Japanese specimens). . . . . . . . . . . . . .3
3 Do sal su ace wi h dis inc and deep coa se punc u es, and wi h hick yellowish se ae (Fig. 2). P ono um wi h dis inc punc-
u es and ows o deep punc u es along midline. [Male]: s e ni e VIII (Fig. 24) poin ed iangula ly. Median lobe o aedeagus
(Figs. 26–27): apex o apical lobe sha ply poin ed in la e al iew; lagellum sho , bu basal pla e o lagellum long and la ge;
scle i e Z small, wi h s aigh a achmen poin ed owa d apex. In Japan known only om Hokkaidô. [Female]: p oximal po -
ion o spe ma heca (Fig. 29) wi h hick coils and widely ex ended la e ally. . . . . . . . . . . . . . . . . A. (C.) bino a a K aa z, 1856
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FIGURES 14–21. Te minalia o Aleocha a (Cop ocha a) e na. 14. e gi e VIII o male; 15. e gi e VIII o emale; 16. s e ni e
VIII o male; 17. s e ni e VIII o emale; 18. male geni alia: median lobe o aedeagus in la e al iew; 19. male geni alia: median
lobe o aedeagus in en al iew; 20. apical lobe o pa ame i e, la e al iew; 21. emale geni alia: spe ma heca.
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FIGURES 22–29. Te minalia o Aleocha a (Cop ocha a) bino a a. 22. e gi e VIII o male; 23. e gi e VIII o emale; 24.
s e ni e VIII o male; 25. s e ni e VIII o emale; 26. male geni alia: median lobe o aedeagus in la e al iew; 27. male geni alia:
median lobe o aedeagus in en al iew; 28. apical lobe o pa ame i e, la e al iew; 29. emale geni alia: spe ma heca.
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FIGURES 30–33. Mou h pa s o Aleocha a (Cop ocha a) squali ho ax o male. 30. labium; 31. maxilla; 32. lab um; 33.
men um.
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FIGURES 34–41. Te minalia o Aleocha a (Cop ocha a) squali ho ax. 34. e gi e VIII o male; 35. e gi e VIII o emale; 36.
s e ni e VIII o male; 37. s e ni e VIII o emale; 38. male geni alia: median lobe o aedeagus in la e al iew; 39. male geni alia:
median lobe o aedeagus in en al iew; 40. apical lobe o pa ame i e, la e al iew; 41. emale geni alia: spe ma heca.
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Ma sue-shi, Shimane-ken, 17. VII. 2010, S. Yamamo o leg. (unde d y lo sam on sandy beach; cYam); 1
♂
(GT), 1
♀
(GT), 5 sex? (GT), Taki-chô, Hikawa-gun, Shimane-ken, 2. VI. 1980, S.-I. Naomi leg. (KUM). [Kyûshû]: 5
♂♂
(GT), 1
♂
(YT), 2
♀♀
(GT), 1
♀
(YT), 122 sex? (GT), 17 sex? (YT), Wa a i, Fuku su-shi, Fukuoka-ken, 21. IV.
2012, S. Yamamo o, M. Ma uyama, T. Kanao leg. ( om d y o we seaweed on huge sandy beach; KUM); 5 sex?
(GT), 3 sex? (YT), Wa a i, Fuku su-shi, Fukuoka-ken, 21. IV. 2012, S. Yamamo o, M. Ma uyama, T. Kanao leg.
( om d y o we seaweed on small sandy beach; KUM); 1
♂
(YT), 1
♀
(YT), 4 sex? (GT), 1 sex? (YT), Mi oma,
Higashi-ku, Fukuoka-shi, Fukuoka-ken, 3. VI. 2001, S. Oga a leg. (KUM); 2 sex? (GT), Ka suma (Shikano-shima),
Higashi-ku, Fukuoka-shi, Fukuoka-ken, 21. III. 2012, S. Yamamo o leg. ( om seaweed on huge sandy beach;
cYam); 1 sex? (GT), Ka a su-shi, Saga-ken, 7. VII. 1975, H. Ohishi leg. (KUM); 2 sex? (GT), Minamiôsumi-chô
(Sa a-misaki), Kimo suki-gun, Kagoshima-ken, 28. V. 1958, Y. Miyake leg. (cHay); 1
♀
(GT), 11 sex? (GT),
Naka ane-chô (Tanega-shima), Kumage-gun, Kagoshima-ken, 8-13. V. 1996, M. Ma uyama leg. (KUM).
Redesc ip ion. Body (Figs. 3–4): hick and spindle shaped, especially o small indi iduals (Fig. 4), small o
medium sized, ca. 3.48 mm in body leng h (3.14–3.83 mm, N = 15), and 1.88 mm in o e body leng h (1.68–2.15
mm, N = 15), no mally mode a e in size; en i e body hea ily obus ; do sal su ace ma ; o ebody co e ed wi h
g anula mic os uc u es. Colo (Figs. 3–4, 11–13): g ound colo g ay o blackish b own; legs, especially a sal
segmen s, and mou h pa s da k b own o eddish b own; ely a g ay in mos cases as Fig. 3, bu some indi iduals
wi h huge yellowish po ion excep o an e io and la e al ma gins as Fig. 4; an ennae eddish b own o da k
b own, su ace densely pubescen wi h minu e yellowish se ae. Head (Figs. 3–4): ci cula (head leng h = 0.42–0.56
mm (mean = 0.48 mm), head wid h = 0.46–0.56 mm (mean = 0.51 mm), N = 15), sligh ly ans e se (head wid h /
head leng h = 1.08 (mean), N = 15), wides a eyes; su ace en i ely co e ed wi h hexagonal e icula ions, lacking
punc a ion, bu co e ed wi h sho and hick se ae spa sely. An ennae (Figs. 3–4): monili o m, obus and
hickened apically, sligh ly longe han head leng h (an enna leng h = 0.57–0.84 mm (mean = 0.73 mm), N = 15);
segmen I, abou 2.1 imes as long as b oad, segmen II sligh ly sho e han I, segmen III appa en ly sho e han
II, segmen s IV o VI mo e o less sphe ical, as long as wid h excep o s em o each segmen , segmen VII
mode a ely wide han long, segmen s VIII o IX clea ly ans e se, segmen X s ongly ans e se, segmen XI
hick and conical, nea ly 1.2 imes as long as wid h, as same leng h as segmen I, app oxima e ela i e leng h o
segmen s om basal o apex: 10.5: 10.0: 6.0: 4.5: 4.5: 4.5: 4.5: 4.5: 4.5: 4.5: 10.5. Mou h pa s (Figs. 30–33):
mandibles sligh ly asymme ic, le one wi h one oo h nea apex. Clypeus ounded apically. Lab um (Fig. 32)
ans e se, abou 1.3 imes as wide as long, an e io ma gin sligh ly ema gina ed medially, basal hal semi-
anspa en , apodeme oundly p oduced pos e o-medially; su ace wi h nume ous pseudopo es sca e ed andomly
in apical pa . Ligula (Fig. 30) bilobed, wi h each lobe ounded apically. Labial palpus (Fig. 30) wi h segmen I
hicke and longe han II, segmen II weakly diala ed, segmen III sho e han II. Men um (Fig. 33) nea ly
apezoidal, much wide han long, abou 2.0 imes as wide as long; an e io ma gin s ongly ema gina ed; en i e
su ace wi h nume ous pseudopo es sca e ed andomly. Maxilla (Fig. 31): maxilla y palpus wi h hick and long
segmen II and III, segmen II as long as III, segmen IV sho , less han hal leng h o segmen III; galea hick and
sho , clea ly sho e han segmen II o maxilla y palpus; lacinia wi h nume ous se ae and wi h dozens o hick
spines pec ina ely. Tho ax: p ono um (Figs. 3–4) ans e se (p ono um leng h = 0.60–0.76 mm (mean 0.69 mm),
p ono um wid h = 0.77–0.94 mm (mean = 0.88 mm), p ono um wid h / p ono um leng h = 1.27 (mean), N = 15),
clea ly wide han head (p ono um wid h / head wid h = 1.71 (mean), N = 15), wides a ound basal 1/3, weakly
na owing apically; su ace uni o mly co e ed wi h la ge hexagonal e icula ions, hick-sho se ae bu wi hou
dis inc punc a ion; midline longi udinally ele a ed weakly abo e, glab ous, lacking punc a ion. In e coxal p ocess
o meso en i e (Fig. 13) wi h s ongly de eloped ca ina along midline. In e coxal p ocess o me a en i e (Figs.
12–13) na ow and long, appa en ly poin ed apically. Me a en i e (Fig. 13) ough, densely pubescen like ely a.
Ely a (Figs. 3–4, 11) wide han long (ely a leng h = 0.62–0.78 mm (mean = 0.68 mm), ely a wid h = 0.90–1.13
mm (mean = 1.04 mm), N = 15), ugose, mode a ely co e ed wi h yellowish hick and sho se ae; pos e io
ma gins deeply no ched la e ally (Fig. 11); [G ay ype]: do sal su ace en i ely g ay, bu [Yellow ype]: wi h a pai
o la ge yellowish po ions, ex ending om apical o pos e io excep o an e io ma gins and shoulde s close o
p ono um as well as la e al ma gins o ely a and ely al su u e (pos e io ma gins colo ed). Legs (Figs. 3–4): sho
(hind ibia leng h = 0.49–0.65 mm (mean = 0.57 mm), N = 15) and hick, dozens o hick spines on ibia, especially
on o e and mid ibia; ela i e leng hs o a sal segmen s om basal o apical: 5.0: 3.5: 3.5: 3.0: 8.0 in o e a sus,
8.0: 5.0: 5.0: 4.5: 14.5 in mid a sus, 17.5: 10.0: 7.0: 7.0: 14.5 in hind a sus. Abdomen (Figs. 3–4): ogose, excep
o an e io ma gin o each e gi e and pos e io ma gin o e gi e VI o VII; e gi e VIII (Figs. 34–35) wi h dozens
o hick spines sca e ed in pos e io hal .
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[Male]: pos e io ma gin o e gi e VIII (Fig. 34) ounded, wi h app oxima ely 7 mac ose ae on each side.
S e ni e VIII (Fig. 36) wi h one mac ose a (di icul o iden i y); pos e io ma gin weakly poin ed. Median lobe o
aedeagus as in Figs. 38–39: compac ly elonga ed and na owed owa d apex; lagellum long, clea ly longe han
median lobe o aedeagus; scle i e Y la ge; scle i e Z la ge wi hou a achmen a apex; apical lobe in la e al iew
sligh ly ben , na owing subapically, median lobe in en al iew elonga ed, weakly na owing owa d apex; basal
swelling o median lobe la ge and o al; apical lobe looks sho isosceles. Apical lobe o pa ame i e (Fig. 40) sho
and hick, wides a subapical po ion, bea ing 4 se ae.
[Female]: e gi e VIII (Fig. 35) wi h ounded pos e io ma gin like male, wi h app oxima ely 4 mac ose ae.
Pos e io ma gin o s e ni e VIII (Fig. 37) weakly poin ed, wi h one mac ose a (di icul o con i m). Spe ma heca
(Fig. 41): apical in agina ion o spe ma heca iny and shallow; spe ma hecal head used wi h spe ma hecal neck;
a achmen o spe ma hecal duc p ominen ; (sn) sho , sligh ly na owing owa d basal po ion o spe ma hecal
s em; coiled po ion ex emely long, composed wi h coun less imes o mode a ely hick coils, no ex ended
la e ally, wa ing owa d memb anous po ion o spe ma hecal duc ; coils di ec ly connec ed wi h hick (sm); each
pa o spe ma heca excep o (sm) en i ely and s ongly scle o ized; inne wall o (sh) coa sely and ha o (sb) in
apical po ion also coa sely s ia e wi h ough mesh s ia e in mid o pos e io pa o (sb).
Diagnosis. Aleocha a squali ho ax can be easily dis inguished om he o he Cop ocha a in Japan by a
combina ion o he ollowing cha ac e s a es: body hick and spindle shaped; su ace ma wi h g anula
mic os uc u es, co e ed wi h hick and sho se ae coa sely bu wi hou punc a ion on head and p ono um (Figs. 3–
4); p ono um clea ly wide han head, wi hou ows o punc u es along midline; ely a wi h pos e io ma gin deeply
no ched, some imes wi h yellowish po ion on do sal su ace (YT: Figs. 4, 11); in e coxal p ocess o me a en i e
na ow and sha ply poin ed apically (Fig. 13); e gi e and s e ni e VIII wi h hick spines in pos e io hal (Figs. 34–
35). [Male]: lagellum long; scle i e Z la ge wi hou a achmen (Figs. 38–39); apical lobe o pa ame i e sho ,
hick, wi hou nume ous minu e se ae (Fig. 40). [Female]: coun less coiled po ion wi h spe ma heca (Fig. 41).
Con i med dis ibu ion by he p esen s udy. [JAPAN]: Hokkaidô, Honshû, Kyûshû, Tanega-shima.
O he locali ies in he li e a u e. [EAST ASIA]: Sou h Ko ea (Ahn e al., 2000; Pa k & Ahn, 2004).
Rema ks. Aleocha a squali ho ax is a a he common species in Japan bu has a limi ed dis ibu ion globally.
Un il now, his axon has only been eco ded om Japan and Sou h Ko ea (Sme ana, 2004; F ank & Ahn, 2011). A
ecen s udy p o ided he i s eco d o A. squali ho ax om Sou h Ko ea and b ie ly edesc ibed he species and
p o ided a igu e o he median lobe o he aedeagus o he male, based on Ko ean specimens (Ahn e al., 2000).
The dis ibu ion o A. squali ho ax in Japan is wide, om no heas e n Hokkaidô (Nemu o) o sou hwes e n
Kyûshû (Tanega-shima). We p o ide new dis ibu ional eco ds om mainland Kyûshû.
This species is conside ed a highly de i ed axon o he subgenus Cop ocha a and i has a ac ed special
a en ion om esea che s (Be nhaue & Schee pel z, 1926; Assing, 1995; Maus, 1998). Thus, he line d awings o
mou h pa s, which a e impo an cha ac e s in Aleocha inae ha ha e ne e been desc ibed, a e also p o ided in
his pape .
His o ical no es. This species was o iginally desc ibed as “Aleocha a squali ho ax” om “Hagi [Hagi-shi,
wes e n end o Honshû, Japan] (Fau el), Hakoda e [Hakoda e-shi, Hokkaidô, Japan] (Lewis)” by Sha p (1888).
A e wa d, Fenyes (1920) included i wi hin he peculia subgenus Eucha ina Casey, 1906. A ew yea s la e ,
Be nhaue and Schee pel z (1926) ans e ed A. squali ho ax o he newly es ablished subgenus Skenocha a
Be nhaue & Schee pel z, 1926 as he ype species, and i was he only species assigned o Skenocha a. Assing
(1995) aised his subgenus o gene ic ank (s ill only one species emained in he axon). This pape was especially
impo an because A. squali ho ax was edesc ibed wi h igu es and he male lec o ype was designa ed. The
sys ema ic posi ion o Skenocha a was unclea , al hough Assing (1995) en a i ely ea ed i as a dis inc genus.
Maus and Ashe (1998) and Maus (1998) de ec ed Skenocha a wi h all apomo phies ha cha ac e ize he
subgenus Cop ocha a; i.e., he exis ence o do sal longi udinal ows on he p ono um, a s ongly coiled
spe ma heca in he emale, and a comple ely ca ina e meso en i e. The e o e, Skenocha a was synonymized unde
he subgenus Cop ocha a by Maus (1998). In addi ion o mo phological e idence, molecula in o ma ion
suppo ed his ea men (Maus e al., 2001; see Phylogeny o A. squali ho ax).
Bionomics. SY and MM collec ed he species a se e al di e en loca ions in Hokkaidô, Honshû and Kyûshû
ha aced bo h he Paci ic Coas and he Sea o Japan (Fig. 45). All o he loca ions we e sandy beaches, none o
he a eas we e e y small, and hey gene ally had good en i onmen al condi ions (Fig. 43). Indi iduals we e ound
om comple ely d y seaweed masses on beaches ha we e a om he sho eline om si es ha we e and close o
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coas al ege a ion a Hamama su and U ayako an in Hokkaidô; specimens we e also collec ed om somewha we
seaweed a Wa a i, Kyûshû (Fig. 43). Adul s we e obse ed and collec ed om ea ly sp ing (Ma ch) o mid- all
(Oc obe ), wi h he highes numbe o indi iduals being obse ed be ween Ap il and May.
Hos eco ds. No in o ma ion is a ailable o A. squali ho ax.
Phylogeny. Phylogene ic ees in Maus e al. (2001) show ela ionships among A. e na, A. bino a a, and A.
squali ho ax. All h ee species we e included in he Cop ocha a clade. Aleocha a squali ho ax o med a sis e
g oup wi h li o al A. (C.) sulcicollis Manne heim, 1843 om No h Ame ica. Mic osculp u e is p esen in
Empleno a, Maseocha a, Polys omo a, and A. squali ho ax, bu none o he analyses suppo ed a monophyle ic
clade o hese axa (Maus e al., 2001).
Colo a ia ion in ely a. Two pa e ns o ely al colo a ia ion we e ound. The g ay ype (Fig. 3) is
appa en ly he ypical o m and he yellow ype (Figs. 4, 11) is much a e . We examined bo h ypes
mo phologically, bu no di e ences we e ound be ween he wo ypes. Thus, he au ho s concluded ha he
di e ence ep esen s colo a ia ion wi hin A. squali ho ax. The yellow ype accoun ed o only 12.6% o he o al
numbe o indi iduals collec ed (N=470). The a io also showed conside able a ia ion among loca ions. Fo
example, no yellow- ype indi iduals we e ound among 55 specimens a San iga-hama in Hokkaidô. In con as ,
he yellow ype accoun ed o 14.1% in Wa a i (da a combined ac oss small and la ge beaches), Kyûshû (N=156),
and he highes a io, 45.9%, was eco ded in Ka suu a, Honshû (N=37).
FIGURES 42–43. Habi a s o he subgenus Cop ocha a o he genus Aleocha a in Japan. 93. he summi o Sobo-san (loca ed
in bounda ies be ween Ôi a and Miyazaki P e ec u es; only obse ed; A. (C.) e na: a ow); 43. Wa a i (Fuku su, Fukuoka
P e ec u e; A. (C.) squali ho ax: a ow).
Discussion
Does Aleocha a bipus ula a occu in Japan?
As a yellow-spo ed Aleocha a species, Aleocha a (Cop ocha a) bipus ula a (Linnaeus, 1760) was hough o be a
e y common species in Japan. The i s eco d o A. bipus ula a in Japan was gi en by Sha p (1888) as A. ni ida
G a enho s , 1802, which was la e synonymized unde A. bipus ula a. Adachi (1957) ecognized A. bipus ula a in
his ca alog o Japanese o e bee les. Naomi (1989) and Sme ana (2004) ollowed his ea men . Howe e , we
examined mo e han 250 Japanese specimens ha ha e hi he o been iden i ied as A. bipus ula a, and no specimen
o A. bipus ula a was ound. These specimens we e iden i ied as A. e na Say, 1833 o A. bino a a K aa z, 1856. In
conclusion, we exclude A. bipus ula a om he Japanese insec auna.
The con usions o A. bipus ula a in No h Ame ica add essed in Hemachand a e al. (2005) and add essed in
Eu ope in Welch (1997) illus a e some o he axonomic di icul ies ha ha e been encoun e ed wi h he species.
Maus (1998) iden i ied he dis ibu ion pa e n o A. bipus ula a, which includes all o Eu ope h ough No h A ica
o Pakis an and India. The e o e, he Sou h Ko ean eco d o A. bipus ula a in Cho and Ahn (2001) is highly
doub ul. Pa k e al. (2011) eco ded Aleocha a e na o he i s ime om Sou h Ko ea; he e o e, he Sou h
Ko ean eco d o A. bipus ula a migh be a misiden i ica ion o A. e na. Fu he s udy in Eas Asia is needed o
e alua e he ue di e si y o he subgenus Cop ocha a in he egion.
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FIGURES 44–45. Collec ing locali ies o Aleocha a (Cop ocha a) spp. based on specimens examined in Japan. 44. Aleocha a
(Cop ocha a) e na (Indica ed by ed ci cle) and A. (C.) bino a a (Indica ed by yellow squa e); 45. A. (C.) squali ho ax
(Indica ed by blue iangle).
Examined specimen da a o Aleocha a bipus ula a. AUSTRIA: [Bu genland]: 1
♂
, 1
♀
, Illmi z, VI. 1995, K.
Peschke leg. (wi hou collec ing da e; de . D . C. Maus, 1996; KUM).
Geni al pa s o bo h sexes we e examined. This species is dis inguished om A. e na and A. bino a a by a
much smalle coiled po ion (1–5 coils) o he emale spe ma heca and he shape o a scle i e Z inside he median
lobe o he aedeagus in he male (Maus, 1998).
Di e si y and dis ibu ion
We ecognized only h ee species as Japanese congene s o he subgenus Cop ocha a. This numbe is much
smalle han he species coun s in USA (se en species: Klimaszewski, 1984) and Sou h A ica (six species:
Klimaszewski & Jansen, 1994). Howe e , i is e y close o he numbe o he species in UK ( ou species: Welch,
1997) and Sou h Ko ea ( h ee species, al hough A. bipus ula a is doub ul: Ahn e al., 2000; Cho & Ahn, 2001;
Pa k e al., 2011).
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Two species, A. e na and A. bino a a, ha e wide global dis ibu ion anges, bu A. squali ho ax is con ined o
Eas Asia. Aleocha a e na is a Hola c ic species and i s p esence in Japan is no su p ising. In con as , he inding
o A. bino a a om Japan is qui e unexpec ed. This species has mainly been eco ded om Eu ope. P e iously, he
eas e nmos eco d o his species was om eas Sibe ia (Sme ana, 2004). Fo ha eason, he Japanese specimens
ep esen he eas e nmos eco ds o A. bino a a. All o he Japanese specimens o his species we e collec ed om
Hokkaidô loca ed in no heas e n Japan, which has a con inen al clima e (Fig. 44).
Acknowledgemen s
SY co dially hank associa e p o esso D . Sa oshi Kami ani in Kyushu Uni e si y o con inual guidance and
suppo o he s udy in a ious ways. We exp ess ou since e g a i ude o D . Roge Boo h (BMNH), M . Hi o umi
Mo iya (SCM) and D . Akiko Sai ô (CBM) o loan o he ins i u ional ma e ials. Ou hanks also due o he
colleagues indica ed in he ma e ial sec ion o kindly p o iding us oppo uni ies o examine he specimens. We
hank D . Al ed F. New on J . and D . Ma ga e K. Thaye in he Field Museum o Na u al His o y, Chicago o
kindly o e ing us li e a u e e e ences. Finally, we deeply indeb ed o anonymous e iewe s o use ul commen s
on he d a . This s udy is a con ibu ion om he En omological Labo a o y, Facul y o Ag icul u e, Kyushu
Uni e si y, Fukuoka (se . 6, no. 117).
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APPENDIX
APPENDIX 1. Species lis o Japanese names o Aleocha a (subgenus Cop ocha a).
Subgenus Cop ocha a Mulsan & Rey, 1874
Scien i ic name Japanese name
Aleocha a (Cop ocha a) e na Say, 1833 Fu amon-higebu o-hanekakushi
A. (C.) bino a a K aa z, 1856 Kibane-higebu o-hanekakushi
A. (C.) squali ho ax Sha p, 1888 Fu o- suyakeshi-higebu o-hanekakushi
