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Ngirhaphium Evenhuis & Grootaert from southern Thailand (Diptera: Dolichopodidae) with the description of a new species

  • Princess Maha Chakri Sirindhon Natural History Museum (PSUNHM), Prince of Songkla University, Songkhla, Thailand, 90110

Abstract and Figures

The genus Ngirhaphium Evenhuis & Grootaert, 2002 is reported for the first time from Thailand in particular from mangroves on the coast of the Andaman Sea in southern Thailand. Three species were found: N. murphyi Evenhuis & Grootaert, 2002, N. sivasothii Grootaert & Puniamoorthy, 2014 and N. chutamasae sp. nov. The latter species is described and illustrated and a key to all four known species is provided. COI barcode data showed that the new species is most closely related to N. murphyi with a genetic distance of 7%. The distance with the other species is 11 to 12%.
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Accepted by B. Sinclair: 12 Feb. 2015; published: 8 Apr. 2015
ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 3946 (1): 125
Ngirhaphium Evenhuis & Grootaert from southern Thailand
(Diptera: Dolichopodidae) with the description of a new species
Department of Biology, Faculty of Science, Prince of Songkhla University, Hat Yai campus, Kho Hong, Hat Yai, Songkhla, Thailand,
90112. E-mail:;
Entomology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B-1000 Brussels, Belgium.
Corresponding author
The genus Ngirhaphium Evenhuis & Grootaert, 2002 is reported for the first time from Thailand in particular from man-
groves on the coast of the Andaman Sea in southern Thailand. Three species were found: N. murphyi Evenhuis & Groot-
aert, 2002, N. sivasothii Grootaert & Puniamoorthy, 2014 and N. chutamasae sp. nov. The latter species is described and
illustrated and a key to all four known species is provided. COI barcode data showed that the new species is most closely
related to N. murphyi with a genetic distance of 7%. The distance with the other species is 11 to 12%.
Key words: Dolichopodidae, Ngirhaphium, new species, mangrove, Thailand
The genus Ngirhaphium Evenhuis & Grootaert, 2002 is a genus of large dolichopodid species that occurs only in
the front mangrove and along creeks in mangroves (Grootaert & Puniamoorthy 2014). Hitherto three species were
known exclusively from Singapore and it is the first time that the genus is reported from another country.
Here we report on three species found in mangroves along the coast of the Andaman Sea in southern Thailand:
N. murphyi Evenhuis & Grootaert, 2002, N. sivasothii Grootaert & Puniamoorthy, 2014 and a new species for
science that is described, illustrated and barcoded.
Material and methods
Study sites and sampling techniques. The present study is based on a survey of the marine dolichopodids in
southern Thailand done by the first author (AS). Both Malaise traps and sweep netting techniques were used to
collect fresh specimens in various mangroves in the provinces of Nakhon Si Thammarat, Songkhla, Pattani and
Satun (Tammalang subdistrict) and Tarutao Island, all in southern Thailand. Terminology following Grootaert &
Puniamoorthy (2014).
Specimen storage. The holotype and paratypes of the new species and other species are preserved in 70%
ethanol to prevent the degradation of DNA and deposited in the collections of the Princess Maha Chakri Sirindhon
Natural History Museum of the Prince of Songkhla University, Hat Yai, Thailand (PSU). Voucher specimens are
also stored in the collection of the Royal Belgian Institute of Natural Sciences, Brussels (RBINS).
Genetic analysis. Total DNA was extracted from a pair of middle legs. The remaining portions of the
specimens sampled were kept as voucher in PSU Natural History Museum. The sample tissues were placed into 1.5
ml sterile tubes and pulverized by adding 50 µl of tissue lysis buffer; eventually incubated at 65°C for 30 minutes.
Subsequently, 2 µl of Proteinase K was added and incubated at 60°C overnight (24 hr.). The mixture was agitated
with 7 µl of 8M potassium acetate for 5 minutes and incubated at -20°C for 30 mins before extraction of the
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aqueous supernatant. This procedure was repeated once before an equal volume of 95% ethanol was added to
precipitate the DNA pellet. The pellet was washed subsequently in 70% ethanol and 30 µl of TE buffer was applied
to dissolve the DNA before storage at -20°C.
Universal primers amplifying portion of the mitochondrial cytochrome c oxidase I (COI) gene (LCOI1490 and
HCO2198, Folmer et al. 1994) were as previously used by Grootaert & Puniamoorthy (2014). Thermocycling
conditions were as follows: initial denaturation at 95°C for 3 mins, followed by 40 cycles of denaturation at 94°C
for 1 min., annealing at 48°C for 1 min., and extension at 72°C for 1.5 min. A final extension of 5 mins at 72°C was
used. Gel electrophoresis was performed to verify the PCR success in a 1% agarose gel using 5 µl of the reaction
mix. All PCRs were purified and sequenced commercially by First BASE Sequencing Company (Malaysia).
The evolutionary history was inferred using the Neighbor-Joining method (MEGA6, Tamura et al. 2013). The
optimal tree with the sum of branch length = 0.20749235 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. The analysis involved 13 nucleotide sequences. Codon positions included were
1st+2nd+3rd+Noncoding. All positions containing gaps and missing data were eliminated. There were a total of
494 positions in the final dataset.
Genus Ngirhaphium Evenhuis & Grootaert
Ngirhaphium Evenhuis & Grootaert, 2002: 310. Type species by original designation: Ngirhaphium murphyi Evenhuis &
Grootaert, 2002.
Diagnosis. Medium to large sized species (4.5–8 mm) with a metallic green or blue ground-colour. Antenna very
long in males, a little shorter in females. Arista apical, basal article long. Rostrum in male small with well-
developed labellae. Large rostrum in female. Vertex excavated (cf. Sciapodinae).
Mid and hind coxae without exterior bristle. Femora with inconspicuous bristling. All tibiae with strong
bristles. Fore leg in male with tarsomere 4 bearing an asymmetrical, apical dorsal forked protuberance (absent in
females); terminal segment with a pair of normal claws and a thickened claw-like structure beneath the posterior
claw. Females with the claws as usual, but the terminal segment bears a long dorsal protuberance. Mid and hind
legs with tarsomeres 1–4 with an apical comb of spinules ventrally.
Wing with tip of M
sharply bent upwards just before reaching the wing border and ending near tip of R
Ngirhaphium chutamasae sp. nov.
(Figs 1–6)
Diagnosis. A large species differing from the other Ngirhaphium species mainly in the structure of the male
genitalia. Cercus in lateral view slightly shorter than dorsal surstylus. Cercus brown, tip pointed bearing a single
yellow bristle. Dorsal surstylus brown, bordered with short, stout yellow bristles. Outer branch of apical fork on the
fore tarsomere 4 slightly longer than inner branch. M
with a short stub on apical bend.
Material examined. HOLOTYPE ♂, labelled: “THAILAND: Satun prov., Tammalang (6°32'21.05"N,
100°04'9.42"E), 3.x.2014 (reg. 34030, leg. P. Grootaert)” (PSU); PARATYPE: 1 ♂, Tammalang (6°32'21.05"N,
100°04'9.42"E), 6.viii.2014 (leg. A. Samoh) (RBINS).
Etymology. The species is dedicated to Associate Professor Dr. Chutamas Satasook, director of the Princess Maha
Chakri Sirindhon Natural History Museum of the Prince of Songkhla University, Hat Yai as a token for her
dynamic support of our research.
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FIGURE 1. Ngirhaphium chutamasae sp. nov., male habitus; inset: apical tarsomeres on fore leg, showing the large fork-like
extensions on tarsomere 4 and the additional claw-like structure on tarsomere 5 (photo: J. Brecko). Scale = 1 mm.
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FIGURES 2–6. Ngirhaphium chutamasae sp. nov., holotype male genitalia: 2. Left ventral surstylus; 3. Lateral view of genital
capsule with left ventral surstylus removed; 4. Cerci dorsally; 5. Dorsal surstylus in ventral view; 6. Ventral view of genital
capsule. Abbreviations: ae: aedeagus; c: cercus; ds: dorsal surstylus; f: foramen; hy: hypandrium; sp: sperm pump; vs: ventral
surstylus. Scale = 0.1 mm.
Description. Male. Length body: 7 mm; wing: 5.6 mm. Head. Frons shining metallic green (not dusted). Face
greenish brown in ground-colour; apex of face and clypeus yellowish brown in ground-colour, wide, nearly as wide
as front of frons, parallel-sided, grey dusted with very short clypeus (less than 0.1 length of face). Eyes pass beyond
border of face; eyes densely set with white hairs. Ocellar callus globular protruding from frons with 2 very long
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ocellars, directed backward, divergent. Vertical bristles long, half as long as ocellars, rather anteriad on frons at
level of ocellar callus, close to eye border, long, black, directed forward and cruciate. Pair of long black
postverticals directed backward and crossing. Postoculars above strong, black in single row, below white and
mixed with very long white hairs below mouth; postcranium greenish in ground-colour but grey dusted. Palpus
long, strap-shaped, yellowish with few short black hairs, no bristles. Labella brown with black hairs. Antenna very
long, completely black. First segment long, 3X as long as second segment; second segment short, apically with
crown of short black bristles. Third segment very long strap-shaped, laterally flattened, about 6X as long as width
at base. Arista apical, apical article longer than basal article, gradually tapered towards tip. Length of scape: 0.52;
pedicel: 0.13; postpedicel: 0.95; basal aristal article: 0.34; apical aristal article: 0.4 (in mm). Thorax and scutellum
dark metallic green in ground-colour (bluish when seen from in front), covered with fine grey dusting. All hairs and
bristles black. Pleura more densely grey dusted than mesonotum. Acrostichals biseriate, about 7 pairs, rows
widening slightly behind. Presutural dorsocentrals multiseriate; 6 postsutural dorsocentrals: 4 short and 2 long
prescutellars; 1 pair of strong scutellars. One long humeral with shorter bristle in front; 1 strong posthumeral, 2
strong notopleurals, 1 postsutural, 1 supraalar, 1 very strong postalar. Propleural bristles black, 6 short upper and 2
longer lower propleural bristles (lower one twice as long as upper). Legs (Fig. 1) yellow, all bristles black. All
coxae greenish black in ground-colour, covered with fine greyish dusting. All trochanters brown. Tip of hind tibia
annulated brown at tip. All tarsi yellowish, becoming darker towards tip. Apical tarsomeres completely black. Fore
leg. Coxa with short black bristles. Fore femur slightly swollen on basal half; row of minute posteroventrals in
apical half. Short preapical posterior bristle and 1 stronger anterior preapical bristle directed forward. Fore tibia
with 4 strong ad, 4 strong pd and crown of 4 apicals. All tarsal segments densely set with black hairs and short
black bristles. Tarsomere 4 with dorsal asymmetrical fork, extended over tarsomere 5; fork about 1.5X length of
tarsomere; outer branch of fork slightly longer than inner branch, tips pointed (Fig. 1, inset). Terminal segment
with pair of long normal claws and thicker claw-like structure beneath posterior claw. Two well-developed pulvilli
and empodium present. Length of femur, tibia and tarsal segments (in mm): 2 : 1.96 : 0.98 : 0.56 : 0.28 : 0.28 : 0.28.
Mid leg. Coxa with short bristles anteriorly; no exterior bristle. Mid femur as wide as fore femur; ventrally with
inconspicuous bristles; 1 strong anterior preapical and 2 tiny posterior preapicals. Tibia with 6 ad, 6 pd (might be
considered as dorsal), 8 longer av and crown of long apicals. Tarsomeres 1–4 ventrally at tip with pair of short
spine-like bristles as well as comb of shorter black spinules. Apical tarsomere dorsally thickly set with long black
squamiform bristles. Length of femur, tibia and tarsal segments (in mm): 2 : 3.08 : 1.68 : 0.84 : 0.77 : 0.35 : 0.42.
Hind leg. Coxa bare. Hind femur thicker than mid femur, as wide as fore femur; ventrally almost bare; 1 strong
anterior preapical, 1 fine posterior preapical. Tibia stronger bristled than mid tibia with 7 long av, 7 ad, 7 pd and
crown of long apicals. Tarsomeres 1–4 ventrally at tip with pair of short spine-like bristles as well as comb of
shorter black spinules. Length of femur, tibia and tarsal segments (in mm): 2 : 3.78 : 1.68 : 0.98 : 0.77 : 0.42 : 0.35.
Wing mostly tinged brownish, but anteriorly between costa and R
with yellowish brownish tinge. Tp brown
seamed. Veins dark brown, yellowish at base. M
sharply bent upwards and ending in costa closely near tip of
. Tp straight, about as long as apical part of M
. Anal vein reaching wing border. Halter with white knob.
Squama white with long white cilia. Abdomen shiny dark metallic green; tips and sides of tergites with greyish
dusting. Sternites greyish dusted. Tergites densely set with quite long black bristles; hind-marginal bristles slightly
longer than other bristles. Only tergite 5 with very long marginal bristles. Sternites with very short hairs except for
longer marginals on sternite 4. Genital capsule black. Cercus brown, slightly shorter than dorsal surstylus (Fig. 3).
Ventrally at base with black sclerotisation. Both cerci fused for almost entire length, only tips free (Fig. 4). Tip of
cercus pointed, with single yellow apical bristle, dorsally set with long black bristles. Dorsal surstyli brown, much
enlarged, forming clasper transverse on cercus, bordered with short, stout yellow bristles, shorter than surstylus is
wide (Fig. 3). Ventrally with dark spur-like apex (Fig. 5). Ventral and dorsal surstyli not fused. Ventral surstylus
yellowish, large, rounded (Fig. 2); tip with short hair-like bristles; inner carina running parallel to dorsal border.
Hypandrium dorsally with large rounded black protuberance set with spinules (Fig. 3). Female. Indistinguishable
from females of N. sivasothii (see Remarks section).
Remarks. The new species is morphologically almost identical to N. sivasothii, except for the larger forked
extension on fore tarsomere 4 in male and the very different male genitalia. The outer branch of the apical fork on
fore tarsomere 4 is slightly longer than the inner branch, the outer branch is slightly shorter than the inner branch in
N. sivasothii.
The wing is brownish tinged and only the Tp is dark seamed. In N. sivasothii the wing is darker and the
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longitudinal veins as well as Tp are generally black seamed. Vein M
includes a short stub on the apical bend in
the new species. Such a stub was never observed in the large populations of N. sivasothii in Singapore.
Presently females of the new species are indistinguishable from females of N. sivasothii. Both species have
been collected together. For this reason, no females were included in the material examined section.
FIGURES 7–12. Lateral view genital capsule: 7. Ngirhaphium sivasothii; 8. N. caeruleum; 9. N. murphyi; dorsal view cerci:
10. N. sivasothii; 11. N. caeruleum; 12. N. murphyi. Abbreviations: c: cercus, ds: dorsal surstylus (modified after Grootaert &
Puniamoorthy 2014). Scale = 0.1 mm.
Ngirhaphium murphyi, Evenhuis & Grootaert
(Figs 9, 12)
Ngirhaphium murphyi Evenhuis & Grootaert, 2002: 310. Type locality: SINGAPORE: Kranji mangrove.
N. murphyi: Grootaert & Puniamoorthy, 2014: 147 (figs 1–3, 4, 5, 18).
Diagnosis. A large species (5.7–7.3 mm), generally with clear wings. Mesonotum and tergites metallic green.
Apical aristal article quite thick, nearly half as long as basal aristal article. Male with cerci longer than surstyli and
thus the tips are visible outside the surstyli (Figs 9, 12).
Material examined. THAILAND: 1 ♂, 4 ♀, Satun province, Tarutao Island, Talo Wao bay (6°36'58.7"N
99°40'43.1"E), 11.viii.2014 (leg. A. Samoh) (PSU)
Remarks. The pedicel is yellowish-brown in females and black in males. The pedicel is always black in both
sexes in Singapore populations (Grootaert & Puniamoorthy 2014).
Ngirhaphium sivasothii Grootaert & Puniamoorthy
(Figs 7, 10)
Ngirhaphium sivasothii Grootaert & Puniamoorthy, 2014: 150 (figs 6–8, 9–10, 17). Type locality: SINGAPORE: Semakau
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Diagnosis. A medium-sized species (4.5–5.5 mm), generally with dark infuscate wing and with longitudinal veins
and Tp (posterior cross vein) brownish seamed. Mesonotum and tergites metallic green. Apical aristal article
shorter, but nearly as long as basal article. Male with dorsal surstylus half as long as cerci, with a rectangular bend,
set with very long bristles (Fig. 7). Cercus much longer than dorsal surstylus, tip wide, rounded, set with many long
yellow bristles (Fig. 10). Outer branch of apical fork on the fore tarsomere 4 slightly shorter than inner branch.
Material examined. THAILAND: Satun province: 1 ♂, 2 ♀, Tammalang (6°32'21.05" N, 100°04'9.42" E); 4
7 ♀, 6.viii.2014 (leg. A. Samoh); 1 , 2 ♀, 3.x.2014 (reg. 34030, leg. P. Grootaert & A. Samoh); 7 ♂, 20 ♀,
Tarutao Island, Talo Wao bay (6°36'58.7"N 99°40'43.1"E), 12.viii.2014 (leg. A. Samoh); 2 ♂, 1 ♀, Tanjong Po
(6°36'57.43" N, 99°57'25.66" E), 3.x.2014 (leg. A. Samoh) (PSU).
Remarks. Some specimens had quite clear wings without the brown of black seams along the longitudinal
veins and the Tp (posterior cross vein).
Key to males of Ngirhaphium
1 Mesonotum and tergites metallic blue. Antenna with apical aristal article filiform and much longer than basal article (Singa-
pore). Genitalia as in Figures 8 and 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .N. caeruleum Grootaert & Puniamoorthy
- Mesonotum and tergites mainly metallic green. Antenna with apical aristal article shorter or about half as long as apical article
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Cerci in lateral view nearly as long as dorsal surstyli (Figs 3, 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. chutamasae sp. nov.
- Cerci in lateral view longer than dorsal surstyli (Figs 7, 9) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Dorsal surstylus elongate, digitiform with truncate apex (Figs 9, 12) slightly shorter than cercus. Cerci with narrow apex, set
with 2 apical setae (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. murphyi Evenhuis & Grootaert
- Dorsal surstylus with very wide apex (Fig. 7) much shorter than cercus. Cerci with expanded apex set with many yellow setae
(Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. sivasothii Grootaert & Puniamoorthy
FIGURE 13. Neighbour-Joining tree of the COI barcodes of Ngirhaphium with bootstrap values indicated at the nodes. Scale
of genetic distance is 1%.
With three species, Ngirhaphium appears quite diverse on the coast along the Andaman Sea in southern Thailand.
Although several mangroves along the coast of the South China Sea were investigated, we failed to collect any
specimens. This might be due to differences in microhabitat that we failed to recognize, or simply that the genus is
not present there.
Zootaxa 3946 (1) © 2015 Magnolia Press
Ngirhaphium murphyi and N. sivasothii also occur in Singapore. The geographic distance between the
Singaporean populations and those in the Satun province (Tammalan, Tarutao Island) is about 800 km and the
genetic distance between the two populations is less than 1% for N. murphyi and about 1% for N. sivasothii (Fig.
13). This is very low compared to another mangrove species, Teuchophorus simplicissimus Grootaert & Meuffels,
which differed by 6.5% for COIb with a geographic distance of only 240 km between Singapore and Pulau Tioman
(Lim et al. 2009).
Ngirhaphium chutamasae sp. nov. is most closely related to N. murphyi and they cluster with a bootstrap of 94
while the genetic distance is 7% (Fig. 13) The genetic distance between the new species and N. sivasothii and N.
caeruleum is 11 % and 12 % respectively.
The authors are very grateful for the support of Associate Prof. Dr. Chutamas Satasook (Prince of Songkhla
University, Hat Yai). The staff of the Nature Reserve at Tammalang is thanked for their help in the field. Our thanks
also to Mr. Rueangrit Promdam and the entomology research unit members, Department of Biology, Faculty of
Science, PSU for assistance with fieldwork, to Mr. Phuripong Meksuwan, Ms Sakiyah Morlor and Ms Bongkot
Wichachoochert for assistance with the molecular work and highly appreciated suggestions, and to the Princess
Maha Chakri Sirindhorn Natural History Museum staff for their help. This study was in part supported by funds
from the National Research Universities grant (NRU), Government of Thailand. We also thank Dr. Brad Sinclair
and two anonymous referees for many crucial comments.
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial
cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3,
Evenhuis, N. & Grootaert, P. (2002) Annotated checklist of the Dolichopodidae (Diptera) of Singapore, with new records and
descriptions of new species. Raffles Bulletin of Zoology, 50, 301–316.
Grootaert, P. & Puniamoorthy, J. (2014) Revision of Ngirhaphium (Insecta: Diptera: Dolichopodidae), with the description of
two new species from Singapore’s mangroves. Raffles Bulletin of Zoology, 62, 146–160.
Lim, G.S., Hwang, W.S., Kutty, S., Meier, R. & Grootaert, P. (2009) Mitochondrial and nuclear markers of Oriental species
support the monophyly of Dolichopodidae and suggest a rapid origin of the subfamilies (Diptera: Empidoidea). Systematic
Entomology, 35, 59–70.
Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis
version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
... Recent intensive sampling efforts in Thailand (reviewed in part by Plant et al. 2011) and increased taxonomic study have begun to reveal the empidoid diversity of this tropical country (Barták & Kubík 2008;Barták et al. 2013;Daugeron & Grootaert 2003Daugeron et al. 2011;Grootaert & Kiatsoonthorn 2001;Grootaert & Meuffels 2001;Grootaert & Shamshev 2003Horvat 2002;Meuffels & Grootaert 2004;Plant 2009aPlant , 2009bPlant , 2009cPlant , 2010aPlant , 2010bPlant , 2013Samoh et al. 2015Samoh et al. , 2017Shamshev & Grootaert 2004Shamshev et al. 2006;Sinclair & Plant 2017). Although sampling has been geographically and temporally biased, taxonomic workup of the samples uneven (taxa variously assigned to species, operational taxonomic units or merely to higher taxa), and metadata of variable quality (Plant 2015a), analyses are beginning to reveal much valid and useful information on the geographical and habitat associations of distinct Empidoidea communities, their types of distribution, possible biogeographic affinities and locations of hotspots of diversity and endemism (Plant 2009b(Plant , 2013(Plant , 2014Plant et al. 2011Plant et al. , 2012. ...
This study is based on more than 25,000 specimens of the superfamily Empidoidea (Diptera) collected throughout a full year on a 2000 m elevational habitat succession gradient along a 21 km transect on Doi Inthanon, the highest mountain in Thailand. The samples were sorted to 58 genera and 458 morphospecies (Empididae, 73; Hybotidae, 203; Dolichopodidae, 179; Brachystomatidae, 3). The data were used to prepare the first thorough taxon-focussed description of how diversity of a major group of Diptera is structured in tropical forest biotopes. We found significant spatial (elevation / habitat) and temporal (seasonal) variations in richness (α-diversity) and abundance at family-level. α-Diversity of the four families was maximal in damp evergreen forests at higher elevation (1500–2500 m), but Dolichopodidae also had a major subsidiary peak in lowland dry evergreen forest at 500–1000 m. Genus-, tribe- and subfamily level α-diversity / elevation profiles were varied, indicating that overall family-level richness is a composite of many taxa that contribute low, high or mid-elevation specialisms. We provide a detailed analysis of these specialisms for each of the 58 genera. Adult phenology was correlated with the monsoon and had three characteristic phases: (i) pre-monsoon commencement during the latter part of the hot dry season, (ii) a ‘flush’ of maximal richness during the early-monsoon, and (iii) a secondary richness maximum associated with the late-monsoon. Maximum α-diversity occurred in phases (i) and (ii) but communities in phase (iii) had characteristically low evenness in which a few abundant species were dominant. Cluster analysis and ordination resolved three well-founded communities with different species-abundance distributions, high levels of species-level specialism and habitat-fidelity associated with moist hill evergreen forest (MHE) at >2000 m; mid elevation evergreen forests (EM) at 1000–2000 m and dry lowland forest (DL) at <1000 m. The three forest types with which these communities are associated are widespread and typical of northern Thailand and the diversity characteristics of each habitat are likely scalable to larger geographic areas. The transition from lowland DL through to upper montane MHE communities was generally characterised by increasing abundance, lower evenness (higher dominance), slower temporal turnover of community composition (relaxation of seasonality), longer periods of adult flight activity and rare species contributing less to species richness. Oriental biogeographic influences are strong at lower elevations but Palaearctic influences are increasingly important at higher elevations. The mixing of Oriental and Palaearctic elements in MHE forests is thought to explain the greater phylogenetic complexity at higher elevation (as measured by taxonomic distinctness).
... Thinophilus in Southeast Asia thrives in marine habitats (Grootaert & Meuffels 2001;Evenhuis & Grootaert 2002;Samoh et al. 2015Samoh et al. , 2017Grootaert 2018), where they are often found on rocky shores, but most species are found in front mangroves or follow the creeks inside the mangrove. A few species are non-marine, such as T. mekongensis Grootaert, 2017, and the very common and widespread T. nitens Grootaert & Meuffels, 2001 and T. setiventris Grootaert & Meuffels, 2001. ...
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Three new species of Thinophilus Wahlberg, 1844 are described from marine sites on the Andaman Sea coasts in peninsular Thailand: T. inaequalis Samoh & Grootaert sp. nov. belonging to the ‘Thinophilus simplex-group’ sensu Grootaert, 2018, T. plektron Samoh & Grootaert sp. nov., an unplaced species characterised by a long flattened apical spur on the hind tibia, and T. subapicalis Samoh & Grootaert sp. nov. belonging to the ‘Thinophilus spinatus-group’ sensu Grootaert, 2018. New records are given for following species occurring in peninsular Thailand: T. apicatus Grootaert, 2018, T. chaetulosus Grootaert, 2018, T. clavatus Zhu, Yang & Masunaga, 2006, T. evenhuisi Grootaert, 2018, T. lenachanae Grootaert, 2018, T. longicilia Evenhuis & Grootaert, 2002, T. nigrilineatus Grootaert, 2018, T. simplex Grootaert, 2018, T. superbus Grootaert, 2018, and T. yeoi Grootaert, 2018.
... Oldroyd (Insecta, Diptera, Dolichopodidae) from rocky shores in southern Thailand with the description of a new species INTRODUCTION The present study forms part of a survey of the marine dolichopodid flies occurring in peninsular Thailand (Samoh et al., 2015;2017). During this study a number of Cymatopus and Thambemya specimens were occasionally collected on rocky shores. ...
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Four species of the genus Cymatopus Kertèsz, 1901 and one species of Thambemyia Oldroyd, 1956 were found in southern Thailand. A key is given for all five species and new data of their distribution are provided. Cymatopus mayakunae sp. nov. is described as new for science. COI barcodes (313 base pairs) seem to be good indicators for delimitation of the species but do not support a phylogeny.
... In a previous survey (Grootaert & Meuffels 2001) 15 species belonging to seven genera of Dolichopodidae were found. Samoh et al. (2015) added the genus Ngirhaphium Evenhuis & Grootaert, 2002 with three species, resulting in 18 known species from southern Thailand to date. Only three species of Thinophilus have been recorded from peninsular Thailand until now: T. nitens Grootaert & Meuffels, 2001, T. parmatus Grootaert & Meuffels, 2001 and T. setiventris Grootaert & Meuffels, 2001. ...
... In a previous survey (Grootaert & Meuffels 2001) 15 species belonging to seven genera of Dolichopodidae were found. Samoh et al. (2015) added the genus Ngirhaphium Evenhuis & Grootaert, 2002 with three species, resulting in 18 known species from southern Thailand to date. Only three species of Thinophilus have been recorded from peninsular Thailand until now: T. nitens Grootaert & Meuffels, 2001, T. parmatus Grootaert & Meuffels, 2001 and T. setiventris Grootaert & Meuffels, 2001. ...
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Eight new species of marine dolichopodid fl ies from southern Thailand belonging to the genus Thinophilus Wahlberg, 1844 are described and illustrated: Thinophilus boonrotpongi sp. nov., T. langkawensis sp. nov., T. minutus sp. nov., T. parmatoides sp. nov., T. parvulus sp. nov., T. spinatus sp. nov., T. spinatoides sp. nov. and T. variabilis sp. nov. A key is provided to the species of the Thai- Malay Peninsula.
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A review is given of the Ngirhaphium species of Southeast Asia and two new species are described from Thailand. Ngirhaphium meieri Samoh & Grootaert, new species is described from a mangrove in Takua Pa district, Phang Nga Province, peninsular Thailand. NGS-barcodes (313 bp) are congruent with traditional taxonomic delimitation. Ngirhaphium caeruleum Grootaert & Puniamoorthy sensu lato seems to consist of a species-complex with 32 haplotypes and has a wide distribution in the southern part of the South China Sea. There is a 4.2 % genetic difference between the Thai/Cambodian population and the Singapore/Brunei populations in N. caeruleum: despite the many haplotypes and differences in male terminalia, the Thai and Cambodian haplotypes formed a haplotype group, while the Singapore and Bruneian specimens formed another haplotype group. Nonetheless, the Thai population is morphological distinct enough to be considered a morpho-species: it is here described as N. thaicum Samoh & Grootaert, new species. An updated distribution of the six Southeast Asian Ngirhaphium species is given with additional taxonomic notes, a key to the species and a phylogenetic diagram. We find that the Andaman Sea coast to be the most diverse with four species while the South China Sea region has only two species both belonging to the caeruleum species complex. N. sivasothii Grootaert & Puniamoorthy is common and occurring from Singapore up to the Thai coast of the Andaman Sea, with 28 haplotypes. Remarkable in all species is the limited distribution of the haplotypes, where most seem endemic to a limited area and thus provide information about the origin of the distribution.
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The type-species of Ngirhaphium Evenhuis & Grootaert, N. murphyi is re-described. Two new species are described from mangroves in Singapore: N. sivasothii, new species, and N. caeruleum, new species. A key to the three known species of the genus is given and male genitalia are illustrated. COI barcodes are provided for the three species, as well as data on distribution and phenology. Genetic distances of at least 10% suggest that the species have been genetically separated for a long time.
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We announce the release of an advanced version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which currently contains facilities for building sequence alignments, inferring phylogenetic histories, and conducting molecular evolutionary analysis. In version 6.0, MEGA now enables the inference of timetrees, as it implements our RelTime method for estimating divergence times for all branching points in a phylogeny. A new Timetree Wizard in MEGA6 facilitates this timetree inference by providing a graphical user interface (GUI) to specify the phylogeny and calibration constraints step-by-step. This version also contains enhanced algorithms to search for the optimal trees under evolutionary criteria and implements a more advanced memory management that can double the size of sequence data sets to which MEGA can be applied. Both GUI and command-line versions of MEGA6 can be downloaded from free of charge.
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The Dolichopodidae is a species-rich dipteran group with almost 7000 described species. The monophyly of the subfamilies and their relationships remain largely unknown because the polarities of key morphological characters are unclear and molecular data are available only for 9 of the 19 proposed subfamilies. Here we test whether molecular data from two nuclear (18S, 28S) and four mitochondrial (12S, 16S, Cytb, COI) genes can resolve the higher-level relationships within the family. Our study is based on 76 Oriental species from 12 dolichopodid subfamilies and uses eight species of Empididae and Hybotidae as outgroups. Parsimony and likelihood analyses confirm the monophyly of the Dolichopodidae, as well as the monophyly of five of the ten subfamilies represented by more than two species [Sympycninae, Sciapodinae, Dolichopodinae, Hydrophorinae (excluding tribe Aphrosylini), Neurigoninae]. There is strong support for restoring the tribe Aphrosylini as a separate subfamily Aphrosylinae. The monophyly of Medeterinae, Peloropeodinae and Diaphorinae is dependent on which tree reconstruction technique is used, how indels are coded, and whether the fast-evolving sites are excluded. Overall, we find that our sample of Oriental species is largely compatible with the subfamily concepts that were developed for the northern temperate fauna. However, our data provide little support for relationships between the subfamilies. Branch lengths, saturation, and distance plots suggest that this is probably the result of the rapid origin of dolichopodid subfamilies over a relatively short time. We find that genera that are difficult to place into subfamilies based on morphological characters are generally also difficult to place using molecular data. We predict that a dense, balanced taxon sample and protein-encoding nuclear genes will be needed to resolve the higher-level relationships in the Dolichopodidae.
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We describe "universal" DNA primers for polymerase chain reaction (PCR) amplification of a 710-bp fragment of the mitochondrial cytochrome c oxidase subunit I gene (COI) from 11 invertebrate phyla: Echinodermata, Mollusca, Annelida, Pogonophora, Arthropoda, Nemertinea, Echiura, Sipuncula, Platyhelminthes, Tardigrada, and Coelenterata, as well as the putative phylum Vestimentifera. Preliminary comparisons revealed that these COI primers generate informative sequences for phylogenetic analyses at the species and higher taxonomic levels.
Forty-four species of Dolichopodidae are listed from the island of Singapore, with 28 of these constituting new records for the country. One new genus: Ngirhaphium; and five new species: Thinophilus murphyi, Thinophilus asiobates, Thinophilus longicilia, Ngirhaphium murphyi, and Scotiomyia singaporensis, are described and illustrated.