(Coleoptera: Staphylinidae: Aleocharinae:
Aleocharini) and its systematic position
637: 621 – 637 (2004) Munetoshi Maruyama
Department of Zoology, National Science Museum, Hyakunin-chô, Shinjuku-ku, Tôkyô-to, Japan, 169-0073
The Canadian Entomologist 136: 621 – 637 (2004)
Abstract—The genus Creochara Cameron is redescribed, and one species, C. brevipennis(Bernhauer), is recognized in the genus. The systematic position of the genus is discussed, and it is placed in the Tetrasticta genus group in the tribe Aleocharini. The genus group is characterized by the presence of a sternopleural su- ture and its monophyly is supported by the elongate antennomere I, the development of the metasternal process, and the wide separation of the mesocoxal cavities. Tetrasticta bicolor(Cameron) comb. nov. and Tetrasticta bryanti (Cameron) comb. nov. are excluded from the genus Creochara and redescribed.
Maruyama M. 2004. Nouvelle description et position systématique du genre Creochara (Co- leoptera : Staphylinidae : Aleocharinae : Aleocharini). The Canadian Entomologist 136 : 621–637.
Résumé—Le genre Creochara Cameron est décrit de nouveau et une espPce, C. brevipennis (Bernhauer), est retenue dans le genre. La position systématique du genre est analysée et celui-ci est placé dans le groupe de genres de Tetrasticta dans la tribu des Aleocharini. Le groupe est caractérisé par la présence de la suture ster- nopleurale et sa monophylie est appuyée par l’allongement des antennomPres I, le développement du processus du métasternum et la séparation large des cavités mésocoxales. En outre, Tetrasticta bicolor (Cameron) comb. nov. et Tetrasticta bryanti(Cameron) comb. nov. sont exclus de Creochara et décrits de nouveau.
[Traduit par la Rédaction]
Introduction
The genus Creochara was originally described by Cameron (1939) based on a single species, Myrmedonia brevipennis Bernhauer, 1903, known from Vietnam, India, Thailand, Indonesia, and Japan. Cameron (1943) added two species to the genus: C. bicolor and C. bryanti, from Malaysia. Consequently, at one time there were three species from the Orient in this genus. After examining the type series of these three species, I have transferred the latter two species to the genus Tetrasticta Kraatz.
Although Ashe and Maus (1997) recognized the genus Creochara as a member of the subtribe Aleocharina in the tribe Aleocharini, the systematic relationships between the genera in the subtribe remain uncertain. The first purpose of this study was to clar- ify the systematic position of the genus. In the course of a close examination of the ex- ternal structures of the specimens and comparisons with material and descriptions of Aleocharini genera, I found a synapomorphy supporting a sister-group relationship among some genera of the subtribe. The second purpose was to redefine the genus Creocharabased on the single species in the genus, C. brevipennis (Bernhauer), which is redescribed. In addition, I redescribe C. bicolor and C. bryanti, which are transferred to the genus Tetrasticta. The original descriptions by Bernhauer (1903) and Cameron
621
(1925, 1939, 1943) do not include a level of detail sufficient to adequately distinguish these taxa.
Material and methods
The technical procedures adopted here generally follow Naomi and Maruyama (1997). The terminology of the microstructures largely follows Sawada (1972), Seevers (1978), and Klimaszewski (1984). The numbers of setae and pores in the descriptions are confined to one side of the body, except for the medial pseudopores on the prementum.
The depositories of the material examined are abbreviated as follows: BMNH The Natural History Museum, London, United Kingdom
FMNH Field Museum of Natural History, Chicago, Illinois, United States DEI Deutsches Entomologisches Institut, Eberswalde Finow, Germany NMW Naturhistorisches Museum Wien, Wien, Austria
The material for which depositories are not specified is deposited in the personal collection of the author.
All measurements in the text are maximum length and given in millimetres, and the following abbreviations are used: AL, antennal length; BL, body length; EL, elytral length; EW, elytral width; FTL, foretibial length; HL, head length; HTL, hind tibial length; HW, head width; L, length; MTL, midtibial length; PL, pronotal length; PW, pronotal width; W, width.
Taxonomy
Genus Creochara Cameron, 1939
CreocharaCameron, 1939: 653 [description]; Blackwelder 1952, p 111 [list]; Ashe and Maus 1997 [genus list of Aleocharina].
Type species: Myrmedonia brevipennis Bernhauer, 1903, by monotypy.
Diagnosis
This genus can be distinguished from the other genera of the Tetrasticta group by a combination of the following nine character states: (1) secondary sexual dimorphism obvious, especially in body sculpture and shape of abdominal segment VIII; (2) mentum transverse, 2.25 times as wide as long; (3) mentum with deeply emarginated anterior margin; (4) antennomeres II–X wider than long; (5) pronotum somewhat wider than long (PW/PL = approximately 1.24); (6) pronotum subtrapezoidal; (7) pronotal disc with three pairs of small depressions; (8) each elytron obliquely truncated on pos- terior margin; (9) legs long, midtibia slightly longer than metasternum.
Redescription
Body: large, stout, slender (Fig. 1). Head: almost circular, moderately convex above; apical margin of clypeus almost truncated; frons produced in V shape; eyes large, prominent, 0.62 times as long as head. Antennae with antennomere I long, longer than combined length of antennomeres II–IV; antennomere II slightly shorter than III, dilated apically; antennomere III much shorter than half of length of I, dilated apically;
antennomere IV much wider than long, short, one third as long as III; antennomeres V– X wider than long; antennomere XI oval, longer than wide (Fig. 7). Mouthparts: labrum slightly wider than long (W/L = 1.4–1.5); anterior margin rounded but narrowly emarginate medially; lateral margins weakly sclerotized, semitransparent (Fig. 2). Maxillary palpus with segment II long, as long as III, and slightly curved (Fig. 3). Ga- lea slightly curved and densely pubescent apically (Fig. 3). Lacinia with thick setae along inner margin and densely pubescent around the setae (Fig. 3). Mentum much wider than long (W/L = 2.25), and with several pseudopores around anterior margin; anterior margin largely emarginate (Fig. 4). Prementum with 1 setal pore, 3 or 4 real pores, and about 30 medial pseudopores; real pores small and poorly differentiated from
FIGURE1. Creochara brevipennis: habitus, male, dorsal view. Scale bar = 1.0 mm.
FIGURES 2–6. Creochara brevipennis: 2, labrum, dorsal view; 3, maxilla, ventral view; 4, mentum, ventral view; 5, labium, ventral view (palpus eliminated and hypopharynx indicated on right side); 6, meso- and metathoraces, ventral view. Scale bars = 0.1 mm (Figs. 2–5) and 0.5 mm (Fig. 6).
pseudopores; anterior margin deeply emarginated; posteromedial margin with two long projections (Fig. 5). Labial palpus with segment I slightly longer than II, dilated api- cally (Fig. 5). Ligula bilobed; each lobe with four setulae apically (Fig. 5). Thorax: pronotum subtrapezoidal, slightly wider than long; disc with three pairs of small depres- sions (two medially, one laterally) (Fig. 8). Mesosternal process absent (Fig. 6). Metasternal process broad and long, two thirds as long as mesocoxal cavity, and trun- cated apically (Fig. 6). Scutellum with anterior margin rounded; posterior margin slightly pointed. Elytra: short, not reaching posterior margin of metanotum, much wider than long, rounded on posterior margin; surface rugosely punctured and covered with yellow setae. Hind wing: entire. Legs: long, midtibia slightly longer than metasternum; tibiae with apical spur. Abdomen: narrowed posteriorly, widest around segments III and IV.
Male: dorsal surface of head and pronotal disc roughly reticulated, matted. Ante- rior margin of clypeus deeply emarginate. Tergite VII with surface sparsely covered with minute granules. Tergite VIII large, longer than sternite VIII; posterior margin rounded and crenated; surface moderately covered with granules and each granule bear- ing seta (Fig. 9).
Female: dorsal surface of head and pronotal disc slightly reticulated, glossy. An- terior margin of clypeus slightly emarginate or truncate. Tergite VII with surface smooth. Tergite VIII short, much shorter than sternite VIII; posterior margin truncated; surface moderately covered with setae, and base of each seta slightly rugose (Fig. 11).
Creochara brevipennis (Bernhauer, 1903) (Figs. 1–15, 37, 38)
Myrmedonia brevipennisBernhauer, 1903: 21 [original description]. Myrmedonia (?Pella) montana Cameron, 1925: 46 [original description].
Creochara brevipennis (Bernhauer, 1903): Cameron 1939, p 654 [redescription]; Blackwelder 1952, p 111 [list]; Scheerpeltz 1965, p 121 [new records from West- ern Sunda Islands and Thailand]; Pace 1985, p 190 [additional record from India]; Hayashi 1994, p 4 [new record from Japan].
Type material
Myrmedonia brevipennis. Syntypes. VIETNAM: 1 % (Fig. 37), “Tonkin, Montes Mauson, April,Mai 2–3000′, H. Fruhstorfer” / “brevipennis, Bernh. Type, Fruhstorfer” /
“Chicago NHMus, M. Bernhauer, Collection”; 2 &, “Tonkin, Montes Mauson, April,Mai 2–3000′, H. Fruhstorfer” / “Myrmedonia, brevipennis, det. Klima.” / “Chi- cago NHMus, M. Bernhauer, Collection” (FMNH).
Myrmedonia montana. Syntypes. INDIA: 3 %, 11 &, “Matiana 7900, Simla Hills.” / “Dr. Cameron, 11-IX. 1921.” / “M. Cameron., Bequest., B.M.1955–147.” (one
% (Fig. 38) has the label with “Zyras, montanus, Cam, TYPE”). INDONESIA: 1 %,
“Sumatra, SI-Rambe, XII.90-III.91, E. Modigliani” / “Museo Civ., Genova” / “Zyras, montanus, Cam” / “M. Cameron., Bequest., B.M.1955–147.” (BMNH). All syntypes were examined.
Other material
VIETNAM: 1 %, “Hoa Binh, Tonkin” / “Cotype” / “Zyras, brevipennis, det. Dr. Rambousek” / “M. Cameron., Bequest., B.M.1955–147” (BMNH). INDIA: 1 %,
“Arni Gad., Mussoorie” / “Dr. Cameron, 16–10–1921” / “M. Cameron., Bequest., B.M.1955–147”; 1 %, “Dehra Dun., Dr. Cameron, 6–2–1922” / “M. Cameron., Be- quest., B.M.1955–147” (BMNH). INDONESIA: 1 %, “Mte. Singalang, Lugio 1878,
O. Beccari” (NMW). JAPAN: 2 &, Tenkawa-mura, Nara-ken, 9.vii.1990, Saito A; 180 sex?, Hirai (Experimental Forest of Hokkaidô University), Kozagawa-chô, Wakayama- ken, Honshû, 12–15.vii.1999, Maruyama M; 1 %, Kumosô-san, Kisawa-mura, Tokushima-ken, Shikoku, 13–15.vii.2001, Sugimura A; 1 %, Yasuda-gawa, Kishira, Uchinoura-chô, Kagoshima-ken, Kyûshû, 5.v.1991, Haga K (at a dog carcass on sandy seashore of river mouth); 7 %, 12 &, Shiratani (700 m), Yaku-shima, Kagoshima-ken, 28–29.viii.1990, Ôhara M; 2 %, Yodogawa-tozanguchi, Yaku-machi, Yaku-shima, Kagoshima-ken, 2–5.viii.1998, Matsumoto K; 1 %, Yuwan-dake, Amami-Ôshima, Kagoshima-ken, 17.vii.1963, Yamasaki T; 1 %, Nishinakama, Amami-Ôshima, Kagoshima-ken, 5.vi.1969, Maeda Y (used in Hayashi 1994); 1 &, Yona, Kunigami- Son, Okinawa-jima, Okinawa-ken, 9.iii.1991, Kishimoto T. TAIWAN: 3 %, 3 &, Paleng, Taoyuan Hsien, 11.vi.1992, Kanô R; 1 %, Nanshanchi, Puli, Nantou-hsien, 13.iv.1986, Ôhara M; 1 %, Liukuei, Kaosiung-hsien, 26.v.1999, Chen W-L.
FIGURES 7–11. Creochara brevipennis: 7, antennomeres I–V (right, dorsolateral view); 8, pronotum, dorsal view; 9, male tergite VIII, dorsal view; 10, male sternite VIII, ventral view; 11, female tergite VIII, dorsal view. Scale bars = 0.5 mm (Figs. 7, 8) and 0.2 mm (Figs. 9–11).
Redescription
Colour: black; mouthparts, legs, sometimes abdominal segments II and III red- dish brown. Head: surface sparsely covered with minute, yellow setae. Antennae much longer than combined length of head and pronotum; relative lengths of antennomeres from base to apex: 51:19:20:5:10:11:10:10:10:10:26. Thorax: pronotum with surface of disc sparsely covered with yellow, minute setae, and with four or five small bristles lat- erally; three pairs of depressions on disc, each bearing a small bristle; scutellum with surface rugosely punctured and moderately covered with minute setae (Fig. 8). Elytra: surface rugosely punctured and covered with yellow setae, and with 10 small bristles laterally. Abdomen: abdominal tergites III–VII with surface sparsely covered with small setae; tergite VIII with 4 bristles; sternite VIII with posterior margin somewhat truncate, and with about 20 bristles (Fig. 10); number of bristles of tergites III–VII: 2, 2, 2, 2, 2; tergite VIII with 1 bristle laterally.
Male: median lobe of aedeagus much narrowed apically in lateral view; inner sac with flagellum long, coiled seven times (Figs. 12, 13); apical lobe of paramerite slightly dilated apically (Fig. 14).
Female: spermatheca gently curved, weakly sclerotized around junction of basal and apical portions; basal portion dilated apically, as long as apical portion, with
FIGURES 12–15. Creochara brevipennis: 12, median lobe of aedeagus, lateral view; 13, apical lobe of median lobe of aedeagus, dorsal view; 14, apical lobe of paramerite, lateral view; 15, spermatheca. Scale bars = 0.1 mm.
opening of spermathecal gland near apex; apical portion constricted at middle, inner wall of apical half moderately striate (Fig. 15).
Measurements: BL, 5.33–7.78 (7.08 ± 0.73); HL, 0.94–1.09 (1.06 ± 0.08); HW, 1.04–1.23 (1.20 ± 0.07); AL, 2.05–2.58 (2.41 ± 023); PL, 1.04–1.29 (1.19 ± 0.09); PW, 1.25–1.62 (1.48 ± 0.11); EL, 0.96–1.23 (1.12 ± 0.10); EW, 1.43–1.85 (1.75 ± 0.14); FTL, 0.88–1.89 (1.04 ± 0.09); MTL, 1.18–1.86 (1.49 ± 0.13); HTL, 1.40–1.86 (1.63 ± 0.13); HW/HL, 1.04–1.24 (1.14 ± 0.06); PW/PL, 1.16–1.28 (1.24 ± 0.04). n = 10.
Bionomics
In Japan and Taiwan, adult specimens were collected from April to July exclu- sively in laurel forests. The type series of M. montana (the material listed in Cameron 1925, 1939) and the specimens examined from Japan and Taiwan were all collected at the carcasses of vertebrates (e.g., birds and deer).
In the Experimental Forest of Hokkaidô University (Wakayama, Japan), traps baited with rotten chicken attracted numerous adults, which were observed only in the evening until midnight; no individuals were found during the day. By contrast, during the day, the traps attracted Aleochara curtula (Goeze, 1777) and Aleochara lata Gravenhorst, 1802, which are also attracted to carrion and are roughly the same size as C. brevipennis. This may be an example of temporal habitat segregation of beetles oc- cupying similar habitats.
The adults were observed licking the rotten chicken or liquid from the chicken on the asphalt road.
Remarks
The aedeagus of C. brevipennis is characterized by the long, coiled flagellum. The female spermathecal duct is long and almost the same length as the male flagellum. The apex of the flagellum is shaped like the head of a nail. Therefore, it is most likely that the flagellum is used for spermatophore displacement during mating, as already re- ported in A. curtula by Gack and Peschke (1994).
Distribution
Vietnam (Bernhauer 1903); India (Cameron 1925, 1939; Pace 1985); Indonesia: Sumatra (Cameron 1939) and Western Sunda Islands (Scheerpeltz 1965); Thailand (Scheerpeltz 1965); Malaysia (Scheerpeltz 1965); Japan: Honshû*, Shikoku*, Kyûshû*, Yakushima*, Amami-Ôshima (Hayashi 1994), and Okinawa-jima*; Taiwan* (asterisks indicate new localities).
Species to be excluded from Creochara
Genus Tetrasticta Kraatz, 1857
TetrastictaKraatz, 1857: 54 [original description]; Kraatz 1859, p 19 [list], plate 1 [fig- ures of whole body, maxillary palpus, and labium]; Fenyes 1918, p 25 [list]; Fenyes 1921, p 417 [redescription]; Bernhauer and Scheerpeltz 1926, p 798 [list]; Cameron 1939, p 659 [redescription]; Blackwelder 1952, p 111 [list].
Type species: Tetrasticta polita Kraatz, 1857, by monotypy.
Diagnosis
This genus can be distinguished from the other genera of the Tetrasticta group by the combination of the following nine character states: (1) secondary sexual dimor- phism unclear in general appearance; (2) mentum transverse, 1.6–1.9 times as wide as long; (3) mentum slightly emarginate or truncate on anterior margin; (4) antennomeres IV–X wider than long; (5) pronotum much wider than long (PW/PL > 1.40); (6) pronotum slightly elliptical; (7) disc with three pairs of small depressions; (8) posterior margin of elytron horizontally truncated in dorsal view; (9) legs short, midtibia shorter than metasternum.
The genus Tetrasticta is distinguished from the genus Creochara by a combina- tion of character states 1, 2, 3, 5, 8, and 9. Characters of the two species described
FIGURES16–21. Tetrasticta bicolor: 16, mentum, dorsal view; 17, antennomeres I–V, right, dorsolateral view; 18, pronotum, dorsal view; 19, male tergite VIII, dorsal view; 20, male sternite VIII, ventral view; 21, female tergite VIII, apical part, dorsal view. Scale bars = 0.1 mm (Fig. 16), 0.2 mm (Figs. 17, 19–21), and 0.5 mm (Fig. 18).
below were not consistent with the diagnostic characters of the genus Creochara but are consistent with those of the genus Tetrasticta.
The type series of these species and the type species of the genus, Tetrasticta polita Kraatz, 1857 (DEI), were compared.
Tetrasticta bicolor (Cameron, 1943), comb. nov. (Figs. 16–26, 39)
Creochara bicolorCameron, 1943: 183 [original description].
Type material
Creochara bicolor. Syntypes. MALAYSIA: 1 &, “Mt. Matang, W. Sarawak, G. E. Bryant., 1. 2. 14, 1000 m” / “G 58” / “n. gn.? n. sp., Myrmedonia” / “C. bicolor, Cam, TYPE” / “M. Cameron., Bequest., B.M.1955–147.”; 1 % (Fig. 39), “Penang, G. E. Bryant., 1. XI. 13” / “M. Cameron., Bequest., B.M.1955–147.”; 1 &, “Penang, G. E. Bryant., 14. XI. 13” / “M. Cameron., Bequest., B.M.1955–147.” (BMNH). All syntypes were examined.
FIGURES22–26. Tetrasticta bicolor: 22, median lobe of aedeagus, lateral view; 23, apical lobe of median lobe of aedeagus, dorsal view; 24, apical lobe of paramerite, lateral view; 25, 26, spermathecae (Sarawak and Penang, respectively). Scale bars = 0.1 mm.
Redescription
Body: broad, shining (Fig. 39). Colour: blackish brown; antennomeres I–IV, mouthparts, and legs reddish yellow; elytra mainly yellow, but black around posterolateral corners. Head: somewhat pentangular, moderately convex above, slightly wider than long; apical margin of clypeus rounded; frons produced, V shaped; surface sparsely covered with minute, yellow setae; eyes prominent, length 0.65 times that of head. Antennae slightly shorter than combined length of head and pronotum; antennomere I long, as long as combined length of antennomeres II–IV; antennomere II small, half as long as III, slightly dilated apically; antennomere III dilated apically, about half as long as I; antennomere IV short, much wider than long, one quarter as long as III; antennomeres V–X wider than long; antennomere XI oval, longer than wide; relative lengths of antennomeres from basal to apical: 26:7:14:3.5:5:7:7:6:7:7:17 (Fig. 17). Mouthparts: labrum with anterior margin truncated. Mentum somewhat hex- agonal, much wider than long (W/L = 1.9); anterior margin emarginated; anterolateral corner with several pseudopores. (Fig. 16). Prementum with 1 setal pore, 3 real pores, 4 or 5 pseudopores among the real pores, and about 30 medial pseudopores. Thorax: pronotum much wider than long; disc with three pairs of small depressions (two pairs medially, one pair laterally); surface sparsely covered with yellow setae, and with six bristles along lateral margin; each depression bearing a small bristle (Fig. 18). Scutellum rounded posteriorly; surface rugosely punctured and moderately covered with small setae. Elytra: wider than long, subparallel-sided, rounded posterolaterally; sur- face moderately punctured and covered with yellow setae, and with four bristles later- ally. Legs: moderate in length; relative lengths of tarsomeres from basal to apical: 4.5:4.5:5:4.5:7 in foretarsus; 10:8:8:6:14 in midtarsus; 20:11:10:9:19 in hind tarsus. Ab- domen: subparallel-sided, widest around segments IV and V; tergites III–VII sparsely covered with small setae; tergite VIII with 4 bristles; sternite VIII generalized in shape, semicircular in dorsal view, with about 10 bristles (Fig. 20).
Male: tergite VIII with posteromedial margin roundly protruded; surface coarsely punctured (Fig. 19). Median lobe of aedeagus narrowed apically in lateral view; inner sac with flagellum of copulatory piece coiled seven times (Figs. 22, 23); apical lobe of paramerite somewhat dilated apically (Fig. 24).
Female: tergite VIII generalized in shape; surface smooth (Fig. 21). Spermatheca: gently curved; border between basal and apical portions membranous; basal portion di- lated apically, shorter than apical portion, with opening of spermathecal gland, which is prominent; apical portion elliptical, inner wall of apical part three-quarters to four-fifths densely striate (Figs. 25, 26).
Measurements: BL, 5.5–5.7 (5.63 ± 0.06); HL, 0.82–0.83 (0.83 ± 0.01); HW, 0.95–0.99 (0.98 ± 0.02); AL, 1.5–1.58 (1.55 ± 0.04); PL, 0.9–0.96 (0.93 ± 0.03); PW, 1.33–1.39 (1.37 ± 0.03); EL, 0.87–0.9 (0.88 ± 0.02); EW, 1.41–1.51 (1.47 ± 0.05); FTL, 0.56–0.71 (0.64 ± 0.08); MTL, 0.86–0.9 (0.88 ± 0.02); HTL, 0.98–1.06 (1.01 ± 0.04); HW/HL, 1.16–1.19 (1.18 ± 0.02); PW/PL, 1.45–1.48 (1.47 ± 0.02). n = 3.
Distribution
Malaysia: Borneo, Penang.
Tetrasticta bryanti (Cameron, 1943), comb. nov. (Figs. 27–36, 40)
Creochara bryantiCameron, 1943: 183 [original description].
Type material
Creochara bryanti. Syntypes. MALAYSIA: 1 & (Fig. 40), “Mt. Matang, W. Sarawak, G. E. Bryant., 1. 2. 14, No. 16. 14” / “G 61” / “C. bryanti, Cam, TYPE” /
“M. Cameron., Bequest., B.M.1955–147.”; 1 &, “Mt. Matang, W. Sarawak, G.E. Bryant., 1. 2. 14, No. 16. 14” / “G 61” / “M. Cameron., Bequest., B.M.1955– 147.”; 3 %, 2 &, “Mt. Matang, W. Sarawak, G. E. Bryant., XII. 1913” / “M. Cameron., Bequest., B.M.1955–147.” (BMNH). All syntypes were examined.
Redescription
Body: slender, shining (Fig. 40). Colour: blackish brown; antennomeres I–III, mouthparts, legs, and abdominal segment III yellow; elytra mainly brown, but black around posterolateral corners. Head: almost circular, moderately convex above, slightly wider than long; apical margin of clypeus slightly emarginated; surface sparsely cov- ered with minute, yellow setae; frons produced, V shaped; eyes large, prominent, length 0.65 times that of head. Antennae slightly longer than combined length of head and
FIGURES27–32. Tetrasticta bryanti: 27, mentum, dorsal view; 28, antennomeres I–V, right, dorsolateral view; 29, pronotum, dorsal view; 30, male tergite VIII, dorsal view; 31, male sternite VIII, ventral view; 32, female tergite VIII, apical part, dorsal view. Scale bars = 0.1 mm (Fig. 27), 0.2 mm (Figs. 28, 30–32), and 0.5 mm (Fig. 29).
pronotum; antennomere I long, as long as combined length of antennomeres II–IV; antennomere II as long as III, slightly dilated apically; antennomere III much dilated apically; antennomere IV short, much wider than long, half as long as III; antennomeres V–X wider than long; antennomere XI oval, longer than wide; relative lengths of antennomeres from basal to apical: 25:10:10:5:6:6:6:5.5:6:6:15.5 (Fig. 28). Mouth- parts: labrum wider than long; anterior margin slightly emarginated. Mentum subtrapezoidal, much wider than long (W/L = 1.8); anterior margin deeply emarginated; several pseudopores around anterolateral corner (Fig. 27). Prementum with 1 setal pore, 4 real pores, and about 20 medial pseudopores. Thorax: pronotum much wider than long; disc with three pairs of small depressions (two medially, one laterally); surface sparsely covered with yellow setae, and with seven or eight bristles laterally; each de- pression bearing a small bristle (Fig. 29). Scutellum rounded posteriorly; surface rugosely punctured and moderately covered with small setae. Elytra: wider than long, subparallel-sided, rounded posterolaterally; surface moderately punctured and covered with yellow setae, and with four or five bristles laterally. Legs: moderate in length; rel- ative lengths of tarsomeres from basal to apical: 4:3:3:4:6 in foretarsus; 8:6:5.5:5:11 in midtarsus; 15:9:6:5:14 in hind tarsus. Abdomen: subparallel-sided, widest around seg- ments IV and V; tergites III–VII sparsely covered with small setae; tergite VIII with six
FIGURES33–36. Tetrasticta bryanti: 33, median lobe of aedeagus, lateral view; 34, apical lobe of median lobe of aedeagus, dorsal view; 35, apical lobe of paramerite, lateral view; 36, spermatheca. Scale bars = 0.1 mm.
FIGURES 37–40. Habitus: 37, syntype of Myrmedonia brevipennis, male (Vietnam); 38, syntype of Myrmedonia montana, male (India); 39, syntype of Creochara bicolor, male (Sarawak); 40, syntype of Creochara bryanti, male (Sarawak).
bristles; sternite VIII generalized in shape, semicircular in ventral view, with eight or nine bristles (Fig. 31).
Male: tergite VIII with medial part of posterior margin slightly protruded (Fig. 30). Median lobe of aedeagus much narrowed apically in lateral view; inner sac with flagellum of copulatory piece coiled 11 times (Figs. 33, 34); apical lobe of paramerite dilated apically (Fig. 35).
Female: tergite VIII generalized in shape (Fig. 32); surface smooth. Spermatheca: curved at middle; border between basal and apical portions broadly membranous; basal portion dilated apically, longer than apical portion, with opening of spermathecal gland, which is prominent; apical portion oval, constricted near base, inner wall moderately striate apically (Fig. 36).
Measurements: BL, 3.0–4.4 (3.47 ± 0.47); HL, 0.53–0.63 (0.58 ± 0.04); HW, 0.63–0.78 (0.69 ± 0.05); AL, 1.15–1.4 (1.26 ± 0.11); PL, 0.53–0.64 (0.57 ± 0.05); PW, 0.75–0.91 (0.83 ± 0.06); EL, 0.56–0.68 (0.61 ± 0.04); EW, 0.86–1.15 (0.99 ± 0.1); FTL, 0.41–0.53 (0.46 ± 0.04); MTL, 0.44–0.66 (0.55 ± 0.07); HTL, 0.59–0.81 (0.67 ± 0.08); HW/HL, 1.14–1.24 (1.2 ± 0.04); PW/PL, 1.43–1.52 (1.48 ± 0.03). n = 7.
Distribution
Malaysia: Borneo.
Discussion
When Cameron (1939) established the genus Creochara, he placed it in the tribe Aleocharini in the subfamily Aleocharinae. The tribe Aleocharini is subdivided into three subtribes: the Aleocharina, Compactopediina, and Hodoxenina (Newton and Thayer 1992). None of these is well defined. Ashe and Maus (1997) listed the genus Creocharaas a member of the subtribe Aleocharina without any explanation.
The tribe Aleocharini has been well characterized by the presence of pseudo- segments of the maxillary and labial palpi; these character states are unique to the tribe and are apomorphic. In addition to these character states, Aleocharini is characterized by the filiform lateral lobe of the premental apodeme (Fig. 5, llpa). This state is unique to the tribe and apomorphic; i.e., it is considered to be an autapomorphy of the tribe. The genus Creochara is well characterized by these states and thus placed in Aleocharini.
The genus Creochara is closely allied to the aleocharine genera Tetrasticta, Paraleochara Cameron, 1920, Cratoacrochara Pace, 1986, Ilarochara Pace, 1993, AleonictusKistner, 1997, and Formicaenictus Kistner, 1997, and these genera form the Tetrastictagroup. This group of genera is characterized by a combination of the follow- ing five character states: (1) antennomere I long, longer than the combined length of antennomeres II–IV; (2) mesosternal process short; (3) metasternal process well devel- oped, broad, much longer than the mesosternal process; (4) mesocoxal cavities widely separated; and (5) copulatory piece with a long flagellum, which is more than 5 times as long as its basal part. Moreover, I confirm the presence of the sternopleural suture (Fig. 6, sps) in the genera Tetrasticta, Paraleochara, Creochara, Aleonictus, and Formicaenictus. Although I could not examine specimens or drawings of the mesosterna of Cratoacrochara and Ilarochara, these genera are expected to have this character state because of their close resemblance to the genera Tetrasticta and Creochara, in that their mesocoxal cavities are widely separated, the aedeagus is de- pressed laterally, and the copulatory piece has a coiled flagellum.
The elongate antennomere I, the development of the metasternal process, and the wide separation of the mesocoxal cavities are unique to the Tetrasticta group in the
tribe Aleocharini and are considered apomorphic. The subfamily Aleocharinae is repre- sented by 51 tribes (Newton and Thayer 1992), and most of the tribes, including the primitive “lower aleocharines” (Ashe 1994), lack these character states. The similar states are also observed in the tribes Lomechusini and Athetini (Aleocharinae), but the tribes are probably distantly related to Aleocharini. Though the systematic position of Lomechusini and Athetini among aleocharine tribes is unknown, these tribes are proba- bly allied not to Aleocharini but to the other aleocharine tribes, e.g., Falagrini and Homalotini, mainly based on the aedeagal structures. Therefore, the elongate antennomere I, the development of the metasternal process, and the wide separation of the mesocoxal cavities can be regarded as autapomorphies of the Tetrasticta group.
The presence of the sternopleural suture in the Tetrasticta group is apparently plesiomorphic and is not observed in the other genera of Aleocharini. On the basis of this character state and the monophyly of Aleocharini, disappearance of the sterno- pleural suture is an autapomorphy of the tribe, excluding the members of the Tetrasticta group. The genera of the subtribe Aleocharina are recognized by the generalized body shape, but this is characteristic of the tribe Aleocharini, except for members of the subtribes Compactopediina and Hodoxenina, which have a highly specialized body shape because of their termitophily. Actually, no character states supporting monophyly of the subtribe Aleocharina have been found. Therefore, Aleocharina can be regarded as a non-monophyletic group. A subtribal revision of Aleocharini will be needed, but the members of all the genera of the tribe were not available for examination in the present study. A phylogenetic analysis based on all the known genera of Aleocharini will estab- lish the systematic position of the Tetrasticta group in the tribe.
Acknowledgments
I thank M Brendell (BMNH), K Haga (Saitama), Y Hayashi (Hyôgo), T Kishimoto (Japan Wildlife Research Centre, Tôkyô), S-I Naomi (Natural History Mu- seum and Institute, Chiba), A Newton (FMNH), M Ôhara (Hokkaidô University Mu- seum, Sapporo), H Schillhammer (NMW), H Sugaya (Systematic Entomology, Hokkaidô University, Sapporo; SEHU), A Sugimura (Aichi), and L Zerche (DEI) for material, and I Kawashima (Yokosuka) for preparing the habitus figure of C. brevipennis. Thanks are also due to K Yoshizawa (SEHU) for reading an earlier ver- sion of the manuscript and the staff of the Experimental Forest of Hokkaidô University, Wakayama, for their kind assistance during the collecting trip in Kozagawa-chô. This study was supported by a Research Fellowship of the Japan Society for the Promotion of Science for Young Scientists (Postdoctoral Fellowship) to the author.
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(Received: 7 October 2003; accepted: 9 June 2004)
