Research Article |
Corresponding author: Patrick Grootaert ( patrick.grootaert@naturalsciences.be ) Academic editor: Marija Ivković
© 2018 Chufei Tang, Ding Yang, Patrick Grootaert.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Tang C, Yang D, Grootaert P (2018) Revision of the genus Lichtwardtia Enderlein in Southeast Asia, a tale of highly diverse male terminalia (Diptera, Dolichopodidae). ZooKeys 798: 63-107. https://doi.org/10.3897/zookeys.798.28107
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In the present paper the Oriental species of the genus Lichtwardtia Enderlein, 1912 are revised based on the type material of known species and new material from Singapore and Cambodia. A re-description and illustration of the holotype female of Lichtwardtia ziczac (Wiedemann, 1824) is given but since it has been described on the basis of a female only and its provenance India Orientalis is only a vague indication of its type locality, it is considered as a nomen dubium. All the species put as junior synonyms by
Although Lichtwardtia is a common genus in Southeast Asia it is generally not abundant locally. It is often found in anthropogenic disturbed habitats only. Four species are recorded from Singapore while eight species are sympatric and very abundant at the locality of Siem Reap in Cambodia.
Lichtwardtia , new species, Oriental, revision
With its bayonet-like vein M and long soft hairs on the apical segment of the arista-like stylus, the genus Lichtwardtia Enderlein is easily recognised as such, but assigning specimens to the correct species is another story. Even the validity of the genus Lichtwardtia itself has been uncertain since its establishment (
There are 22 species known in the world: 16 from the Afrotropical Region (
The taxonomy of Lichtwardtia suffered from the description of species based on females only. Pairing males to these females became almost impossible as females of Lichtwardtia barely differ without careful observation and dissection, which also raises the concern of synonyms. Typically, many species were set to be the synonym of the iconic species L. ziczac (Wiedemann) that itself was described on the basis of a female. Its provenance “India orientalis” is vague since in the past this area extended from Pakistan in the West to New Guinea in the East. In addition, the single type of L. coxalis Kertész, 1901 is probably destroyed (
Techniques to study ancient DNA might one day elucidate its status and that of related species. Wing interference patterns (WIPs) might be a tool which would help pairing the specimens. It has been proved useful on the taxonomy of Campsicnemus, which is another genus of Dolichopodidae. However, this could not fully support the identification independently. Often the pattern arrangements between females and males are still at ‘similar’ level, not exactly the same. Here we do not consider this a reliable method for identification because there is no standard at this stage. Meanwhile, the WIPs of Lichtwardtia between different species seem not as distinguishably different as those of Campsicnemus. The level of similarity that could confirm the pairing to the level of species should not only be determined by the study of one single genus but extended to other genera.
In the present paper we revise the known Oriental fauna and add eight species new for science. With illustrations of the male terminalia we hope to provide a framework to recognise well-defined morpho-species. In addition, some (COI) barcodes (600 bp) and NGS barcodes (313 bp) are provided for all the species with fresh specimens available (uploaded to GenBank, with accession number MH536852–MH536856). In contrast to what we originally thought, the genus Lichtwardtia is as diverse in the Oriental region as it is in the Afrotropical region (
In addition to the type material of the previously described species, new Lichtwardtia specimens were collected during a two-year survey with Malaise traps at the temple site of Preah Khan, a temple at the Angkor site (Siem Reap, Cambodia) and in the garden of the Sam Vesna Centre in Siem Reap. The site of Preah Khan was situated along a path bordering a secondary forest (Figure
Morphological terminology for adult structures mainly follows McAlpine (1981), and the structures of the male genitalia follow Cumming and Wood (2009). Photos were taken with a Canon EOS 600D camera at the Royal Belgian Institute of Natural Sciences and then stacked by Helicon Focus 6.0 or with the Visionary Digital BK Plus Lab System at the Laboratory for Evolutionary Biology (NUS).The type material is deposited at the Royal Belgian Institute of Natural Sciences (
COI barcoding was done following the techniques described in Lim et al. (2010) and by
As for the other old type material seen in this study, we preferred not to dissect the specimens for examination as the features on the body are enough for identification and the specimens might be examined in the near future with less destructive techniques. Abbreviations in the text are as follows:
a anterior;
acr acrostichal bristle(s);
ad anterodorsal bristle(s);
av anteroventral bristle(s);
C costal vein;
d dorsal bristle(s);
dc dorsocentral bristle(s);
hm postpronotal bristle(s);
npl notopleural bristle(s);
p posterior bristle(s);
pa postalar bristle(s);
pd posterodorsal bristle(s);
pm presutural supraalar bristle(s);
pv posteroventral bristle(s);
sa postsutural supraalar bristle(s);
sc scutellar bristle(s);
sr presutural intraalar bristle(s);
v ventral bristle(s).
Abbreviations on the figures are as follows:
ap apical bristle(s);
bvl basoventral epandrial lobe;
dsur dorsal surstylus;
eb epandrial bristles;
hy hypandrium;
L left hand side;
ph phallus;
pg postgonite;
R right hand side;
vsur ventral surstylus;
sur surstylus.
Lichtwardtia Enderlein, 1912: 406. Type species: Lichtwardtia formosana Enderlein, 1912 (original designation).
Vaalimyia Curran, 1926: 398. Type species: Vaalimyia violacea Curran, 1926 [= Dolichopus angularis Macquart, 1842] (original designation).
The generic synonymy list is as given by Yang et al. (2005) except that Lichtwardtia formosana Enderlein, 1912 was considered to be a junior synonym of L. ziczac (
Small to medium-sized species (3.5–5.0 mm). Head: overall dark metallic green including vertex, with thick pale pollinosity; face wide, slightly narrow at middle, slightly raised from lateral view, frons and face both with thick pale pollinosity that hiding the ground colour, face parallel sided, but slightly narrowed near mid-length. Hairs and bristles on head black but lower postocular bristles pale. Vertex flat. Ocellar tubercle distinct. With pairs of strong vertical and diverging ocellar bristles present, with weak postvertical bristles which are approx. a half-length of vertical bristles. Antenna wholly or mostly yellow; scape with short dorsal bristles, longer than pedicel; arista-like stylus dorsal, two-segmented, with feather-like long hairs on apical segment. Eyes dichoptic, with hairs between facets. Clypeus long and wide.
Thorax: dark metallic blue-black. Metapleuron with one narrow black stripe. Hairs and bristles on thorax black. Acr biseriate, hair-like. Five strong pairs of dc. With one pa, two sa, one sr, one hm, one pm, two npl. Propleuron with long curved bristle just above base of fore coxa. With hairs anterior to spiracle. Scutellum with two pairs sc, apical pair long and strong, basal pair short and weak. Mid and hind coxae with outer bristle(s). Fore femur without distinct bristle. Mid and hind femora each with preapical bristle. Hind femur thick, 5.0 times longer than wide. Hind tarsomere I with strong dorsal bristles, shorter than hind tarsomere II.
Wing: usually clear, sometimes with faint brown clouding around distal vein M and dm-cu crossvein, occasionally partly smoky. Vein costa sometimes widened at the joint with R1, with various callus. M with fading M2, M1 with one short subvein. Crossvein dm-cu straight. Vein M joining margin just before apex. CuAx around 1.0.
Abdomen: metallic green (in Oriental), nearly 1.5 times longer than wide, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia: Epandrium distinctly longer than wides. Epandrial lobe with three long pale bristles Cercus nearly triangular, margin rounded, usually pale with broad black outer margin, with weak digitations around outer margin, with black simple or specialised marginal bristles on digitations. Hypandrium and phallus various, often with various denticles.
The cercus is more or less triangular, bordered by some strong marginal bristles that are strongly flattened or truncate. All cercus have one or two large inside bristles near the dorsal border. The postgonite is broad and the tip is bifid. The tip of the phallus is ventrally denticulate and the hypandrium is a simple tube.
Holotype male (coll.
Wing entirely hyaline; with a slight swelling of the costa where R1 merges with costa. Postpedicel mainly dark yellow but blackish on apically half, nearly as long as wide. Mid coxa with a dark brown stripe anteriorly, a paler brown band posteriorly. Hind coxa entirely yellow. Cercus rounded, marginal bristles black, not flattened nor on tubercles. Hypandrium simple and smooth, with no denticle; phallus with double rows of spinules on ventral half.
Male.Body length 3.4–3.7 mm, wing 3.4–3.5 × 1.0 mm.
Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black but lower postocular bristles pale. Antenna dark yellow; postpedicel blackish on apical half, blunt apically, nearly as long as wide arista-like stylus dark brown, nearly as long as width of head, feather-like, with long pubescence, basal portion 0.2 times as long as apical portion. Proboscis dark yellow, with black hairs; palpus dark yellow, with1 short black apical bristle.
Thorax dark green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of long acr. Scutellum with two pairs sc, apical pair long and strong, basal pair short and weak. Legs mainly yellow; but mid coxa with two large brown square spots laterally: a dark brown stripe anteriorly, a paler brown band posteriorly; mid and hind tarsomeres I–II blackish on tip, mid and hind tarsomere III blackish at apical half, mid and hind tarsomeres IV–V wholly black. Fore and mid coxae anteriorly with rows of bristle-like hairs, fore coxa anteriorly with three long strong marginal bristles, mid coxa with two outer bristles and two marginal bristles, hind coxa with two outer bristles, basal one strong, apical one smaller. Mid trochanter with three short ap, all dorsal. Hind trochanter with two weak ap, one dorsal, one posteroventer. Fore femur with one short weak av. Mid femur with one strong pd and one preapical pv in normal strength. Hind femur with one strong pd at apical quarter. Fore tibia with four ad (of which one strong and preapical), one av and three ap. Mid tibia with two ad, three pd, one pv, and four ap. Hind tibia with four ad, four pd, two pv and three ap. Fore tarsomere I with one short av at base. Hind tarsomere I with one strong ad and two short av. Relative lengths of tibia and five tarsomeres of legs LI : 7.0 : 5.0 : 2.5 : 2.0 : 1.2 : 1.0; LII : 15.0 : 7.5 : 3.8 : 3.0 : 2.0 : 1.5; LIII : 16.2 : 7.0 : 7.5 : 5.0 : 2.2 : 1.5. Wing nearly hyaline, veins brown. Costa slightly widened at the joint with R1. M with fading M2, M1 with one short subvein. Crossvein dm-cu straight. CuAx ratio 1.0. Lower calypter pale with black hairs. Haltere pale.
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia (Figure
Female. Similar to male in size and morphology except for the male terminalia.
This new species is named after the country (Cambodia) where it was found.
Lichtwardtia cambodiensis sp. n. belongs to the L. nodulata group and thus is related to L. dentalis and L. semakau sp. n. The latter two species both have distinctly darkened and flattened marginal bristles on the cercus while L. cambodiensis sp. n. only has thin, and weak marginal bristles on the cercus. Further, L. cambodiensis has a weak swelling on the joint point of wing vein R1 and costa, whereas the swelling is distinct in L. dentalis, and absent in L. semakau.
Lichtwardtia dentalis Zhang, Masunaga & Yang, 2009: 198, figs 1–4.
CAMBODIA (all coll.
THAILAND: 1 male (coll.
Costa swollen before and at the level where R1 joins the costa (Figure
We refer to the detailed description of L. dentalis in
The specimens from northern Thailand and central Cambodia correspond entirely to the description and figures given by
L. dentalis is very closely related to L. semakau sp. n. from Singapore and we refer to the comments under the latter species.
Holotype male (coll. LKCNHM): SINGAPORE: Semakau, 30 August 2012, sweep netting (leg. Jayanthi Puniamoorthy & P Grootaert). Paratypes (all coll. LKCNHM): SINGAPORE: 1 male, Pulau Ubin, 14 July 2012, sweep netting in mangrove (leg. Jayanthi Puniamoorthy & P Grootaert); 9 males, 14 females, Kranji nature trail, 27 July 2005, sweep netting in park (leg. P Grootaert); 1 male, Semakau, 13 December 2012, sweep netting in halophilous vegetation on sandy beach (leg. P Grootaert).
Basal 7/8 of mid coxa and basal half of hind coxa brown, mid coxa with one black band at middle. Vein R1 with one oval thickness at where connect to C. Phallus with two rows of black denticles on ventral half, anterior row only with four small sparse denticles, posterior row with eight dense denticles.
Male.Body length 3.6–3.8 mm, wing 3.2–3.3 × 1.0 mm.
Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black except posteroventral hairs pale. Antenna dark yellow; postpedicel nearly triangular, blunt apically, nearly as long as wide, with short black pubescence; arista-like stylus dorsal, inserted at basal half of postpedicel, nearly as long as head width, black, feather-like, with long black pubescence, basal segment 0.3 times as long as apical portion of arista-like stylus. Proboscis brown, with black hairs; palpus dark brown, with one short black apical bristle.
Thorax dark metallic green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of acr. Scutellum with two pairs sc, apical pair long strong, basal pair short and weak. Legs mainly dark yellow, but base of fore coxa, basal 7/8 of mid coxa and basal half of hind coxa brown, mid coxa with one black band at middle. Fore coxa anteriorly with three strong bristle at apical half, mid coxa with cluster of anterior hairs three marginal bristles anteriorly and one long strong outer bristle, hind coxa with one long strong outer bristle. Mid trochanter with two weak apical bristle dorsally. Fore femur without distinct bristle. Mid and hind femora each with one preapical pv. Fore tibia with one row of two ad, two pd, one pv and three ap. Mid tibia with three ad, four pd, two pv and four ap. Hind tibia with four ad and three ap. Mid tarsomere I with one short strong pd. Hind tarsomere I with one short strong ad at middle and one av at basal quarter. Relative lengths of tibia and five tarsomeres of legs LI : 6.6 : 3.3 : 1.6 : 1.3 : 1.0 : 1.0; LII : 10.6 : 5.0 : 2.0 : ? : ? : ? (mid tarsus partly lost); LIII : 10.0 : 4.0 : 5.0 : 3.3 : 2.0 : 1.3. Wing nearly hyaline, tinged brown; veins brown. R1 with one oval thickness at where connect to C. M with fading M2, M1 with one short subvein. Crossvein dm-cu somewhat arched. CuAx ratio 1.4. Lower calypter pale with black hairs. Haltere pale.
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male genitalia: Epandrium 1.7 times longer than wide; epandrial lobe with three pale ap. Ventral surstylus and dorsal surstylus both with five short nail-like ap. Cercus nearly triangular, pale except the black ring, with weak digitations around outer margin, with long strong black marginal bristle on digitations, with one long strong bristles on digitation at apical half. Hypandrium simple. Phallus with two rows of black denticles on ventral half, anterior row only with four small sparse denticles, posterior row with eight dense denticles.
Female. Similar to male in size and morphology except for the male terminalia.
The name refers to the swelling of the costa.
The new species should be compared to L. hirsutiseta since there is a node on the costa that is however distinctly before R1 joins the costa (Figure
Holotype male (coll. LKCNHM): SINGAPORE: Semakau, back mangrove in the old mangrove (1°12'19.9"N, 103°45'34.1"E), 3 April 2012, sweep netting along path (leg. P Grootaert). Paratype (coll. LKCNHM): 1 female, same provenance as holotype.
Antenna entirely dark yellow. Arista-like stylus feathered. Wing clear, faintly brownish tinged. No thickening of the costa where R1 joins the costa. Fore and hind coxae entirely yellow. Mid coxa with a rectangular brown spot anteriorly, posteriorly pale brownish. Cercus yellow, brownish seamed with margin distinctly digitated and bearing five thickened marginal bristles near the tip. Hypandrium simple, lacking protuberances. Phallus ventrally with at least twelve black denticles.
Male.Body length 3.9 mm, wing 3.5 × 1.2 mm. Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black but lower postocular hairs pale. Antenna dark yellow (Figure
Thorax dark green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of long acr. Scutellum with two pairs sc, apical pair long strong, basal pair short and weak. Legs mainly yellow, but mid coxa with a dark brown rectangular spot anteriorly and a pale brown posteriorly. Fore and mid coxae anteriorly with rows of bristle-like hairs, fore and mid coxae anteriorly with three long strong preapical bristles and rows of anterior bristles, mid coxa with one outer bristles at apical third, hind coxa with two outer bristles, basal one strong, apical one relatively weak. Mid trochanter with two ap dorsally. Hind trochanter with one outer bristle at middle. Fore femur with one strong pv at middle. Mid femur with one strong pd and one preapical pv. Hind femur with one strong pd at apical quarter. Fore tibia with one ad, three pd, one av and three ap. Mid tibia with two ad, three pd, one pv and four ap. Hind tibia with three ad, three pd (one preapical), two weak pv and three ap. Fore tarsomere I with one short av at base. Hind tarsomere I with one strong ad at middle and two short apical bristle. Relative lengths of tibia and five tarsomeres of legs LI : 7.0 : 3.2 : 1.2 : 1.2 : 1.0 : 1.0; LII : 6.0 : 4.0 : ? : ? : ? : ? (mid tarsus partly lost); LIII : 10.4 : 4.0 : ? : ? : ? : ?. Wing nearly hyaline, nearly pale; veins brown. No thickening of the costa where R1 joins the costa. M with fading M2, M1 with one short subvein. Crossvein dm-cu straight. CuAx ratio 1.2. Lower calypter pale with black hairs. Haltere pale.
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia (Figure
Female. Body length 4.3 mm, wing 3.4 × 1.3 mm. Resembling the male except for the terminalia and face wider.
The name refers to the type locality Semakau, an island on the southern coast of Singapore.
L. semakau sp. n. resembles very much L. dentalis
L. semakau sp. n. should be compared also to L. cambodiensis sp. n. that differs in the male also by a thickening of a costa and the marginal bristles on the cercus that are all thin and paler.
Holotype male (coll. LKCNHM): SINGAPORE: West Coast Park, 7 December 2003, shrubs along sandy beach (leg. P Grootaert), sweeping. Paratype (coll. LKCNHM): 1 female; collecting information same to the holotype
Wings clear, but having the cross veins brownish seamed. Mid coxa brown with one anterior dark brown stripe and one lateral band, both dark brown. Hind coxa yellow. Hypandrium simple. Phallus with one row of five big clear regular black denticles ventrally, with one blunt denticle hidden in hypandrium.
Male.Body length 3.7 mm, wing 3.4 × 1.1 mm.
Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black except posteroventral hairs pale. Antenna dark yellow; postpedicel nearly triangular, blunt apically, 1.2 times as long as wide, with pale pubescence; arista-like stylus dorsal, inserted at basal half of postpedicel, nearly as long as head width, dark yellow, feather-like, with long black pubescence, basal segment 0.3 times as long as apical portion of arista-like stylus. Proboscis dark yellow, with black hairs; palpus dark yellow, with one black apical bristle.
Thorax dark green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of strong acr. Scutellum with two pairs sc, apical pair long strong, basal pair short and weak. Legs mainly yellow, but fore coxa somewhat brownish at base, mid coxa brown with one anterior dark brown stripe and one lateral band, both dark brown, hind coxae dark yellow, lightly brownish at basal half; mid and hind tarsi brown from tip of tarsomere II to tarsomere V. Fore and mid coxae anteriorly with row of thin bristles and four strong bristles at apical half, hind coxa with two short outer bristles. Fore trochanter with three weak outer bristle at middle. Mid trochanter with two short weak outer bristles. Fore femur with one short weak av. Mid femur with one strong pd at apical 1/6 and one preapical pv. Hind femur with one strong pd at apical quarter. Fore tibia with two pd, one av, and one pv and three short ap. Mid tibia with two ad, four pd (basal 1st relatively weak), one pv at middle and four ap. Hind tibia with four ad, four pd, two pv and three ap. Fore tarsomere I with one short av at base. Hind tarsomere I with one short strong ad at middle, one pv at basal fifth and two short apical bristle. Relative lengths of tibia and five tarsomeres of legs LI : 6.0 : 3.0 : 1.6 : 1.2 : 1.0 : 1.0; LII : 8.0 : 5.0 : 2.0 : 1.6 : 1.2 : 1.0; LIII : 10.0 : 4.0 : 4.0 : 3.0 : 2.0 : 1.4. Wing nearly hyaline, cross veins seamed brownish; veins brown. M with fading M2, M1 with one short subvein. Crossvein dm-cu straight. CuAx ratio 1.0. Lower calypter pale with black hairs. Haltere pale.
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia: Epandrium 1.8 times longer than wide; epandrial lobe with three pale ap. Surstylus with five short nail-like ap. Cercus nearly triangular, pale, covered by thin pale bristles, with weak digitations around outer margin, with long strong pale marginal bristle on digitations. Hypandrium simple. Phallus with one row of five big clear regular black denticles ventrally, with one blunt denticle hidden in hypandrium.
Female. Body length 3.6–4.0 mm, wing length 3.1–3.2 × 1.1 mm. Very similar to male, but postpedicel as long as wide. Wing hyaline with anterior border faintly brownish and cross veins brownish seamed. No swelling of the costa before or at the point where R1 joins the costa. The ratio of the proximal section of M1, and the distal section is 0.4/0.6 (Figure
This species is comparable to L. infuscata sp. n. but the latter has quite brownish wings also with the cross veins seamed brown. Both species have also the hind coxa entirely yellow and the phallus bears ventrally rather strong spines. In L. singaporensis sp. n. there are five strong spines while in L. infuscata there is a double row of ten denticles that are smaller than in L. singaporensis sp. n. The epandrium has a pointed tip while its tip in L. infuscata sp. n. is truncated. The size of the cercus is larger in L. infuscata sp. n. L. infuscata has a pale ventral protuberance with a black seam on the epandrium on which the phallus-hypandrial complex is resting. This protuberance is absent in L. singaporensis sp. n. The brownish seams along the cross veins suggest that L. singaporensis sp. n. represents the enigmatic L. ziczac. However we refrain from given the description of the male of L. singaporensis sp. n. as the true L. ziczac. See comments under L. ziczac.
The hypandrium has a bifurcated tip and a curved appendage. The phallus has a preapical ventral process.
Holotype male (coll.
Antenna nearly entirely dark yellow. Wing brownish tinged, a distinct swelling of the costa present just before R1 joins the costa. Mid coxa with two light brown bands laterally. Hind coxa entirely yellow. Cercus with elongate tip bearing a longer bristle than the other marginals; hypandrium with bifurcate tip and a black curved appendage on the right-hand side. Phallus wide, with preapical ventral process.
Male.Body length 3.7 mm, wing 3.5 × 1.1 mm.
Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black; upper four postocular bristles black but all lower postoculars pale. Antenna dark yellow except portion at level of arista-like stylus slightly brownish. Postpedicel nearly triangular, blunt apically, with brown pubescence, nearly as long as wide; arista-like stylus dark brown, nearly as long as width of head, feather-like, with long pubescence, basal portion 0.2 times as long as apical portion. Proboscis dark yellow, with black hairs; palpus dark yellow, with one short weak black apical bristle.
Thorax dark green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of long acr. Scutellum with two pairs sc, apical pair long and strong, basal pair very short and weak. Legs mainly yellow, but mid coxa with two light brown bands laterally, hind coxa entirely yellow. Fore and mid coxae anteriorly to laterally with rows of bristle-like hairs and six marginal bristles; mid coxa with one outer bristles at apical third, hind coxa with two outer bristles, basal one strong, apical one weak. Mid trochanter with two ap. Hind trochanter with one outer hair at middle. Fore femur without distinct bristles. Mid femur with one strong ad and one preapical pv. Hind femur with one strong anterior on apical fourth. Fore tibia with three ad, one av and three ap (of which one posterodorsal very long). Mid tibia with two ad, five pd, two pv, and four ap. Hind tibia with three ad, two pd, one pv and four ap. Fore tarsomere I with one short av at base. Hind tarsomere I with one strong dorsal beyond middle, two short av close to base and two short apical bristle. Relative lengths of tibia and five tarsomeres of legs LI : 10.0 : 4.0 : 2.5 : 2.0 : 1.5 : 1.5; LII : 16.0 : 7.5 : 4.0 : 3.0 : 1.5 : 1.0; LIII : 18.0 : 7.5 : 7.5 : 6.0 : 4.0 : 2.5. Wing brownish tinged, veins brown. M with fading M2, M1 with one short subvein. Crossvein dm-cu straight. CuAx ratio 1.3. Lower calypter pale with long black hairs. Haltere white.
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia (Figure
Female. Unknown.
This species resembles superficially L. polychroma in having a swelling of the costa and by the similar elongate cercus and the complicated structure of the tip of the hypandrium. However, L. polychroma possesses a large brown tooth at the right side. L. conspicabilis sp. n. has a curved black extension of the left side and the tip of the phallus is deeply indented.
The name conspicabilis refers to the remarkable structure of the hypandrium.
Cambodia.
Holotype male (coll.
Postpedicel of antenna mostly brownish yellow. Wing faintly tinged brownish, no swelling of the costa present. Mid coxa with two brown bands laterally, one brown, one light brown. Hind coxa entirely yellow. Tip of hypandrium bifurcate with a black narrow ventral arm that is strongly toothed and a broad arm dorsally yellowish at base and black at tip; apex rounded, bordered with teeth. Phallus yellowish with a very strong yellowish brown dorsal hook near middle resting in the ventral cavity of the epandrium; apical half of phallus, with a ventral curved followed by a strong dorsal curved; tip pointed.
Male.Body length 4.0 mm, wing 3.8 × 1.3 mm. Head with frons shining metallic green; face slightly raised, as wide as postpedicel, frons and face covered with a thick white dusting. Hairs and bristles on head black; upper five postoculars black, lower postocular bristles pale. Antenna mainly yellow, postpedicel largely brownish yellow; postpedicel nearly triangular, blunt apically, with yellow pubescence, 1.2 times as long as wide; arista-like stylus dark yellow, nearly as long as width of head, feather-like, basal portion 0.5 times as long as apical portion. Proboscis dark yellow, with black hairs; palpus dark yellow, with one short weak black apical bristle.
Thorax dark metallic green, with a fine pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of acr half as long as dc. Scutellum with two pairs sc, apical pair long strong, basal pair short and weak. Legs mainly yellow, but mid coxa with two brown bands laterally, one brown, one light brown. Fore and mid coxae anteriorly to laterally with rows of bristle-like hairs and eight preapical bristles; mid coxa with one strong outer bristles at apical third, hind coxa with two outer bristles, basal one strong, apical one relatively weak. Mid trochanter with two dorsal ap. Hind trochanter with one outer bristle at middle. Fore femur without distinct bristles. Mid femur with one strong ad and a weak pd. Hind femur with one strong ad at apical quarter. Fore tibia with two ad, two pd, one av and four ap. Mid tibia with two ad, three pd, one pv and four ap. Hind tibia with four ad, four d and four ap; ventrally in basal third with a row of small erect hairs. Fore and mid tarsomere I without distinct bristles. Hind tarsomere I with one strong d at apical third. Relative lengths of tibia and five tarsomeres of legs LI : 8.2 : 5.0 : 2.0 : 1.6 : 1.0 : 1.0; LII : 13.2 : 6.6 : 3.3 : 2.6 : 2.0 : 1.3; LIII : 13.2 : 6.0 : 6.0 : 4.3 : 3.3 : 1.6. Wing nearly hyaline, tinged brown; veins brown. M with fading M2, M1 with one short subvein. Crossvein dm-cu straight. CuAx ratio 1.0. Lower calypter pale with black hairs. Haltere pale.
Abdomen metallic green, with fine pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia (Figure
Female. Unknown.
The species name alludes to the monstrous appendages on the male terminalia.
The male is easily recognised having these huge extensions on the terminalia. The very strong dorsal tooth on the phallus resembles superficially the strong tooth present on the tip of the hypandrium of L. polychroma and L. zhangae. However the origin of these structure is different: in L. monstruosa sp. n., it is on the phallus, while in the others it is on the hypandrium.
The cercus is more or less triangular, with some strong flattened marginals and a dorsal marginal that shifted to the inside. The postgonite is tubiform. The tip of the phallus has no denticle. The hypandrium has a strong subapical dorsal hook.
Rhagoneurus
polychromus
Loew, 1864: 346, Fig.
There is a single male conserved in the collection of Becker (MfN, Berlin) bearing the label “Rhaconeurus polydromus m” in the handwriting of Loew (Figure
The information on the locality is new because Loew did not give a precise locality in his description. Rambodda (nowadays cited as Ramboda) is a small village in Sri Lanka known for its famous waterfalls. Johannes (John) Nietner (died 1874) was a German naturalist chiefly interested in botany and entomology. Born in Potsdam, he was a plantation owner in Rambodda, Ceylon and described many new insect species from the island. Having a special interest in insects, he made large collections and sent specimens for study by experts abroad. Collections from him are in the Deutschen Entomologischen Institut, the Museum für Naturkunde in Berlin, in the Naturhistorischen Museum in Vienna and the Natural History Museum in London.
We designate the male as lectotype since Loew did not designate a holotype. A female was included in the description but we failed to find it. Stacked images of the lectotype male were provided by the courtesy of Mr. Bernhard Schurian and Sven Marotske (MfN, Berlin).
CAMBODIA (all coll.
Antenna largely yellow, legs yellow (Figs
Male.Body length 4.2 mm, wing 3.8 × 1.3 mm.
Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black except postocular bristles yellow. Antenna yellow; postpedicel with extreme tip and dorsal margin brownish; elongate triangular, blunt at tip, nearly as long as wide; arista-like stylus with long densely set hairs. Proboscis dark yellow, with short black hairs; palpus, dark yellow with one black apical bristle.
Thorax dark green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of strong acr, with dense short strong bristles at anterior portion. Scutellum with two pairs sc, apical pair long strong, basal pair short and weak. Legs mainly yellow. Fore and hind coxa entirely yellow, but mid coxa with a black band anteriorly and a broad band posteriorly. Fore coxa anteriorly at base with a few short erect bristles, anteriorly densely covered with short black bristle-like hairs, four very long ap and a few shorter bristles. Mid coxa anteriorly densely covered with short black hairs, with a long outer bristle at the tip of the blackish band; hind coxa with two outer bristles, basal one strong, apical one short and weak. Mid and hind trochanters both with several short weak hairs. Fore femur lacking ventral bristles. Mid femur with one preapical pv. Hind femur with one strong ad at apical quarter. Fore tibia with two ad, two pd, one av, and three ap. Mid tibia with two ad, three pd, one av, and four ap; all long strong. Hind tibia with two ad, four pd, one pv, and three ap; all long. Hind tarsomere I with one strong ad at middle, one short strong ad at basal third and two short apical bristle. Relative lengths of tibia and five tarsomeres of legs LI : 9.0 : 6.0 : 2.4 : 1.2 : 1.0 : 1.0; LII : 16.0 : 8.0 : 5.0 : 4.0 : 2.4 : 1.6; LIII : 18.0 : 6.0 : 4.0 : 4.0 : ? : ?. Wing nearly hyaline, tinged brownish, veins brown. M with fading M2, M1 with one short subvein. Crossvein dm-cu straight. CuAx ratio 1.1. Lower calypter pale with black hairs. Haltere pale.
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia. Epandrium 1.9 times longer than wide (Figure
Female. Unknown.
Cambodia, Sri Lanka.
Lichtwardtia ziczac (Wiedemann, 1824) sensu Zhang, Masunaga & Yang, 2009: 199, figs 11–14. Re-description.
This species is dedicated to Dr. Lili Zhang of the IOZ Museum in Beijing who re-described and illustrated Lichtwardtia ziczac (Wiedemann) for the first time with detailed drawings.
Costa without swelling. Hypandrium with a large brown dorsal preapical tooth. Phallus smooth.
For a full description we refer to
In having the cross veins clouded, Lichtwardtia ziczac (Wiedemann) is distinctly different from L. ziczac sensu
Bali (Indonesia).
Lichtwardtia formosana Enderlein, 1912: 407. Type locality: Taiwan (China).
Lichtwardtia taiwanensis Zhang, Masunaga & Yang, 2009: 199, figs 6–10. Type locality: Taiwan (China).
Lichtwardtia formosana Enderlein, 1912, in Selivanova, Negrobov & Yang, 2010: 144, re-description, figs 1–5.
CAMBODIA (all coll.
SINGAPORE (all coll. LKCNHM): 1 male, Semakau, 3 April 2012, sweeping along path in back mangrove (leg. P Grootaert). 1 male, Semakau, 12 July 2012, sweeping along path in mangrove (leg. Jayanthi Puniamoorthy & P Grootaert). 1 male, 2 females, West Coast Park, 7 December 2003; sweeping nets along shrubs along sandy beach (leg. P Grootaert). 1 male, 5 female, Sungei Buloh, 1–6 June 2005, malaise trap in mangrove (leg. P Grootaert). 5 female, same provenance the previous, 20–27 July 2005. 1 male, 2 females, Clementi Woods, sweeping nets along drains in park (leg. P Grootaert). 1 male, 1 female, Labrador Park, 3 April 2005, sweeping along drains (leg. P Grootaert).
Antenna entirely dark yellow. Mid coxa entirely darkened, hind coxa anteriorly with a rectangular brown spot. No thickening of the costa present. Wing nearly hyaline, lightly tinged brown. Apex of phallus looking smooth though microscopic small denticles are present.
Male. Body length 3.8 mm, wing 3.5 × 1.2 mm.
Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black except postocular bristles pale. Antenna dark yellow; postpedicel nearly triangular, blunt apically, 1.4 times as long as wide, covered by short brown pubescence; arista-like stylus dorsal, black to dark yellow onwards, inserted at basal half of postpedicel, nearly as long as head width, black, feather-like, with long black pubescence, basal segment 0.5 times as long as apical portion of arista-like stylus. Proboscis brown, with black hairs; palpus yellow, with one short black apical bristle.
Thorax dark metallic green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of strong acr. Scutellum with two pairs sc, apical pair long strong, basal pair short and weak. Legs mainly yellows, except basal 2/3 of mid coxa brown, hind coxa with one brown spot, tip of hind tibia brown, tarsi yellow to brown onwards. Fore coxa anteriorly with five strong bristles at apical half, mid coxa anteriorly with four strong bristles, hind coxa with two short outer bristles. Fore and hind trochanters with one weak outer bristle at middle. Mid trochanter with two short weak outer bristles. Fore femur without distinct bristle. Mid and hind femora each with one preapical pv. Fore tibia with two short ad, two pd, one av, and three ap. Mid tibia with two ad, three pd and four ap. Hind tibia with six ad, four pd, two pv, and three ap. Hind tarsomere I with one short strong ad at apical third, one av at basal fifth and one short apical bristle. Relative lengths of tibia and five tarsomeres of legs LI : 2.5 : 1.1 : 0.5 : 0.4 : 0.3 : 0.3; LII : 3.0 : 2.0 : 1.0 : 0.8 : 0.5 : 0.3; LIII : 4.0 : 1.5 : 1.5 : 1.0 : 0.8 : 0.5. Wing nearly hyaline, lightly tinged brown; veins brown. M with fading M2, M1 with one short subvein. Crossvein dm-cu almost straight. CuAx ratio 1.0. Lower calypter pale with black hairs. Haltere pale.
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia: Epandrium 2.0 times longer than wide (Figure
Female. Has the same characteristics as the male: large rectangular black spot on the hind coxa and the tip of the hind tibia brown.
Lichtwardtia formosana is the only Oriental species known at the moment with a rectangular black (dark brown) patch on the hind coxa. In all other species the hind coxa is entirely yellow. Lichtwardtia formosana looks different but has a double row of microscopic denticles ventrally on the tip of the phallus. Otherwise the shape of the postgonite is identical to the L. nodulata group. The cercus lacks a strong dorsal bristle at the inside.
China (Taiwan), Cambodia, Singapore.
Rhagoneurus hirsutisetus de Meijere, 1916: 229. Male. Type locality Batavia (= Jakarta, Indonesia).
Holotype male Batavia, August, 1907, leg. Jacobson (Figure
A larger species (body length 4.5 mm; wing length 4.0 mm). Postpedicel mainly dark yellow, but blackish on dorsum and tip. Wing brown with cross veins not brownish seamed and costa with a distinct swelling well before R1 reaches the costa (Figure
As marked in the diagnosis L. hirsutiseta is quite unique in having a largely darkened postpedicel while other Lichtwardtia have generally an entirely yellow postpedicel. The swelling of the costa is also unique. It is well separated and before R1 reaches the costa. The complex male terminalia resemble L. monstruosa sp. n. but not L. polychroma.
Holotype male (coll.
Wing anteriorly brown, grey below Cu and M1. Costa lacking a swelling. Postpedicel about as long as wide with rounded tip, apex brownish yellow, ventrally yellowish. Mid coxa anteriorly with a black stripe, posteriorly brown. Hind coxa entirely yellow. Phallus with strong black spines, phallus and hypandrium resting on a ventral protuberance of the epandrium. Cercus seamed with long, fine black and pale bristles.
Male. Body length 4.1 mm, wing 3.2 × 1.1 mm. Head dark metallic green, with thick pale pollinosity; face slightly raised, frons and face both with thick pale pollinosity, gradually narrowed downward. Hairs and bristles on head black but lower postocular bristles yellowish. Antenna yellow except but apical half of postpedicel brown; postpedicel nearly triangular, 1.2 times as long as wide; arista-like stylus black, almost as long as head width, feather-like, with long regular pubescence. Proboscis dark yellow, with black hairs; palpus very small, dark yellow, with one short black apical bristle.
Thorax dark green, with pale grey pollinosity. Hairs and bristles on thorax black; five strong dc, ten pairs of strong acr. Scutellum with two pairs sc, apical pair long strong, basal pair short and weak. Legs mainly yellow. Mid coxa anteriorly with a black stripe, posteriorly brown. Hind coxa entirely yellow. Fore tarsomere I brown, apical fifth of each hind tarsomere brownish. Fore and mid coxae anteriorly with rows of bristle-like hairs, and four strong ap, hind coxa with two outer bristles, basal one strong, apical one relatively weak. Mid and hind trochanters both with four short weak ap. Fore femur with one weak preapical av. Mid femur with one strong pd and one preapical av. Hind femur with one strong pd at apical quarter. Fore tibia with two ad, three pd, one av and three ap. Mid tibia with two ad, four pd and four ap; all long strong. Hind tibia with four ad, four pd (of which one preapical), one d, two pv and three ap. Fore tarsomere I with one short av at base. Hind tarsomere I with one strong ad at apical third, one short ad at base and two short apical bristle. Relative lengths of tibia and five tarsomeres of legs LI : 7.5 : 4.5 : 1.8 : 1.3 : 1.0 : 1.0; LII : 11.2 : 6.3 : 3.0 : 2.5 : 1.5 : 1.3; LIII : 12.5 : 5.0 : 5.2 : 3.8 : 2.5 : 1.5. Wing anteriorly brownish infuscate (Figure
Abdomen metallic green, with pale pollinosity. Hairs and bristles on abdomen black.
Male terminalia: Epandrium 2.0 times longer than wide; epandrial lobe with three long pale bristles. Ventral surstylus with five ap, of which ventral most apical bristle rod-like and others spinous; dorsal surstylus both with five short nail-like ap and one long digitation close to dorsal margin which with one long spinous bristle. Phallus with strong black spines, phallus, and hypandrium resting on a ventral protuberance of the epandrium. Cercus nearly triangular, pale except the thick black ring, with weak digitations around outer margin, with long pale marginal bristles on digitations. Hypandrium simple. Phallus strong with strong black denticles (Figure
Female. Unknown.
From Latin infuscāre to darken.
It is not clear if L. infuscata sp. n. belongs to the L. dentalis –group though it has the tip of the phallus ventrally denticulate and a simple hypandrium. The cercus is more elongate and lacks a dorsal bristle at the inside. Others see comments under L. ziczac.
Cambodia.
Rhagoneurus coxalis Kertész, 1901: 411. Female. Type locality: Singapore.
The type was probably conserved at the Hungarian Museum for Natural History. It is no longer in their collections and probably destroyed (
Hind coxa largely blackish brown. Hind femur without preapical bristle and hind basal tarsomere lacking long bristles. Cross veins not brownish seamed.
It is likely that this species is one of the four species that we actually recorded in Singapore. It is not L. singaporensis sp. n. since it has the cross veins brownish seamed and the hind coxa yellowish. L. nodulata sp. n. has a broad swelling where R1 joins the costa and the hind coxa is yellowish (Figure
Since it was described on the basis of a female and that the holotype seems to be lost, we think it is not appropriate to sink L. formosana as a junior synonym having its holotype conserved. The confusion is bigger since de Meijere (1912) quotes L. coxalis (as Rhagoneurus) from “Neu Guinea” but this is a misreading because Kertész clearly says that he obtained a female from Singapore by the courtesy of his friend Biro. Meanwhile, the description by Kertész is too simple to acquire any further comparison. Therefore, at this moment, L. coxalis is considered as a nomen dubium.
Dolichopus ziczac Wiedemann, 1824: 40. Female. Type locality: India Orientalis
Dolichopus ziczac Wiedemann, 1830: 232
Dolichopus
zickzack
Wiedemann, 1824. Male in
Lichtwardtia ziczac (Wiedemann, 1824) sensu Zhang, Masunaga & Yang, 2009: 199, figs 11–14.
Holotype female, India Orientalis, on pin in collection ZMUC (Copenhagen).
Female. A medium-sized species (body: 4 mm; wing: 3.2 mm). Wing hyaline with anterior border faintly brownish and cross veins brownish seamed. No swelling of the costa before or at the point where R1 joins the costa. The ratio of the proximal section of M1, and the distal section is 0.4/0.6 (Figure
Dolichopus ziczac was described by
Brownish seams along the cross veins are not very common in Lichtwardtia and actually only known in L. ziczac and L. singaporensis sp. n. The ratio of the proximal section of M1, and the distal section is however 0.435/0.564. Thus the distal section is not as long as in L. ziczac. We do not consider both species as conspecific for the moment because in the near future it might be possible to extract ancient DNA from the holotype without using destructive techniques. This can be decisive about the status of both species. We propose to consider L. ziczac (Wiedemann) from terra incognita as a nomen dubium and not to complicate again the taxonomy of Lichtwardtia by applying the name ziczac to the male of singaporensis without genetic information. Remarkable is that among the more than 200 Lichtwardtia specimens belonging to six species that we found in Seam Reap, no specimens with brownish seamed cross veins were found.
1 | Hind coxa with anteriorly half or basal half brownish | 2 |
– | Hind coxa entirely yellow | 3 |
2 | Anterior half of hind coxa with a dark brown rectangular area that bears the two black exterior bristles; cross veins not seamed brownish. Apex of phallus looking smooth though microscopic small denticles are present (Figure |
formosana Enderlein |
– | Hind coxa with basal half brownish and cross veins brownish seamed. Phallus ventrally with strong denticles, hypandrium unarmed. (Figs |
singaporensis sp. n. |
3 | Wing with a colour pattern (Figure |
infuscata sp. n. |
– | Wing uniformly hyaline, greyish or brownish tinged but without pattern. Male terminalia various | 4 |
4 | Costa with a swelling before or at the point where R1 joins the costa | 5 |
– | Costa without swelling | 10 |
5 | A distinct brown swelling of the costa where R1 joins the costa (Figure |
nodulata sp. n. |
– | Costa widened before R1 reached costa (Figure |
6 |
6 | Costa widened distinctly before R1 joins costa (Figure |
hirsutiseta de Meijere |
– | Costa widened just before tip of R1 (Figure |
7 |
7 | Tip of phallus with black ventral denticles (Figure |
8 |
– | Tip of phallus lacking ventral denticles; tip of hypandrium with a large brown tooth (Figure |
9 |
8 | Marginal bristles on cercus strong, black (Figure |
dentalis Zhang et al. |
– | Marginal bristles on cercus weak, pale. Swelling of costa weak. Hypandrium simple and smooth, with no denticle; phallus with double rows of spinules on ventral half (Figure |
cambodiensis sp. n. |
9 | Tip of hypandrium with a single large brown preapical tooth (Figs |
polychroma (Loew) |
– | Tip of hypandrium dorsally curved, bifurcate; with a long black appendage near middle of hypandrium (Cambodia) | conspicabilis sp. n. |
10 | Hypandrium simple (Figure |
semakau sp. n. |
– | Hypandrium with tooth-like extensions | 11 |
11 | Hypandrium with a large preapical brown tooth (Bali) | zhangae sp. n. |
– | Hypandrium with ventral and dorsal black saw-toothed extensions (Figure |
monstruosa sp. n. |
With twelve species known now in the Oriental Realm, the genus Lichtwardtia is apparently quite species rich especially in view of the poor local sampling. Here we added eight species new for science. Interestingly, the external non-genitalic morphology is not very diverse but the male terminalia are distinctly different. It is perhaps too early to place all species into species groups but some closely related species can be distinguished.
A first large species group is the L. nodulata group with cambodiensis sp. n., L. dentalis Zhang et al., L. nodulata sp. n., L. semakau sp. n., and L. singaporensis sp. n.
The sister-group L. polychroma and L. zhangae are probably related to the nodulata-group in having a similarly shaped cercus with some strong flattened marginals and a dorsal marginal that shifted to the inside. However the postgonite is tubiform while it is broad with bifid tip in the L. nodulata group. Moreover the tip of the phallus is not ventrally denticulate and the hypandrium has a strong subapical dorsal hook. Provisionally we put both species in the polychroma-group.
Lichtwardtia conspicabilis sp. n. and L. monstruosa sp. n. seem to be related in having a forked hypandrium and a forked phallus, thus lacking the ventral denticles on the tip of the phallus in the nodulata group. Provisionally we place both species in the conspicabilis-group.
It is not clear if L. infuscata sp. n. belongs to the L. nodulata group though it has the tip of the phallus ventrally denticulate and a simple hypandrium. The cercus is more elongate and lacks a dorsal bristle at the inside. L. formosana looks different but has a double row of microscopic denticles ventrally on the tip of the phallus (Figure
At the moment we could only provide barcodes of the species from Singapore (uploaded to GenBank, with accession number MH536852-MH536856). The material from Cambodia was collected in ethanol of poor quality and therefore not suitable for sequencing. The optimal tree with the sum of branch length = 0.27350170 is shown. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test (1000 replicates) are shown next to the branches. The tree is drawn to scale, with branch lengths in the same units as those of the evolutionary distances used to infer the phylogenetic tree. As can be seen in Figure
Variation in morphology of hypandrium and phallus in Oriental Lichtwardtia species. Scale bars: 0.1 mm. A formosana-group: L. formosana: phallus slender, hidden in hypandrium. Tip of phallus ventrally with a double row of microscopic denticles B nodulata-group: L. nodulata: tip of phallus ventrally with strong denticles. Tip of phallus never retracted in the hypandrium C nodulata-group: L. infuscata: dorsal spine present at base of exposed part of phallus D polychroma-group: L. polychroma hypandrium with a large dorsal preapical tooth resting on a large basoventral epandrial lobe. Phallus robust, forked E conspicabilis-group: L. conspicabilis: tip of hypandrium membranous and with a right extension. Phallus robust with a forked tip F conspicabilis-group: L. monstruosa tip of forked hypandrium with denticles. Tip of phallus slender but with a large dorsal tooth near middle of phallus.
As can be seen on Figure
We thank Dr Zoltán Soltész (Hungarian Natural History Museum, Budapest) who checked the collection in Budapest for the type of Rhagoneurus coxalis Kertész and for communicating the report about the loss of the Diptera collection in Budapest. Dr Thomas Pape (Copenhagen) helped us with the photos of the holotype of Lichtwardtia ziczac (Wiedemann). Mr Ben Brugge (Naturalis, the Netherlands) sent us the photos of the holotype of L. hirsutiseta (de Meijere). Sven Marotzke (MfN, Berlin) recovered the lectotype of L. polychroma and provided us with photos made by Bernhard Schurian. Prof Rudolf Meier helped with the sequencing of the specimens of Singapore. Finally Dr Lili Zhang commented on the species that she described. Ms Maimon Hussin (LKCNHM, NUS) Singapore made a number of stacked photos.
Thanks for the help of Dr Torsten Dikow and Dr Norman Woodley (Smithsonian Institution, Washington DC, USA) we could trace the holotype of L. polychroma (Loew).
The sampling in Siem Reap (Cambodia) coordinated by Mr Jérôme Constant (
Last but not least, P Grootaert acknowledges a grant from the National Parks Board in Singapore and especially thanks Dr Lena Chan, senior Director at the Biodiversity Centre, who stimulated the long-term surveys. The research was also funded by the National Natural Science Foundation of China (31772497).