Accep ed by J. Klimaszewski: 24 Jul. 2012; published: 15 Oc . 2012
ZOOTAXA
ISSN 1175-5326 (p in edi ion)
ISSN 1175-5334 (online edi ion)
Copy igh © 2012 · Magnolia P ess
Zoo axa 3517: 1–52 (2012)
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A icle
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u n:lsid:zoobank.o g:pub:F832C768-A8CA-4FEE-8C3B-BD933247FA6E
Re ision o he Seasho e-dwelling Subgene a Empleno a Casey and T iocha a
Be nhaue (Coleop e a: S aphylinidae: genus Aleocha a) om Japan
SHÛHEI YAMAMOTO1, 2 & MUNETOSHI MARUYAMA2, 3
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
3Co espoding au ho : E-mail: dend [email protected]
Abs ac
The Japanese species o he seasho e-dwelling subgene a Empleno a Casey and T iocha a Be nhaue o he genus
Aleocha a G a enho s a e e ised. Fi e species a e ecognised in Empleno a, o which h ee a e desc ibed as new
species: Aleocha a (Empleno a) seg ega a n. sp., A. (E.) hayamai n. sp. and A. (E.) yama o n. sp. The emaining known
species A. (E.) ucicola Sha p and A. (E.) pue zi (Assing) a e edesc ibed. Th ee species ecognised in T iocha a,
Aleocha a (T iocha a) isulca a Weise, A. (T.) ze chei (Assing) and A. (T.) nubis (Assing) a e edesc ibed. All species a e
keyed. Fo some species ecological da a a e epo ed. The phylogene ic ela ionships o he Japanese species a e
discussed, and he dis ibu ions o all species a e mapped.
Key wo ds: biodi e si y, coas al en i onmen , iden i ica ion key, Palaea c ic, edesc ip ion, sup a idal zones, sympa ic
species, axonomy
In oduc ion
Recen s udies ha e e ealed he wo ldwide coas al s aphylinid di e si y (Moo e & Legne , 1976; Hammond,
2000; F ank & Ahn, 2011), and he sub amily Aleocha inae is ep esen ed by 187 species h oughou he wo ld,
ep esen ing he la ges numbe o coas al s aphylinid bee les (F ank & Ahn, 2011). The genus Aleocha a
G a enho s , 1802 is ep esen ed by 16 coas al species belonging o ou subgene a (F ank & Ahn, 2011).
Aleocha a comp ises mo e han 450 species, and is dis ibu ed wo ldwide, excep o An a c ica (e.g.,
Be nhaue & Schee pel z, 1926; Klimaszewski, 1984; Maus e al., 2001). Mos species a e ound nea ly-in es ed
habi a s such as ca ion, animal d oppings, o decaying plan ma e ial. Mos Aleocha a species a e cha ac e ised by
unusual li e his o ies, i. e., he pa asi oid la ae use cyclo hapheous ly pupa ia as hos s. Thus, hey ac as
impo an na u al enemies o many dip e an species (e.g., Klimaszewski, 1984; Klimaszewski & Jansen, 1993;
Maus e al., 2001). Because o hei impo ance in biological con ol, qui e a ew s udies ha e been conduc ed on
some species o cla i y hei biology (Maus e al., 2001). The e a e, howe e , se e al issues s ill emaining o be
sol ed wi h ega d o he axonomy, sys ema ics, phylogeny and li e his o y o Aleocha a. Nume ous unpublished
synonyms, lack o adequa e keys o speci ic iden i ica ion, use o supe icial, and o en useless, ex e nal
cha ac e is ics, and a lack o pic u es o illus a ions pose majo di icul ies (Klimaszewski, 1984). Recen s udies
cla i ied he auna o Eas Asian Aleocha a pa ially such as he subgenus Xenocha a Mulsan & Rey, 1874 in
Sou h Ko ea (Pa k & Ahn, 2010), he subgenus Aleocha a G a enho s , 1802 in mainland China (Luo & Zhou,
2012), and some li o al subgene a (Assing, 1995; Ahn e al., 2000; Pa k & Ahn, 2004). In spi e o hese e o s, he
cu en si ua ion is s ill a om adequa ely wo ked ou in Asia. Knowledge o he Japanese Aleocha a auna is
also incomple e; o da e, only 22 species ha e been eco ded (Sme ana, 2004; Yamamo o & Ma uyama, 2009).
Fu he mo e, many p oblems emain in he axonomy o he Japanese Aleocha a, such as doub ul in e p e a ion o
mos species due o he absence o mode n e isions o ype ma e ial.
YAMAMOTO & MARUYAMA
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The subgene a o Empleno a Casey, 1884 and T iocha a Be nhaue , 1901 a e known as seasho e-dwelling axa
ha inhabi decaying seaweed, deb is, unde d i wood, and some imes ca ion on beaches (e. g., Klimaszewski,
1984; F ank & Ahn, 2011). Thei la ae a e ec opa asi oids o he pupae o dip e an species (Peschke & Fuldne ,
1977; Klimaszewski, 1984; Assing, 1995; Yamazaki, 2008, 2012), as a e membe s o he o he subgene a o
Aleocha a. The subgene a add essed he e a e conside ed as sis e g oups, based on mo phological and molecula
e idence (Assing, 1995; Maus e al., 1998a, 2001), and hey sha e a unique cha ac e s a e: a pai o subapico-
en al p ojec ions on he median lobe o he male aedeagus (Assing, 1995). Empleno a and T iocha a ha e
occasionally been ea ed as dis inc gene a (e. g., See e s, 1978; Lohse, 1984, 1985, 1989; Assing, 1995), bu
ecen s udies ha e con i med ha hese gene a a e phylogene ically nes ed wi hin he genus Aleocha a (Maus e
al., 2001).
The subgenus Empleno a includes eigh species om Eu ope, No h A ica, Eas Asia, and No h Ame ica on
bo h he A lan ic and Paci ic coas s (Klimaszewski, 1984; Assing, 1995; Maus & Ashe, 1998a; Sme ana, 2004;
F ank &Ahn, 2011). All o he h ee known species o T iocha a a e, on he o he hand, es ic ed o he Paci ic
coas o Eas Asian coun ies, namely, Russia (Fa Eas ), Japan, Sou h Ko ea, No h Ko ea, and Hong Kong
(Assing, 1995; Maus & Ashe, 1998c; Ahn e al., 2000; de Rougemon , 2001; Pa
ś
nik, 2001; Ma uyama, 2002; Pa k
& Ahn, 2004; Sme ana, 2004; F ank & Ahn, 2011). Taxonomic and aunal in o ma ion on bo h subgene a has been
g adually accumula ing.
S udy o he Japanese s aphylinid auna began wi h a se ies o pape s by Sha p in he la e 19 h cen u y, and he
i s Japanese Empleno a species, A. (E.) ucicola, was desc ibed as Aleocha a ucicola om “Amakusa and
Iwosima, nea Nagasaki” (Amakusa-shi, Kumamo o-ken (o Iô-jima, Nagasaki-shi, Nagasaki-ken), Kyûshû)
(Sha p, 1874). In he same decade, Weise (1877) desc ibed wo species, Aleocha a isulca a and A. a iolosa (as
genus Homalo a Manne heim, 1830), bo h om “Hagi” (Hagi-shi, Yamaguchi-ken, wes e n end o Honshû).
Be nhaue (1901b) es ablished he subgenus T iocha a o A. isulca a. La e , Fenyes (1920) ecognised
addi ional i e species om Aus alia in T iocha a, bu Be nhaue & Schee pel z (1926) subsequen ly excluded
hese i e species. As a esul , only A. isulca a was con i med in he subgenus a ha ime. On he o he hand, A.
ucicola was placed in he subgenus Empleno a in Be nhaue and Schee pel z (1926). Sawada (1971) edesc ibed
A. (T.) isulca a and A. (E.) ucicola wi h emphasis on he cha ac e s a es o mou h pa s. The Palaea c ic species
o Empleno a and T iocha a we e e ised by Assing (1995) (as dis inc gene a), and h ee addi ional species we e
desc ibed om he Russian Fa Eas , namely A. (E.) pue zi, A. (T.) ze chei, and A. (T.) nubis, which we e la e
eco ded om Japan (Ma uyama, 2002). Assing (1995) ega ded Homalo a a iolosa as a synonym o A. (E.)
ucicola. Recen ly, Ahn e al. (2000) and Pa k and Ahn (2004) published eco ds o Empleno a and T iocha a om
Sou h Ko ea.
In he p esen s udy, we cla i y he Japanese auna o Empleno a and T iocha a in he i s e isiona y
con ibu ion o he Japanese species o Aleocha a. We examined mo e han wo housands o specimens, which
we e collec ed om all o e Japan and adjacen egions, so ha he esul s p o ide us wi h mo e eliable aunis ic
in o ma ion on he subgene a. In iew o he ongoing des uc ion o he coas al and seasho e en i onmen s o
Japan, i is c ucial o comp ehensi ely in es iga e he auna in hese a eas. Iden i ica ion o Empleno a and
T iocha a is di icul e en o skilled axonomis s. Hence, we also es ablish an easie iden i ica ion me hod. The
species o Empleno a and T iocha a a e ex emely simila in gene al appea ance e en in he geni al s uc u es.
The e o e, an easy- o-use key allowing an accu a e iden i ica ion o he species is p o ided.
Ma e ials and me hods
Ma e ial om ins i u ions and p i a e collec ions
Mos specimens used we e supplied om ins i u ional o pe sonal collec ions. In addi ion, we collec ed o he
ma e ials. The deposi o ies o he ma e ial examined a e abb e ia ed as ollows (all collec ions, excluding he
Na u al His o y Museum, London, and he Field Museum o Na u al His o y, Chicago, a e in Japan):
Ins i u ions:
BMNH The Na u al His o y Museum, London (R. Boo h).
CBM The Na u al His o y Museum and Ins i u e, Chiba (A. Sai ô).
Zoo axa 3517 © 2012 Magnolia P ess · 3
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
EEEU Labo a o y o En i onmen al En omology, Ehime Uni e si y, Ma suyama (H. Yoshi omi).
FMNH The Field Museum o Na u al His o y, Chicago (M. K. Thaye ).
HUM The Hokkaido Uni e si y Museum, Sappo o (M. Ôha a).
KUM The Kyushu Uni e si y Museum, Fukuoka (M. Ma uyama).
SCM The Sagamiha a Ci y Museum, Kanagawa (H. Mo iya).
P i a e Collec ions:
cHayam Takeshi Hayama (Okinawa-ken).
cHayas Yasuhiko Hayashi (Hyôgo-ken).
cI ô Ta eo I ô (Kyô o- u).
cKaw Yasuko Kawakami (Ôsaka- u).
cMa Mune oshi Ma uyama (Fukuoka-ken).
cOno Hi oki Ono (Chiba-ken).
cWa Takashi Wa anabe (Kanagawa-ken).
cYam Shûhei Yamamo o (Fukuoka-ken).
Mos o he ype se ies including holo ypes a e deposi ed in he collec ion o KUM. Some pa a ypes o he new
species o be desc ibed in his pape and iden i ied specimens will be dis ibu ed o he collec ions abo e.
We did no examined he ype specimens o he species desc ibed by Weise (1877) because hey we e ecen ly
e ised by Assing (1995).
Collec ing me hods
The subgene a Empleno a and T iocha a inhabi nea ly he same en i onmen , he sup a idal zones o sandy o
pebbly beaches, and sal ma shes. Specimens we e usually hand-picked o aspi a ed om he si ed ma e ial o
decaying seaweed. Collec ed bee les we e ans e ed o a ube wi h a piece o co on pape and a small amoun o
e hyl ace a e. Some specimens we e p ese ed in a small ube illed wi h e hanol.
Dissec ing, d awing and edi ing images
The echnical p ocedu es used we e gene ally as desc ibed by Ma uyama (2006), wi h some di e ences. Fi s ,
he specimens we e so ened wi h a small amoun o wa e o de ach abdominal e minalia, hen soaked in
po assium chlo ide (KOH) solu ion, bo h conduc ed a oom empe a u e (abou 20–25°C). Second, he soaking
imes o each me hod di e ed (15–30 min o wa e so ening o he en i e body; 1 day in KOH o he emo ed
pa s; cleaning in 80% e hanol o 10 min; dehyd a ion in 99% e hanol o 10 min). Holo ypes we e dissec ed and
moun ed wi h Eupa al, ollowing Ma uyama (2004b).
A ligh mic oscope (Olympus BH-2, Olympus, Tôkyô, JAPAN, wi h a d awing de ice) was used o
obse a ion and d awing o body pa s. Also, a high magni ica ion s e eomic oscope (Nikon SMZ 1500, Nikon,
Tôkyô, Japan, wi h ligh de ice) was used o obse a ion o gene al appea ance.
Pic u es o specimens we e aken using a digi al came a (Canon EOS 7D, Canon, Tôkyô, JAPAN) wi h an
ex eme mac o lens (Canon MP-E 65 mm F2.8 1–5×, Canon) and a mac o lash (Mac o Twin Li e MT-24EX Flash,
Canon). Then, ocus s acking was conduc ed using he au omon age so wa e Combine ZM (Alan Hadley, UK,
h p://www.hadleyweb.pwp.blueyonde .co.uk/). All digi al images we e edi ed using Adobe Pho oshop Elemen s
6.0 (Adobe, San Jose, CA, USA).
Te minology and abb e ia ions
The e minology o he mo phology used in his s udy la gely ollows: Blackwelde (1936), Sawada (1972),
Klimaszewski (1984), Thaye (2005), and Ma uyama (2006). The e minology o he geni alia used by Welch
(1997) was adop ed. A ew male geni alia e ms we e adop ed om Klimaszewski (1984). The numbe o se ae and
po es in he desc ip ions we e con ined o one side o he body, excep o he mac o se ae and senso y se ae o
abdominal pa s and o he medial pseudopo es on he labium.
All measu emen s in he ex a e maximum leng h and gi en in millime es, as minimum leng h–maximum
leng h (mean ± SD).
The ollowing abb e ia ions a e used: AL, an enna leng h; BL, body leng h (app oxima e whole leng h); EL,
leng h o ely a om end o pos e io ma gin o p ono um o ely al hind ma gin; EW, ely al wid h; FBL, o e
body leng h, om apex o clypeus o he pos e io ma gin o ely a; HL, head leng h, om apex o pos clypeus o
YAMAMOTO & MARUYAMA
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he pos e io ma gin o head capsule, excep o memb anous an eclypeus and p eoccipu ; HTL, hind ibial leng h;
HW, head wid h; PL, p ono al leng h; and PW, p ono al wid h. Geni al pa s 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; sh, head o spe ma heca; sm, memb anous po ion o spe ma hecal duc ; sn, neck o
spe ma heca; and ss, scle o ised po ion o spe ma hecal s em (a e Welch, 1997, excep o (sc): we used (ss)
ins ead o (sc) o he same e m; see, Figs. 18, 22).
O he in o ma ion
The loca ions eco ded in his pape we e adop ed om he la es e sion o he Wo ld Geode ic Sys em (WGS
84). Some specimens ha e li le in o ma ion on he collec ion si e. In his case, bo h la i ude and longi ude we e
es ima ed. We ma ked he es ima ed locali ies wi h an as e isk, and oughly poin hem ou on maps o beaches
(Figs. 101–105). Some specimens wi h da a label o longi ude and la i ude da a we e con e ed in o decimal
deg ees.
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 old and his o ically
impo an ma e ial. The o iginal spellings on he label a ached o he specimen we e adop ed in such cases.
Japanese place names gene ally ollow he same spelling used in Japan excep o some English nouns. Wo ds
a e nea ly co esponded as ollows: -chôme (ci y block o i egula size); -chô, -machi ( own); -ku (wa d); -shi
(ci y); -gun (coun y); - o, - u, -ken (p e ec u e); -gawa, -kawa ( i e , s eam); -jima, -shima, -Ôshima, - ô (island); -
kaigan, -hama (seasho e beach); -sho ô (islands, a chipelago); -misaki, -zaki (cape); -numa (ma sh).
Collec ing me hods and condi ions o specimen labels a e abb e ia ed as ollows: FIT, ligh in e cep ion ap;
YPT, yellow pan ap. O he abb e ia ions: BRL, blue ound label pinned by a cu a o ; HW, hand w i en; PRL,
pu ple ound label pinned by a cu a o ; RRL, ed ound label pinned by a cu a o ; YRLS, yellow ound label by D .
Da id Sha p.
Taxonomy o he o e bee les
T ibe Aleocha ini Fleming, 1821
Aleocha idae Fleming, 1821: 49 ( ype genus: Aleocha a G a enho s , 1802)
Sub ibe Aleocha ina Fleming, 1821
Aleocha idae Fleming, 1821: 49 ( ype genus: Aleocha a G a enho s , 1802)
Piocha diae Fenyes, 1918: 20 ( ype genus: Piocha dia Heyden, 1870)
Diagnosis. The sub ibe is dis inguished by he ollowing cha ac e is ics: 5-5-5 a sal o mula (4-5-5 in Tino us
Sha p, 1833) (Hanley, 2002; Klimaszewski e al., 2002); maxilla y palpus 4-segmen ed, plus apical pseudosegmen
(looks 5-segmen ed); labial palpus 3-segmen ed, plus apical pseudosegmen (looks 4-segmen ed); ligula bilobed
apically, sho e han he segmen I o he labial palpus; la e al p ojec ions o labial apodeme elonga ed
(Ma uyama, 2004); median lobe o he aedeagus wi hou do sal apodeme (Ma uyama, 2004); a leas some species
ha e a e icula ed elum on he pa ame i e o he male aedeagus (See e s, 1978; Ashe & Maus, 1998).
Commen s. This sub ibe is composed o 21 gene a and is dis ibu ed h oughou all zoogeog aphic egions.
The sub ibe Aleocha ina is easily dis inguished om he o he wo sub ibes comp ised o e mi ophiles, namely
Compac opediina Kis ne , 1970, and Hodoxenina Kis ne , 1970, in he ibe Aleocha ini, by a a he gene alized
body shape (wi h some excep ions). Recen ly, howe e , a my mecoid genus o he sub ibe Aleocha ina,
My mecos ica Ma uyama, 2011, was desc ibed om Bo neo (Ma uyama e al., 2011).
The phylogene ic ela ionships wi hin he sub ibe emain unce ain. The phylogene ic concep o his sub ibe
is e y uns able due o unce ain limi s o he genus Aleocha a and ela ed axa (Ashe & Maus, 1998). The
monophyly o Aleocha ina is s ill no es ablished (Ashe & Maus, 1998), and no au apomo phy has been de ec ed
o i . Kanao e al. (2011) showed ha Compac opediina belongs o he Te as ic a K aa z, 1857 genus g oup o
Aleocha ina (sensu Ma uyama, 2004a) and sugges ed non-monophyly o Aleocha ina.
In Japan, 5 gene a, 6 subgene a (all subgene a belong o Aleocha a), and 27 species o his sub ibe ha e
hi he o been eco ded.
Zoo axa 3517 © 2012 Magnolia P ess · 5
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
Genus Aleocha a G a enho s , 1802
Aleocha a G a enho s , 1802: 67 (o iginal desc ip ion; ype species: S aphylinus cu ulus Goeze, 1777).
See, Klimaszewski (1984), Sme ana (2004) and Gouix & Klimaszewski (2007) o u he synonymic in o ma ion.
Redesc ip ion. Body: small o la ge, medium o la ge in many cases; o e all shape di e se, no mally mo e o
less b oad: compac and obus , mo e o less usi o m o cylind ical and a ely slende and na ow (Ashe & Maus,
1998). Colou : uni o mly eddish b own o black in mos cases, some imes ely a ligh e o wi h macula ions.
Head: head capsule somewha ci cula o mo e o less ans e se. An ennae: 11-segmen ed, scape somewha
cla a ed, no mally longe o as long as pedicel (Klimaszewski, 1984). Mou hpa s: lacinia wide, mul ispinose;
galea wide, as long as lacinia (Klimaszewski, 1984). Tho ax: meso en i e a iable, wi hou ca ina o wi h
comple e ca ina. Legs: no mally medium leng h; mesocoxae na owly sepa a ed, wi h long p ocess o
meso en i e. Abdomen: simple and no mally usi o m; e gi e o segmen s III–V imp essed ans e sely a base.
Geni alia: median lobe o aedeagus in male, usually wi h lagellum inside. Spe ma heca o emale, which a e
di e en ly modi ied in di e en subgene a (Klimaszewski & Jansen, 1993), bu wi h some excep ions. De ail
desc ip ions a e in Klimaszewski (1984).
Commen s. Adul s p eda e upon eggs, la ae, and pupa ia o lies; mos known la ae a e in e nal pa asi oids
o cyclo haphous lies (Peschke & Fuldne , 1977; See e s, 1978; Klimaszewski, 1984; Maus e al., 1998ab, 2001;
Yamazaki, 2008, 2012). Thus, Aleocha a species a e conside ed economically impo an as hos agen s. In con as
o he accumula ion o ecological knowledge o some pa icula species, he axonomy a he subgene ic le el and
he e olu ion and phylogeny o his g oup a e poo ly known.
S udies o Japanese species o Aleocha a a e also s ill incomple e and only 22 species in 6 subgene a ha e
been eco ded. Re isions o each subgenus o he Japanese species a e also equi ed.
Key o he li o al subgene a o he genus Aleocha a in Japan
1. Body hick, a he spindle shaped; head and p ono um co e ed wi h g anula mic os uc u es; p ono um somewha
uni o mly ele a ed; e gi e VIII s ongly ounded, bea ing wi h nume ous la ge o al p ojec ions on mid o pos e io
su ace.
(1) Do sal su ace ma and no shining a all. P ono um wides a ound base and mode a ely con ex abo e; clea ly smalle han
head; lacking pubescence along midline. Ely a wi h pos e io ma gins deeply no ched la e ally; su ace coa sely punc u ed,
densely co e ed wi h sho bu obus se ae. Meso en i e comple ely ca ina e. Do sal su ace o abdomen ugosy wi h nume -
ous pubescence. [Male]: pos e io ma gin o s e ni e VIII mode a ely poin ed. Median lobe wi hou subapico- en al p ojec-
ions; lagellum inside e y long. [Female]: basal po ion o spe ma heca coiling coun less imes . . . . . subgenus Cop ocha a
Mulsan & Rey, 1874 [Aleocha a (Cop ocha a) squali ho ax Sha p, 1888] (see, Assing, 1995 o he edesc ip ion, and Maus,
1998 o in o ma ion).
- Body slende o na owly elonga ed; p ono um almos la ened; meso en i e wi hou o wi h sho (incomple e)
ca ina; do sal su ace o abdomen shining; su ace o e gi e VIII simple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Head (Figs. 53, 57, 61) wi h wo longi udinal u ows along midline; p ono um (Figs. 54, 58, 62) wi h longi udinal u -
ows o h ee-dimensional pa e ns.
(2) Body (Figs. 53, 57, 61) shining o ma , o ebody spa sely co e ed by somewha hick and sho se ae. P ono um (Figs. 53–54,
57–58, 61–62) sligh ly la ge han head. Meso en i e (Figs. 55–56, 59–60, 63–64) wi hou ca ina. Do sal su ace o abdomen
(Figs. 53, 57, 61) shining. [Male]: e gi e VIII (Figs. 71, 79, 86) wi h mo e o less unca ed pos e io ma gin. Median lobe
(Figs. 75–76, 83–84, 90–91): a pai o poin ed o o al subapico- en al p ojec ions p esen s; lagellum sho , much sho e
han median lobe. [Female]: e gi e VIII (Figs. 72, 80, 87) simila o ha o male. Spe ma heca (Figs. 78, 85, 92) wi hou coil-
ing po ion and colla (see, example o Empleno a: Fig. 22) . . . . . . . . . . . . . . . . . . . . . . . subgenus T iocha a Be nhaue , 1901
- Head and p ono um lack o any sulcus.
An ennae (Figs. 1–5, 8) long and slende , much longe han p ono um wid h. Head (Fig. 6) impunc a e on medial line. Mes-
o en i e (Fig. 7) wi h a e y sho o mode a e leng h o ca ina ion. Te gi e VIII (e. g., Figs. 14–15) almos unca e, ounded
o weakly ema gina ed, wi h a ow o hick senso y se ae (e.g., Fig. 14: a ow) a pos e io ma gin. [Male]: pos e io ma gin o
s e ni e VIII (Figs. 16, 25, 32, 40, 48) mode a ely o e y s ongly poin ed, a ied g ea ly among species. Median lobe (Figs.
18–19, 27–28, 34–35, 43–44, 50–51): a pai o cu ed o s aigh subapico- en al p ojec ions p esen s in la e al iew; lagel-
lum long o ex emely long. [Female]: basal po ion o spe ma heca (Figs. 22, 29, 37, 45, 52) simple bu wi h colla (see, Fig.
22), lacking coiling pa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subgenus Empleno a Casey, 1884
Rema ks. The e is no eco d o he li o al subgenus Polys omo a Casey, 1906 in Japan and adjacen egions.
The e o e, i was excluded om he key abo e. See, Assing (1995) o de ail in o ma ion o Polys omo a.
YAMAMOTO & MARUYAMA
6 · Zoo axa 3517 © 2012
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FIGURES 1–7. Japanese species o Aleocha a (Empleno a) (Figs. 1–5, habi us). 1. Aleocha a (Empleno a) ucicola; 2. A. (E.)
pue zi; 3. A. (E.) hayamai; 4. A. (E.) yama o; 5. A. (E.) seg ega a; 6. do sal head o A. yama o; 7. meso en i e and me a en i e
o A. ucicola in en al iew.
Zoo axa 3517 © 2012 Magnolia P ess · 7
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
Subgenus Empleno a Casey, 1884
(Figs. 1–52, 93–97, 101–103)
Empleno a Casey, 1884: 17 (o iginal desc ip ion; ype species: Empleno a ma i ima Casey, 1884: 17, by o iginal
designa ion and mono ypy); 1906: 131 (as genus; key o gene a o sub ibe “Aleocha ae”), 172 (as genus; his o ical
e iew; diagonos ic key o species o A lan ic coas ); Eichelbaum, 1909: 245 (as subgenus; gene ic ca alogue o
wo ld S aphylinidae); Fenyes, 1920: 399 (as subgenus; key o wo ld subgenge a o Aleocha a), 1921: 415 (as
subgenus; diagnosis; ca alogue o wo ld species); Schee pel z, 1925: 447 (as subgenus; ca alogue o Palaea c ic
species o Aleocha inae); Be nhaue & Schee pel z, 1926: 795 (as subgenus; ca alogue o wo ld species o
Aleocha inae); To enham, 1949: 404 (as subgenus; gene ic ca alogue o B i ish S aphylinidae); Blackwelde ,
1952: 147 (as subgenus; gene ic ca alogue o wo ld S aphylinidae); Ha ch, 1957: 137 (as subgenus; key o
Aleocha a subgene a o Paci ic No hwes ), 140 ( edesc ip ion o species o Paci ic No hwes ); Liko sky, 1974:
294 (as subgenus; key o subgene a o middle Eu opa); Moo e & Legne , 1975: 328 (as subgenus; ca alogue o
Nea c ic species); See e s, 1978: 59 (as genus; key o Nea c ic gene a o Aleocha inae), 138 (diagnosis);
Klimaszewski, 1984: 9, 10 (phylogene ic ela ionships o Nea c ic Aleocha a), 95 (as subgenus; e ision o
Nea c ic Aleocha a; edesc ip ion o subgenus), 96 (key; edesc ip ions o Nea c ic species); Lohse, 1984: 148 (as
genus; diagnosis); Lohse, 1985: 328 (as genus; key o middle Eu opean species o gene a Empleno a and
Polys omo a); Lohse, 1989: 239 (as genus; key o gene a o middle Eu ope), 240 (as genus; ca alogue o middle
Eu opean species); Assing, 1995: 219 (as genus; diagnos ic key o gene a), 219 (no es on genus), 220 (key o
Palaea c ic species), 220 (desc ip ions o each species); Welch, 1997: 8 (his o ical e iew o B i ish Empleno a and
ela ed axa); Maus & Ashe, 1998a (online) (as subgenus; checklis o subgenus o wo ld; diagnosis o subgenus;
bionomics; discussion o phylogene ic ela ionships); Ashe, 2001: 360 (as subgenus; ca alogue o Nea c ic
species); Sme ana, 2004: 356 (as subgenus; ca alogue o Palaea c ic Aleocha inae); Dauphin, 2005: 47 (as genus;
key o gene a o Empleno a and Polys omo a); Gouix & Klimaszewski, 2007: 26 (as subgenus; ca alogue o
Aleocha ine species o Canada and Alaska).
Polys oma S ephens, 1833: 91 (o iginal desc ip ion; ype species: Aleocha a obscu ella G a enho s , 1806:
159; ixed by S ephens, 1833: 91, by mono ypy); S ephens, 1835: 430 (as genus; edesc ip ion o genus); Thomson,
1861: 47 (as genus; edesc ip ion o genus), 48 (key o Scandina ian species); Mulsan & Rey, 1874: 169 (as
genus; de ailed edesc ip ion o genus as Polys ome [misspelling]), 172 (key o species o F ance), 173
( edesc ip ions o species o F ance); Fowle , 1888: 21 (as subgenus; diagnosis; diagnos ic key o species o B i ish
Islands); Casey, 1894: 289 (as genus; as “Polis oma” [misspelling]: see, Casey, 1906: 272; la e ci ed by some
au ho s as synonym o Empleno a); Ganglbaue , 1895: 45 (as subgenus; edesc ip ions o middle Eu opean
species); Be nhaue , 1901a: 504 (as subgenus; edesc ip ions o Palaea c ic species); Johansen, 1914: 31 (as
subgenus; ca alogue and key o species o Denma k); Fenyes, 1921: 415 (as synonym o Empleno a); Schee pel z,
1925: 447 (as synonym o Empleno a); Be nhaue & Schee pel z, 1926: 795 (as synonym o Empleno a);
To enham, 1949: 404 (as synonym o Empleno a; no e); Blackwelde , 1952: 147, 318 (as synonym o Empleno a);
Palm, 1972: 443 (as subgenus; ca alogue; edesc ip ions o species o Sweden); Moo e & Legne , 1975: 328 (as
synonym o Empleno a); Klimaszewski, 1984: 95 (as synonym o Empleno a); Lohse, 1985: 328 (as synonym o
Empleno a); Sme ana, 2004: 356 (as synonym o Empleno a); Gouix & Klimaszewski, 2007: 26 (as synonym o
Empleno a).
Polys omo a Casey, 1906: 136 (o iginal desc ip ion; ype species: Aleocha a g isea K aa z, 1856: 96; ixed by
Casey, 1906: 136, by o iginal designa ion and mono ypy); Eichelbaum, 1909: 245 (as genus; gene ic ca alogue o
wo ld S aphylinidae); Fenyes, 1921: 415 (as synonym o Empleno a); Schee pel z, 1925: 447 (as synonym o
Empleno a); Be nhaue & Schee pel z, 1926: 796 (as synonym o Empleno a); Blackwelde , 1952: 147, 319 (as
synonym o Empleno a); Moo e & Legne , 1975: 328 (as synonym o Empleno a); See e s, 1978: 138 (as synonym
o Empleno a); Lohse, 1985: 328 (as synonym o Empleno a); Lohse, 1989: 239 (as genus; key o gene a and
species o middle Eu ope); Assing, 1995: 219 (key o Asian coas al gene a), 226 (as genus; key o Palaea c ic
species), 229 (no es on species); Welch, 1997: 9 (his o ical commen ); Maus & Ashe, 1998b (online) (wo ld
checklis o subgenus; diagnosis; bionomics; phylogene ic ela ionships); Sme ana, 2004: 357 (as subgenus;
ca alogue o Palaea c ic species); Dauphin, 2005: 47 (as genus; key o gene a, Empleno a and Polys omo a); F ank
& Ahn, 2011: 20 (as subgenus; checklis o coas al S aphylinidae o wo ld).
Polycha ina Rei e , 1909: 28 (o iginal desc ip ion; ype species: Aleocha a g isea K aa z, 1856: 96; ixed by
Rei e , 1909: 28, by mono ypy); Fenyes, 1921: 415 (as synonym o Empleno a); Schee pel z, 1925: 447 (as
synonym o Empleno a); Be nhaue & Schee pel z, 1926: 796 (as synonym o Empleno a); Blackwelde , 1952:
YAMAMOTO & MARUYAMA
8 · Zoo axa 3517 © 2012
Magnolia P ess
147, 317 (as synonym o Empleno a); Moo e & Legne , 1975: 328 (as synonym o Empleno a); Lohse, 1985: 328
(as synonym o Empleno a).
FIGURES 8–13. Body pa s o Aleocha a (Empleno a) ucicola o male. 8. igh an enna; 9. labium; 10. maxilla; 11. men um;
12. lab um; 13. igh o eleg in en al iew.
Zoo axa 3517 © 2012 Magnolia P ess · 9
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
Polys oma ia Rei e , 1909: 28 (o iginal desc ip ion; ype species: Aleocha a obscu ella G a enho s , 1806:
159; ixed by Rei e , 1909: 28); Schee pel z, 1925: 447 (as synonym o Empleno a); Fenyes, 1921: 415 (as
synonym o Empleno a); Be nhaue & Schee pel z, 1926: 796 (as synonym o Empleno a); Blackwelde , 1952:
147, 319 (as synonym o Empleno a); Moo e & Legne , 1975: 328 (as synonym o Empleno a); Klimaszewski,
1984: 95 (as synonym o Empleno a); Lohse, 1984: 148 (as genus; diagnosis, as Polys oma ia Casey); Lohse,
1985: 328 (as synonym o Empleno a); Sme ana, 2004: 356 (as synonym o Empleno a); Gouix & Klimaszewski,
2007: 26 (as synonym o Empleno a).
Redesc ip ion. Body (Figs. 1–5): mode a ely la ened, slende and subpa allel sided; whole leng h a ying
om 2.5 o 7.0 mm, no mally a ound 3.5–4.5 mm; do sal su ace co e ed wi h minu e and dense hexagonal
mic os uc u es; su ace o head, p ono um and ely a co e ed wi h somewha dense, hin hai s. Do sal su ace o
abdomen shining, bu head, p ono um and ely a ma (uni o mly hexagonal- e icula ed). Colou (Figs. 1–5):
uni o mly blackish b own o black some imes wi h ligh e colou in ely a. Head: almos ci cula o somewha o al
(HW/HL
≈
1.14), mode a ely con ex abo e, wides a ound middle; do sal su ace gen ly ele a ed medially, bu
some imes wi h unique ele a ion pa ially; impunc a e on medial line (e.g., Fig. 6). Eyes small, no s ongly
p o uding la e ally. An ennae (Figs. 1–5, 8): mo e o less ili o m, en i ely obus and long, as same as combined
leng h o head and p ono um; segmen I s ou and apically dila ed; and segmen XI clea y long. Mou h pa s:
mandibles asymme ic, le one wi h one oo h nea apex. Clypeus ounded apically. Lab um (e. g., Fig. 12) abou
1.7 imes as wide as long; an e io ma gin mode a ely ema gina ed medially; basal hal semi- anspa en . Labial
palpus (Fig. 9) wi h segmen I clea ly longe han II; segmen II diala ed. Men um (Fig. 11) much wide han long,
nea ly apezoidal; an e io ma gin s ongly ema gina ed; nume ous pseudopo es sca e ed andomly. Maxilla y
palpus (Fig. 10) dis inc ly segmen ed; segmen I long and hick; segmen II sho e and na owe han I; segmen III
sligh ly sho e and na owe han II; pseudosegmen much sho e han o he segmen s; lacinia wi h nume ous
hai s and wi h abou 11 hick spines pec ina ely. Tho ax: p ono um o al and somewha ans e se (PW/PL
≈
1.24),
longe han head leng h (PL/HL
≈
1.25), somewha b oade han head (PW/HW
≈
1.36), wides a ound middle,
mode a ely cons ic ed nea base; ou e ma gin a ound middle wi h a pai o ela i ely long blackish mac ose ae;
su ace la and e enly pubescen bu some imes wi h deep and dis inc punc u es on do sal su ace nume ously.
Hypome a ully isible in la e al iew. In e coxal ca i ies o meso en i e (e.g., Fig. 7) na owly o mode a ely
sepa a ed. Meso en i e wi h sho ca ina (Fig. 7: a ow), o no ca ina e. Ely a: widened owa d pos e io ma gin,
much b oade han long (EW/EL
≈
1.56), and somewha wide han p ono um (EW/PW
≈
1.28); pos e io ma gin o
each ely a nea ly unca e bu gen ly ounded owa d pos e io ma gins; su ace somewha ugosely punc u ed
bo h hexagonal mic osculp u es and p ominen dis inc punc u es in some species; en i e su ace densely co e ed
wi h b own se ae; an e io ma gin wi h one ela i ely long blackish b is le. Legs (e. g., Fig. 13): sho and slende
wi h sho hind a si; nume ous spines and b is les on do sal su ace o each ibia, especially on o e ibia (Fig. 13);
mid ibia as long as me a en i e; each a sal segmen almos same wid h (segmen V excluding claws); hind a si
sho (hind a sal leng h/mid a sal leng h
≈
1.31). Hind wings: en i e; eins weakly scle o ized and e y obscu e;
pos e io ma gin wi h a ow o minu e whi e hai s. Abdomen: elonga ed and sligh ly na owed owa d pos e io
segmen s, na owing ab up ly a ound apex, wides a ound segmen III–IV; a leas e gi e III–VI (Figs. 1–5)
ans e sely imp essed a base. Su ace o e gi e VIII and s e ni e VIII simple and smoo h. Pos e io ma gin o
e gi e VIII (Fig. 14–15, 23–24, 30–31, 38–39, 46–47), bea ing hick and sho se e al senso y se ae (see, Fig. 14:
a ow).
[Male]: pos e io ma gin o s e ni e VIII (Figs. 16, 25, 32, 40, 48) no mally poin ed, and i s shape a ies
among species. Median lobe (Figs. 18–19, 27–28, 34–35, 43–44, 50–51) elonga ed, wides a ound base. Median
lobe wi h a pai o subapico- en al p ojec ions (see, Fig. 18) on median lobe; p ominen inne sac wi h long
p ojec ing lagellum (e. g., Figs. 18–19, 43–44) a leas almos as long as median lobe o aedeagus. Pa ame i e
longe han median lobe o aedeagus; apical lobe o pa ame i e (Fig. 21) sho and poin ed, bea ing wi h ou se ae.
[Female]: pos e io ma gin o e gi e VIII, almos unca e, ounded o sligh ly ema gina ed; su ace o e gi e
VIII (Figs. 15, 24, 31, 39, 47) simila o ha o male. Pos e io ma gin o s e ni e VIII (Figs. 17, 26, 33, 41, 49)
sligh ly o mode a ely poin ed. Spe ma heca (Figs. 22, 29, 37, 45, 52; see, Fig. 22) wi h apical in agina ion o
spe ma heca (ai) sha ow; sphe ical head o spe ma heca (sh), as long as o longe han apical po ion o
spe ma hecal s em (sa); neck o spe ma heca (sn) and (sa) o ming mo e o less in igh angle; basal po ion o
spe ma hecal s em (sb), na owing owa d base; dis inc colla loca ed be ween (sa) and (sb); scle o ized po ion o
spe ma hecal s em (ss) sho and e ec in mos cases; memb aneous po ion o spe ma hecal duc (sm) mode a e o
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16 · Zoo axa 3517 © 2012
Magnolia P ess
pai o subapico- en al p ojec ions gen ly cu ed in la e al iew (Fig. 18); basal swelling o aedeagus (bs),
ci cula ly ounded in la e al iew (Fig. 18); apical lobe o median lobe clea ly isosceles shape in en al aspec
(Figs. 19–20); lagellum sligh ly sho e han he whole leng h o median lobe (Figs. 18–19).
[Female]: pos e io ma gin o e gi e VIII (Fig. 15) ounded, wi h a ound 7 mac ose ae. S e ni e VIII (Fig.
17) wi h 3 la ge mac ose ae and a ound 4 hinne mac ose ae; pos e io ma gin sligh ly poin ed. Spe ma heca
(Fig. 22): spe ma hecal head mo e o less as long as apical po ion o spe ma hecal s em; basal po ion o
spe ma hecal s em (sb) sligh ly na owing owa d base, lacking any ben a ound middle and usually a base;
scle o ized po ion o spe ma hecal s em e ec , connec ed wi h (sb) almos ho izon ally; each pa o
spe ma heca excep o memb aneous po ion o spe ma hecal duc (sm) en i ely and mode a ely scle o ized;
(sm) mode a e in leng h.
Measu emen s (male: n=10): BL, 3.58–4.59 (4.02±0.32); FBL, 1.16–2.03 (1.83±0.16); HL, 0.43–0.60
(0.51±0.06); HW, 0.46–0.66 (0.59±0.06); AL, 0.79–1.27 (1.08±0.15); PL, 0.48–0.73 (0.64±0.07); PW, 0.50–0.90
(0.77±0.12); EL, 0.54–0.83 (0.67±0.09); EW, 0.77–1.21 (1.02±0.13); HTL, 0.45–0.73 (0.61±0.08).
Measu emen s ( emale: n=10): BL, 3.36–5.18 (4.23±0.55); FBL, 1.63–2.12 (1.90±0.16); HL, 0.47–0.57
(0.53±0.03); HW, 0.49–0.65 (0.60±0.05); AL, 0.87–1.23 (1.08±0.12); PL, 0.56–0.73 (0.66±0.05); PW, 0.66–1.04
(0.83±0.11); EL, 0.58–0.80 (0.69±0.08); EW, 0.86–1.18 (1.06±0.11); HTL, 0.48–0.71 (0.61±0.08).
Diagnosis. This species can be dis inguished om he o he species o Empleno a by a combina ion o he
ollowing cha ac e s a es: no mally medium body sized; do sal su ace o o ebody wi h nume ous dis inc
punc u es (Fig. 1); impunc u ed a ea o do sal su ace o head sca cely swollen (almos la ened); meso en i e
wi h sho ca ina abou 0.41 imes as long as meso en i e (Fig. 7). [Male]: s e ni e VIII (Fig. 16) somewha
weakly poin ed owa d apex; subapico- en al p ojec ions on median lobe gen ly cu ed in la e al iew (Fig. 18);
basal swelling o median lobe (bs), ci cula ly ounded in la e al iew (Fig. 18); median lobe elonga ed py i o m
(Fig. 19) and apical lobe isosceles shape in en al aspec (Fig. 20). [Female]: basal po ion o spe ma hecal s em
(sb) (Fig. 22) simple, and gen ly o mode a ely cu ed wi hou ben a middle and base, wi h scle o ized and e ec
scle o ized po ion o spe ma hecal s em (ss).
Con i med dis ibu ion by p esen s udy. [JAPAN]: Honshû, Shikoku, Kyûshû, Hachijô-jima, Awaji-shima,
Tsu-shima, Amakusa-sho ô, Kamikoshiki-jima. See, Fig. 101 o Japanese dis ibu ion.
O he locali ies in li e a u e. [SOUTH KOREA]: Chungnam P o ince, Gyeongnam P o ince, Jeonnam
P o ince, Jeju P o ince (Pa k & Ahn, 2004); [NORTH KOREA]: Hamgyong P o ince (Pa
ś
nik, 2001); [CHINA]:
Heilongjiang (Li & Chen, 1990, 1993: bo h doub ul eco ds), Anhui (Li & Wang, 1993), Hong Kong (de
Rougemon , 2001); [RUSSIA]: Fa Eas (Ahn e al., 2000: no o iginal ci a ion; locali y doub ul); [FRANCE]:
Région Aqui aine (Dauphin, 2005).
Rema ks. This species was o iginally desc ibed “unde seaweed a Amakusa and Iwosima, nea Nagasaki”,
(Kyûshû, Japan: Sha p, 1874) and la e edesc ibed by Sawada (1971), Assing (1995), and Pa k and Ahn (2004).
Sawada (1971) emphasised he chae o axy o mou h pa s, while Assing (1995) paid special a en ion o he se al
condi ion o he do sal su ace o p ono um. Pa k and Ahn (2004) p o ided a sho diagnosis wi h igu es.
Ahn e al. (2000) sho ly edesc ibed Aleocha a ucicola, bu Pa k and Ahn (2004) ega ded i as a
misiden i ica ion o A. pue zi (Assing, 1995) and edesc ibed i again. Howe e , acco ding o hei illus a ions o
he aedeagus, i is pa o a new species being desc ibed in he p esen pape , A. seg ega a, al hough he igu e o
he spe ma heca in Ahn e al. (2000) seems o belong o A. ucicola.
The dis ibu ion o A. ucicola is e y wide. Al hough eco ds ha e been es ic ed in Eas Asia o a long ime,
Dauphin (2005) eco ded he species om no h-wes e n F ance (Région Aqui aine, Gi onde). This amazing eco d
seems accu a e, as D . V. Assing iden i ied he specimens. Dauphin (2005) also men ioned ha he eco ds migh
ha e been unin en ional in asions om o he egions, and special a en ion needs o be paid o his axon wi h
ega d o i s dis ibu ion.
The eco ds om mainland China adop ed in Li and Chen (1990, 1993) a e highly doub ul, because his
species’ eco d locali y (Heilongjiang) is inland.
Assing (1995) synonymised Homalo a a iolosa Weise, 1877 wi h A. ucicola, and designa ed a lec o ype o
he ype se ies o each species.
Yamazaki (2008) e ealed one o he hos species o A. ucicola as Fucellia apicalis Ke ész, 1908 (Dip e a,
An homyiidae). Yamazaki (2012) epo ed ha he kelp ly, Coelopa igida (Fab icius, 1805) (Dip e a,
Coelopidae) was an addi ional ly hos . The a icle also no ed he pa asi ized mon hs (Ap il, No embe , Decembe )
by A. ucicola.
Zoo axa 3517 © 2012 Magnolia P ess · 17
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
FIGURES 38–45. Te minalia o Aleocha a (Empleno a) hayamai. 38. e gi e VIII o male; 39. e gi e VIII o emale; 40.
s e ni e VIII o male; 41. s e ni e VIII o emale; 42. igh an enna o male; 43. male geni alia: median lobe o aedeagus in
la e al iew; 44. male geni alia: median lobe o aedeagus in en al iew; 45. emale geni alia: spe ma heca.
YAMAMOTO & MARUYAMA
18 · Zoo axa 3517 © 2012
Magnolia P ess
Aleocha a (Empleno a) pue zi (Assing, 1995)
(Figs. 2, 23–29, 95–96, 103)
Empleno a pue zi Assing, 1995: 220 (diagnos ic key o species o Palaea c ic Empleno a), 225 (o iginal desc ip ion; ype
locali y: “RUSSIA, Sakhalin, Ko sako dis ., Ismenshy oye lake”); Naomi e al., 2000: 107 ( eco d om Pa amushi Is.,
Ku il Islands); Pa
ś
nik, 2001: 232 ( eco d om No h Ko ea).
Aleocha a (Empleno a) pue zi (Assing, 1995); Maus & Ashe, 1998a (online) (checklis o subgenus o wo ld; phylogene ic
ela ionship); Ma uyama, 2002: 18 ( eco d om Japan (Hokkaidô)); Sme ana, 2004: 356 (ca alogue o Palaea c ic
Aleocha inae); F ank & Ahn, 2011: 10 ( igu e o li ing indi idual; misiden i ica ion o A. seg ega a?), 20 (checklis o
coas al S aphylinidae o wo ld).
Type specimens. No examined.
Non- ype specimens. JAPAN: [Hokkaidô]: 1
♂
, 1
♀
, Goyômai, Nemu o-shi (43.363N, 145.795E), 14 VI
1999, Ôha a-M. (43°21
′
47
″
N, 145°47
′
43
″
E; unde seaweed; HUM); 1
♂
, 3
♀
, Makkayô-misaki, Habomai,
Nemu o-shi (43.339N, 145.747E), 14 VI 1999, Ôha a-M. (43°20
′
19
″
N, 145°44
′
48
″
E; unde seaweed (Lamina ia
sp., Zos e a sp.); HUM); 2
♂
, 4
♀
, Hamama su, Nemu o-shi (43.206N, 145.529E), 26 IX 2000, Ma uyama-M.
(cMa ); 1
♂
, Ochiishi, Nemu o-shi, (43.198N, 145.530E*), 24 VIII 1999, Ma uyama-M. (unde seaweed on sandy
beach; cMa ); 1
♂
, 2
♀
, Ochiishihigashi, Nemu o-shi (43.189N, 145.511E), 10 VI 2010 (21:00), Yamamo o-S.
( om seaweed (Zos e a sp.) on sandy beach; cYam); 1
♂
, Ochiishinishi, Nemu o-shi, (43.171N, 145.503E*), 4 V
1995, Naka ani-M. (KUM); 1
♂
, 1
♀
, Mochi ippu-numa, Hamanaka-chô, Akkeshi-gun (43.022N, 145.019E), 11
VI 2010, Yamamo o-S. (collec ed om seaweed (Lamina ia sp., Sa gassum sp.) on sandy ma sh nea coas ;
cYam); 1
♀
, Shunkuni ai, Numu o-shi (43.281N, 145.425E), 24 VIII 2001, Ma uyama-M. (cMa ); 6
♂
, 1
♀
,
Todowa a (No suke-zaki peninsulae), No suke, Be sukai-chô, No suke-gun (43.597N, 145.301E*), 6 VII 1986,
Nomu a-S. (KUM); 1
♂
, 1
♀
, Moko o, Abashi i-shi (43.970N, 144.322E), 2 VII 1994, Haga-K. (unde seaweed;
KUM); 2
♂
, 1
♀
, Fu a su-iwa, Abashi i-shi (44.049N, 144.258E*), 6 VI 1989, Ka ô-T. (KUM); 1
♂
, Kimuaneppu
cape, Hamasa oma, Sa oma-chô, Toko o-gun (44.108N, 143.912E), 10 VII 1986, Nomu a-S. (KUM); 1
♀
,
Komuke-gensei-kaen pa k, Komukai, Mombe su-shi (44.296N, 143.441E), 23 VII 2009, Ôha a-M. (44°17
′
47
″
N,
143°26
′
26
″
E; unde seaweed; HUM: HK-09-MO-054/SA); 1
♀
, Chikube su, Habo o-chô, Tomamae-gun
(44.419N, 141.739E), 22 VII 2009, Ôha a-M. (44°25
′
09
″
N, 141°44
′
22
″
E; unde seaweed (Lamina ia sp.); HUM:
HK-09-MO-046/SA); 3
♂
, Maehama (Teu i- ô), Habo o-chô, Tomamae-gun (44.432N, 141.336E), 14 VI 2008,
Ôha a-M. (44°25
′
55
″
N, 141°20
′
08
″
E; unde seaweed; HUM: TE-08-MO-006); 1
♂
, 1
♀
, Aikage (Teu i- ô),
Habo o-chô, Tomamae-gun (44.419N, 141.321E), 15 VI 2008, Ôha a-M. (44°25
′
07
″
N, 141°19
′
16
″
E; unde
seaweed; HUM: TE-08-MO-007); 1
♂
, 1
♀
, Ayo o cape, Kojôhama, Shi aoi-chô, Shi aoi-gun (42.453N,
141.206E), 27 VIII 2009, Ôha a-M. (N42°27
′
11
″
, E141°12
′
21
″
; unde seaweed (Sa gassum sp., Lamina ia sp.), eel
g ass (Zos e a sp.); HUM: HK-09-MO-092/RO, SA); 1
♀
, Yamasedoma i (Okushi i- ô), Okushi i-chô, Okushi i-
gun (42.201N, 139.540E), 12 VII 2008, Ôha a-M. (42°12
′
03
″
N, 139°32
′
24
″
E; unde seaweed; HUM: OK-08-MO-
018); 1
♀
, Okushi i (Okushi i- ô), Okushi i-gun (42.170N, 139.517E), 12 VII 2008, Ôha a-M. (42°10
′
12
″
N,
139°31
′
00
″
E; unde seaweed; HUM: OK-08-MO-014); 1
♂
, Tomisa o (Okushi i- ô), Okushi i-chô, Okushi i-gun
(42.072N, 139.471E), 12 VII 2008, Ôha a-M. (42°04
′
18
″
N, 139°28
′
17
″
E; unde seaweed; HUM: OK-08-MO-
015); 3
♀
, Hakoda e-shi (41.760N, 140.693E*), 13 V 1971, Hi ano-Y. (KUM); 1
♀
, Esan-misaki, Esanmisaki-chô,
Hakoda e-shi (41.812N, 141.183E), 15 VII 2009, Ôha a-M. (41°48
′
42
″
N, 141°10
′
59
″
E; unde seaweed (Lamina ia
sp.); HUM: HK-09-MO-042/CO); 1
♀
, Tachimachi-misaki, Sumiyoshi-chô, Hakoda e-shi (41.745N, 140.721E*),
15 IX 2008, Nishikawa-M (KUM); 3
♀
, Shi akami-misaki, Shi akami, Ma sumae-chô, Ma sumae-gun (41.398N,
140.199E*), 12 VII 2009, Ôha a-M. (42°23
′
52
″
N, 140°11
′
56
″
E: mis yping?; unde seaweed (Sa gassum sp.);
HUM: HK-09-MO-033/SI).
O he specimens. RUSSIA: Ku il Islands: [Pa amushi Is.]: 5
♂
, 5
♀
, Med ezhiy Wa e all, Shelekho o
(50.367N–50.378N, 155.611E–155.656E), 18 VII 1997, Sai ô-A. (50°22.012
′
N–50°22.694
′
N,
155°36.677
′
E–155°39.380
′
E; al . 0–10m; CBM: CBM-ZI 81521(-81530)); 1
♀
, B ynkhano o bay, sou h end o he
island (50.021N, 155.405E), 1 VIII 1996, Ôha a-M. (50°01
′
17
″
N, 155°23
′
79
″
E; by hand pick up; unde seaweed;
IKIP; HUM). [U up Is.]: 1
♂
, 1
♀
, Vese aya i e nea mou h o Na ally bay (46.094N, 150.142E), 6 VIII 1995,
Ôha a-M. (46°05
′
38
″
N, 150°08
′
33
″
E; by hand pick up; IKIP; HUM: UR-95-MO-006); 1
♀
, same da a, bu 7 VIII
1995, (IKIP: wi hou collec ion numbe ); 1
♀
, en i ons o Vs echnyi i e , inland coas al ma gin o Negodnaya
Bay (45.951N, 150.181E), 29 VIII 1995 (14:30–16:30), Ôha a-M. (45°56
′
63
″
N, 150°10
′
52
″
E; by hand wi h
Zoo axa 3517 © 2012 Magnolia P ess · 19
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
aspi a o ; al . 1m; unde seaweed along sandy coas line; IKIP; HUM: UR-95-MO-070); 1
♀
, O k y yybay
(45.864N, 149.793E), 4 VIII 1995, Ôha a-M. (45°51
′
49
″
N, 149°46
′
95
″
E; by hand pick up; unde logs and ocks on
sho e; IKIP; HUM: UR-95-MO-002); 1
♀
, same da a, bu (45.851N, 149.770E), 5 VIII 1995, (N45°51
′
04
″
,
E149°46
′
12
″
; unde seaweed along seasho e; IKIP; HUM: UR-95-MO-005).
Redesc ip ion. Body (Fig. 2): small o ela i ely la ge sized, no mally medium sized; somewha obus ,
na owly elonga ed; su ace o p ono um and ely a co e ed wi h nume ous sho hai s a he densely; dis inc bu
inconspicuous punc u es on do sal su ace o head and p ono um, and shallow dis inc punc u es on ely a. Colou
(Fig. 2): gland colou blackish b own o da k g ay; legs, especially a sal segmen s, b own o eddish b own:
maxilla y and labial palpi blackish b own o yellowish b own; an ennae blackish b own and pa ly eddish b own.
Head: op o do sal su ace la ened, wi hou unique ele a ion; su ace spa sely co e ed wi h sho se ae.
An ennae: hick and obus ; sligh ly sho e han combined leng h o head and p ono um; segmen I, nea ly 2.6
imes as long as b oad; segmen II clea ly sho e han I; segmen III as same leng h as II; segmen s IV and V
somewha sphe ical, as long as wid h; segmen s VI o X clea ly ans e se; segmen XI hick and subconical wi h
ounded apical ma gin, 1.2 imes as long as b oad; ela i e leng h (wid h) o segmen s om basal o apical: 9(3.5):
5(3): 5(3): 3(3): 3(3.5): 2.5(4): 3(5): 3 (5): 3(5): 3(5): 6(5). Tho ax: p ono um ela i ely wide han long (PW/PL
=1.26), a li le b oade han head (PW/HW =1.36); su ace wi h minu e hexagonal e icula ions and inconspicuous
dis inc punc u es. Me a en i e, abou 1.6 imes as long as meso en i e. In e coxal p ocess o meso en i e
sha ply poin ed, bu wi h ounded apex, and ha ing sho ca ina along midline, abou 0.4 imes as long as
meso en i e. In e coxal p ocess o me a en i e b oad and sho , one hi d as long as mesocoxal ca i y. Legs:
hind ibia sho , 0.9 imes as long as ely a; ela i e leng hs o a some es om basal o apical: 5: 4.5: 4.5: 4.5: 10.5
in o e a sus, 7.5: 6: 5: 5: 13.5 in mid a sus, 9: 7: 7: 7.5: 16 in hind a sus.
[Male]: pos e io ma gin o e gi e VIII (Fig. 23) nea ly unca e, wi h a ound 8 mac ose ae. S e ni e VIII (Fig.
25) wi h abou 7 mac ose ae and a ound 12 hin mac ose ae; pos e io ma gin poin ed somewha s ongly. Median
lobe o aedeagus (Figs. 27–28) elonga ed and na owed owa d apex, elonga ed py i o m in en al iew (Fig. 28);
a pai o subapico- en al p ojec ions almos s aigh and somewha obus in la e al iew (Fig. 27); basal swelling
o aedeagus (bs), poin ed sha ply owa d o amen mediale in la e al iew (Fig. 27); apical lobe clea ly isosceles
shape in en al iew (Fig. 28); lagellum much sho e han he whole leng h o median lobe (Figs. 27–28); apical
pa o copula o y piece scle o ized aiangula ly in la e al iew (Fig. 27).
[Female]: e gi e VIII (Fig. 24) mode a ely ounded owa d pos e io , wi h a ound 7 mac ose ae. S e ni e VIII
(Fig. 26) wi h 8 mac ose ae, bu hickness o mac ose ae a ying wi hin species; pos e io ma gin sligh ly poin ed
bu much weake han male. Spe ma heca (Fig. 29): spe ma hecal head longe han spe ma hecal neck (sn); basal
po ion o spe ma hecal s em na owing owa ds base, wi h ben a ound middle, some imes a base; scle o ized
po ion o spe ma hecal s em e ec ; each pa o spe ma heca excep o memb aneous po ion o spe ma hecal duc
(sm) en i ely and mode a ely scle o ized; (sm) mode a e in leng h.
Measu emen s (male: n=10): BL, 3.29–4.65 (3.77±0.44); FBL, 1.76–2.28 (2.02±0.17); HL, 0.52–0.74
(0.64±0.07); HW, 0.61–0.79 (0.69±0.06); AL, 0.97–1.33 (1.17±0.10); PL, 0.66–0.85 (0.74±0.07); PW, 0.82–1.05
(0.92±0.08); EL, 0.66–0.89 (0.75±0.09); EW, 0.92–1.35 (1.15±0.12); HTL, 0.48–0.78 (0.67±0.09).
Measu emen s ( emale: n=10): BL, 3.20–4.73 (4.01±0.51); FBL, 1.81–2.43 (2.10±0.18); HL, 0.50–0.77
(0.62±0.08); HW, 0.59–0.81 (0.69±0.07); AL, 0.99–1.18 (1.08±0.07); PL, 0.63–0.84 (0.74±0.08); PW, 0.84–1.07
(0.95±0.08); EL, 0.58–0.82 (0.72±0.07); EW, 0.86–1.37 (1.18±0.16); HTL, 0.52–0.79 (0.64±0.07).
Diagnosis. This species is simila o he o he Japanese Empleno a species, bu can be dis inguished om hem
by he ollowing poin s: a ying om small o la ge size (no ex emely la ge); dis ibu ion (see, Fig. 103)
es ic ed o no he n egion (Hokkaidô, Japan; Russian Fa Eas ; No h Ko ea); punc u es on do sal su ace o
head, p ono um and ely a inconspicuous (Fig. 2); impunc u ed a ea o head la ened; an ennae hick;
me a en i e, abou 1.62 imes as long as meso en i e o which wi h sho ca ina, abou 0.37 imes as long as
meso en i e. [Male]: s e ni e VIII (Fig. 25) somewha s ongly poin ed owa d pos e io ma gin; apex o scle i e
inside he median lobe, iangula ly scle o ized in la e al iew (Fig. 27); subapico- en al p ojec ions on median
lobe clea ly s aigh in la e al iew (Fig. 27); basal swelling o median lobe, sha ply poin ed owa d en al ma gin
in la e al iew (Fig. 27). [Female]: basal po ion o spe ma hecal s em wi h ben a ound middle pa (Fig. 29), and
wi h e ec scle o ized po ion o spe ma hecal s em.
Con i med dis ibu ion by au ho s. [JAPAN]: Hokkaidô, Teu i- ô, Okushi i- ô (See, Fig. 103); [RUSSIA]:
Ku il Islands: Pa amushi Is., U up Is.
YAMAMOTO & MARUYAMA
20 · Zoo axa 3517 © 2012
Magnolia P ess
O he locali ies in li e a u e. [RUSSIA]: Fa Eas : Sakhalin, Kamcha ka Peninsula, P imo sky K ai (Assing,
1995); [NORTH KOREA]: Chagang P o ince (Pa
ś
nik, 2001).
Rema ks. Aleocha a pue zi was o iginally desc ibed om Russian e i o y by Assing (1995). The esul s o
he Biological Expedi ion o he Na u al His o y Museum and Ins i u e, Chiba, Japan, o he No h Ku il Islands
p o ided us wi h he dis ibu ional in o ma ion o his species, and o he eco ds o he species come om
Pa amushi Island (o he Ku il Islands) (Naomi e al., 2000). He e, we p esen he i s eco d om U up Island
om his same island g oup. These specimens we e collec ed in Russia unde he su ey o he In e na ional Ku il
Island P ojec (IKIP), which is an in e na ional collabo a ion o Ame ican, Russian, and Japanese scien is s o
su ey he auna and lo a o he Ku il A chipelago (Takahashi & Ôha a, 2004).
Pa
ś
nik (2001) epo ed his species om No h Ko ea, and la e Ma uyama (2002) eco ded i o he i s ime
om Japan (no heas e n pa o Hokkaidô). Pa k and Ahn (2004) eco ded A. pue zi om Sou h Ko ea and also
co ec ed he misiden i ica ion o his species epo ed as “A. ucicola” in he p e ious pape (Ahn e al., 2000).
Howe e , he iden i ica ion o he species in bo h pape s is inco ec , and we conside i a new species (A.
seg ega a) acco ding o hei illus a ions.
Aleocha a (Empleno a) seg ega a Yamamo o & Ma uyama n. sp.
(Figs. 5, 30–37, 93–94, 97, 102)
“Aleocha a (Empleno a) ucicola Sha p, 1874”; Ahn e al., 2000: 243 (diagnosis wi h igu es; la e ega ded as
misiden i ica ion o “A. pue zi” by Pa k & Ahn, 2004, bu ac ually misiden i ica ion o A. seg ega a; e-examina ion o
specimens is needed).
“Aleocha a (Empleno a) pue zi Assing, 1995”; Pa k & Ahn, 2004: 195 (key o li o al Aleocha a species o Ko ea), 196
( edesc ip ion, bu misiden i ica ion o A. seg ega a; e-examina ion o specimens is needed).
Type se ies. Holo ype,
♂
, JAPAN: Ikinoma suba a-1chôme, Nishi-ku, Fukuoka-shi, Fukuoka-ken, Kyûshû
(33.580N, 130.293E), 28 VI 2009, Yamamo o-S. (collec ed by an aspi a o ; om d a ed seaweed on ine sandy
beach in day ime; KUM).
Pa a ypes, JAPAN: [Hokkaidô]: 1
♀
, Ma ue (Okushi i- ô), Okushi i-chô, Okushi i-gun (42.072N,
139.471E), 12 VII 2008, Ôha a-M. (42°04
′
18
″
N, 139°28
′
17
″
E; unde seaweed; 5
♂
, 6
♀
, Sawa a, Mo i-machi,
Kayabe-gun (42.123N, 140.647E), 15 VII 2009, Ôha a-M. (42°07
′
23
″
N, 140°38
′
50
″
E; unde seaweed (Sa gassum
sp.); HUM: HK-09-MO-040/SA); HUM: HK-09-MO-032/SA); 2
♂
, Esashi-chô, Hiyama-gun (41.857N,
140.125E*), 4 VI 1977, Nishikawa-N. (KUM); 1
♀
, Hakoda e-shi (41.760N, 140.693E*), 13 V 1971, Hi ano-Y.
(KUM); 1
♂
, 1
♀
, Hon-chô, Kikonai-chô, Kamiiso-gun (41.676N, 140.439E), 12 VII 2009, Ôha a-M.
(41°40
′
33
″
N, 140°26
′
22
″
E; unde seaweed (Lamina ia sp., Sa gassum sp.) and eel g ass (Zos e a sp.); HUM: OK-
08-MO-015). [Honshû]: 1
♂
, Rikuchû-Yagi, Taneichi, Hi ono-chô, Kunohe-gun (40.349N, 141.764E), 28 VI
2010, Ôha a-M. (40°20
′
56
″
N, 141°45
′
49
″
E; unde seaweed (Lamina ia sp., Sa gassum sp.) and eel g ass (Zos e a
sp.) on sandy beach: HUM: HN-10-MO-044/SA); 2
♀
, Raga, Tanoha a-mu a, Shimohei-gun, Iwa e-ken (39.939N,
141.940E*), 30 V 1998, Wa anabe-T. (cWa ); 1
♂
, Jôdoga-hama, Hi achihama-chô, Miyako-shi, Iwa e-ken
(39.649N, 141.981E), 29 V 1998, Wa anabe-T. (cWa ); 3
♀
, Isoha a, Ki aiba aki-shi, Iba aki-ken (36.790N,
140.752E), 24 V 1998, Ohmomo-S. (KUM); 1
♂
, Majima, Mu akami-shi, Niiga a-ken (38.273N, 139.449E*), 5 V
2000, no collec o ’s name (KUM); 1
♀
, Ben en-misaki (Sadoga-shima), Higashikowashimizu, Sado-shi, Niiga a-
ken (38.008N, 138.546E), 4 V 1998, Kinoshi a-T. (cWa ); 1
♂
, same da a, 5 V 1998, Tsuyuki-S. (cWa ); 1
♂
,
Hamaogi-kaigan, Hamaogi, Kamogawa-shi, Chiba-ken (35.118N, 140.144E), 24 IV 1990, Takeda-T. (KUM); 1
♂
,
1
♀
, same da a, bu 6 IV 1993; 2
♂
, 1
♀
, Heisau a, Sakai, Ta eyama-shi, Chiba-ken (34.948N, 139.801E), 1 IV
1997, Ma uyama-M. (cMa ); 1
♀
, Shinmaiko-hama, Yawa a, Fu su-shi, Chiba-ken (35.250N, 139.866E), 11 VI
1995, Emo o-K. (cWa ); 5
♂
, 4
♀
, Shibasaki-bashi b idge, Kanaya, Fu su-shi, Chiba-ken (35.174N, 139.819E), 14
VI 2009, Ono-H. (cOno); 1
♂
, 1
♀
, Kaneda swimming beach, Kaneda, Minamishi au a-machi, Miu a-shi,
Kanagawa-ken (35.167N, 139.660E), 19 III 2001, Wa anabe-T. (cWa ); 2
♂
, 2
♀
, Jôgashima (Jôga-shima), Misaki-
machi, Miu a-shi, Kanagawa-ken (35.132N, 139.621E), 10 IV 2003, Wa anabe-T. (cWa ); 1
♀
, Mi o beach, Hasse-
machi, Miu a-shi, Kanagawa-ken (35.177N, 139.620E), 29 V 2007, Wa anabe-T. (cWa ); 5
♂
, 4
♀
, same da a, bu
5 VI 2007; 1
♂
, 1
♀
, same da a, bu 14 V 2009 (collec ed wi h Aleocha a (E.) hayamai); 3
♂
, 1
♀
, same da a, bu
12 VI 2009; 3
♂
, 1
♀
, A asaki, Nagai, Yokosuka-shi, Kanagawa-ken (35.194N, 139.600E*), 12 VI 2009,
Zoo axa 3517 © 2012 Magnolia P ess · 21
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
Wa anabe-T. (cWa ); 1
♂
, 3
♀
, Shônan-kaigan, Kanagawa-Ken (35.315N, 139.351E*), 2 VII 1976, Niiza o
(KUM); 1
♂
, Eno-shima, Fujisawa-shi, Kanagawa-ken (35.300N, 139.485E*), 11 VI 2001, Wa anabe-T. (cWa ); 1
♂
, 3
♀
, Mi suishi, Manazu u, Manazu u-shi, Ashiga ashimo-gun, Kanagawa-ken (35.138N, 139.163E), 15 VII
1989, Wa anabe-T. (cWa ); 2
♂
, Shi a-hama, Shi ahama, Shimoda-shi, Shizuoka-ken (34.689N, 138.973E), 13 III
1989, Oda-H. (KUM); 33
♂
, 18
♀
, Tsumeki-zaki, Shimoda-shi, Shizuoka-ken (34.659N, 138.987E), 21 IV 2003,
Ma uyama-M. (cMa ); 1
♂
, 1
♀
, i e mou h o Kushida-gawa, Ma sunase-chô, Ma suzaka-shi, Mie-ken
(34.605N, 136.581E), 29 IV 2002, Kawakami-Y. (cKaw); 1
♂
, Maizu u, Kyô o- u (35.514N, 135.387E*), 3 V
1980, Hayashi-Y. (cHayas); 1
♂
, Shimakage, Miyazu-shi, Kyô o- u (35.576N, 135.250E*), 31 III 2000, Yasukawa-
K. (KUM); 1
♀
, Shionomisaki, Kushimo o-chô, Higashimu o-gun, Wakayama-ken (33.445N, 135.756E*), 17 V
1964, Yasui-M. (cHayas); 1
♀
, Kada-kaigan, Kada, Wakayama-shi, Wakayama-ken (34.280N, 135.074E), 4 IX
2003, Wada-T. (KUM); 1
♀
, Ni iga-hama, Isonou a, Wakayama-shi, Wakayama-ken (34.259N, 135.094E), 15 V
2007, Kawakami-Y. (cKaw); 1
♀
, i e mou h o Ashiya-gawa, Minamihama-chô, Ashiya-shi, Hyôgo-ken
(34.708N, 135.309E), 27 V 1998, Kawakami-Y. (cKaw); 1
♂
, Sumau a-kaigan, Sumau adô i, Suma-ku, Kôbe-shi,
Hyôgo-ken (34.637N, 135.103E), 16 VI 1998, Kawakami-Y. (cKaw); 1
♀
, same da a, bu 15 IV 2004; 4
♂
, same
da a, bu 27 V 2010; 2
♂
, 1
♀
, Maiko-hama, Ta umi-ku, Kôbe-shi, Hyôgo-ken (34.638N, 135.027E), 16 VI 1998,
Kawakami-Y. (cKaw); 2
♂
, Nada-kaigan (Awaji-shima), Nada, Minamiawaji-shi, Hyôgo-ken (34.207N,
134.812E*), 1 IV 1977, Kaneda (KUM); 1
♀
, same locali y, bu 3 X 1972, Tomokuni-M. and Sakai-M. (EEEU); 1
♂
, Gunge-hama (Awaji-shima), Gunge, Awaji-shi, Hyôgo-ken (34.475N, 134.845E), 5 V 2000, Kawakami-Y.
(cKaw); 2
♂
, 1
♀
, Sumo o coas (Awaji-shima), Sumo o-shi, Hyôgo-ken (34.343N, 134.902E*), 23 V 1998,
Kawakami-Y. (cKaw); 1
♀
, i e mou h o Akashi-gawa, Akashi, Akashi-shi, Hyôgo-ken (34.644N, 134.975E), 9
IV 2002, Kawakami-Y. (cKaw); 1
♀
, Yagi, Ôkubo-chô, Akashi-shi, Hyôgo-ken (34.670N, 134.933E), 9 V 1999,
Kawakami-Y. (cKaw); 1
♂
, Ma oga a beach, Ma oga a, Ma oga a-chô, Himeji-shi, Hyôgo-ken (34.769N,
134.746E), 16 V 2010, Kawakami-Y. (KUM); 1
♂
, Fukudoma i, Ma oga a-chô, Himeji-shi, Hyôgo-ken (34.769N,
134.737E), 2 VI 2000, Kawakami-Y. (cKaw); 1
♀
, Shi ahama, Shi ahama-chô, Himeji-shi, Hyôgo-ken (34.777N,
134.708E), 2 VI 2000, Kawakami-Y. (cKaw); 1
♂
, 3
♀
, i e mou h o Chigusa-gawa, Onzaki, Akô-shi, Hyôgo-
ken (34.731N, 134.393E), 15 VI 1999, Kawakami-Y. (cKaw); 1
♂
, 1
♀
, Kugui, Higashi-ku, Okayama-shi,
Okayama-ken (34.584N, 134.086E*), 31 III 2003, Fuji ani-Y. (KUM); 5
♂
, 2
♀
, same da a, bu 9 V 2003 (unde
seaweed on beach); 3
♂
, 2
♀
, same da a, bu 28 V 2003; 1
♂
, Inazumi, Sugeu a, Mihonoseki-chô, Ma ue-shi,
Shimane-ken (35.564N, 133.166E*), 1-2 VI 2008, Hayama-T. (sandy beach; cHayam); 2
♂
, Okidoma i, Tako,
Shimane-chô, Ma ue-shi, Shimane-ken (35.604N, 133.096E*), 3 VI 2008, Hayama-T. ( om seaweed; cHayam); 2
♂
, Konami-kaigan, Nonami, Shimane-chô, Ma ue-shi, Shimane-ken (35.588N, 133.098E), 6 V 2008, Hayama-T.
(cHayam); 1
♀
, Kakanokuke o, Shimane-chô, Ma sue-shi, Shimane-ken (35.578N, 133.050E), 9 VI 2009,
Hayama-T. (cHayam); 1
♀
, same da a, bu 19 VII 2010, Yamamo o-S. (shingle beach; unde lo som; cYam); 41
♂
,
56
♀
, Kou a-kaigan, Kou a, Kashima-chô, Ma ue-shi, Shimane-ken (35.5201N, 132.975E), 8-10 VI 2009,
Hayama-T. (FIT; cHayam); 2
♀
, same da a, bu 3-5 VII 2009; 12
♂
, 36
♀
, same da a, bu 18-20 VI 2009; 1
♂
, 1
♀
,
same da a, bu 16-18 VII 2009; 3
♂
, 1
♀
, same locali y, bu 17 VII 2010, Yamamo o-S. (unde lo sam on sandy
beach; cYam); 1
♂
, same da a, bu 18 VII 2010; 2
♀
, Sakau a-kaigan, Sakau a-chô, Izumo-shi, Shimane-ken
(35.508N, 132.861E), 4 VII 2008, Hayama-T. (shingle beach wi h sho e ee s; cHayam); 1
♂
, same da a, bu 15 IV
2009 ( om seaweed); 1
♂
, 4
♀
, same da a, bu 20 IV 2009; 1
♂
, i e mou h o Sakau a-gawa, Sakau a-chô,
Izumo-shi, Shimane-ken (35.508N, 132.861E), 24 III 2009, Hayama-T. (shingle beach; cHayam); 1
♂
, i e mou h
o Mi su-gawa, Mi su-chô, Izumo-shi, Shimane-ken (35.498N, 132.826E), 30 IV 2008, Hayama-T. (cHayam); 2
♀
,
i e mou h o Inome-gawa, Inome-chô, Izumo-shi, Shimane-ken (35.441N, 132.710E), 21 III 2008, Hayama-T.
(shingle beach; cHayam); 5
♂
, Owashi-hama, Nakayama, Taisha-chô, Izumo-shi, Shimane-ken (35.433N,
132.634E), 10 IV 2009, Hayama-T. (cHayam); 2
♂
, Akaishihana, Hinomisaki, Taisha-chô, Izumo-shi, Shimane-
ken (35.411N, 132.650E*), 2 IV 2009, Hayama-T. (cHayam); 2
♂
, same da a, bu 2-8 V 2009 (FIT; cHayam); 1
♀
,
So ozono-kaigan, Sashiumi, Ko yô-chô, Izumo-shi, Shimane-ken (35.333N, 132.664E), 12-13 IV 2009, Hayama-
T. (YPT; cHayam); 2
♂
, Kumu a-kaisuiyokujô swimming beach, Kumu a, Taki-chô, Izumo-shi, Shimane-ken
(35.296N, 132.637E), 5 X 2008, Hayama-T. (cHayam); 2
♂
, 1
♀
, Kumu a, Taki-chô, Izumo-shi, Shimane-ken
(35.296N, 132.637E*), 18 IV 2009, Hayama-T. (cHayam); 1
♂
, Kuchi aki, Taki-chô, Izumo-shi, Shimane-ken
(35.271N, 132.581E), 18 IV 2009, Hayama-T. (cHayam); 1
♂
, 1
♀
, Iwami-kaigan, Kushi o-chô, Hamada-shi,
Shimane-ken (34.955N, 132.127E*), 16-17 VI 2008, Hayama-T. (YPT; cHayam); 1
♂
, 1
♀
, Kiami-chô, Masuda-
shi, Shimane-ken (34.676N, 131.750E*), 15 VI 2008, Hayama-T. (cHayam); 2
♂
, 1
♀
, same da a, bu 15-16 VI
YAMAMOTO & MARUYAMA
22 · Zoo axa 3517 © 2012
Magnolia P ess
2008 (FIT); 1
♂
, 1
♀
, same da a, bu 10 V 2009; 7
♀
, same da a, bu 10-11 V 2009 (FIT); 7
♂
, 11
♀
, Yoshimi,
Shimonoseki-shi, Yamaguchi-ken (34.064N, 130.905E*), 1 VI 1980, Naomi-S.I. (KUM). [Shikoku]: 3
♂
, 5
♀
,
A iake-hama, Mu omo o-chô, Kanonji-shi, Kagawa-ken (34.139N, 133.642E), 9 IV 2011, Fujimo o-H. (sandy
beach; KUM); 21
♂
, 10
♀
, Iwagi (Akahone-jima), Kamijima-chô, Ochi-gun, Ehime-ken (34.234N, 133.159E*), 2
V 2009, Sa ô-Y. (EEEU); 3
♀
, same locali y, bu 7 V 2009, Senda-Y. (EEEU); 1
♀
, Hôjô suji, Ma suyama-shi,
Ehime-ken (33.974N, 132.770E), 16 VI 2008, Senda-Y. (EEEU). [Kyûshû]: 3
♂
, 4
♀
, Wa a i, Fuku su-shi,
Fukuoka-ken (33.786N, 130.454E), 21 IV 2012, Yamamo o-S., Ma uyama-M., Kanao-T. ( om seaweed on small
sandy beach wi h Aleocha a (T iocha a) isulca a; KUM); 4
♂
, 6
♀
, Wa a i, Fuku su-shi, Fukuoka-ken (33.790N,
130.447E), 21 IV 2012, Yamamo o-S., Ma uyama-M., Kanao-T. ( om seaweed on huge sandy beach wi h A. (T.)
isulca a; KUM); 1
♀
, Mi oma, Higashi-ku, Fukuoka-shi, Fukuoka-ken (33.703N, 130.417E), 3 VI 2001, Oga a-
S. (KUM); 2
♂
, 1
♀
, Shikano-shima, Higashi-ku, Fukuoka-shi, Fukuoka-ken (33.675N, 130.312E*), 24 VIII 1980,
Naomi-S.I. (KUM); 3
♂
, 1
♀
, Shimomaga-hama, Ka suma, Higashi-ku, Fukuoka-shi, Fukuoka-ken (33.684N,
130.290E), 15 V 2011 (11:30), Yamamo o-S. (seaweed on sandy beach oge he wi h A. (T.) isulca a; cYam); 25
♂
, 56
♀
, same da a, bu 9 VII 2011 (15:00-16:00; om seaweed and lo sam on beach); 28
♂
, 39
♀
, same da a, bu
21 III 2012 (14:00); 3
♂
, 7
♀
, Odo-kôen pa k, Odo-2chôme 6, Nishi-ku, Fukuoka-shi, Fukuoka-ken (33.596N,
130.311E), 28 VI 2009, Yamamo o-S. ( om decaying seaweed along sandy seasho e; cYam); 24
♂
, 26
♀
,
Ikinoma suba a-1chôme, Nishi-ku, Fukuoka-shi, Fukuoka-ken (33.580N, 130.293E), 28 VI 2009, Yamamo o-S.
( om seaweed on sandy beach; same da a as holo ype; cYam); 3
♂
, 6
♀
, Noko (Nokono-shima), Nishi-ku,
Fukuoka-shi, Fukuoka-ken (33.608N, 130.299E), 20 III 2009, Yamamo o-S. ( om a seaweed mass on sandy beach
wi h A. (T.) isulca a and A. (T.) ze chei; cYam]; 1
♀
, Hongô (Saki o-jima), Saki o-chô, Saikai-shi, Nagasaki-ken
(33.003N,129.550E*), 23 X 1977, Imasaka-S. (KUM). [Ryûkyû]: 1
♀
, Naka ane-chô (Tanega-shima), Kumage-
gun, Kagoshima-ken (30.515N, 130.980E), 8-13 V 1996, Ma uyama-M. (cMa ).
Desc ip ion. Body (Fig. 5): medium o ela i ely la ge size, no mally medium sized. Uni o mly densely hai ed
excep o head along midline; su ace o p ono um and ely a co e ed wi h nume ous somewha long hai s
densely; head, p ono um and ely a wi h dis inc punc u es on do sal su ace densely. Colou (Fig. 5): gland colou
blackish b own o da k g ay; legs, especially a sal segmen s, eddish b own o da k b own: maxilla y and labial
palpi eddish b own o yellowish b own; an ennae blackish b own and pa ly eddish b own. Head: head capsule
somewha ec angle; su ace en i ely a he la ened, bu impunc u ed a ea along midline longi udinally swollen
(weakly ca ina e); su ace spa sely co e ed wi h hin and somewha long se ae excep o swollen a ea; uni o mly
obse ed compa a i ely la ge hexagonal e icula ions on do sal su ace. An ennae: mode a ely hick and long;
segmen I, nea ly 2.1 imes as long as b oad; segmen II clea ly sho e han I; segmen III as same leng h as II;
segmen IV sphe ical o sho ba el shape, as long as wid h; segmen s V and VI sho ba el shape; sebmen s VII
o X clea ly ans e se; segmen XI, abou 1.6 imes as long as b oad; ela i e leng h (wid h) o segmen s om
basal o apical: 10.5(5): 7(4): 7(4): 4(4): 4.5(5): 4.5(5): 4(5): 3.5(5): 4(6): 4.5(6): 9(5.5). Tho ax: p ono um
somewha wide han long (PW/PL =1.24), a li le b oade han head (PW/HW =1.36); su ace ma wi h la ge
hexagonal e icula ions wi h somewha shallow dis inc punc u es. Me a en i e, abou 1.5 imes as long as
meso en i e. Apex o in e coxal p ocess o meso en i e sha ply poin ed wi h e y sho and inconspicuous ca ina
along midline, abou 0.3 imes as long as meso en i e. In e coxal p ocess o me a en i e e y b oad and sho ,
less han one hi d as long as mesocoxal ca i y, and ounded apically. Legs: ela i e leng hs o a some es om
basal o apical: 5.5: 3: 4.5: 4.5: 10.5 in o e a sus, 7.5: 5.5: 5.5: 5.5: 13.5 in mid a sus, 9.5: 6.5: 6.5: 7.5: 16.5 in
hind a sus.
[Male]: ou e ma gin o e gi e VIII (Fig. 30) nea ly unca e, wi h a ound 7 mac ose ae. S e ni e VIII (Fig. 32)
wi h abou 5 mac ose ae and a ound 10 hin mac ose ae; pos e io ma gin s ongly poin ed wi h ounded apex.
Median lobe o aedeagus (Figs. 34–35) elonga ed and sligh ly na owing owa d apex in la e al iew (Fig. 34),
subpa allel sided in en al iew (Fig. 35); a pai o subapico- en al p ojec ions gen ly cu ed in la e al iew (Fig.
34); basal swelling o aedeagus, ci cula ly poin ed in la e al iew (Fig. 34); apical lobe sho and ab up ly
na owing owa d apex in en al iew (Figs. 35–36); lagellum clea ly sho e han he whole leng h o median
lobe (Figs. 34–35).
[Female]: e gi e VIII (Fig. 31) wi h mode a ely ounded ma gin, wi h a ound 7 mac ose ae. S e ni e VIII (Fig.
33) wi h abou 7 mac ose ae; pos e io ma gin sligh ly poin ed. Spe ma heca (Fig. 37): en i ely hick; spe ma hecal
head abou wo o h ee imes longe han apical po ion o spe ma hecal s em; spe ma hecal neck sho e han (sh);
basal po ion o spe ma hecal s em sho , sligh ly na owing owa d base, lacking any ben a ound middle and base;
Zoo axa 3517 © 2012 Magnolia P ess · 23
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
lack o scle o ized po ion o spe ma hecal s em; each pa o spe ma heca en i ely and mode a ely scle o ized
excep o memb aneous po ion o spe ma hecal duc (sm); (sm) mode a e in leng h.
Measu emen s (male: n=10): BL, 3.28–4.96 (4.11±0.58); FBL, 1.57–2.35 (1.92±0.23); HL, 0.44–0.66
(0.55±0.08); HW, 0.52–0.71 (0.62±0.06); AL, 0.95–1.23 (1.13±0.09); PL, 0.56–0.78 (0.68±0.07); PW, 0.71–1.00
(0.86±0.09); EL, 0.57–0.84 (0.67±0.08); EW, 0.93–1.27 (1.11±0.12); HTL, 0.53–0.75 (0.63±0.07).
Measu emen s ( emale: n=10): BL, 3.25–4.63 (4.18±0.49); FBL, 1.64–2.15 (1.92±0.16); HL, 0.42–0.65
(0.57±0.07); HW, 0.56–0.71 (0.64±0.05); AL, 0.92–1.26 (1.10±0.10); PL, 0.60–0.80 (0.70±0.06); PW, 0.75–0.99
(0.85±0.07); EL, 0.56–0.85 (0.70±0.09); EW, 0.96–1.22 (1.12±0.09); HTL, 0.54–0.72 (0.63±0.06).
Dis ibu ion. [JAPAN]: Hokkaidô, Honshû, Shikoku, Kyûshû, no he n Ryûkyû, Okushi i- ô, Sadoga-shima,
Awaji-shima, Tanega-shima. See, Fig. 102.
O he locali ies in li e a u e. [SOUTH KOREA]: Gangwon (Kangwon) P o ince, Chungnam P o ince,
Gyeongnam P o ince, Jeonnam P o ince, Jeju P o ince (Ahn e al., 2000; Pa k & Ahn, 2004; bo h as “A. pue zi”;
e-examina ions a e needed).
Diagnosis. This new species is simila o he o he species, especially A. ucicola and A. pue zi, also in male
geni alia. We ound se e al dis inguishable cha ac e s a es om o he Empleno a species below: a ying om
medium o ela i ely la ge size (no e y small o e y la ge); en i e body pubescen wi h weak bu dense punc u es
on do sal su ace o head, p ono um and ely a (Fig. 5); impunc u ed a ea along midline o head longi udinally and
p ominen ly swollen abo e, o ming ca ina, bu o he a eas la ened compa ed o A. ucicola; an ennae ela i ely
hick and obus , wi h sho ba el o sphe ical shape o segmen IV; meso en i e wi h qui e sho ca ina, abou
0.27 imes as long as meso en i e; in e coxal ca i y o meso en i e sha ply poin ed. [Male]: s e ni e VIII (Fig.
32) s ongly poin ed wi h ounded apex; subapico- en al p ojec ions on median lobe gen ly ounded in la e al
iew (Fig. 34); median lobe subpa a ell sided in en al iew (Fig. 35); apex o median lobe sho in la e al iew
(Fig. 34); apical lobe isosceles shape, bu na owing ab up ly a base in en al aspec (Figs. 35–36): basal po ion
o median lobe poin ed oblongly, owa ds o en al ma gin o median lobe in la e al aspec (Fig. 34). [Female]:
spe ma heca (Fig. 37) qui e hick and wide, wi hou e ec and scle o ized s em (ss).
E ymology. The species is named a e he La in adjec i e seg ega us meaning sepa a ed. Because o he close
simila i y o he ex e nal appea ances among A. seg ega a, A. ucicola, and A. pue zi, some esea che s ha e o en
con used hese axa (e.g., Ahn e al., 2000; Pa k & Ahn, 2004).
Rema ks. This new species is e y simila o Aleocha a (Empleno a) ucicola and A. (E.) pue zi in gene al
appea ance. The e o e, he species has gene a ed some p oblems ega ding he axonomy o Empleno a species
occu ing in adjacen egions o Japan (see, ema ks o A. ucicola and A. pue zi).
Ca ina like longi udinal ele a ion on he do sal su ace o he head cha ac e ises his species in mo phology,
bu obse a ion unde a mic oscope is needed o con i m he cha ac e s a e.
The species has a wide- anging dis ibu ion in Japan, om he no he n (Hokkaidô) o he sou he n islands
(Tanega-shima, no he n Ryûkyû). In addi ion, his species may be he mos common Empleno a species in Japan,
because la ge numbe s o specimens ha e been collec ed by many esea che s. P e ious eco ds o he Asian
Empleno a specimens need o be eexamined o be con i med, especially hose eco ded om China and he
Ko ean peninsula.
Aleocha a (Empleno a) hayamai Yamamo o & Ma uyama n. sp.
(Figs. 3, 38–45, 93–94, 97, 103)
Type se ies. Holo ype,
♂
, JAPAN: Kou a-kaigan, Kou a, Kashima-chô, Ma ue-shi, Shimane-ken, Honshû
(35.521N, 132.975E), 18 VII 2010, Yamamo o-S. (by hand wi h aspi a o ; unde lo sam on sandy beach in he ho
day ime; KUM).
Pa a ypes, JAPAN: [Honshû]: 2
♂
, Mi o beach, Hasse-machi, Miu a-shi, Kanagawa-ken (35.177N,
139.620E), 14 V 2009, Wa anabe-T. (cWa ); 1
♂
, Inugu (Dôgo Is., Oki-sho ô), Okinoshima-chô, Oki-gun,
Shimane-ken (36.219N, 133.372E*), 10 XI 2009, Hayama-T. (cHayam); 4
♂
, 3
♀
, Kakanokuke o, Shimane-chô,
Ma sue-shi, Shimane-ken (35.578N, 133.051E), 1 IV 2009, Hayama-T. (shingle beach; cHayam); 2
♂
, 2
♀
, same
da a, bu 9 VI 2009; 2
♂
, same da a, bu 18 VII 2010, Yamamo o-S. (cYam); 1
♂
, same da a, bu 19 VII 2010; 1
♂
,
Kaka (Ka su a-jima), Shimane-chô, Ma sue-shi, Shimane-ken (35.566N, 133.052E), 19 VII 2010, Yamamo o-S.
YAMAMOTO & MARUYAMA
24 · Zoo axa 3517 © 2012
Magnolia P ess
(by hand pick; sandy beach; unde seaweed a ound e ening; cYam); 18
♂
, 4
♀
, Kou a-kaigan, Kou a, Kashima-
chô, Ma ue-shi, Shimane-ken (35.521N, 132.975E), 8-10 VI 2009, Hayama-T. (FIT; cHayam); 31
♂
, 26
♀
, same
da a, bu 18-20 VI 2009; 3
♂
, same da a, bu 3-5 VII 2009; 4
♂
, 8
♀
, same da a, bu 16-18 VII 2009; 8
♂
, 4
♀
,
same da a, bu 17 VII 2010, Yamamo o-S. (by hand pick up wi h aspi a o ; unde lo sam on sandy beach; KUM);
1
♂
, 2
♀
, same da a, bu 18 VII 2010 (same da a as holo ype); 2
♂
, Sakau a-kaigan, Sakau a-chô, Izumo-shi,
Shimane-ken (35.508N, 132.861E), 20 IV 2009, Hayama-T. (cHayam); 1
♀
, same da a, bu 15 V 2009; 1
♀
, i e
mou h o Sakau a-gawa, Sakau a-chô, Izumo-shi, Shimane-ken (35.508N, 132.861E), 13 IV 2008, Hayama-T.
(cHayam); 1
♂
, same da a, bu 31 IV 2008; 1
♂
, same da a, bu 17 V 2008; 1
♂
, 31 VIII 2008; 1
♂
, same da a, bu
5 VIII 2009; 2
♂
, 1
♀
, Owashi-hama, Nakayama, Taisha-chô, Izumo-shi, Shimane-ken (35.433N, 132.634E), 24
III 2009, Hayama-T. (shingle beach; cHayam); 1
♂
, 1
♀
, Akaishihana, Hinomisaki, Taisha-chô, Izumo-shi,
Shimane-ken (35.411N, 132.650E*), 2 IV 2009, Hayama-T. (cHayam); 7
♀
, same da a, bu 2-8 V 2009 (FIT); 1
♂
,
Kiami-chô, Masuda-shi, Shimane-ken (34.676N, 131.747E), 10-11 V 2009, Hayama-T. (FIT; cHayam). [Kyûshû]:
1
♂
, Odo-kôen pa k, Odo-2chôme 6, Nishi-ku, Fukuoka-shi, Fukuoka-ken (33.596N, 130.311E), 28 VI 2009,
Yamamo o-S. ( om seaweed on sandy beach wi h Aleocha a (E.) seg ega a; cYam); 2
♂
, Sa o (Kamikoshiki-
jima), Sa o-machi, Sa sumasendai-shi, Kagoshima-ken (31.844N, 129.919E), 11-12 IV 1973, Fu uki-Y. (EEEU).
Desc ip ion. Body (Fig. 3): small o medium sized, slende body; su ace o p ono um and ely a somewha
densely hai ed by a he sho and hin se ae; o e-body (head, p ono um and ely a) weakly shining due o
ex emely minu e hexagonal mic os uc u es. Colou (Fig. 3): gland colou comple e black; legs da k b own o
eddish b own, a sal segmen s o ange b own o eddish b own: maxilla y and labial palpi blackish b own o
yellowish b own; an ennae blackish b own and pa ly eddish b own. Head: impunc u ed a ea along midline no
uniquely ele a ed a all. An ennae (Fig. 42): en i ely slende ; sligh ly longe han combined leng h o head and
p ono um; segmen I, nea ly 2.8 imes as long as b oad; segmen II somewha sho e han I; segmen III as same
leng h as II; segmen s IV and V long ba el shape, clea ly longe han wid h; segmen s VI o VII sho ba el shape;
segmen s VIII o X ans e se; las segmen somewha long, abou 1.7 imes as long as b oad; ela i e leng h
(wid h) o segmen s om basal o apical: 11(4): 7(3): 6.5(3): 4(3): 4.5(3.5): 4(4): 4(4): 5(5): 6(5): 4.5(5): 10(6).
Tho ax: p ono um somewha s ongly con ex abo e, wi h mode a ely ounded pos e io ma gin; sligh ly wide
han long (PW/PL =1.23), clea ly b oade han head (PW/HW =1.38), wides jus behind middle; su ace wi h
ex emely minu e hexagonal e icula ions wi h inconspicuous punc u es. Me a en i e, abou 1.6 imes as long as
meso en i e. In e coxal p ocess o meso en i e sha ply poin ed wi h e y sho ca ina along midline, abou 0.3
imes as long as meso en i e. In e coxal p ocess o me a en i e b oad and somewha long, and apical ma gin
poin ed mode a ely. Legs: ela i e leng hs o a some es om basal o apical: 4.5: 3: 3: 3: 10 in o e a sus, 6: 5: 5:
5: 13 in mid a sus, 8: 6: 6: 6: 16 in hind a sus.
[Male]: pos e io ma gin o e gi e VIII (Fig. 38) e y weakly ema gina ed medially, wi h a ound 7 o 8
mac ose ae. S e ni e VIII (Fig. 40) wi h abou 12 mac ose ae; pos e io ma gin e y s ongly poin ed owa d apex,
o ming iangula . Median lobe o aedeagus (Figs. 43–44) C-cu ed (sponge-gou d shape) in la e al iew (Fig.
43); mode a ely na owed apically in en al iew (Fig. 44); a pai o subapico- en al p ojec ions hick, obus and
s aigh in la e al iew (Fig. 43); basal swelling o aedeagus, ci cula ly ounded in la e al iew (Fig. 43); apical
lobe o median lobe e y sho isosceles shape in en al aspec (Fig. 44); lagellum ex emely longe han he
whole leng h o median lobe (Figs. 43–44).
[Female]: pos e io ma gin o e gi e VIII (Fig. 39) sligh ly ounded, bu e y weakly ema gina ed medially,
wi h abou 7 mac ose ae. Pos e io ma gin o s e ni e VIII (Fig. 41) ounded o weakly poin ed, wi h a ound 8
mac ose ae. Spe ma heca (Fig. 45): spe ma hecal head as long as apical po ion o spe ma hecal s em; basal
po ion o spe ma hecal s em e y sho and cu ed; scle o ized po ion o spe ma hecal s em hin and e ec ; each
pa o spe ma heca excep o memb aneous po ion o spe ma hecal duc (sm) en i ely and e y weakly
scle o ized; (sm) ex emely long in leng h.
Measu emen s (male: n=10): BL, 2.94–4.65 (4.00±0.49); FBL, 1.66–2.07 (1.87±0.15); HL, 0.46–0.68
(0.56±0.07); HW, 0.53–0.69 (0.60±0.05); AL, 1.11–1.51 (1.30±0.14); PL, 0.59–0.78 (0.69±0.06); PW, 0.60–0.92
(0.83±0.06); EL, 0.57–0.77 (0.67±0.06); EW, 0.94–1.27 (1.08±0.10); HTL, 0.57–0.79 (0.66±0.07).
Measu emen s ( emale: n=10): BL, 3.23–4.64 (4.04±0.38); FBL, 1.50–2.11 (1.91±0.20); HL, 0.44–0.62
(0.55±0.05); HW, 0.51–0.66 (0.60±0.04); AL, 1.03–1.32 (1.18±0.09); PL, 0.53–0.73 (0.66±0.07); PW, 0.68–0.91
(0.83±0.07); EL, 0.62–0.77 (0.69±0.05); EW, 0.87–1.18 (1.07±0.10); HTL, 0.55–0.78 (0.68±0.07).
Zoo axa 3517 © 2012 Magnolia P ess · 25
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
FIGURES 46–52. Te minalia o Aleocha a (Empleno a) yama o. 46. e gi e VIII o male; 47. e gi e VIII o emale; 48.
s e ni e VIII o male; 49. s e ni e VIII o emale; 50. male geni alia: median lobe o aedeagus in la e al iew; 51. male geni alia:
median lobe o aedeagus in en al iew; 52. emale geni alia: spe ma heca.
Dis ibu ion. [JAPAN]: Honshû, Kyûshû, Dôgo Is., Kamikoshiki-jima. See, Fig. 103.
Diagnosis. Aleocha a hayamai is a peculia species and can be easily disc imina ed om o he Japanese
Empleno a species by ollowing cha ac e s a es: small o medium size body (no la ge species); gland colou
YAMAMOTO & MARUYAMA
32 · Zoo axa 3517 © 2012
Magnolia P ess
Tsuyuki-S. (cWa ); 2 sex?, Ika ashi-hama, Ika ashi-2no-chô, Niiga a-shi, Niiga a-ken (37.871N, 138.927E), 29 V
1991, Hayashi-M. (sandy beach; cKaw); 1
♂
, Naga-hama, Nagai-2chôme 13, Yokosuka-shi, Kanagawa-Ken
(35.191N, 139.615E), 7 IV 1975, Tao-M. (KUM); 1 sex?, Tsumeki-zaki, Shimoda-shi, Shizuoka-ken (34.659N,
138.987E), 21 IV 2003, Ma uyama-M. (cMa ); 2 sex?, Mukaiawagasaki, Uchinada-machi, Kahoku-gun, Ishikawa-
ken (36.640N, 136.622E), 12 IV 1948, Takaba-S. (KUM); 2 sex?, Obama-shi, Fukui-ken (35.485N, 135.719E*), 6
VI 1980, Naomi-S.I. (KUM); 1
♀
, Maizu u, Kyô o- u (35.514N, 135.387E*), 3 V 1980, Hayashi-Y. (cHayas); 1
sex?, i e mou h o Onosa o-gawa, Ta ui, Sennan-shi, Ôsaka- u (34.377N, 135.250E), 15 IV 2001, Kawakami-Y.
(cKaw); 3
♂
, 24 sex?, Kugui, Higashi-ku, Okayama-shi, Okayama-ken (34.584N, 134.086E*), 31 III 2003,
Fuji ani-Y. (KUM); 2
♀
, 31 sex?, same da a, bu 9 V 2003 (unde seaweed on beach); 18 sex?, same da a, bu 28 V
2003; 1
♂
, 11 sex?, Inazumi, Sugeu a, Mihonoseki-chô, Ma ue-shi, Shimane-ken (35.564N, 133.166E*), 1-2 VI
2008, Hayama-T. (sandy beach; cHayam); 70 sex?, Konami-kaigan, Nonami, Shimane-chô, Ma ue-shi, Shimane-
ken (35.589N, 133.098E), 2-3 VI 2008, Hayama-T. (cHayam); 1 sex?, Kou a-kaigan, Kou a, Kashima-chô, Ma ue-
shi, Shimane-ken (35.520N, 132.975E), 16-18 VII 2009, Hayama-T. (FIT; cHayam); 47 sex?, So ozono-kaigan,
Sashiumi, Ko yô-chô, Izumo-shi, Shimane-ken (35.333N, 132.664E), 8 III 2007, Hayama-T. (unde seaweed;
cHayam); 37 sex?, same da a, bu 12-13 IV 2009; 26 sex?, Ko oga-hama, Maji, Nima-chô, Ôda-shi, Shimane-ken
(35.131N, 132.389E), 2 V 2009, Hayama-T. (YPT on sup a idal zone; cHayam); 6 sex?, Ku oma su-chô, Gô su-
shi, Shimane-ken (35.057N, 132.310E), 2 V 2009, Hayama-T. ( om seaweed; cHayam); 1 sex?, Iwami-kaigan,
Kushi o-chô, Hamada-shi, Shimane-ken (34.955N, 132.127E), 16-17 VI 2008, Hayama-T. (YPT; cHayam); 1 sex?,
Nagashima (Naga-shima), Kaminoseki-chô, Kumage-gun, Yamaguchi-ken (33.827N, 132.091E), 1 V 2008,
Mo iguchi-M. (KUM). [Shikoku]: 3 sex?, A iake-hama, Mu omo o-chô, Kanonji-shi, Kagawa-ken (34.139N,
133.642E), 9 IV 2011, Fujimo o-H. (sandy beach; KUM); 23 sex?, Iwagi (Akahone-jima), Kamijima-chô, Ochi-
gun, Ehime-ken (34.234N, 133.159E*), 2 V 2009, Sa ô-Y. (EEEU); 5 sex?, same da a, bu 7 V 2010, Senda-Y.; 2
♂
, 2
♀
, 2 sex?, Ôu a, Ma suzaka-shi, Ehime-ken (33.996N, 132.771E), 15 IV 2010, Kawakami-Y. (cKaw).
[Kyûshû]: 9 sex?, Wa a i, Fuku su-shi, Fukuoka-ken (33.786N, 130.454E), 21 IV 2012, Yamamo o-S., Ma uyama-
M., Kanao-T. ( om seaweed on small sandy beach wi h Aleocha a (Empleno a) seg ega a; KUM); 10 sex?,
Wa a i, Fuku su-shi, Fukuoka-ken (33.790N, 130.447E), 21 IV 2012, Yamamo o-S., Ma uyama-M., Kanao-T.
( om seaweed on huge sandy beach wi h A. (E.) seg ega a; KUM); 2 sex?, Mi oma, Higashi-ku, Fukuoka-shi,
Fukuoka-ken (33.703N, 130.417E), 3 VI 2001, Oga a-S. (KUM); 13
♂
, 23 sex?, Shimomaga-hama, Ka suma,
Higashi-ku, Fukuoka-shi, Fukuoka-ken (33.684N, 130.290E), 15 V 2011 (11:30), Yamamo o-S. (seaweed on sandy
beach wi h A. (E.) seg ega a; cYam); 113 sex?, same da a, bu 21 III 2012; 2 sex?, Noko (Nokono-shima), Nishi-
ku, Fukuoka-shi, Fukuoka-ken (33.608N, 130.299E), 20 III 2009, Yamamo o-S. ( om a seaweed mass on sandy
beach wi h A. (E.) seg ega a and A. (T.) ze chei; cYam); 1 sex?, Take o-kaigan, Kayaki-machi, Nagasaki-shi,
Nagasaki-ken (32.638N, 129.799E), 29 III 1992, Kusui-Y. (cI ô); 2 sex?, Kin (Tsu-shima), Kami sushima-chô,
Tsushima-shi, Nagasaki-ken (34.568N, 129.469E), 5 V 2009, Yamamo o-S. ( om decaying seaweed on sandy
beach wi h A. (E.) ucicola; cYam); 1
♂
, 3
♀
, 8 sex?, Higashimochida, Ai a-chô, Ai a-gun, Kagoshima-ken
(31.724N, 130.640E*), 22 V 1984, Ôha a-M. (KUM). [Ryûkyû]: 1
♂
, 8
♀
, 5 sex?, Naka ane-chô (Tanega-shima),
Kumage-gun, Kagoshima-ken (30.515N, 130.980E), 8-13 V 1996, Ma uyama-M. (cMa ); 1
♀
, Akaogi (Amami-
Ôshima), Ta sugou-chô, Ôshima-gun, Kagoshima-ken (28.415N, 129.627E), 25 III 1978, Naomi-S.I. (KUM); 2
sex?, Kame su (Tokuno-shima), Tokunoshima-chô, Ôshima-gun, Kagoshima-ken (27.716N, 129.018E*), 24 IV
1954, Kuma a-T. (SCM); 2 sex?, China-chô (Okinoe abu-jima), Ôshima-gun, Kagoshima-ken (27.396N,
128.560E*), 24 III 1966, I ô-T. (cHayas).
O he specimens. [JAPAN]: 1 sex?, Japan/Mus. Ge m. [HW]// isulca a We. [HW]/de . Be nhaue // Chicago
NHMus/M. Be nhaue /Collec ion. [FMNH]. [CHINA, Hong Kong]: 1 male, Tai Long Wan, Lan au Island
(22.220N, 113.884E*), 30 III 1997, de Rougemon G. M. (BMNH); 1 sex?, Hong Kong/Waeken Coll/93-58 [HW]/
/China/B i . Mus. [HW]// isulca a/Wse. [HW]//Chicago NHMus/M. Be nhaue /Collec ion. [FMNH].
Redesc ip ion. Body (Fig. 53): medium o somewha la ge, no mally medium sized; ex emely obus and
hea ily scle o ized; e y na owly subpa allel sided; en i e body shining, bu punc u ed a eas o o ebody,
especially ely a, ma . Colou (Figs. 53–56): gland colou black o blackish b own wi h some imes ligh e colou in
ely a; legs, especially a sal segmen s, b own o eddish b own; maxilla y and labial palpi eddish b own o
b ownish b own; an ennae da k b own and pa ly eddish b own. Head (Fig. 53): longi udinal deep u ows on
each side o midline, connec ed by deep ans e se sulci a base. An ennae (Fig. 65): hick and qui e obus ; clea ly
sho e han combined leng h o head and p ono um; segmen I, nea ly 2.2 imes as long as b oad; segmen II
Zoo axa 3517 © 2012 Magnolia P ess · 33
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
p ominen ly sho e han I; segmen III p ominen ly sho e han II; segmen IV no mally sphe ical in ce ain angle;
segmen s V o X s ongly ans e se; segmen XI, abou 1.3 imes as long as b oad; ela i e leng h (wid h) o
segmen s om basal o apical: 10(4.5): 6.5(3): 4.5(3): 2(3): 2(4): 2(4): 2(4): 2(4.5): 2(4.5): 2(4.5): 6(4.5). Tho ax:
p ono um (Figs. 53–54) sligh ly wide han long (PW/PL =1.19), a li le b oade han head (PW/HW =1.26);
su ace blackishly shining wi hou hexagonal e icula ions and shallow dis inc punc u es, bu some deep-dis inc
punc u es and p ominen longi udinal h ee u ows along midline o ming h ee-dimensional pa e ns;
impunc u ed a eas la gely simila o hal -moon shape (wi hou excep ion, and ne e cu in o comple ely sepa a ed
pieces; yellow colou ed: Fig. 54), su ace o hal -moon shining, and loca ed a each side o midline; pa ly co e ed
wi h sho and hick b own-se ae along u ows and deep punc u es (whi e line: Fig. 54); u ows o each side o
midline sligh ly ex ended owa d an e io (blue line: Fig. 54). In e coxal p ocess o meso en i e (Figs. 55–56)
ex emely sha p. Su ace o meso en i e (Figs. 55–56) ough. In e coxal p ocess o me a en i e (Fig. 55)
iangula and poin ed abo e. Legs (Fig. 70): hind ibia qui e sho , abou 0.7 imes as long as ely a (measu ed
along midline); ela i e leng hs o a some es om basal o apical: 4: 3: 3: 3: 6 in o e a sus, 6: 4: 4: 4: 8 in
mid a sus, 8: 5: 5: 5: 10 in hind a sus. Abdomen: pos e io ma gin o e gi e VIII (Figs. 71–72), wi h a ow o hick
and sho se e al senso y se ae.
[Male]: pos e io ma gin o e gi e VIII (Fig. 71) nea ly unca e, wi h a ound 8 mac ose ae. S e ni e VIII (Fig.
73) wi h abou 4 mac ose ae and a ound 5 hin mac ose ae; pos e io ma gin p oduced weakly and medially.
Median lobe o aedeagus (Figs. 75–76) compac ly elonga ed, mode a ely na owed apically, elonga ed py i o m in
en al iew (Fig. 76); a pai o ci cula subapico- en al p ojec ions in la e al iew (Fig. 75); apical lobe o
median lobe ex emely sho -isosceles shape in en al iew (Fig. 76); lagellum sho e han he whole leng h o
median lobe (Figs. 75–76).
[Female]: pos e io ma gin o e gi e VIII (Fig. 72) gen ly ounded o almos unca e, wi h a ound 7
mac ose ae. S e ni e VIII (Fig. 74) wi h 4 la ge mac ose ae and a ound 3 hinne mac ose ae; pos e io ma gin
mode a ely poin ed and no big di e ences in compa ed wi h male. Spe ma heca (Fig. 78): spe ma hecal head la ge,
nea ly wice longe han spe ma hecal s em (sa); spe ma hecal neck clea ly sho e han (sa); basal po ion o
spe ma hecal s em dis o ed, wi h some ben s.
Measu emen s (male: n=10): BL, 3.12–3.91 (3.62±0.24); FBL, 1.68–1.98 (1.84±0.09); HL, 0.45–0.62
(0.52±0.05); HW, 0.54–0.69 (0.63±0.04); AL, 0.74–0.91 (0.81±0.06); PL, 0.58–0.74 (0.68±0.05); PW, 0.68–0.86
(0.79±0.05); EL, 0.57–0.68 (0.62±0.03); EW, 0.82–0.99 (0.91±0.05); HTL, 0.37–0.54 (0.47±0.05).
Measu emen s ( emale: n=10): BL, 2.84–4.01 (3.34±0.42); FBL, 1.29–1.81 (1.62±0.17); HL, 0.38–0.51
(0.46±0.04); HW, 0.46–0.61 (0.55±0.05); AL, 0.59–0.83 (0.72±0.07); PL, 0.42–0.64 (0.58±0.07); PW, 0.55–0.79
(0.70±0.08); EL, 0.47–0.66 (0.58±0.06); EW, 0.66–0.89 (0.80±0.08); HTL, 0.27–0.48 (0.41±0.05).
Con i med dis ibu ion by au ho s. [JAPAN]: Honshû, Shikoku, Kyûshû, Ryûkyû, Sadoga-shima, Tsu-
shima, Tanega-shima, Amami-Ôshima, Tokuno-shima, Okinoe abu-jima (See, Fig. 104 o Japanese dis ibu ion);
[CHINA]: Hong Kong.
O he eco ds in li e a u e. [SOUTH KOREA]: Chungnam P o ince, Jeonnam P o ince, Jeju P o ince
(Pa k & Ahn, 2004); [RUSSIA]: Fa Eas (Ahn e al., 2000; no o iginal ci a ion; doub ul eco d).
Diagnosis. Aleocha a isulca a can be easily dis inguished om he o he T iocha a species by a combina ion
o he ollowing cha ac e s a es: do sal su ace o head and p ono um shining; longi udinal subpa allel u ows on
head connec ed wi h a deep u ow a basal head (no sha ed cha ac e in A. nubis); impunc u ed a eas on p ono um
simila o hal -moon shape (ne e cu in o comple ely sepa a ed pieces) and loca ed a each side o midline (de ails
in Fig. 54); u ows on p ono um deep and p ominen , ne e cu in o pieces o wa y (Fig. 54); pos e io ma gin o
e gi e VIII (Figs. 71–72) wi h a ow o hick senso y se ae (dis inguishable cha ac e om A. ze chei and A. nubis).
[Male]: a pai o subapico- en al p ojec ions on median lobe o aedeagus ci cula in la e al iew (Fig. 75).
[Female]: spe ma hecal s em (bs) (Fig. 78) mode a ely cu ed wi h some ben s which is making somewha simila
o M-shape.
Rema ks. This species is qui e simila o A. ze chei, so ha econside a ion o he eco ds in he li e a u e om
Japan and adjacen egions is needed. Fo example, Ahn e al. (2000) eco ded A. isulca a wi h a sho
edesc ip ion and illus a ions o geni alia o bo h sexes, howe e , his u ned ou o be a misiden i ica ion o A.
ze chei by Pa k and Ahn (2004). In he la e pape , he p esence o sho longi udinal ca ina on e gi e segmen s IV
o V is men ioned as an impo an cha ac e s a e o A. isulca a, bu we could no con i m his cha ac e . They
may ha e misiden i ied he cha ac e s a es o misjudged hem.
YAMAMOTO & MARUYAMA
34 · Zoo axa 3517 © 2012
Magnolia P ess
FIGURES 71–78. Te minalia o Aleocha a (T iocha a) isulca a. 71. e gi e VIII o male; 72. e gi e VIII o emale; 73.
s e ni e VIII o male; 74. s e ni e VIII o emale; 75. male geni alia: median lobe o aedeagus in la e al iew; 76. male geni alia:
median lobe o aedeagus in en al iew; 77. apical lobe o pa ame i e, la e al iew; 78. emale geni alia: spe ma heca.
Old eco ds in Hokkaidô and he sou he n Ku il Islands a e possibly misiden i ica ion o Aleocha a
(T iocha a) ze chei and A. (T.) nubis. We did no con i m any specimen o ue A. isulca a in such egions.
Zoo axa 3517 © 2012 Magnolia P ess · 35
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
Yamazaki (2008) eco ded Fucellia apicalis Ke ész, 1908 (Dip e a, An homyiidae) as one o he hos species
o A. (T.) isulca a and A. (E.) ucicola. In o ma ion on he pa asi ized mon hs (Ap il, May) is p esen ed in
Yamazaki (2012). Pupa ion akes place in he pupa ium o i s hos ly (Yamazaki, 2008, 2012).
Aleocha a (T iocha a) ze chei (Assing, 1995)
(Figs. 57–60, 79–85, 105)
T iocha a ze chei Assing, 1995: 230 (diagnos ic key o species o T iocha a), 231 (o iginal desc ip ion; ype locali y:
“RUSSIA: P imo skiy K ay, Ryazano ka, 14km SW Sla yanka, 42.48 N 131.12 E”); Pa
ś
nik, 2001: 232 ( eco d om
No h Ko ea).
Aleocha a (T iocha a) ze chei (Assing, 1995); Maus & Ashe, 1998c (online) (wo ld checklis o subgenus); Ma uyama, 2002:
18 ( eco d om Japan (Hokkaidô)); Pa k & Ahn, 2004: 196 ( eco d om Sou h Ko ea); Sme ana, 2004: 358 (ca alogue o
Palaea c ic species o Aleocha inae); F ank & Ahn, 2011: 10 ( igu e o li ing indi idual), 20 (checklis o coas al
S aphylinidae o wo ld).
“Aleocha a (T iocha a) isulca a (Weise, 1877)”; Ahn e al., 2000: 245 ( edesc ip ion; as A. isulca a; misiden i ica ion o A.
ze chei, co ec ed by Pa k & Ahn (2004)).
Type specimens. No examined.
Non- ype specimens. JAPAN: [Hokkaidô]: 1
♀
, 1 sex?, Yanbe su (nea Niku u-numa), Koshimizu-chô,
Sha i-gun (43.918N, 144.538E), 27 VII 1993, Haga-K. (unde seaweed on sandy beach; KUM); 4 sex?,
Hamakoshimizu, Koshimizu-chô, Sha i-gun (43.935N, 144.450E), 3 X 1996, Ka ô-T. (KUM); 1
♂
, Moko o (nea
Moko o-eki ailway s a ion), Abashi i-shi (43.970N, 144.322E), 2 VII 1994, Haga-K. (unde seaweed; KUM); 1
sex?, Abashi i, Abashi i-shi (43.979N, 144.304E*); 24 VII 2009, Ôha a-M. (45°58
′
45
″
N, 144°18
′
13
″
E:
mis yping?; unde seaweed (Sa gassum sp.), eel g ass (Zos e a sp.); HUM: HK-09-MO-056/SA); 2 sex?, Koi oi,
Shi anuka-chô, Shi anuka-gun (42.982N, 144.163E), 19 VIII 2005, Kishimo o-K. (KUM); 1 sex?, Ko an, A su a-
ku, Ishika i-shi (43.355N, 141.427E), 9 VII 2009, Ôha a-M. (43°21
′
18
″
N, 141°25
′
37
″
E; unde seaweed on sandy
beach; HUM: HK-09-MO-029/SA); 1
♂
, 2
♀
, Zenibako, O a u-shi (43.144N, 141.161E), 30 V 1977, Nishikawa-
N. (KUM); 1 sex?, Mina o-machi, Rankoshi-chô, Isoya-gun (42.880N, 140.364E), 23 IV 1999, Sasaki-K. (KUM);
1 sex?, Kojô-hama, Shi aoi-chô, Shi aoi-gun (42.462N, 141.218E), 30 VII 1953, Shu o-S. (SCM); 1 sex?, Nanae-
hama, Nanaehama, Hoku o-shi (41.815N, 140.702E), 15 VIII 1977, Nishikawa-N. (KUM). [Honshû]: 1 sex?,
Kodoma iaoiwa, Nakadoma i-machi, Ki a suga u-gun, Aomo i-ken (41.165N, 140.329E*), 15 IV 2004, Abe-A.
(KUM); 1 sex?, Sichi inaga-hama (nea Benzu-numa), Kizuku i a eoka, Tsuga u-shi, Aomo i-ken (40.867N,
140.286E), 1 VII 2006, Hayashi-M. (sandy beach; cKaw); 1 sex?, Rikuchû-Yagi, Taneichi, Hi ono-chô, Kunohe-
gun, Iwa e-ken (40.349N, 141.764E), 28 VI 2010, Ôha a-M. (40°20
′
56
″
N, 141°45
′
49
″
E; sandy beach; unde
seaweed (Lamina ia sp., Sa gassum sp.) and eel g ass (Zos e a sp.); HUM: HN-10-MO-044/SA); 1 sex?, Raga,
Tanoha a-mu a, Shimohei-gun, Iwa e-ken (39.939N, 141.940E*), 30 V 1998, Wa anabe-T. (cWa ); 1 sex?, Omo o,
Iwaizumi-chô, Shimohei-gun, Iwa e-ken (39.850N, 141.974E), 27 VI 2010, Ôha a-M. (39°50
′
59
″
N, 141°58
′
27
″
E;
sandy beach; HUM: HN-10-MO-042/SA); 2 sex?, Idou a bay, Fujizuka, Wakabayashi-ku, Sendai-shi, Miyagi-ken
(38.181N, 140.962E), 8 VIII 2009, Ozaki-T. (KUM); 5 sex?, Isoha a, Ki aiba aki-shi, Iba aki-ken (36.790N,
140.752E), 24 V 1998, Ohmomo-S. (KUM); 1 sex?, Heba a, Ka uu a-shi, Chiba-ken (35.165N, 140.334E), 12 V
1991, Takeda-T. (KUM); 1
♂
, 1
♀
, 1 sex?, Uba a, Ka suu a-shi, Chiba-ken (35.137N, 140.275E), 6 IV 1993,
Takeda-T. (KUM); 1 sex?, Ama su-kaigan, Ama su, Kamogawa-shi, Chiba-ken (35.122N, 140.154E), 3 XI 1990,
Takeda-T. (KUM); 1 sex?, Tôjô-kaigan (in on o an aqua ium: Kamogawa Seawo ld), Higashi-chô, Kamogawa-
shi, Chiba-ken (35.115N, 140.121E), 27 IX 1987, Haga-K. (unde lo sam on sandy beach; KUM); 1 sex?,
Heisau a, Sakai, Ta eyama-shi, Chiba-ken (34.948N, 139.801E), 1 IV 1997, Ma uyama-M. (cMa ); 1
♂
, 1
♀
, 20
sex?, same place, bu 26 XII 2007, Ono-H. (cOno); 4 sex?, Miu a-shi, Kanagawa-ken (35.180N, 139.655E*), 26 III
1966, Ki yû-R. (SCM); 1 sex?, Ku ihama, Yokosuka-shi, Kanagawa-ken (35.225N, 139.713E*), 25 IV 1965,
Ki yû-R. (SCM); 1
♀
, Naga-hama swimming beach, Nagai-2chôme 13, Yokosuka-shi, Kanagawa-Ken (35.190N,
139.615E), 7 IV 1975, Tao-M. (KUM); 8 sex?, Hi a suka beach, Hi a suka-shi, Kanagawa-ken (35.315N,
139.355E), 28 IX 1958, Siba a-Y. (SCM); 1
♂
, Shônan-kaigan, Kanagawa-Ken (35.315N, 139.351E*), 2 VII 1976,
Niiza o (KUM); 6 sex?, Tsumekizaki, Shimoda-shi, Shizuoka-ken (34.659N, 138.987E), 21 IV 2003, Ma uyama-
M. (cMa ); 2 sex?, Mukaiawagasaki, Uchinada-machi, Kahoku-gun, Ishikawa-ken (36.640N, 136.622E*), 12 IV
1948, Takaba-S. (cHayas); 1
♂
, 3 sex?, Uchinada-machi, Kahoku-gun, Ishikawa-ken (36.656N, 136.636E*), 15 VI
YAMAMOTO & MARUYAMA
36 · Zoo axa 3517 © 2012
Magnolia P ess
1961, Hayashi-Y. (cHayas); 1
♂
, 1 sex?, Kanaiwa, Kanazawa-shi, Ishikawa-ken (36.607N, 136.589E), 15 VII
1961, Hayashi-Y. (cHayas); 27 sex?, Echizen-kaigan, Fukui-ken (36.036N, 136.012E*), 17 IX 2001, Hoshina-H.
(sandy beach; unde wood logs; KUM); 1 sex?, i e mou h o Kushida-gawa, Ma sunase-chô, Ma suzaka-shi,
Mie-ken (34.605N, 136.582E), 28 IV 2002, Kawakami-Y. (cKaw); 2 sex?, Nagusano-hama, Kemi, Wakayama-shi,
Wakayama-ken (34.165N, 135.182E), 14 V 2002, Kawakami-Y. (cKaw); 1 sex?, Kada-kaigan, Kada, Wakayama-
shi, Wakayama-ken (34.280N, 135.074E), 19 IV 2003, Kawakami-Y. (cHayas); 1
♀
, Takashino-hama, Takaishi-
shi, Ôsaka- u (34.531N, 135.432E), 4 IX 1958, Kimu a-Y. (cHayas); 1
♂
, 2
♀
, 4 sex?, Hamade a, Nishi-ku, Sakai-
shi, Ôsaka- u (34.543N, 135.439E), 5 V 1965, I ô-T. (cHayas); 1
♀
, 1 sex?, sho es o Sakai, Ôsaka- u (34.582N,
135.465E*), 20 VI 1956, Sawada-K. (cHayas); 1
♀
, 12 sex?, Mina o-ku, Ôsaka-shi, Ôsaka- u (34.667N,
135.447E), 7 IV 1957, Shiba a-T. (cHayas); 1
♂
, 7 sex?, Nishinomiya, Hyôgo-ken (34.724N, 135.331E), 14 IV
1957, Shiba a-T. (cHayas); 2 sex?, Kôshien-hama, Kôshienhama-1chôme, Nishinomiya-shi, Hyôgo-ken (34.712N,
135.349E), 27 V 1998, Kawakami-Y. (cKaw); 1 sex?, same da a, bu 29 IV 2003; 2 sex?, Sumau a-kaigan,
Sumau adô i, Suma-ku, Kôbe-shi, Hyôgo-ken (34.637N, 135.103E), 19 IV 1998, Kawakami-Y. (cKaw); 1
♂
, same
da a, bu 27 V 2010; 1 sex?, Gunge-hama (Awaji-shima), Gunge, Awaji-shi, Hyôgo-ken (34.475N, 134.845E), 5 V
2000, Kawakami-Y. (cKaw); 1 sex?, Ma u ahama coas (Awaji-shima), Amanishi-machi, Minamiawaji-shi,
Hyôgo-ken (34.201N, 134.732E), 13 V 1998, Kawakami-Y. (cKaw); 1 sex?, Ma ue-kaigan, Ma sue, Akashi-shi,
Hyôgo-ken (34.654N, 134.955E), 10 V 1998, Kawakami-Y. (cKaw); 3 sex?, Yagi, Ôkubo-chô, Akashi-shi, Hyôgo-
ken (34.670N, 134.933E), 9 V 1999, Kawakami-Y. (cKaw); 4 sex?, i e mou h o Kako-gawa, You a, Onoe-chô,
Kakogawa-shi, Hyôgo-ken (34.737N, 134.811E), 1 VI 1999, Kawakami-Y. (cKaw); 4
♂
, 1
♀
, Ma oga a-kaigan,
Ma oga a, Ma oga a-chô, Himeji-shi, Hyôgo-ken (34.769N, 134.746E), 16 V 2010, Kawakami-Y. (cKaw); 2 sex?,
Kugui, Higashi-ku, Okayama-shi, Okayama-ken (34.584N, 134.086E*), 31 III 2003, Fuji ani-Y. (KUM); 13 sex?,
same da a, bu 9 V 2003 (unde seaweed on beach); 13 sex?, same da a, bu 28 V 2003; 2
♂
, 2
♀
, 1 sex?, Kogushi,
Minami-ku, Okayama-shi, Okayama-ken (34.591N, 134.037E), 11 III 1977, Wa anabe-A. (KUM); 1
♂
, Inazumi,
Sugeu a, Mihonoseki-chô, Ma ue-shi, Shimane-ken (35.564N, 133.166E*), 1-2 VI 2008, Hayama-T. (sandy beach;
cHayam); 4 sex?, So ozono-kaigan, Sashiumi, Ko yô-chô, Izumo-shi, Shimane-ken (35.333N, 132.664E), 12-13
IV 2009, Hayama-T. (unde seaweed; cHayam); 1 sex?, Ku oma su-chô, Gô su-shi, Shimane-ken (132.310N,
35.057E), 2 V 2009, Hayama-T. ( om seaweed; cHayam); 11 sex?, Kiami-chô, Masuda-shi, Shimane-ken
(34.676N, 131.750E*), 15-16 VI 2008, Hayama-T. (FIT on sup a idal zone; cHayam); 1 sex?, same locali y, bu 10
V 2009, Hayama-T. (cHayam); 1 sex?, Chûdô coas , Aiohigashi, Yamaguchi-shi, Yamaguchi-ken (33.996N,
131.450E), 4 V 2003, Kawakami-Y. (cKaw). [Shikoku]: 13 sex?, Tosadoma iu a (Ôge-jima), Na u o-chô, Na u o-
shi, Tokushima-ken (34.232N, 134.639E), 29 IX 1972, Tomokuni-M. and Sakai-M. (EEEU); 1 sex?, i e mou h o
Yoshino-gawa (be ween Nada-bashi b idge and Daijû-zeki sluice), Tokushima-shi, Tokushima-ken (34.089N,
134.561E), 3 V 1999, Kawakami-Y. (cKaw); 3 sex?, es ua y o Yoshino-gawa, Tokushima-shi, Tokushima-ken
(34.074N, 134.595E*), 15 V 2007, Changdo-H. (EEEU); 1 sex?, Ôu a, Ma suyama-shi, Ehime-ken (33.996N,
132.771E), 15 IV 2010, Kawakami-Y. (cKaw); 1 sex?, Takahama-machi, Ma suyama-shi, Ehime-ken (33.898N,
132.709E*), 14 V 2002, Ku iha a-T. (sandy beach; EEEU); 1 sex?, Mo i, Iyo-shi, Ehime-ken (33.740N,
132.678E), 20 III 2001, Sakai-M. (EEEU). [Kyûshû]: 10 sex?, Noko (Nokono-shima), Nishi-ku, Fukuoka-shi,
Fukuoka-ken (33.608N, 130.299E*), 13 V 1998, Fujimo o-H. (KUM); 1 sex?, same locali y, bu 4 V 2008, Ôkawa-
A. (KUM); 1 sex?, same place name (33.608N, 130.299E), bu 20 III 2009, Yamamo o-S. ( om a seaweed mass on
sandy beach wi h Aleocha a (Empleno a) seg ega a and A. (T.) isulca a; cYam).
O he specimens. JAPAN: 2 sex? (specimens on he same pape ca d), Japan./G Lewis./1910-320.//
Aleocha a/ isulca a/Weise [HW]//Chicago Na . His . Mus. (ex. D. Sha p Colln./by exchange wi h/B i . Mus. Na .
His .) [FMNH]; 1 sex?, Japan/mus./ge m. [HW]// isulca a Ws [HW]/de . Be nhaue //Chicago NHMus/M.
Be nhaue /Collec ion. [FMNH]; 1 sex?, isulca a/Wse. Japan/S andinge [HW]//Chicago NHMus/M. Be nhaue /
Collec ion. [FMNH]; 1 sex?, aleocha a/ isulca a Wse/Japan./V. S andinge [HW]//Chicago NHMus/M. Be nhaue /
Collec ion. [FMNH]; 3 sex?, BEPPU/NO.-KIUSHIU [Beppu-shi, Ôi a-ken, Kyûshû]//Japan/Rei e .//Chicago
NHMus/M. Be nhaue /Collec ion. [FMNH]; 1 sex?, BEPPU/NE-KYUSHU [Beppu-shi, Ôi a-ken, Kyûshû]//Japan/
Rei e // isulca a Ws. [HW]/de . Be nhau//Chicago NHMus/M. Be nhaue /Colle ion. [FMNH]; 2 sex?, BEPPU/
NE-KYUSHU//Japan/Rei e .//Chicago NHMu/M. Be naue /Collec ion. [FMNH]. RUSSIA: Ku il Islands:
[U up Is.]: 1
♂
, 3
♀
, 5 sex?, nea i e mou h o Bys aya, No o-Ku ylisk bay (46.223N, 150.319E), 8 VIII 1995,
Ôha a-M. (46°12
′
84
″
N, 150°18
′
69
″
E; hand pick up; eces o cows; IKIP; HUM: UR-95-MO-8); 1
♂
, O k y yy bay
(45.864N, 149.793E), 4 VIII 1995, Ôha a-M. (N45°51
′
49
″
, E149°46
′
95
″
; hand pick up; unde logs and ocks on
Zoo axa 3517 © 2012 Magnolia P ess · 37
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
sho e; HUM: UR-95-MO-002); 1
♂
, O k y yy bay (45.851N, 149.770E), 5 VIII 1995, Ôha a-M. (45°51
′
04
″
N,
149°46
′
12
″
E; unde logs and ocks on sho e; HUM: UR-95-MO-005).
FIGURES 79–85. Te minalia o Aleocha a (T iocha a) ze chei. 79. e gi e VIII o male; 80. e gi e VIII o emale; 81. s e ni e
VIII o male; 82. s e ni e VIII o emale; 83. male geni alia: median lobe o aedeagus in la e al iew; 84. male geni alia: median
lobe o aedeagus in en al iew; 85. emale geni alia: spe ma heca.
YAMAMOTO & MARUYAMA
38 · Zoo axa 3517 © 2012
Magnolia P ess
Redesc ip ion. Body (Fig. 57): small o somewha la ge, no mally medium sized; s ongly scle o ized;
na owly subpa allel sided bu mode a ely hick; en i e body shining somewha s ongly, bu some indi iduals
shining weakly due o sc a ches o ough su ace en i ely. Colou (Figs. 57–60): gland colou ligh black o g ayish
b own wi h almos same (some imes ligh e ) colou in ely a; legs, especially a sal segmen s, ligh b own o
eddish b own: maxilla y and labial palpi eddish b own o b ownish b own; an ennae da k b own o eddish
b own. Head (Fig. 57): longi udinal deep u ows on each side o midline, connec ed by somewha deep ans e se
imp ession (o u ow) a base. An ennae: mode a ely hick and qui e obus ; clea ly sho e han combined leng h
o head and p ono um; segmen I, abou 1.9 imes as long as b oad; segmen II clea ly sho e han I; segmen III
p ominen ly sho e han II; segmen IV sphe ically ans e se; segmen V o X s ongly ans e se; segmen XI,
abou 1.5 imes as long as b oad; ela i e leng h (wid h) o segmen s om basal o apical: 9.5(5): 6(4): 5.5(3.5):
3(3.5): 3(4): 3(5): 3(5): 3(5.5): 3(5.5): 3.5(6): 7.5(5). Tho ax: p ono um (Figs. 57–58) sligh ly wide han long
(PW/PL =1.17), a li le b oade han head (PW/HW =1.32); su ace blackishly shining in mos cases, bu
some imes nea ly ma because o nume ous sc a ches o many la ge punc u es, o oo many undula ions making
ough su ace in somecases; do sal su ace lack o hexagonal e icula ions as well as shallow dis inc punc u es, bu
some deep-dis inc punc u es and shallow-wa ing u ows o ming h ee-dimensional pa e ns; pa e ns e y
di e se and a ying g ea ly wi hin he species, [Typical ype (see, Fig. 58)]: impunc u ed a ea la gely simila o
hal -moon shape on each side, bu wi h cu ing o by an oblique ow o se ae o punc u es a ound middle o each
hal -moon pa e n, o ming wo pai s o impunc u ed a eas (yellow colou ed: Fig. 58), [O he ypes]: pa e ns
i egula and each impunc u ed a ea small and inconspicuous, o lack o impunc u ed a ea and ha ing ough
su ace en i ely in some cases; longi udinal h ee u ows along midline, some imes weak, wa y and inconspicuous
(blue line: Fig. 58). In e coxal p ocess o meso en i e (Figs. 59–60) elonga ed and subpa allel sided wi h ounded
apex. Su ace o meso en i e (Figs. 59–60) ma and weakly shining. In e coxal p ocess o me a en i e (Fig. 59)
qui e sho wi h ounded an e io ma gin, abou 1/4 as long as mesocoxal ca i y. Legs: mid ibia as long as
me a en i e; hind ibia sho , abou 0.8 imes as long as ely a; ela i e leng hs o a some es om basal o apical:
6: 3: 3: 3: 7 in o e a sus, 7: 4: 4: 4: 10 in mid a sus, 8: 5: 5: 5: 11 in hind a sus. Abdomen: pos e io ma gin o
e gi e VIII (Figs. 79–80), wi h a ow o hin and long se e al senso y se ae.
[Male]: pos e io ma gin o e gi e VIII (Fig. 79) nea ly unca e o wa y, wi h a ound 8 mac ose ae. S e ni e
VIII (Fig. 81) wi h abou 5 la ge mac ose ae and a ound 4 hin mac ose ae; pos e io ma gin p oduced weakly and
medially. Median lobe o aedeagus (Figs. 83–84) compac ly elonga ed, elonga ed py i o m in en al iew (Fig.
84); a pai o subapico- en al p ojec ions sha ply poin ed in la e al iew (Fig. 83); apical lobe o median lobe
ex emely sho -isosceles shape in en al iew (Fig. 84); lagellum nea ly as same leng h as he whole leng h o
median lobe (Figs. 83–84).
[Female]: e gi e VIII (Fig. 80) gen ly ounded owa d pos e io and sligh ly ema gina ed medially, wi h
a ound 3 la ge mac ose ae and a ound 5 hin mac ose ae. S e ni e VIII (Fig. 82) wi h 4 la ge mac ose ae and a ound
3 hinne mac ose ae; pos e io ma gin e y weakly poin ed. Spe ma heca (Fig. 85): spe ma hecal head la ge, abou
wice longe han apical po ion o spe ma hecal s em (sa); spe ma hecal neck inconspicuous, clea ly sho e han
(sa); basal po ion o spe ma hecal s em simple.
Measu emen s (male: n=10): BL, 3.19–3.91 (3.62±0.25); FBL, 1.55–2.00 (1.79±0.14); HL, 0.44–0.58
(0.51±0.04); HW, 0.52–0.67 (0.61±0.05); AL, 0.77–0.92 (0.83±0.05); PL, 0.61–0.78 (0.70±0.06); PW, 0.70–0.89
(0.81±0.07); EL, 0.55–0.78 (0.64±0.07); EW, 0.89–1.02 (0.95±0.05); HTL, 0.44–0.55 (0.51±0.04).
Measu emen s ( emale: n=10): BL, 3.19–4.47 (3.79±0.49); FBL, 1.58–2.09 (1.82±0.18); HL, 0.43–0.57
(0.49±0.05); HW, 0.51–0.67 (0.60±0.05); AL, 0.69–0.83 (0.75±0.05); PL, 0.58–0.75 (0.67±0.05); PW, 0.70–0.89
(0.79±0.06); EL, 0.55–0.73 (0.66±0.06); EW, 0.82–1.09 (0.93±0.09); HTL, 0.43–0.55 (0.49±0.04).
Con i med dis ibu ion by au ho s. [JAPAN]: Hokkaidô, Honshû, Shikoku, Kyûshû, Awaji-shima (See, Fig.
105 o Japanese dis ibu ion); [RUSSIA]: Ku il Islands: U up Is.
O he locali ies in li e a u e. [SOUTH KOREA]: Gangwon (Kangwon) P o ince, No h Gyeongsang
P o ince, Jeju P o ince (Ahn e al., 2000: as A. isulca a; Pa k & Ahn, 2004); [NORTH KOREA]: Chagang
P o ince (Pa
ś
nik, 2001); [RUSSIA]: Fa Eas : Sakhalin, P imo sky K ai (Assing, 1995).
Diagnosis. This species is ex emely simila o he o me species, Aleocha a isulca a, in many poin s
especially in ex e nal appea ance. Howe e , ou s udy e ealed some impo an cha ac e s a es o dis inguish A.
ze chei om o he wo species as ollows: longi udinal subpa allel u ows on head connec ed wi h a somewha
deep imp ession (no by a clea ly sha ow and incospicuous u ow); impunc u ed a ea on p ono um (Fig. 58)
Zoo axa 3517 © 2012 Magnolia P ess · 39
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
la gely simila o a hal -moon shape on each side, bu wi h cu ing o by an oblique ow o se ae a ound middle o
each hal -moon pa e n, o lacking impunc u ed a eas and ha ing ough su ace en i ely; longi udinal sulci on
do sal su ace o p ono um shallow, some imes mo e o less making wa y lines (Fig. 58); pos e io ma gin o
e gi e VIII (Figs. 79–80) lack o a ow o hick senso y se ae bu ins ead ha e hin and long se ae. [Male]:
pos e io ma gin o s e ni e VIII (Fig. 81) weakly poin ed; a pai o subapico- en al p ojec ions on median lobe
na owly and sha ply poin ed in la e al iew (Fig. 83). [Female]: base o spe ma hecal s em (sb) (Fig. 85) e y
simple and no s ongly cu ed.
Rema ks. This species is sympa ically dis ibu ed wi h A. isulca a in mos locali ies in Japan, al hough A.
ze chei was known only om Hokkaidô by he p e ious s udy (Ma uyama, 2002). Ahn e al. (2000) misiden i ied
A. ze chei as A. isulca a and his was la e co ec ed by Pa k and Ahn (2004) as A. ze chei in combina ion wi h
eal eco ds o A. isulca a. Aleocha a ze chei and A. isulca a ha e o en been con used, and e-examina ion o
he ma e ial used in he p e ious s udies is needed.
Aleocha a (T iocha a) nubis (Assing, 1995)
(Figs. 61–64, 86–92, 98–100, 104)
T iocha a nubis Assing, 1995: 230 (diagnos ic key o species o genus), 232 (o iginal desc ip ion; ype locali y: “RUSSIA,
Sakhalin, Ko sako dis ., Ismenhy oye lake”); Naomi e al., 2000: 107 ( eco d om Pa amushi Is., Ku il Islands).
Aleocha a (T iocha a) nubis (Assing, 1995); Maus & Ashe, 1998c (online) (wo ld checklis o subgenus; bionomics);
Ma uyama, 2002: 18 ( eco d om Japan (Hokkaidô)); Sme ana, 2004: 358 (ca alogue o Palaea c ic species o
Aleocha inae); F ank & Ahn, 2011: 20 (checklis o coas al S aphylinidae o wo ld).
Type specimens. No examined.
Non- ype specimens. JAPAN: [Hokkaidô]: 3
♂
, 1
♀
, 4 sex?, Hamama su, Nemu o-shi (43.206N, 145.528E),
14 VI 2010, Yamamo o-S. (unde seaweed on sandy beach; cYam); 2 sex?, Ochiishi, Nemu o-shi (43.607N,
145.286E*), 24 VIII 1999, Ma uyama-M. (unde seaweed on sandy beach; cMa ); 1 sex?, No suke (No suke-zaki
peninsulae), Be sukai-chô, No suke-gun (43.607N, 145.286E), 14 IX 2009, Ôha a-M., Yamamo o-H. and Fu u a-
M. (43°36
′
26
″
N, 145°17
′
08
″
E; unde seaweed (Lamina ia sp.) and eel g ass (Zos e a sp.); HUM: HK-09-MO-100/
SI); 1
♂
, 2
♀
, Shibe su, Nemu o-shi (43.656N, 145.136E*), 18 VII 1977, Naomi-S.I. (KUM); 1 sex?, Koi oi,
Shi anuka-chô, Shi anuka-gun, (42.982N, 144.163E), 19 VIII 2005, Kishimo o-K. (KUM); 1 sex?, Ô sumina o-
machi, Toyoko o-chô, Nakagawa-gun (42.667N, 143.628E), 26 VII 2009, Ôha a-M. (42°40
′
02
″
N, 143°37
′
41
″
E;
unde seaweed (Sa gassum sp.); HUM: HK-09-MO-059/SA); 1 sex?, Ôkishi (nea Ôkishi unnel), Toyou a-chô,
Abu a-gun (42.587N, 140.675E*), 15 IX 1997, Ôha a-M. (unde seaweed on beach; KUM); 1 sex?, Sawa a, Mo i-
machi, Kayabe-gun (42.123N, 140.647E), 15 VII 2009, Ôha a-M. (42°07
′
23
″
N, 140°38
′
50
″
E; unde seaweed
(Sa gassum sp.); HUM: HK-09-MO-040/SA).
O he specimens. RUSSIA: Ku il Islands: [Pa amushi Is.]: 4
♂
, 8
♀
, Med ezhiy Wa e all, Shelekho o
(50.367N-50.378N, 155.611E-155.656E), 18 VII 1997, Sai ô-A. (50°22.012
′
N -50°22.694
′
N, 155°36.677
′
E -
155°39.380
′
E; al . 0-10m; CBM: CBM-ZI 81532(-81543)). [Kha imko an Is.]: 3 sex?, Se e gine bay (49.181N,
154.466E), 8 VIII 1996, Ôha a-M. (49°10
′
51
″
N, 154°27
′
59
″
E; hand picking; unde pebble and logs on sandy
beach; IKIP; HUM: KH-96-MO-022B). [K a e naya Ysnkicha Is.]: 5
♂
, 2
♀
, 25 sex?, a en ance o he island
(47.515N, 152.817E), 14 VIII 1995, Ôha a-M. (47°30
′
54
″
N, 152°49
′
00
″
E; IKIP; HUM: US-95-MO-021).
[Simshi Is.]: 1
♀
, Malaya bay (47.087N, 152.131E), 18 VIII 1995, Ôha a-M. (N47°05
′
14
″
, E152°07
′
51
″
; hand
pick up; unde logs and ocks on sho e; IKIP; HUM: SI-95-MO-032). [Chi upoi Is.]: 1
♂
, 1 sex?, Peschanaya bay
(46.548N, 150.906E), 23 VIII 1995, Ôha a-M. (46°32
′
53
″
N, 150°54
′
22
″
E; unde seaweed on sandy beach; IKIP;
HUM: CH-95-MO-048). [U up Is.]: 1 sex?, inland coas al ma gin o Na alie bay, en isions o Obzhi aya i e , in
small ca e o ock ace nea i e mou h a eas end o co e (46.101N, 150.175E), 7 VIII 1995, U bain-B.K.
(46°05
′
85
″
N, 150°09
′
91
″
E; by hand wi h o ceps; in, on, and unde ocks in and a ound small ca e along coas ; al .
1m; IKIP; HUM: UR-95-BKU-028); 1
♂
, nea mou h o Vese aya i e , Na alii bay (46.094N, 150.142E), 6 VIII
1995, Ôha a-M. (46°05
′
38
″
N, 150°08
′
33
″
E; hand pick up; unde ocks; IKIP; HUM: UR-95-MO-006); 1
♂
, inland
coas al ma gin o O k y yi bay; en i ons o Shabalina i e (45.864N, 149.794E), 4 VIII 1995 (15:00-16:00),
U bain-B.K. (45°51
′
51
″
N, 149°46
′
99
″
E; by hand; along coas al beach wi h ine sand; unde logs; al . 2m; sand
empe a u e: 15 deg ee celcius; IKIP; HUM: UR-95-BKU-014); 1
♂
, 1
♀
, O k y yy bay (45.864N, 149.793E), 4
YAMAMOTO & MARUYAMA
40 · Zoo axa 3517 © 2012
Magnolia P ess
VIII 1995, Ôha a-M. (45°51
′
49
″
N, 149°46
′
95
″
E; unde logs and ocks on sho e; IKIP; HUM: UR-95-MO-002); 1
sex?, Toko an, O k y yy bay (45.864N, 149.793E), 4 VIII 1995, Ôha a-M. (45°51
′
49
″
N, 149°46
′
95
″
E; unde logs
and ocks on sho e; IKIP; HUM: UR-95-MO-001); 1
♂
, 6 sex?, O k y yy bay (45.851N, 149.770E), 5 VIII 1995,
Ôha a-M. (45°51
′
04
″
N, 149°46
′
12
″
E; unde seaweed along seasho e; IKIP; HUM: UR-95-MO-005).
Redesc ip ion. Body (Fig. 61): small o la ge size, no mally somewha la ge; ex emely obus and s ongly
scle o ized; somewha hick bu na owly elonga ed; do sal su ace o en i e body excep o abdomen ma due o
la ge hexagonal mic os uc u es. Colou (Figs. 61–64): gland colou g ayish black o da k g ay wi h almos same
(some imes ligh e ) colou in ely a; legs, especially a sal segmen s, b own o eddish b own: maxilla y and labial
palpi eddish b own o b ownish b own; an ennae da k b own o eddish b own. Head (Fig. 61): longi udinal deep
u ows on each side o midline, connec ed by shallow ans e se imp ession a base; do sal su ace wi h la ge
hexagonal e icula ions. An ennae: ela i ely hick and qui e obus ; sho e han combined leng h o head and
p ono um; segmen I, abou 2.3 imes as long as wid h; segmen II p ominen ly sho e han I, and s ongly and
apically dila ed; segmen III p ominen ly sho e han II; segmen IV sligh ly ans e se; segmen V o X
mode a ely ans e se; segmen XI, abou 1.3 imes as long as b oad; ela i e leng h (wid h) o segmen s om
basal o apical: 9(4): 6(3): 4(3): 2.5(3.5): 2.5(4): 3(4.5): 3(4.5): 3(5): 3(5): 3.5(5): 6.5(5). Tho ax: p ono um (Figs.
61–62) sligh ly wide han long (PW/PL =1.20), a li le b oade han head (PW/HW =1.28); somewha cons ic ed
bo h apical and pos e io ma gin; su ace blackishly, bu weakly shining due o la ge hexagonal mic os uc u es on
en i e su ace and shallow punc u es wi h se ae (connec ed wi h whi e line: Fig. 62); shallow punc u es and u ows
o ming h ee-dimensional pa e n; pa e ns di e se and a ying wi hin he species, [Typical ype (see, Fig. 62)]:
each o impunc u ed a ea la gely simila o hal -moon shape and ele a ed like dooms (yellow colou ed: Fig. 62)
and lack o cu ing o by a ow o se ae a ound middle o each hal -moon pa e n, [O he ype]: pa e ns i egula
(no hal -moon shape as in Fig. 62) and sligh ly ele a ed abo e; do sal su ace wi h longi udinal h ee somewha
deep and s aigh u ows along midline (blue line: Fig. 62). In e coxal p ocess o meso en i e (Fig. 63) elonga ed
and somewha sha ply poin ed. Su ace o meso en i e (Figs. 63–64) somewha ough and shining weakly. In e
coxal p ocess o me a en i e (Fig. 63) sho wi h ounded an e io ma gin. Legs: hind ibia sho , abou 0.9 imes as
long as ely a (measu ed along midline); ela i e leng hs o a some es om basal o apical: 6: 4: 4: 4: 8 in
o e a sus, 8: 5: 5: 5: 10 in mid a sus, 10: 6: 6: 6: 12 in hind a sus. Abdomen: pos e io ma gin o e gi e VIII
(Figs. 86–87), wi h a ow o hin and long se e al senso y se ae; su ace o e gi e VIII and s e ni e VIII pubescen .
[Male]: pos e io ma gin o e gi e VIII (Fig. 86) almos unca e o sligh ly ema gina ed medially, wi h a ound
11 mac ose ae. S e ni e VIII (Fig. 88) wi h abou 5 mac ose ae and a ound 8 hin mac ose ae; pos e io ma gin
poin ed iangula ly. Median lobe o aedeagus (Figs. 90–91) elonga ed, mode a ely na owed apically, elonga ed
py i o m in en al iew (Fig. 91); a pai o poin ed la ge and o al subapico- en al p ojec ions in la e al iew (Fig.
90); apical lobe o median lobe isosceles shape in en al iew (Fig. 91); lagellum sligh ly sho e han he whole
leng h o median lobe (Figs. 90–91).
[Female]: pos e io ma gin o e gi e VIII (Fig. 87) nea ly unca e, wi h a ound 3 mac ose ae and a ound 5
hin mac ose ae. Pos e io ma gin o s e ni e VIII weakly poin ed (Fig. 89) wi h 4 la ge mac ose ae and a ound 4
hinne mac ose ae. Spe ma heca (Fig. 92): head (sh) hick and la ge, mo e han wice longe han apical po ion o
spe ma hecal s em; spe ma hecal neck ex emely sho ; basal po ion o spe ma hecal s em na owing owa d base,
wi h s ong ben (<90°).
Measu emen s (male: n=10): BL, 3.72–4.95 (4.39±0.45); FBL, 1.82–2.23 (2.04±0.14); HL, 0.53–0.65
(0.61±0.04); HW, 0.61–0.74 (0.68±0.04); AL, 0.79–1.07 (0.96±0.09); PL, 0.64–0.81 (0.74±0.06); PW, 0.77–0.97
(0.89±0.07); EL, 0.66–0.80 (0.72±0.05); EW, 0.89–1.13 (1.03±0.08); HTL, 0.49–0.65 (0.59±0.05).
Measu emen s ( emale: n=10): BL, 3.34–5.00 (4.12±0.50); FBL, 1.86–2.24 (2.05±0.11); HL, 0.54–0.72
(0.61±0.05); HW, 0.63–0.79 (0.70±0.04); AL, 0.82–1.11 (0.93±0.09); PL, 0.62–0.87 (0.73±0.07); PW, 0.78–1.01
(0.88±0.06); EL, 0.62–0.80 (0.71±0.05); EW, 0.91–1.23 (1.03±0.09); HTL, 0.49–0.70 (0.57±0.06).
Con i med dis ibu ion by au ho s. [JAPAN]: Hokkaidô (see, Fig. 104); [RUSSIA]: Ku il Islands:
Pa amushi Is., Kha imko an Is., K a e naya Ysnkicha Is., Simshi Is., Chi upoi Is., U up Is.
O he locali ies in li e a u e. [RUSSIA]: Fa Eas : Sakhalin, Kamcha ka Peninsula (Assing, 1995).
Diagnosis. Aleocha a (T iocha a) nubis is simila o A. ze chei in many poin s (e.g., pos e io ma gin o e gi e
VIII in bo h sexes wi hou a ow o hick senso y se ae). Al hough hey sha e many cha ac e is ics, we ound ou
some dis inc ions: dis ibu ion es ic ed o no he n a ea (Hokkaidô, Japan (see, Fig. 104); Russian Fa Eas ; No h
Ko ea); usually la ge species, occasionally small; do sal su ace o o ebody ma and no s ongly shining;
Zoo axa 3517 © 2012 Magnolia P ess · 41
REVISION OF JAPANESE EMPLENOTA AND TRIOCHARA
longi udinal subpa allel u ows on head connec ed wi h a shallow imp ession (no by a deep u ow); impunc u ed
a eas (o lacking o ha a eas) on p ono um (Fig. 62) g ea ly a ying wi hin species, bu p ominen mic os uc u es
and hexagonal e icula ions on do sal su ace; longi udinal sulci on do sal su ace o p ono um (Fig. 62) deep,
no mally s aigh lines; pos e io ma gin o e gi e VIII (Figs. 86–87) wi h a ow o hin and long senso y se ae as
in A. ze chei; su ace o bo h e gi e VIII and s e ni e VIII (Figs. 86–89) pubescen . [Male]: pos e io ma gin o
s e ni e VIII (Fig. 88) poin ed iangula ly; subapico- en al p ojec ions on median lobe o aedeagus la ge and
o al, poin ed apical-la e ally in la e al aspec (Fig. 90). [Female]: base o (sb) e y s ongly cu ed (Fig. 92).
Rema ks. Assing (1995) desc ibed his halophilous species om Sakhalin and Kamcha ka Peninsula, Russia,
and Ma uyama (2002) la e eco ded i o he i s ime om Japan (Hokkaidô). Naomi e al. (2000) epo ed A.
nubis om he Ku il Islands, bu only om Pa amushi Island. In his s udy, we p esen se e al new dis ibu ional
eco ds o he Ku il Islands. Ou da a sugges a wide anging dis ibu ion o A. nubis h oughou Fa Eas Russia
o Hokkaidô. These eco ds also show ha A. nubis is dis ibu ed only in he suba c ic zone in Eas Asia. No
specimen has been ound om Honshû, Kyûshû, o Ryûkyû in Japan (see, Fig. 104). Ma ing beha iou (Fig. 100)
was obse ed unde d i ed seaweed a Hamama su (Hokkaidô) (see, bionomics and dis ibu ion o T iocha a).
Key o he Japanese species o he subgenus T iocha a
1. Body no mally la ge (Mean: 4.30 mm). Do sal su ace o o ebody ma because o la ge mic o e icula ions. Collec ed in
no heas e n egions such as he Russian Fa Eas and Hokkaidô, Japan (Fig. 104).
(1) Subpalla el sulci on head, connec ed by a sha ow dep ession a he base o head. In e coxal p ocess o meso en i e some-
wha sha ply poin ed (Fig. 64). Pos e io ma gin o e gi e VIII (Figs. 86–87) wi h a ow o hin and long senso y se ae.
[Male]: subapico- en al p ojec ions on median lobe la ge and o al, poin ed apical-la e ally (Fig. 90). [Female]: basal po ion
o spe ma hecal s em (sb) (Fig. 92) wi h a s ong ben a base. . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleocha a (T iocha a) nubis
- Body small o medium (Mean: 3.59 mm). Su ace mo e o less shining. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Su ace o p ono um di e se, no mally wi h wo pai s o impunc u ed a eas (Fig. 58) o ha ing ough su ace en i ely.
Longi udinal h ee u ows on su ace o p ono um shallow and wa y (Fig. 58).
(2) Apex o in e coxal p ocess o meso en i e subpa a ell sided (Fig. 60). Pos e io ma gin o e gi e VIII (Figs. 79–80) wi h a
ow o hin and long senso y se ae. [Male]: subapico- en al p ojec ions on median lobe o aedagus sha ply poin ed (Fig. 83).
[Female]: basal po ion o spe ma hecal s em (sb) (Fig. 85) simple wi hou some p ominen ben s. . . . . . . . . . .A. (T.) ze chei
- Mic os uc u es on do sal su ace o o ebody minu e (su ace shining). Longi udinal h ee u ows on p ono um deep
and s aigh (sligh ly ex ended an e io ly; Fig. 54)
Longi udinal subpa allel u ows on head connec ed wi h a deep u ow o imp ession. Impunc u ed a ea on p ono um (Fig.
54) la gely simila o a hal -moon shape and si ua ed on each side o midline. In e coxal p ocess o meso en i e sha ply
poin ed. Pos e io ma gin o e gi e VIII (Figs. 71–72) wi h a ow o hick senso y se ae. [Male]: subapico- en al p ojec ions
on median lobe ci cula shape in la e al iew (Fig. 75). [Female]: basal s em o spe ma heca (sb) (Fig. 78) somewha simila o
M-shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. (T.) isulca a
Bionomics and Dis ibu ion
Empleno a. Mos specimens used in he p esen s udy we e collec ed in he sup a idal zones o sandy seasho es,
which a e also inhabi ed by many s aphylinid species. Mos Japanese ep esen a i es o Empleno a, excep A.
yama o, we e no mally ound by seasho e lines on sandy coas s composed o ine o coa se sand and small
amoun s o cobbles ones and pebbles. These en i onmen s con ain only small amoun s o mud and ma ix, making
hem somewha di e en om in e idal zones, which a e p e e ed by mos in e idal s aphylinid bee les including
species o he gene a B yo hinusa Casey, 1904, Diaulo a Casey, 1904 (bo h Aleocha inae), and Thinobius
Kiesenwe e , 1844 (Oxy elinae). Ou su eys o li o al Aleocha a species esul ed in he collec ion o abundan
ma e ial om seaweed, and occasionally om ca ion o unde lo sam, some o which can be in es ed by ly
la ae as hos s o he o e bee les. The i s au ho collec ed hund eds o specimens in many a eas and con i med
ha Empleno a species inhabi a eas close o sho elines. These a eas a e usually mode a ely mois . Howe e , hey
a e no co e ed by seawa e h ough wa e ac ion. A Kou a-kaigan (Figs. 93–94), in Shimane-ken, he i s au ho
ound h ee sympa ic species, namely A. ucicola, A, seg ega a, and A. hayamai in one mass o seaweed.
Ano he habi a , he so-called shingle beach composed o sho e ee , ocks, and cobbles ones, is also sui able
o he bee les, and A. yama o was ound only in ha habi a . O he emainig Empleno a species, oo, p e e shingle
beaches. In addi ion, A. pue zi was ound in mo e he e ogeneous en i onmen s, such as sal ma shes, acco ding o
YAMAMOTO & MARUYAMA
48 · Zoo axa 3517 © 2012
Magnolia P ess
(1984) do no ma ch wi h he Japanese species. Fo example, he Nea c ic species a e ega ded as ha ing he
ollowing cha ac e s: meso en i e no ca ina e, ou h an ennal segmen sphe ical, and spe ma heca o emale wi h
sho and e ec duc . Howe e , hese cha ac e s a e no sha ed by some Japanese species. Also, he mo phological
cha ac e s o he subapico- en al p ojec ions on he median lobe o he aedeagus g ea ly di e be ween Japanese
species and Nea c ic axa. The o me species ha e s aigh o gen ly cu ed subapico- en al p ojec ions, while
he Nea c ic ones ha e clea ly angled p ojec ions (see, Klimaszewski, 1984: Figs. 193, 195). The i s au ho
obse ed some specimens o he Nea c ic species Aleocha a (Empleno a) li o alis (Mäklin, 1853) and Aleocha a
(E.) paci ica (Casey, 1894) and con i med ha hese wo g oups may belong o di e en lineages. The axonomic
s udy o seasho e aleocha ines a e a he complica ed due o he con e gence o many axonomically impo an
mou hpa cha ac e s (Ma uyama, 2011) and he Empleno a complex is no excep ion. The e o e, o es ablish a
p ecise phylogeny, s udies ocusing on DNA and wo ldwide esea ch a e needed o esol e hese p oblems. In
pa icula , DNA da a would help o cons uc a mo e p ecise phylogeny o he subgene a, in pa icula he
ela ionships amongs he eas e n and wes e n Palaea c ic, and Nea c ic egions.
Obse ed specimen da a o Nea c ic species
A. li o alis: 1
♂
, J. H. Eme on/G a on/Y-15-05 MS. [HW]//Aleocha a/(Empleno a)/ma i ima (Csy.)/de . J.
Klimaszewski. [FMNH].
A. paci ica: 1
♂
, Redondo,/Cal./D . A. Fenyes//Sep .//634. [HW]//Aleocha a/(Empleno a)/paci ica (Csy.)/de . J.
Klimaszewski//Chicago NHMus/M. Be nhaue /Collec ion. [FMNH].
Geni al pa s o he abo e specimens we e also examined.
Acknowledgemen s
The coope a ion o he pe sons men ioned abo e by o e ing o lending us in aluable ma e ials used in his pape is
g ea ly app ecia ed. We would like o hank: M . Hi o umi Fujimo o (Kagawa-ken), M . Takeshi Hayama
(Okinawa-ken), M . Yasuhiko Hayashi (Hyôgo-ken), M . Ta eo I ô (Kyô o- u), M . Taisuke Kanao (Kyushu
Uni e si y), M . Toshiyuki Ka ô (Hokkaidô), D . Sa oshi Kami ani (Kyushu Uni e si y), Ms. Yasuko Kawakami
(Ôsaka- u), M . Hi oki Ono (Chiba-ken), M . Toshihi o Ozaki (Aki a-ken), M . Yoshihi o Senda (EEEU), M .
Yasu oshi Shiba a (Tôkyô- o), and M . Takashi Wa anabe (Kanagawa-ken). Special hanks a e also due o M .
Hi o umi Mo iya (SCM), D . Shun-Ichi ô Naomi (CBM), D . Al ed F. New on (FMNH), D . Shûhei Nomu a
(Na ional Science Museum,Tokyo), D . Masahi o Ôha a (Hokkaido Uni e si y), D . Akiko Sai ô (CZM), D . Kôhei
Sawada (Ôsaka- u), D . Ma ga e K. Thaye (FMNH), and o D . Hi oyuki Yoshi omi (EEEU) o suppo in
a ious ways. Finally, we deeply indeb ed o D . Volke Assing (Hanno e , Ge many), D . Jan Klimaszewski
(Na u al Resou ces Canada) and an anonymous e iewe o use ul commen s on he manusc ip . This pape is
con ibu ed om he En omological Labo a o y, Facul y o Ag icul u e, Kyushu Uni e si y, Fukuoka (Se . 6, No.
114).
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APPENDIX 1. Species lis o Japanese names o Aleocha a (subgene a Empleno a and T iocha a).
Subgenus Empleno a
Scien i ic name Japanese name
Aleocha a (Empleno a) ucicola Sha p, 1874 Tsuyakeshi-higebu o-hanekakushi
A. (E.) pue zi (Assing, 1995) Ki a- suyakeshi-higebu o-hanekakushi
A. (E.) seg ega a n. sp. Nise- suyakeshi-higebu o-hanekakushi
A. (E.) hayamai n. sp. Hayama- suyakeshi-higebu o-hanekakushi
A. (E.) yama o n. sp. Ô- suyakeshi-higebu o-hanekakushi
Subgenus T iocha a
Scien i ic name Japanese name
Aleocha a (T iocha a) isulca a Weise, 1877 Hoso-sesuji-higebu o-hanekakushi
A. (T.) ze chei (Assing, 1995) Nise-sesuji-higebu o-hanekakushi
A. (T.) nubis (Assing, 1995) Ezo-sesuji-higebu o-hanekakushi
