Revision of the Seashore-dwelling Subgenera Emplenota Casey and Triochara Bernhauer (Coleoptera: Staphylinidae: genus Aleochara) from Japan
Author
Yamamoto, Shûhei
Author
Maruyama, Munetoshi
text
Zootaxa
2012
3517
1
52
journal article
10.5281/zenodo.282595
3c0b0572-9d83-492a-8656-229c8b2043a3
1175-5326
282595
F832C768-A8CA-4FEE-8C3B-BD933247FA6E
Subgenus
Emplenota
Casey, 1884
(
Figs. 1–52
,
93–97
,
101–103
)
Emplenota
Casey, 1884
: 17
(original description;
type
species:
Emplenota maritima
Casey, 1884
: 17
, by original designation and monotypy); 1906: 131 (as genus; key of genera of subtribe “Aleocharae”), 172 (as genus; historical review; diagonostic key to species of Atlantic coast);
Eichelbaum, 1909
: 245
(as subgenus; generic catalogue of world
Staphylinidae
); Fenyes, 1920: 399 (as subgenus; key of world subgengera of
Aleochara
), 1921: 415 (as subgenus; diagnosis; catalogue of world species);
Scheerpeltz, 1925
: 447
(as subgenus; catalogue of Palaearctic species of
Aleocharinae
);
Bernhauer & Scheerpeltz, 1926
: 795
(as subgenus; catalogue of world species of
Aleocharinae
);
Tottenham, 1949
: 404
(as subgenus; generic catalogue of British
Staphylinidae
);
Blackwelder, 1952
: 147
(as subgenus; generic catalogue of world
Staphylinidae
);
Hatch, 1957
: 137
(as subgenus; key to
Aleochara
subgenera of Pacific Northwest), 140 (redescription of species of Pacific Northwest);
Likovsky, 1974
: 294
(as subgenus; key of subgenera of middle Europa);
Moore & Legner, 1975
: 328
(as subgenus; catalogue of Nearctic species);
Seevers, 1978
: 59
(as genus; key to Nearctic genera of
Aleocharinae
), 138 (diagnosis);
Klimaszewski, 1984
: 9
, 10 (phylogenetic relationships of Nearctic
Aleochara
), 95 (as subgenus; revision of Nearctic
Aleochara
; redescription of subgenus), 96 (key; redescriptions of Nearctic species);
Lohse, 1984
: 148
(as genus; diagnosis);
Lohse, 1985
: 328
(as genus; key of middle European species of genera
Emplenota
and
Polystomota
);
Lohse, 1989
: 239
(as genus; key to genera for middle Europe), 240 (as genus; catalogue of middle European species);
Assing, 1995
: 219
(as genus; diagnostic key to genera), 219 (notes on genus), 220 (key to Palaearctic species), 220 (descriptions of each species);
Welch, 1997
: 8
(historical review of British
Emplenota
and related taxa); Maus & Ashe, 1998a (online) (as subgenus; checklist of subgenus of world; diagnosis of subgenus; bionomics; discussion of phylogenetic relationships);
Ashe, 2001
: 360
(as subgenus; catalogue for Nearctic species);
Smetana, 2004
: 356
(as subgenus; catalogue for Palaearctic
Aleocharinae
);
Dauphin, 2005
: 47
(as genus; key to genera of
Emplenota
and
Polystomota
);
Gouix & Klimaszewski, 2007
: 26
(as subgenus; catalogue of Aleocharine species of
Canada
and Alaska).
Polystoma
Stephens, 1833: 91
(original description;
type
species:
Aleochara obscurella
Gravenhorst, 1806
: 159
; fixed by Stephens, 1833: 91, by monotypy); Stephens, 1835: 430 (as genus; redescription of genus);
Thomson, 1861
: 47
(as genus; redescription of genus), 48 (key to Scandinavian species);
Mulsant & Rey, 1874
: 169
(as genus; detailed redescription of genus as
Polystome
[misspelling]), 172 (key to species of
France
), 173 (redescriptions of species of
France
);
Fowler, 1888
: 21
(as subgenus; diagnosis; diagnostic key to species of British Islands);
Casey, 1894
: 289
(as genus; as “
Polistoma
” [misspelling]: see,
Casey, 1906
: 272
; later cited by some authors as synonym of
Emplenota
);
Ganglbauer, 1895
: 45
(as subgenus; redescriptions of middle European species);
Bernhauer, 1901a
: 504
(as subgenus; redescriptions of Palaearctic species);
Johansen, 1914
: 31
(as subgenus; catalogue and key to species of
Denmark
); Fenyes, 1921: 415 (as synonym of
Emplenota
);
Scheerpeltz, 1925
: 447
(as synonym of
Emplenota
);
Bernhauer & Scheerpeltz, 1926
: 795
(as synonym of
Emplenota
);
Tottenham, 1949
: 404
(as synonym of
Emplenota
; note);
Blackwelder, 1952
: 147
, 318 (as synonym of
Emplenota
);
Palm, 1972
: 443
(as subgenus; catalogue; redescriptions of species of
Sweden
);
Moore & Legner, 1975
: 328
(as synonym of
Emplenota
);
Klimaszewski, 1984
: 95
(as synonym of
Emplenota
);
Lohse, 1985
: 328
(as synonym of
Emplenota
);
Smetana, 2004
: 356
(as synonym of
Emplenota
);
Gouix & Klimaszewski, 2007
: 26
(as synonym of
Emplenota
).
Polystomota
Casey, 1906
: 136
(original description;
type
species:
Aleochara grisea
Kraatz, 1856
: 96
; fixed by
Casey, 1906
: 136
, by original designation and monotypy);
Eichelbaum, 1909
: 245
(as genus; generic catalogue of world
Staphylinidae
); Fenyes, 1921: 415 (as synonym of
Emplenota
);
Scheerpeltz, 1925
: 447
(as synonym of
Emplenota
);
Bernhauer & Scheerpeltz, 1926
: 796
(as synonym of
Emplenota
);
Blackwelder, 1952
: 147
, 319 (as synonym of
Emplenota
);
Moore & Legner, 1975
: 328
(as synonym of
Emplenota
);
Seevers, 1978
: 138
(as synonym of
Emplenota
);
Lohse, 1985
: 328
(as synonym of
Emplenota
);
Lohse, 1989
: 239
(as genus; key to genera and species of middle Europe);
Assing, 1995
: 219
(key to Asian coastal genera), 226 (as genus; key to Palaearctic species), 229 (notes on species);
Welch, 1997
: 9
(historical comment);
Maus & Ashe, 1998b
(online) (world checklist of subgenus; diagnosis; bionomics; phylogenetic relationships);
Smetana, 2004
: 357
(as subgenus; catalogue of Palaearctic species);
Dauphin, 2005
: 47
(as genus; key of genera,
Emplenota
and
Polystomota
);
Frank & Ahn, 2011
: 20
(as subgenus; checklist of coastal
Staphylinidae
of world).
Polycharina
Reitter, 1909
: 28
(original description;
type
species:
Aleochara grisea
Kraatz, 1856
: 96
; fixed by
Reitter, 1909
: 28
, by monotypy); Fenyes, 1921: 415 (as synonym of
Emplenota
);
Scheerpeltz, 1925
: 447
(as synonym of
Emplenota
);
Bernhauer & Scheerpeltz, 1926
: 796
(as synonym of
Emplenota
);
Blackwelder, 1952
: 147
, 317 (as synonym of
Emplenota
);
Moore & Legner, 1975
: 328
(as synonym of
Emplenota
);
Lohse, 1985
: 328
(as synonym of
Emplenota
).
FIGURES 8–13.
Body parts of
Aleochara
(
Emplenota
)
fucicola
of male. 8. right antenna; 9. labium; 10. maxilla; 11. mentum; 12. labrum; 13. right foreleg in ventral view.
Polystomaria
Reitter, 1909
: 28
(original description;
type
species:
Aleochara obscurella
Gravenhorst, 1806
: 159
; fixed by
Reitter, 1909
: 28
);
Scheerpeltz, 1925
: 447
(as synonym of
Emplenota
); Fenyes, 1921: 415 (as synonym of
Emplenota
);
Bernhauer & Scheerpeltz, 1926
: 796
(as synonym of
Emplenota
);
Blackwelder, 1952
: 147
, 319 (as synonym of
Emplenota
);
Moore & Legner, 1975
: 328
(as synonym of
Emplenota
);
Klimaszewski, 1984
: 95
(as synonym of
Emplenota
);
Lohse, 1984
: 148
(as genus; diagnosis, as
Polystomaria
Casey
);
Lohse, 1985
: 328
(as synonym of
Emplenota
);
Smetana, 2004
: 356
(as synonym of
Emplenota
);
Gouix & Klimaszewski, 2007
: 26
(as synonym of
Emplenota
).
Redescription. Body
(
Figs. 1–5
): moderately flattened, slender and subparallel sided; whole length varying from 2.5 to 7.0 mm, normally around
3.5–4.5 mm
; dorsal surface covered with minute and dense hexagonal microstructures; surface of head, pronotum and elytra covered with somewhat dense, thin hairs. Dorsal surface of abdomen shining, but head, pronotum and elytra mat (uniformly hexagonal-reticulated).
Colour
(
Figs. 1–5
): uniformly blackish brown to black sometimes with lighter colour in elytra.
Head
: almost circular or somewhat oval (HW/HL ≈1.14), moderately convex above, widest around middle; dorsal surface gently elevated medially, but sometimes with unique elevation partially; impunctate on medial line (e.g.,
Fig. 6
). Eyes small, not strongly protruding laterally.
Antennae
(
Figs. 1–5
,
8
): more or less filiform, entirely robust and long, as same as combined length of head and pronotum; segment I stout and apically dilated; and segment XI cleary long.
Mouth parts
: mandibles asymmetric, left one with one tooth near apex. Clypeus rounded apically. Labrum (e. g.,
Fig. 12
) about 1.7 times as wide as long; anterior margin moderately emarginated medially; basal half semi-transparent. Labial palpus (
Fig. 9
) with segment I clearly longer than II; segment II dialated. Mentum (
Fig. 11
) much wider than long, nearly trapezoidal; anterior margin strongly emarginated; numerous pseudopores scattered randomly. Maxillary palpus (
Fig. 10
) distinctly segmented; segment I long and thick; segment II shorter and narrower than I; segment III slightly shorter and narrower than II; pseudosegment much shorter than other segments; lacinia with numerous hairs and with about 11 thick spines pectinately.
Thorax
: pronotum oval and somewhat transverse (PW/PL ≈1.24), longer than head length (PL/HL ≈1.25), somewhat broader than head (PW/HW ≈1.36), widest around middle, moderately constricted near base; outer margin around middle with a pair of relatively long blackish macrosetae; surface flat and evenly pubescent but sometimes with deep and distinct punctures on dorsal surface numerously. Hypomera fully visible in lateral view. Inter coxal cavities of mesoventrite (e.g.,
Fig. 7
) narrowly or moderately separated. Mesoventrite with short carina (
Fig. 7
: arrow), or not carinate.
Elytra
: widened toward posterior margin, much broader than long (EW/EL ≈1.56), and somewhat wider than pronotum (EW/PW ≈1.28); posterior margin of each elytra nearly truncate but gently rounded toward posterior margins; surface somewhat rugosely punctured both hexagonal microsculptures and prominent distinct punctures in some species; entire surface densely covered with brown setae; anterior margin with one relatively long blackish bristle.
Legs
(e. g.,
Fig. 13
): short and slender with short hind tarsi; numerous spines and bristles on dorsal surface of each tibia, especially on foretibia (
Fig. 13
); midtibia as long as metaventrite; each tarsal segment almost same width (segment V excluding claws); hindtarsi short (hind tarsal length/mid tarsal length ≈1.31).
Hind
wings
: entire; veins weakly sclerotized and very obscure; posterior margin with a row of minute white hairs.
Abdomen
: elongated and slightly narrowed toward posterior segments, narrowing abruptly around apex, widest around segment III–IV; at least tergite III–VI (
Figs. 1–5
) transversely impressed at base. Surface of tergite VIII and sternite VIII simple and smooth. Posterior margin of tergite VIII (
Fig. 14–15
,
23–24
,
30–31
,
38–39
,
46–47
), bearing thick and short several sensory setae (see,
Fig. 14
: arrow).
[
Male
]: posterior margin of sternite VIII (
Figs. 16
,
25
,
32
,
40
,
48
) normally pointed, and its shape varies among species. Median lobe (
Figs. 18–19
,
27–28
,
34–35
,
43–44
,
50–51
) elongated, widest around base. Median lobe with a pair of subapico-ventral projections (see,
Fig. 18
) on median lobe; prominent inner sac with long projecting flagellum (e. g.,
Figs. 18–19
,
43–44
) at least almost as long as median lobe of aedeagus. Paramerite longer than median lobe of aedeagus; apical lobe of paramerite (
Fig. 21
) short and pointed, bearing with four setae.
[
Female
]: posterior margin of tergite VIII, almost truncate, rounded or slightly emarginated; surface of tergite VIII (
Figs. 15
,
24
,
31
,
39
,
47
) similar to that of male. Posterior margin of sternite VIII (
Figs. 17
,
26
,
33
,
41
,
49
) slightly or moderately pointed. Spermatheca (
Figs. 22
,
29
,
37
,
45
,
52
; see,
Fig. 22
) with apical invagination of spermatheca (ai) sharrow; spherical head of spermatheca (sh), as long as or longer than apical portion of spermathecal stem (sa); neck of spermatheca (sn) and (sa) forming more or less in right angle; basal portion of spermathecal stem (sb), narrowing toward base; distinct collar located between (sa) and (sb); sclerotized portion of spermathecal stem (ss) short and erect in most cases; membraneous portion of spermathecal duct (sm) moderate to long length; inner wall of (sh), (sa) and collar moderately striate; each part of spermatheca except for (sm) weakly
Diagnosis.
The members of this subgenus are similar in habitus to those of the tribe
Athetini
(e.g., genera
Adota
Casey, 1910
;
Psammostiba
Sawada, 1976
) found in Japanese seashore, but are discriminated from them by the following character states: antennae shorter, somewhat shorter or almost same as combined length of head and pronotum; maxillary palpi with four segments and a pseudosegment; labial palpi with three segments and a pseudosegment; tarsal fomula: 5-5-5; [
Male
]: median lobe of aedeagus with a prominent flagellum; a pair of subapico-ventral projections on median lobe (see,
Figs. 18–19
); [
Female
]: spermatheca lack of any coiling part; spermathecal stem with a distinct collar (see, example of
Fig. 22
).
FIGURES 14–22.
Terminalia of
Aleochara
(
Emplenota
)
fucicola
. 14. tergite VIII of male; 15. tergite VIII of female; 16. sternite VIII of male; 17. sternite VIII of female; 18. male genitalia: median lobe of aedeagus in lateral view; 19. male genitalia: median lobe of aedeagus in ventral view; 20. apical lobe of median lobe in ventral aspect; 21. apical lobe of paramerite, lateral view; 22. female genitalia: spermatheca.
As sympatric species of
Aleochara
,
A.
(
Coprochara
)
squalithorax
Sharp, 1888
is also similar in general appearance, but differentiated from it as follows: body clearly flattened; antennae longer and slender, more or less shorter or longer than head and pronotum combined; dorsal surface of pronotum flattened; dorsal surface of pronotum smooth (
A. squalithorax
Sharp, 1888
with rough surface and with elevated pronotum; see, also key of the littoral subgenera); [
Male
]: aedeagus with a pair of subapico-ventral projections (
Figs. 18–19
); [
Female
]: spermatheca lack of coiling portion (e.g.,
Fig. 22
) (
A. squalithorax
, basally coiled numerous times).
Comments.
Some species have a spherical or longitudinal elevation on the dorsal surface of the head base (e.g.,
Fig. 6
: arrow) and these are important character states for distinguishing between each species.
Assing (1995)
noted that different pronotal pubescence patterns can be discriminated between two geographical groups, namely, between those in North
America
and the eastern Palaearctic and those in the western Palaearctic.
Klimaszewski (1984)
redefined the subgenus and noted some important character states, including mesoventrite not carinate and antennae with fourth segment spherical; however, these characters are not important and are shared by only some species in the subgenus.
Welch (1997)
and
Park and Ahn (2004)
already pointed out the presence of a carina on the mesoventrite for some species, including
Aleochara fucicola
.
Emplenota
species are parasitoids on the dipteran families:
Anthomyiidae
,
Coelopidae
, and
Sepsidae
. These dipteran families have hitherto been recorded as host agents (
Peschke & Fuldner, 1977
; Maus
et al
., 1998b;
Yamazaki, 2008
,
2012
). Descriptions of the larvae of the European species,
Aleochara
(
Emplenota
)
obscurella
Gravenhorst, 1806
(as
A. algarum
Fauvel, 1862
) are in
Lesne and Mercier (1922)
and
Paulian (1938
,
1941
: 313). Pupation takes place inside the puparia of flies (
Scott, 1920
;
Yamazaki, 2008
,
2012
).