Zootaxa 1830: 21–36 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition)

New and little-known species of the Tachydromia Meigen (Diptera, ) from Thailand

IGOR SHAMSHEV1 & PATRICK GROOTAERT2 1All-Russian Institute of Plant Protection, shosse Podbel’skogo 3, 188620, St.Petersburg – Pushkin, Russia. E-mail: [email protected] 2Department of Entomology, Royal Belgian Institute of Natural Sciences, Rue Vautier 29, B-1000, Brussels, Belgium. E-mail: [email protected]

Abstract

Six new species of the genus Tachydromia Meigen are described from Thailand: T. doi sp. nov., T. monocercus sp. nov., T. ocellata sp. nov., T. phanensis sp. nov., T. phu sp. nov., T. tigeri sp. nov. A new species group within Tachydromia is distinguished and an updated key to the genus from the Oriental Realm is provided.

Key words: Diptera, Hybotidae, Tachydromia, new species, Thailand, Orient

Introduction

Species of the genus Tachydromia Meigen are likely among the most attractive within the Hybotidae owing to their external appearance and behaviour. They are shining black or blackish brown, usually have wings with brownish or blackish pattern and are found running about quickly on tree-trunks, logs, stones, sand, or leaves of lower herbage resembling in some respects small hymenopterans or ants (Chvála 1970). The group is almost worldwide in distribution (unknown yet from Australia) and currently includes 101 spe- cies, of which 9 species are known from the Oriental Region (Saigusa & Yang 2002; Shamshev & Grootaert 2005; Yang & Grootaert 2006). This paper is a first contribution in the study of the Hybotidae collected during the TIGER project in Thai- land (the Thailand Group for Entomological Research). We describe six new species of Tachydromia from Thailand and provide an updated key to species in the genus for the Oriental Region. Currently, 15 spe- cies of Tachydromia are known from this Region and eight species in Thailand. It should be noted that most specimens examined in this study have been collected at high altitudes and this should not mislead the under- standing of the occurrence of Tachydromia. We still see it as a genus that radiated in a temperate climatic zone such as in the Palaearctic Region and consider the occurrence on mountain tops in Thailand as Palaearctic relicts.

Material and methods

The flies were collected in Malaise traps and transferred to 75% ethanol. Terms used for adult structures pri- marily follow those of McAlpine (1981), although the terminology for the antenna follows Stuckenberg (1999); and for the male terminalia follows Sinclair & Cumming (2006). To facilitate observations, the termi-

Accepted by N. Evenhuis: 23 Jun. 2008; published: 28 Jul. 2008 21 nalia were macerated in cold 10% KOH and hot 85% lactic acid and immersed in glycerine. Drawings of mor- phological features were made with a camera lucida attached to a compound microscope. In descriptions, right and left side of the male terminalia are based on the unrotated position viewed posteriorly, such that in the illustrations the right surstylus appears on the readers left side and vice versa. All male terminalia are fig- ured in their unrotated position. Holotypes and a number of paratypes of new species described here are deposited in the Queen Sirikit Botanic Garden (Chiang Mai), Thailand.

Key to Tachydromia species of the Oriental Region

1. Halter with brown to black knob ...... 2 –. Halter entirely yellow ...... 5 2. Wing with large brown patch on subapical part (Fig. 26) ...... 3 –. Wing brownish infuscate leaving subapical part whitish, brown along anterior margin Figs. 24, 25) .....4 3. Antenna with basal segments yellowish, postpedicel brownish. Fore femur with brown elongate ventral patch basally; hind femur brown on apical 1/4; hind tibia brown on about apical 2/3. Male: fore tibia mostly bare posteriorly but with rather long curved thin setae along sides of bare space (Thailand)...... T. tigeri sp. nov. –. Antenna with basal segments blackish, postpedicel yellowish. Fore and hind femora entirely yellow; hind tibia almost entirely brown, brownish yellow apically. Male: fore tibia unmodified (China, Guangdong)...... T. guangdongensis Yang & Grootaert 4. Fore femur largely brownish; hind femur pale yellow on basal 1/5, otherwise dark brown; hind tibia brown on basal 1/3–1/2. Male: fore tibia unmodified (Thailand) ...... T. phanensis sp. nov. –. Fore femur almost entirely yellowish, indistinctly brownish clouded posteriorly on subapical part; hind femur broadly brownish on middle part; hind tibia dark brown on about apical half. Male: fore tibia slightly curved (viewed dorsally), with ventral spinules becoming stronger on apical half and several spinule-like setulae posteriorly near middle (Thailand) ...... T. phu sp. nov. 5. Wing with 2 broad brownish bands separated along entire length or connected somewhere ...... 6 –. Wing with more or less distinct brownish patch closer to wing apex or (rarer) lacking prominent pattern (Fig. 30) ...... 10 6. Antenna yellow. Hind femur brownish on apical half, hind tibia black brown on apical 1/6–1/5. (Thailand, Singapore)...... T. luang Shamshev & Grootaert –. Antenna dark brown to black (sometimes basal segments somewhat darker). Hind femur and tibia largely blackish brown...... 7 7. Palpus pale yellow (China, Henan)...... T. henanensis Saigusa & Yang –. Palpus dark...... 8 8. Hind femur brown on apical 2/3 (Thailand) ...... T. monocercus sp. nov. –. Hind femur entirely blackish brown ...... 9 [T. papuana Grootaert, the only Tachydromia species known from Australasian Region, belongs here.]

9. Laterotergite with thin finger-like process. Wing with bands connected on cells r1 and r4+5. (China, Henan) ...... T. digitiformis Saigusa & Yang –. Laterotergite unmodified. Wing with 2 broad brown bands separated along the whole length (India) ...... T. latifascipennis Brunetti 10. Hind femur entirely yellow...... 11 –. Hind femur brown on apical part...... 12 11. Fore tarsomeres 4 and 5 brown. Hind tibia brownish on about basal half (Thailand)...... T. doi sp. nov. –. Only fore tarsomere 5 darkened on apical half. Hind tibia entirely yellow (Thailand)......

22 · Zootaxa 1830 © 2008 Magnolia Press SHAMSHEV & GROOTAERT ...... T. terricoloides Shamshev & Grootaert 12. Palpus yellowish brown. Fore tibia almost entirely brownish, extreme base yellow. Wing largely finely infuscate, somewhat deeper on middle part along veins (Thailand) ...... T. thaica Shamshev & Grootaert –. Palpus pale yellow. Fore tibia at most brownish apically. Wing with 1 or 2 brownish patches on subapical part ...... 13

13. Wing with 2 brownish patches on subapical part (Fig. 27); vein R2+3 incomplete (Thailand), male ...... T. ocellata sp. nov. –. Wing with 1 brownish patch on subapical part (female T. ocellata has also only one patch, Fig. 31); vein

R2+3 complete...... 14 14. Fore tarsomeres 4 and 5 brown; mid tibia yellow; hind femur on apical half (except for apex) and hind tibia in basal 2/3 blackish (China, Henan)...... T. longyuwanensis Saigusa & Yang –. Only fore tarsomere 5 darkened on apical part; mid tibia blackish; hind femur darkened on apex; hind tibia yellow (Taiwan)...... T. bistigma (Bezzi)

Systematic account

Chvála (1970, 1975) divided Tachydromia into eight species groups. However, we have already noted that in the Oriental Region this classification has some problems and there are species that cannot be definitely included in any of the groups proposed (Shamshev & Grootaert 2005). This primarily concerns T. luang and four species described here, which would run to the calcanea or terricola groups (sensu Chvála 1970). How- ever, many characters, including male terminalia structure, indicate that these species are more closely related to each other than to the species belonging to these two species groups. To clarify this inconsistency a revision of the relationships within Tachydromia will be necessary which, however, is beyond the scope of our paper. Nevertheless, we provisionally recognised T. phanensis sp. nov., T. phu sp. nov., T. tigeri sp. nov., T. ocellata sp. nov. and T. luang as comprising a separate, hitherto unknown group of species within Tachydromia (T. luang group). In addition, two new species are described in this paper, namely T. monocercus sp. nov. and T. doi sp. nov., that belong to previously described species groups of Tachydromia.

Tachydromia luang group

The group can be distinguished from other species groups by the following combination of characters: palpus yellow (rarely brownish yellow), lacking silvery hairs; prothoracic sclerites finely pollinose (except T. luang); usually 2 notopleurals; vein R2+3 straight. It should be added that the wing pattern is very variable within this group. The presence of a ventral process on the right epandrial lamella of the male terminalia is likely a syna- pomorphy of the T. luang group.

Tachydromia phanensis sp. nov. (Figs 1–3, 24)

Diagnosis. Palpus yellow. Legs with complicated colour pattern; fore femur largely brownish. Wing almost entirely finely brownish infuscate, paler near base and hyaline on subapical space, brown along anterior mar- gin. Halter with brown knob. Male: fore tibia unmodified.

TACHYDROMIA THAILAND Zootaxa 1830 © 2008 Magnolia Press · 23 FIGURES 1–3. Tachydromia phanensis sp. nov. 1. hypopygium, ventral view. 2. the same, dorsal view. 3. left surstylus, lateral view. a: anus; HY: hypandrium; LEL: left epandrial lamella; LS: left surstylus; RC: right cercus; REL: right epan- drial lamella; RS: right surstylus; vp: ventral process of right epandrial lamella. Scale 0.1 mm.

Description. Male. Wing 1.8–1.9 mm. Head black in ground-colour. Eyes extending beyond ocellar tubercle. Frons subshining, slightly widened toward ocellar tubercle, above antennae nearly 1.5 times as broad as anterior ocellus. Ocellars minute. Occiput largely finely pollinose, narrowly shining in upper part beyond eyes; with 2 short strong black setae in upper part, 2 similar setae on middle part laterally and numerous pale hair-like setae near neck and on lower part. Basal antennal segments brownish; postpedicel paler, brownish yellow, short, elongate oval; stylus very long, brownish, short pubescent. Proboscis largely brown, yellow proximally. Palpus unmodified, short, elongate oval, yellow, with 1 long black and 1 subequal but thinner brownish yellow subapical setae and 4–5 brownish yellow short setae basally. Thorax dark brown in ground-colour, largely shining; prominent setae black; prothoracic sclerites (except postpronotal lobe) pollinose. Postpronotal lobe large, lacking conspicuous setae. Mesonotum with 2 notopleu- ral (posterior one longer) and 2 scutellar setae; Acrostichals and dorsocentrals minute, the former arranged in 2–3 irregular rows, lacking on prescutellar depression, the latter uniserial. Legs long, with complicated colour pattern; fore coxa almost entirely brown (yellow apically), mid coxa almost entirely yellow (brownish basally), hind coxa entirely yellow; all trochanters yellow; fore femur brownish near extreme base, broadly in apical half posteriorly and dorsally (except extreme apex), with nar- row brownish anterior patch basally, otherwise yellow; mid femur almost entirely brownish, somewhat paler dorsally and near extreme base and apex; hind femur pale yellow on basal 1/5, otherwise dark brown; fore and mid tibiae brownish, hind tibia brown on basal 1/3–1/2, otherwise yellow; all tarsi brownish (hind basitarsus slightly paler basally). Coxae mostly with pale hair-like setae of different length, hind coxa with 1 strong black curved seta anteriorly; pale hair-like setae present between fore and mid coxae. Fore femur strongly thickened, mostly with minute anteroventral and posteroventral setulae, bearing 4–5 stronger spinule-like

24 · Zootaxa 1830 © 2008 Magnolia Press SHAMSHEV & GROOTAERT anteroventral setulae near middle. Fore tibia slightly swollen, with inconspicuous setation. Mid femur slender, with anteroventral and posteroventral very short (the latter longer) spinule-like setae becoming somewhat longer toward base of femur. Mid tibia slender, lacking prominent bristles with some ventral spinule-like setu- lae. Hind femur slender, with rows of short strong anteroventral and posteroventral setae becoming shorter and thinner basally. Hind tibia slender, lacking prominent bristles, with narrow pointed posterior apical comb. Tarsi of all legs unmodified. Wing (Fig. 24) normally developed, rounded at apex, with typical venation for the genus; largely finely brownish infuscate, paler near base and hyaline on subapical space, brown along anterior margin. One short basal costal seta present. Vein R1 meeting costa beyond wing midway. Proximal section of vein R4+5 consider- ably longer than Rs. Vein R2+3 straight. Veins R4+5 and M1+2 parallel toward wing-apex. Crossveins r-m and bm- cu separated. Cells br and bm extending to wing midway. Halter with pale stem and brown knob. Abdomen with segment 1 pale, otherwise brown, covered mostly with scattered, short, black setulae; seg- ments 7 and 8 with moderately long setae. Terminalia (Figs 1–3) large, dark brown, elongate oval. Right cer- cus long, tapered apically, covered with numerous unmodified setae of different length, bearing 1 very long apical seta. Left cercus about half length of right cercus, pointed apically, covered with scattered short setae. Right epandrial lamella subtriangular (viewed laterally), with large rounded ventral process in subapical part, bearing numerous setae of different lengths. Right surstylus long, narrow, bent inward, with several unmodi- fied setae of different length apically and on inner side. Left epandrial lamella subtriangular, bearing several unmodified setae of different lengths. Left surstylus short, subrectangular, with 1 very long subapical seta, several strong moderately long setae on upper margin and some short setae apically. Female. Fore femur more extensively brownish, lacking stronger anteroventral spinulae near middle. Hind femur lacking prominent anteroventral and posteroventral setae. Otherwise as in male. Cercus brownish, rather short. Material examined. Holotype male: THAILAND Chaiyaphum prov., Pa Hin Ngam NP Tung Dok Gra- jeaw 15°38.208'N 101°23.556'E 720 m Malaise trap 6–12.vii.2006 Kratae Sa-nog & Buakaw Adnafai leg. T331. Paratypes: 1 male, 1 female, THAILAND Sakon Nakhon prov., Phu Phan NP Behind office 17°03.488'N 03°58.497'E 318 m Malaise traps 8–14.vii.2006 Sailom Tongboonchai leg. T199; 1 female, Chiang Mai prov., Doi Inthanon NP Vachiratharn Falls 18°32.311'N 98°36.048'E 700 m Malaise trap 12–19.x.2006 Y. Areeluck leg. T366; 1 female, Sakon Nakhon prov., Phu Phan NP Behind office 17°03.521'N 103°58.450'E 322 m Mal- aise traps 27.vii–2.viii.2006 Winlon Kongnara leg. T207; 1 female, Chaiyaphum prov., Pa Hin Ngam NP creek at Tung Dok Grajeaw 15°38.391'N 101°23.609'E 750 m Malaise trap 19–24.vii.2006 Kratae Sa-nog & Buakaw Adnafai leg. T335. Derivatio nominis. The species is named after one of the National Parks where the species was found: Phu Phan NP in Sakon Nakhon province. Distribution. Thailand. Remarks. The new species is closely related to T. phu sp. nov. sharing similar setation of the left surstylus (1 very long and several strong bristles). The main distinguishing features between these species are given in the key.

Tachydromia phu sp. nov. (Figs 4–6, 25)

Diagnosis. Palpus yellow. Legs largely yellow. Wing largely finely brownish infuscate, hyaline on subapical space, brown along anterior margin. Halter with brown knob. Male: fore tibia slightly curved (viewed dor- sally), with ventral spinules.

TACHYDROMIA THAILAND Zootaxa 1830 © 2008 Magnolia Press · 25 FIGURES 4–6. Tachydromia phu sp. nov. 4. right epandrial lamella, ventral view. 5. epandrium with cerci. 6. left surst- ylus, lateral view. Scale 0.1 mm.

Description. Male. Wing 2.3–2.4 mm. Head black in ground-colour. Eyes extending beyond ocellar tubercle. Frons subshining, slightly widened toward ocellar tubercle, above antennae nearly 1.5 times as broad as anterior ocellus. Ocellars minute. Occiput finely pollinose; with 2 moderately long strong black setae in upper part, 2 similar setae on middle part laterally and numerous pale hair-like setae near neck and on lower part. Basal antennal segments brownish yellow; postpedicel paler, yellow, short, elongate oval; stylus very long, brownish, short pubescent. Proboscis largely yellow, narrowly dark brown proximally. Palpus unmodi- fied, short, elongate oval, yellow, with 1 long black subapical seta and 4–5 yellowish short setae basally. Thorax dark brown in ground-colour, largely shining; prominent setae black; prothoracic sclerites (except postpronotal lobe) pollinose. Postpronotal lobe large, lacking conspicuous setae. Mesonotum with 2 strong notopleural and 2 scutellar setae; Acrostichals and dorsocentrals minute; the former multiserial, lacking on prescutellar depression; the latter uniserial anteriorly, arranged in 3–4 irregular rows and hardly separated from acrostichals posteriorly. Legs long, largely yellow; mid coxa with brownish patch laterally; fore femur indistinctly brownish clouded posteriorly on subapical space; mid tibia brownish on apical 2/3 anteriorly and (less distinct) on basal 2/3 posteriorly; fore tarsomere 4 brownish yellow on apex, tarsomere 5 brown; mid tibia brown basally and becoming gradually paler toward apex; mid tarsomere 4 brownish yellow, tarsomere 5 brown; hind femur broadly brownish on middle part; hind tibia dark brown on about apical half. Coxae mostly with pale hair-like setae of different lengths; several pale hair-like setae present between fore and mid coxae. Fore femur strongly thickened, with tuft of short black spinule-like setae near middle and several spinule-like setulae basally. Fore tibia swollen and slightly curved (viewed dorsally), with ventral spinules over entire length (stronger on apical half) and several spinule-like setulae posteriorly near middle. Fore basitarsus with ventral spinules becoming shorter apically. Mid femur slender, with anteroventral and posteroventral very short (the latter longer) spinule-like setae becoming somewhat longer toward base of femur, bearing 4–5 moderately long strong setae near base. Mid tibia slender, lacking prominent bristles, with some ventral spinule-like setulae. Hind femur slender, with row of short strong anteroventral setae becoming minute basally. Hind tibia slightly thickened on

26 · Zootaxa 1830 © 2008 Magnolia Press SHAMSHEV & GROOTAERT about apical half, lacking prominent bristles, with narrow pointed posterior apical comb. Tarsi of all legs unmodified (hind tarsus missing). Wing (Fig. 25) normally developed, rounded at apex, with typical venation for the genus; largely finely brownish infuscate, hyaline on subapical space, brown along anterior margin. One short basal costal seta present. Vein R1 meeting costa beyond wing midway. Proximal section of vein R4+5 considerably longer than

Rs. Vein R2+3 straight. Veins R4+5 and M1+2 parallel toward wing-apex. Crossveins r-m and bm-cu separated. Cells br and bm extending to wing midway. Halter with pale stem and brown knob. Abdomen with segment 1 pale (viewed dorsally with narrow brownish patch), otherwise brown, subshin- ing, covered mostly with scattered, short, black setulae; segments 7 and 8 with moderately long setae. Termi- nalia (Figs 4–6) large, dark brown, elongate oval. Right cercus rather long, subrectangular, with numerous moderately long marginal setae in subapical part. Left cercus quite long but somewhat shorter than right cer- cus, slightly curved, pointed apically, with numerous unmodified setae of different lengths. Right epandrial lamella subtriangular (viewed laterally), with large subglobular ventral process in subapical part, bearing numerous setae of different lengths. Right surstylus long, narrow, bent inward, with several unmodified setae on inner side and apically. Left surstylus rather subtriangular, with 1 very long bristle on lower margin about middle, 4 strong long bristles on upper margin and some unmodified setae of different lengths apically. Female. Fore coxa sometimes brownish near base anteriorly; hind tarsus brownish. Fore femur with numerous unmodified spinules ventrally; fore tibia evenly thickened, with unmodified ventral spinules; mid femur with short bristles near base; hind femur with hardly prominent anteroventral setae. Otherwise as in male. Cercus brownish, short. Material examined. Holotype male: THAILAND Loei prov., Phu Ruea NP office 17°28.805'N 101°21.242'E 870 m Malaise trap 5–12.vii.2006 Patikhom Tamtip leg. T312. Paratypes: 1 female, THAILAND Chiang Mai prov., Doi Inthanon NP Vachiratharn Falls 18°32.311'N 98°36.048'E 700 m Malaise trap 12–19.x.2006 Y. Areeluck leg. T366; 1 female, Doi Inthanon NP camp- ground pond 18°32.657'N 98°31.482'E 1200 m Malaise trap 2–10.xi.2006 Y. Areeluck leg. T385; 1 female, Doi Inthanon NP Vachiratharn Falls 18°32.311'N 98°36.048'E 700 m Malaise trap 27.ix–5.x.2006 Y. Areeluck leg. T347. Derivatio nominis. The specific epithet alludes to the fact that the species was found on a mountain (“Phu kao” is Thai for mountain though generally only the word “Phu” is used to indicate mountain). Distribution. Thailand. Remarks. The new species is closely related to T. phanensis sp. nov. sharing similar setation of the left surstylus (1 very long and several strong bristles). The main distinguishing features between these species are given in the key.

Tachydromia tigeri sp. nov. (Figs 7–9, 26)

Diagnosis. Postpedicel brownish. Palpus brownish yellow. Legs with complicated colour pattern: largely finely brownish infuscate, with large brown patch on subapical part. Halter with brown knob. Male: fore tibia mostly bare posteriorly but with rather long curved thin setae along sides of bare space. Description. Male. Wing 2.9–3.0 mm. Head black in ground-colour. Eyes extending beyond ocellar tubercle. Frons subshining, slightly widened toward ocellar tubercle, above antennae nearly 1.5 times as broad as anterior ocellus. Ocellars minute. Occiput largely shining, narrowly pollinose behind eyes; with 2 moder- ately long strong black cruciate setae in upper part, 2 similar setae on middle part laterally and numerous pale hair-like setae near neck and on lower part. Basal antennal segments yellow; postpedicel and stylus brownish; postpedicel about 2.5 times longer than wide, with straight upper and convex lower margin; stylus long, short

TACHYDROMIA THAILAND Zootaxa 1830 © 2008 Magnolia Press · 27 pubescent. Proboscis entirely brownish yellow. Palpus unmodified, elongate, brownish yellow, with 1 long black and 1 pale, shorter and thinner subapical setae.

FIGURES 7–9. Tachydromia tigeri sp. nov. 7. right epandrial lamella, ventral view. 8. epandrium with cerci. 9. left sur- stylus, lateral view. Scale 0.1 mm.

Thorax brown in ground-colour, largely shining; prominent setae black; prothoracic sclerites (except post- pronotal lobe) finely pollinose. Postpronotal lobe large, lacking conspicuous setae. Mesonotum with 1 strong notopleural and 2 scutellar setae; acrostichals and dorsocentrals minute, uniserial, lacking on prescutellar depression. Legs long, with complicated colour pattern; fore femur with brown elongate ventral patch basally; fore tibia brown on apical 1/4; fore basitarsus brown on apical 1/3, tarsomeres 2–4 almost entirely (except extreme base), tarsomere 5 entirely brown; mid femur with small brownish subapical patch dorsally; mid tibia entirely brownish; mid tarsomeres 1–2 brownish apically, tarsomeres 3–5 becoming gradually darker brown toward tarsomere 5; hind femur brown on apical 1/4; hind tibia brown on about apical 2/3; hind tarsus almost entirely brownish (except extreme base of tarsomeres 1 and 2); otherwise legs reddish yellow. Coxae with black setae of different lengths; no setae between fore and mid coxae. Fore femur strongly thickened, with minute ventral setulae. Fore tibia spindle-like, mostly bare posteriorly but with rather long curved thin setae along bare space. Mid femur slender, with minute anteroventral and posteroventral spinule-like setulae, bearing 3 black spine- like setae of different lengths near base. Mid tibia with shallow subapical excision, bearing black ventral spinule-like setulae. Hind femur slender, with rows of short anteroventral and posteroventral setae. Hind tibia slender, lacking prominent bristles, with rounded posterior apical comb. Wing (Fig. 26) normally developed, rounded at apex, with typical venation for the genus; largely finely brownish infuscate, with large brown patch near apices of veins R1 and R2+3. One short basal costal seta present. Vein R1 meeting costa beyond wing midway. Proximal section of vein R4+5 considerably longer than

Rs. Vein R2+3 straight. Veins R4+5 and M1+2 parallel toward wing-apex. Crossveins r-m and bm-cu separated. Cells br and bm extending to about wing midway. Halter with yellow stem and brown knob. Abdomen brown, subshining, covered mostly with scattered, short, black setulae; segments 7 and 8 with moderately long setae. Terminalia (Figs 7–9) moderately large, dark brown, elongate oval. Right cercus mod-

28 · Zootaxa 1830 © 2008 Magnolia Press SHAMSHEV & GROOTAERT erately long, rather broad, rounded apically, with numerous unmodified setae of different lengths. Left cercus short, narrowed apically, with several unmodified setae of different lengths. Right epandrial lamella subtrian- gular (viewed laterally), with large subglobular ventral process in subapical part, bearing numerous setae of different lengths. Right surstylus long, narrow, bent inward, with several unmodified setae on inner side and apically. Left surstylus subtriangular, with several long marginal bristles. Female. Fore tibia with unmodified setation; mid tibia unmodified, slender. Otherwise as in male. Cercus brown, long. Material examined. Holotype male: THAILAND Chiang Mai prov., Doi Inthanon NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 9–16.viii.2006 Y. Areeluck leg. T177. Paratypes: 1 male, THAILAND Chiang Mai prov., Doi Inthanon NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 2–8.vii.2006 Y. Areeluck leg. T53; 1 male, Doi Inthanon NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 16–24.viii.2006 Y. Areeluck leg. T184; 1 female, Doi Intha- non NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 2–9.viii.2006 Y. Areeluck leg. T124. Derivatio nominis. The epithet “tiger” alludes to the name of the TIGER Project. Distribution. Thailand. Remarks. The precise relationships of T. tigeri sp. nov. are unclear beyond its inclusion in the T. luang group. The new species can be distinguished from other species known from the Oriental Region as it is out- lined in the key.

Tachydromia ocellata sp. nov. (Figs 10–12, 27, 31)

Diagnosis. Palpus yellow. Legs with complicated colour pattern. Male wing with vein R2+3 incomplete; 2 rounded dark brown spots present on subapical part. Halter yellow.

FIGURES 10–12. Tachydromia ocellata sp. nov. 10. right epandrial lamella, ventral view. 11. epandrium with cerci. 12. left epandrial lamella with surstylus, lateral view. Scale 0.1 mm.

Description. Male. Wing 1.6–1.7 mm. Head black in ground-colour. Eyes extending beyond ocellar tubercle. Frons pollinose, slightly widened toward ocellar tubercle, above antennae nearly 1.5 times as broad

TACHYDROMIA THAILAND Zootaxa 1830 © 2008 Magnolia Press · 29 as anterior ocellus. Ocellars minute. Occiput finely pollinose; with 2 moderately long strong black setae in upper part, 2 similar setae on middle part laterally and numerous pale hair-like setae near neck and on lower part. Basal antennal segments brownish yellow; postpedicel yellow, small, elongate oval; stylus very long, brownish, short pubescent. Proboscis almost entirely yellow, brownish at tip. Palpus unmodified, small, rounded, yellow, with 1 very long black and 1 pale short thin subapical setae. Thorax brown in ground-colour, largely shining; prominent setae black; prothoracic sclerites (except post- pronotal lobe) finely pollinose. Postpronotal lobe very large, lacking conspicuous setae. Mesonotum with 2 notopleural (posterior longer and stronger) and 2 scutellar (shorter than posterior notopleural) setae; acros- tichals and dorsocentrals minute; the former arranged in 1–2 irregular rows, lacking on prescutellar depres- sion; the latter uniserial, 1 prescutellar pair as long as scutellar setae. Legs long, with complicated colour pattern; fore tibia brownish anteriorly and posteriorly; fore tarsomeres 4 and 5 brown; mid femur brownish posteriorly on apical 2/3; mid tibia entirely brownish; mid tarsomeres largely brownish, basitarsus paler basally; hind femur brown on about apical 1/2; hind tibia brown on apical 1/ 4; hind tarsomere 2 brownish on about apical 1/2, tarsomeres 3–5 entirely brownish; otherwise legs yellow. Coxae with yellowish to brownish yellow setae of different length; no setae between fore and mid coxae. Fore femur strongly thickened, with minute brownish yellow, anteroventral and posteroventral spinule-like setulae, bearing 2 longer black spinulae near base. Fore tibia spindle-like, with unmodified setation. Mid femur slen- der, with rows of anteroventral and posteroventral spinule-like setae (the latter black and somewhat longer) and 2 long pale hair-like setae near base. Mid tibia slender, bearing ventral spinule-like setulae. Hind femur slender, with rows of very short anterodorsal and posteroventral setae. Hind tibia slender, lacking prominent bristles, with rounded posterior apical comb.

Wing (Fig. 27) tip deformed, with modified venation, costal vein curved in subapical part, vein R2+3 incomplete, disappearing before meeting of vein R1 with costa; entirely finely brownish infuscate and with 2 rounded dark brown spots on virtual meeting point of vein R2+3 in costa and near wing apex between veins R4+5 and M1+2. Basal costal seta absent. Vein R1 meeting costa near wing midway. Proximal section of vein R4+5 considerably longer than Rs. Veins R4+5 and M1+2 strongly divergent toward wing-apex. Crossveins r-m and bm-cu confluent. Cells br and bm extending to wing midway. Halter yellow. Abdomen brownish (segments 1–2 paler), subshining, covered mostly with scattered, short, black setulae; segments 7 and 8 with moderately long setae. Terminalia (Figs 10–12) rather large, brown, elongate oval. Right cercus bilobed, with both lobes digitiform, curved, bearing some unmodified setae of different lengths; right lobe somewhat longer and narrower. Left cercus quite long, broadened apically and basally, narrow on middle part, with numerous unmodified setae of different lengths. Right epandrial lamella subtriangular (viewed laterally), with large rounded ventral process in subapical part, bearing numerous setae of different lengths. Right surstylus long, narrow, bent inward, with several spine-like sensilla on inner side. Left epan- drial lamella subtriangular, bearing several unmodified setae of different lengths. Left surstylus short, subrect- angular, with several unmodified setae. Female. Body length: 1.75–2 mm; wing: 1.5–1.75 mm.

Wing (Fig. 31) not deformed like in male and vein R2+3 complete, ending in costa; only one rounded dark brown spot present beyond end R2+3. The spot can be variable in colour intensity. The tip of the wing is white. Structure and colour pattern of legs identical to male. Material examined. Holotype male: THAILAND Loei prov., Phu Ruea NP Subhnonghin 17°28.772'N 101°21.308'E 860 m Malaise trap 26.vii–2.viii.2006 Nukoonchai Jaroenchai leg. T319. Paratypes: THAILAND Phetchabun prov., Nam Nao NP Helicopter landing ground 16°43.156'N 01°35.118'E 890 m Malaise trap 1 male, 1 female, 17.vii.2006 – 24.vii.2006 (T276); 1 male, 24–30.vii.2006 (T278); 4 females, 24.vii.2006 – 30.vii.2006 (T279) Noopean Hongyothi leg.

30 · Zootaxa 1830 © 2008 Magnolia Press SHAMSHEV & GROOTAERT 1 female Chaiyaphum prov., Pa Hin Ngam NP, Ecotone between mix deciduous/dry dipterocarp, 15°38.100'N 101°23.857'E, 700m, 18.viii.2006 – 24.viii.2006, Malaise trap, (T446) Katae Sanog & Buakaw Adnafai leg. 2 males, 7 females, Loei prov., Phu Ruea NP, Pan Hin Khan Maak ditch, 17°30.042'N 101°20.474'E, 1219m, 12.viii.2006 – 19.viii.2006, Malaise trap, (T528); 2 females, 26.viii.2006 – 2.ix.2006 (T534), Nuk- oonchai Jaroenchai leg.; 3 males, 1 female, Loei prov., Phu Ruea NP, Huay Taey ditch, 1233m, 12.viii.2006 - 19.viii.2006, Malaise trap, (T529); 3 males, 3 females, 26.viii.2006 – 2.ix.2006 (T535) Nukoonchai Jaroen- chai leg.; 3 males, 5 females, Loei prov., Phu Ruea NP, Nern Wibaak ditch, 19.viii.2006 – 26.viii.2006, 1196m (T530) Nukoonchai Jaroenchai leg.; 1 female, Loei prov., Phu Ruea NP, Pah Lo Noy, 17°30.502'N 101°20.868'E, 1343m, 12.ix.2006 – 19.ix.2006, Malaise trap, (T827) Nukoonchai Jaroenchai leg. 1 female, Sakon Nakhon prov., Phu Phan NP, Kam Hom waterfall at Haew Sin chai, 17°07.415'N 104°01.179'E, 347m, 4.ix.2006 – 10.ix.2006, Malaise trap, (T610) Sailom Tongboonchai leg. Derivatio nominis. The name alludes to the two brownish spots on the tip of the wing. Distribution. Thailand. Remarks. Tachydromia ocellata sp. nov. is closely related to T. luang sharing similar setation of the right surstylus (several spine-like sensilla on inner side). The new species can be readily distinguished from other Tachydromia known from the Oriental Region by its unique venation and pattern of the wing.

FIGURES 13–16. Tachydromia luang Shamshev, Grootaert, 2005. 13. right epandrial lamella, lateral view. 14. epan- drium with cerci. 15. apical part of right surstylus, dorsal view. 16. left epandrial lamella with surstylus, lateral view. Scale 0.1 mm.

TACHYDROMIA THAILAND Zootaxa 1830 © 2008 Magnolia Press · 31 Tachydromia luang Shamshev, Grootaert (Fig. 13–16, 28)

Tachydromia luang Shamshev & Grootaert, 2005: 115.

Material examined. 1 male, THAILAND Loei Phu Ruea NP Subhnonghin 17°28.772'N 101°21.308'E 860 m Malaise trap 26.vii–2.viii.2006 Nukoonchai Jaroenchai leg. T319; 1 male, Phetchabun prov., Nam Nao NP Helicopter landing ground 16°43.184'N 01°35.137'E 875 m Malaise trap 3–10.vii.2006 Noopean Hongyothi leg. T268. Distribution. Singapore, Thailand. Remarks. This species was described from northeaster Thailand (Loei Province) and from mangroves of Singapore (Shamshev & Grootaert 2005). New data show that T. luang is found at middle altitudes of 500-850 m above sea level. Tachydromia luang is closely related to T. ocellata sp. nov. sharing similar setation of the right surstylus (several spine-like sensilla on inner side).

Tachydromia arrogans group

Tachydromia monocercus sp. nov. (Figs 17–19, 29)

Diagnosis. Antenna and palpus brownish. Scutum with 1 pair of long presutural dorsocentrals. Legs with complicated colour pattern; fore tarsomeres 4 and 5, hind tibia (except base) and hind femur on apical 2/3 brown. Wing with 2 broad brownish bands connected on cells r1 and r2+3. Halter pale.

FIGURES 17–19. Tachydromia monocercus sp. nov. 17. right epandrial lamella, ventral view. 18. epandrium with cerci. 19. left surstylus, lateral view. Scale 0.1 mm.

32 · Zootaxa 1830 © 2008 Magnolia Press SHAMSHEV & GROOTAERT Description. Male. Wing 1.8 mm. Head black in ground-colour. Eyes extending beyond ocellar tubercle. Frons shining, slightly widened toward ocellar tubercle, above antennae nearly 1.5 times as broad as anterior ocellus. Two moderately long black ocellars. Occiput shining; with 2 long cruciate black setae in upper part, several shorter setae on middle part and some setulae near neck and on lower part. Antenna entirely brownish; postpedicel small, elongate oval; stylus very long. Proboscis entirely brownish. Palpus unmodified, narrow, elongate oval, brownish, with 1 short black and 1 pale, minute subapical setae. Thorax brown in ground-colour, largely shining; prominent setae black; prothoracic sclerites (except post- pronotal lobe) finely pollinose. Postpronotal lobe large, lacking conspicuous setae. Mesonotum with 2 notopleural (posterior one longer), 1 long postalar and 2 similar scutellar setae; acrostichals lacking; dorso- centrals uniserial, very few in number, lacking on prescutellar depression, 1 presutural pair nearly as long as posterior notopleural. Legs long, with complicated colour pattern; fore femur rather brownish yellow dorsally near apex; fore tibia almost entirely brownish, paler basally; fore tarsomeres 4 and 5 brown; mid leg with similar pattern to fore leg; hind femur brown on apical 2/3; hind tibia almost entirely brownish, brownish yellow basally; hind tarsomeres 2 and 3 apically, tarsomeres 4 and 5 entirely brownish; otherwise legs yellow. Coxae with yellow- ish to brownish yellow setae of different lengths; no setae between fore and mid coxae. Fore femur slightly thickened, with minute ventral unmodified setulae and thin long setae near base. Fore tibia somewhat spindle- like, with unmodified setation. Fore basitarsus with subapical brush of black spinules ventrally. Mid femur slender; with anteroventral and posteroventral short black spinule-like setae on apical 2/3 and several pale set- ulae near base. Mid tibia slender, bearing ventral spinule-like setulae (one subapical stronger). Mid basitarsus with subapical brush of black spinules ventrally. Hind femur slender, with rows of short anteroventral and posteroventral setae. Hind tibia slender, lacking prominent bristles, with rounded posterior apical comb. Wing (Fig. 29) normally developed, rounded at apex, with typical venation for the genus; with 2 broad brownish bands connected on cells r1 and r2+3. One short basal costal seta present. Vein R1 meeting costa before wing midway. Proximal section of vein R4+5 somewhat longer than Rs. Vein R2+3 straight. Veins R4+5 and M1+2 parallel toward wing-apex. Crossveins r-m and bm-cu separated. Cells br and bm extending to 1/3 of wing. Halter pale. Abdomen brownish, subshining, covered mostly with scattered, short, black setulae; segments 7 and 8 with moderately long setae. Terminalia (Figs 17–19) moderately large, dark brown, elongate oval. Right cer- cus rather short, subrectangular, with several setae of different lengths and bearing 5 short subapical spines on inner side. Left cercus hardly prominent. Right epandrial lamella subrectangular, covered with numerous setae of different length, lacking ventral process. Right surstylus broad, rather long, curved inward, with numerous moderately long subapical setae and several spines on inner side. Left surstylus bilobed; upper lobe broad, rather long, with numerous setae of different length; lower lobe small, with several short setae. Female. Unknown. Material examined. Holotype male: THAILAND Sakon Nakhon Phu Phan NP Behind office 17°03.521'N 103°58.450'E 322 m Pan traps 13–14.vii.2006 Winlon Kongnara leg. T195. Derivatio nominis. The name alludes to the greater prominence of the right cercus. Distribution. Thailand. Remarks. Provisionally, the new species could be assigned to the T. arrogans group sensu Chvála (1970) due to possessing pollinose prothoracic sclerites, very long stylus, brownish palpi, wing with two bands and moderately large, elongate oval terminalia. However, the new species has, like the T. connexa group, 2 notopleural bristles. The main distinguishing features between T. monocercus sp. nov. and other species known from the Oriental Region are given in the key.

TACHYDROMIA THAILAND Zootaxa 1830 © 2008 Magnolia Press · 33 Tachydromia terricola group

Tachydromia doi sp. nov. (Figs 20–23, 30)

Diagnosis. Palpus yellow. Legs almost entirely yellow; fore tarsomeres 4–5 entirely and hind tibia on basal half brownish. Wing with large brownish patch on subapical part. Halter yellow. Male: mid femur and mid tibia with subbasal and subapical excision, respectively.

FIGURES 20–23. Tachydromia doi sp. nov. 20. right epandrial lamella with surstylus. 21. right cercus, lateral view. 22. hypopygium, dorsal view. 23. left surstylus. Scale 0.1 mm.

Description. Male. Wing 2.5–2.7 mm. Head black in ground-colour. Eyes extending beyond ocellar tubercle. Frons shining, slightly widened toward ocellar tubercle, above antennae nearly as broad as anterior ocellus. Two pairs of moderately long ocellars. Occiput shining; with 2 long dark setae in upper part, several similar setae on middle part laterally and some pale hair-like setae near neck and on lower part. Basal antennal segments brownish yellow; postpedicel paler, yellowish, short, oval; stylus long, brownish, short pubescent. Proboscis entirely brown. Palpus unmodified, elongate, yellow, with 1 long yellow subapical seta. Thorax brown in ground-colour, largely shining; prominent setae black; prothoracic sclerites (except post- pronotal lobe) pollinose. Postpronotal lobe large, lacking conspicuous setae. Mesonotum with 1 thin notopleu- ral, 1 postalar and 2 scutellar setae; acrostichals lacking; dorsocentrals uniserial, few in number, mostly minute, 1 presutural pair long (as long as scutellars). Legs long, almost entirely yellow; fore tibia brownish yellow anteriorly and posteriorly; fore tarsomeres 4 and 5 brown (except extreme base); mid and hind tarsomere 4 brownish yellow, tarsomere 5 brownish; hind tibia brownish on basal half. Coxae with pale hair-like setae of different length; no setae between fore and mid coxae. Fore femur strongly thickened, with anteroventral and posteroventral short spinule-like setulae, bear- ing 2 long setae near base. Fore tibia evenly swollen, with subapical ventral comb of several black setulae. Fore tarsomere 1 with patch of black ventral spinules on subapical portion. Mid femur slightly thickened, with shallow excavation and small ventral swelling nearer to base, bearing 2 dark spinules sitting on swelling

34 · Zootaxa 1830 © 2008 Magnolia Press SHAMSHEV & GROOTAERT (before excavation) and 1 smaller dark spinule beyond excavation, with 2 long yellowish setae near base and some anteroventral and posteroventral spinules on apical part. Mid tibia with shallow excision on apical part, bearing some ventral spinule-like setulae before excision. Hind femur slender, with rows of short anterodorsal and posteroventral setae. Hind tibia slender, lacking prominent bristles, with narrow pointed posterior apical comb. Wing (Fig. 30) normally developed, rounded at apex, with typical venation for the genus; largely hyaline, with large brownish patch near meeting point of vein R2+3 with costa, which becomes gradually paler toward lower margin of the wing. One short basal costal seta present. Vein R1 meeting costa somewhat before wing midway. Proximal section of vein R4+5 subequal to Rs. Vein R2+3 straight. Veins R4+5 and M1+2 parallel toward wing-apex. Crossveins r-m and bm-cu separated. Cells br and bm extending to 1/3 of wing. Halter yellow. Abdomen brown, subshining, covered mostly with scattered, short, black setulae; segments 7 and 8 with moderately long setae. Terminalia (Figs 20–23) moderately large, dark brown, elongate oval. Right cercus small, deeply concave apically, with scattered short setulae. Left cercus rather short, subrectangular (subtrian- gular viewed laterally), with numerous short setae on subapical part. Right epandrial lamella subglobular (viewed laterally), covered with numerous setae of different lengths. Right surstylus rather short, bilobed; upper lobe bearing long black subapical spine; lower lobe curved inward, of complicated structure. Left surst- ylus as in Fig. 23.

FIGURES 24–31. Wings. 24. Tachydromia phanensis sp. nov. 25. Tachydromia phu sp. nov. 26. Tachydromia tigeri sp. nov. 27. Tachydromia ocellata sp. nov. male. 28. Tachydromia luang Shamshev, Grootaert. 29. Tachydromia monocercus sp. nov. 30. Tachydromia doi sp. nov. 31. Tachydromia ocellata sp. nov. female.

TACHYDROMIA THAILAND Zootaxa 1830 © 2008 Magnolia Press · 35 Female. Mid femur unmodified, with rows of anteroventral and posteroventral thin setae, the latter some- what longer; mid tibia unmodified. Otherwise as in male. Cercus brownish, long. Material examined. Holotype male: THAILAND Chiang Mai prov., Doi Inthanon NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 13–21.ix.2006 Y. Areeluck leg. T252. Paratypes: 2 males, 6 females, THAILAND Chiang Mai prov., Doi Inthanon NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 19–26.x.2006 Y. Areeluck leg. T374; 1 male, 2 females, Doi Inthanon NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 26.x–2.xi.2006 Y. Areeluck leg. T380; 2 males, 5 females, Doi Inthanon NP summit marsh 18°35.361'N 98°29.157'E 2500 m Malaise trap 5– 12.x.2006 Y. Areeluck leg. T362. Derivatio nominis. The name “doi” means mountain in northern Thai language. Distribution. Thailand.

Remarks. On the basis of pollinose prothoracic sclerites, straight vein R2+3, yellow palpi, 1 notopleural bristle and almost entire yellow legs the new species belongs to the T. terricola group sensu Chvála (1970) and it is similar to T. terricoloides Shamshev & Grootaert described from Loei Province of Thailand (Sham- shev & Grootaert 2005). The main distinguishing features between these species are given in the key.

Acknowledgements

The Thailand Insect Group for Entomological Research (TIGER) Project is supported in part by the USA National Science Foundation (DEB-0542864). We thank Brian Brown (Natural History Museum of Los Angeles County, USA), Michael Sharkey (University of Kentucky, USA) and Adrian Plant (National Museum and Galleries of Wales, Cardiff, UK) for arranging the loan of material and Wendy Porras (Instituto Nacional de Biodiversidad, Costa Rica) for pre-sorting samples to family level. The authors are indebted to Jeffrey Cumming (Agriculture and Agri-Food Canada, Ottawa) and Andreas Stark (Halle (Saale)) for valuable com- ments.

References

Chvála, M. (1970) Revision of Palaearctic species of the genus Tachydromia Meig. (= Tachista Loew) (Diptera, Empid- idae). Acta Entomologica Musei Nacionalis Pragae, 38 (1969), 415–524. Chvála, M. (1975) The (Dipt. Empididae) of Fennoscandia and Denmark. Fauna Entomologica Scandi- navica, 3, 1–336. McAlpine, J.F. (1981) Morphology and terminology—Adults [Chapter] 2. In McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Coords.), Manual of Nearctic Diptera, Volume 1. Agriculture Canada Monograph, 27, 9–63. Saigusa, T. & Yang D. (2002) Empididae (Diptera) from Funiu Mountains, Henan, China (I). Studia Dipterologica, 9, 519–543. Shamshev, I. & Grootaert, P. (2005) The genus Tachydromia Meigen (Diptera: Hybotidae) from South East Asia, with the description of three new species. Studia Dipterologica, 12, 109–117. Sinclair, B.J. & Cumming, J.M. (2006) The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera). Zootaxa, 1180, 1–172. Stuckenberg, B.R. (1999) Antennal evolution in the Brachycera (Diptera), with a reassessment of terminology relating to the flagellum. Studia Dipterologica, 6, 33–48. Yang D. & Grootaert, P. (2006) Notes on Tachydromia from China (Diptera: Hybotidae). Transactions of the American Entomological Society, 132, 133–135.

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