European Journal of Taxonomy 112: 1-10
http://dx.doi.org/10.5852/ejt.2015.112
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ISSN 2118-9773
www. europeanj oumaloftaxonomv. eu
2015 • Grootaert P. et al.
Research article
urn:lsid:zoobank.org:pub:A92B8545-3B96-4596-AD29-B36D33E69770
Two new coastal species of Elaphropeza Macquart
(Diptera: Hybotidae) from Bali, Indonesia
Patrick GROOTAERT 1 l Isabella VAN DE VELDE 14 & Igor V. SHAMSHEV 12 5
Entomology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B-1000 Brussels, Belgium.
Email: Patrick.Grootaert@naturalsciences. be and
[email protected]
2 Zoological Institute, Russian Academy of Sciences, Universitetslcaja nab. 1, St. Petersburg 199034,
3 urn:lsid:zoobank.org:author:B80BC556-9087-4D0D-9D69-7FA9BE5779C4
4 urn:lsid:zoobank.org:author:CBlB2335-553C-4A40-BB10-19C62E3875C6
5 urn:lsid:zoobank.org:author:569F41CC-EC2B-4CF0-802A-8D7056C72C93
Abstract. Two new species of Elaphropeza Macquart, 1827 are described from the northeast coast
of Bali (Indonesia): Elaphropeza triseta sp. nov. belonging to the ephippiata- group and E. balinensis
sp. nov. belonging to the biuncinata- group. A COI Neighbour-Joining tree is given for the Southeast
Asian Elaphropeza species showing large genetic distances between the species. The morphological
characterisation of the ephippiata- group and the biuncinata- group sensu Shamshev & Grootaert 2007
is confirmed.
Key words. Elaphropeza , new species, Hybotidae, Southeast Asia.
Grootaert R, Van de Velde I. & Shamshev I. 2015. Two new coastal species of Elaphropeza Macquart (Diptera:
Hybotidae) from Bali, Indonesia. European Journal of Taxonomy 112:1-10. http://dx.doi.org/10.5852/eit.2015.112
Introduction
Shamshev & Grootaert (2007) revised the genus Elaphropeza in the Oriental Realm, reporting 79
species. Apart from the redescription of the holotypes of previously known species, 51 new species
were described mainly from Singapore. These new species were the result of a one-year collecting
campaign with Malaise traps in various biotopes in Singapore, e.g., swamp forest, secondary rain forest
and mangrove. Later, three species were described from Vietnam by Grootaert & Shamshev (2009).
These species were remarkable in having brush-like clusters of long hairs in the intertergal spaces
between tergites 3 and 4 and sometimes 4 and 5 as well. These brushes might help in the evaporation of
pheromones secreted by the glands below the tergites.
Subsequently, Grootaert & Shamshev (2012) published a monograph on the Tachydromiinae of Singapore
which was partly based on a one-month site assessment study performed in 2009, when insects were
collected with Malaise traps in thirteen mangrove fragments all along the coast of Singapore. In that
monograph another eight Elaphropeza species were described, all being coastal. Finally, Wang et al.
(2012) described two new Elaphropeza species from Taiwan (Wang et al. 2012). Thus, 137 species
of Elaphropeza are now known for the Oriental realm, including the two new species described in the
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European Journal of Taxonomy 112: 1-10 (2015)
present paper. This number is considered only a fraction of the number of species that might eventually
be found to occur in the Oriental realm, since most species have only been recorded from the limited
territory of Singapore with an emphasis on fauna restricted to mangroves.
At the end of 2013 the first two authors collected a small number of specimens of Elaphropeza from
shrubs in the supralittoral zone of a shingle beach with black volcanic sands on the northeast coast of
Bali. It is the first record of the genus in Bali and the specimens belong to two species that are new to
science. They are described, illustrated, COI-barcoded and compared with the other species known from
the Oriental realm in order to confirm the separation of the genus Elaphropeza in two species groups
(Shamshev & Grootaert 2007).
Material and methods
Specimens were collected by sweep netting and preserved in 70% ethanol. Holotypes are conserved in
the Museum Zoologici Bogor, Indonesia (MZB). The specimens from which DNA was extracted are
conserved in the tissue collection at the Royal Belgian Institute of Natural Sciences, Brussels (RBINS),
with the cuticular parts still being recognisable. For the barcoding technique we refer to Nagy et al.
(2013). The evolutionary history was inferred using the Neighbor-Joining method MEGA6 (Tamura et
al. 2013). The optimal tree with the sum of branch lengths = 1.87955927 is shown. The percentages
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 evolutionary distances were
computed using the Maximum Composite Tikelihood method and are in the units of the number of base
substitutions per site. The analysis involved 39 nucleotide sequences. Codon positions included were
1 st + 2 nd + 3 rd + Noncoding. All positions containing gaps and missing data were eliminated. There was
a total of 640 positions in the final dataset. Evolutionary analyses were conducted in MEGA6 (Tamura
et al. 2013).
Results
Class Insecta Tinnaeus,1758
Order Diptera Linnaeus, 1758
Superfamily Empidoidea Latreille, 1804
Family Hybotidae Macquart, 1823
Subfamily Tachydromiinae Meigen, 1822
Elaphropeza Macquart, 1827
Elaphropeza Macquart, 1827: 86 (as genus). Type-species: Tachydromia ephippiata Fallen, 1815: 11,
by monotypy.
Ctenodrapetis Bezzi, 1904: 351 (as subgenus of Drapetis ). Type-species: Ctenodrapetis ciliatocosta
Bezzi, 1904 by subsequent designation of Melander (1928: 309).
Elaphropeza triseta sp. nov.
urn:lsid:zoobank.org: act: CD 141962-6947-4418-AB0B-25A25A2F41E7
Figs 1-4
Diagnosis
A species of the E. ephippiata group; recognised by almost entirely yellow thorax, only postnotum
brownish along posterior margin; antenna with scape and pedicel yellow; postpedicel short, about 1.5
times as long as wide, brown; legs yellow, with only tarsomere 5 dark brown, hind tibia with 3 brownish
black anterodorsal bristles; halter yellow.
2
GROOTAERT R et al ., New coastal Elaphropeza (Hybotidae) from Bali
Etymology
The specific epithet refers to the presence of three anterodorsal bristles on the hind tibia.
Type material
Holotype
INDONESIA: $, Bali, Amed, Bunutan, beach, 8°20’ 19.44” S, 115°39’53.39” E, 2 Jan. 2014, leg. P.
Grootaert & I. Van de Velde (MZB).
Paratypes
INDONESIA: 2 $$, 2 §§, Bali, Amed, same provenance as holotype (RBINS); 2 2 5?, Bali,
Arya Amed, Bunutan, 8°20’38.89” S, 115°40’21.81” E, 25 Dec. 2013, leg. P. Grootaert & I. Van de
Velde (RBINS).
Description
Male (Fig. 1)
Length. Body 2.3 mm, wing 2.1 mm.
Head. Occiput entirely black, largely shining, with yellowish setae; inner verticals long inclinate, outer
verticals hardly prominent. Anterior ocellars long proclinate, posterior ocellars minute. Frons subshining,
very narrow, above antennae about as broad as anterior ocellus. Antenna with scape and pedicel yellow,
postpedicel and stylus brownish; pedicel with circlet of subequally short setulae; postpedicel short.
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European Journal of Taxonomy 112: 1-10 (2015)
nearly 1.5 times as long as wide; stylus very long, nearly 4.0 times as long as postpedicel, with short
pubescence. Proboscis brownish yellow. Palpus yellow, small, rounded, silvery pubescent, bearing
scattered setulae, subapical seta short.
Thorax. Almost entirely yellow, only antepronotum with brownish spot dorsally, postnotum brownish
along posterior margin, hypopleuron brownish on extreme apex and metapleuron yellowish brown on
upper part anteriorly; shining, with yellowish setation; anterior spiracle brownish yellow, posterior
spiracle brown. Prothoracic episterna lacking long upturned seta just above fore coxa, with minute
setula on upper part. Postpronotal seta hardly prominent. Mesonotum with 2 long notopleurals, 1 shorter
postsutural supra-alar, 1 long postalar and 4 scutellars (apical pair very long, cruciate; lateral pair short);
acrostichals short, arranged in 4 irregular rows anteriorly but becoming sparser and biserial posteriorly,
distinctly separated from dorsocentrals, lacking on prescutellar depression; dorsocentrals arranged in
3-4 irregular rows, as long as acrostichals, 1 prescutellar pair very long.
Legs. Long, slender, almost entirely yellow, only tarsomere 5 of all legs dark brownish. Coxae and
trochanters with unmodified setation. Fore femur slightly thickened, with rows of minute anteroventral
and posteroventral setae, long seta near base and 1 subapical bristle anteriorly. Fore tibia lacking
prominent bristles (except subapicals). Mid femur with rows of minute anteroventral and posteroventral
spinule-like setae (more numerous near base), 1 long seta near base and 1 long subapical bristle anteriorly.
Mid tibia only with 1 black, short, subapical spine, lacking prominent setae (except subapicals). Hind
femur with short anteroventrals and 3 erect dorsal setae near base. Hind tibia with 3 brownish black
anterodorsal bristles; apical projection small, rounded, brownish. Tarsi of all legs unmodified, with
unmodified setation.
Figs 2-4. Elaphropeza triseta sp. nov. Male terminalia. 2. Right epandrial lamella. 3. Epandrium with
cerci. 4. Left surstylus. c = cerci; Is = left surstylus; rs = right surstylus. Scale bar = 0.1 mm.
4
GROOTAERT R et al ., New coastal Elaphropeza (Hybotidae) from Bali
Wings. Normally developed, finely infuscate, covered with uniform microtrichia; veins mostly yellowish
brown; basal section of M 1+2 , crossveins bm-cu and r-m pale. Costal vein with moderately long, uniform
setulae along anterior margin. Costal bristle long, brown. Costal index: 41/31/31. Vein Rs long, slightly
longer than crossvein bm-cu. Vein R 0+3 evenly bowed. Veins R 4+ . and M |+n somewhat divergent near wing
apex, R 4+5 slightly bowed, M 1+ , straight. Vein CuAj reaching wing margin. Vein A x lacking. Crossvein
bm-cu slightly oblique. Crossvein r-m before middle of cell bm. Squama yellowish, with long, brownish
yellow setae. Halter yellow.
Abdomen. Tergite 1 pale yellow; tergites 2-3 brownish yellow, subtriangular viewed laterally, divided
medially, with scattered unmodified setae; tergite 4 brown, broadest, with squamiform setae; tergite
5 narrow, undivided, with squamiform setae; an ampoule-like structure below tergite 5; tergites 6-7
brownish yellow, tergite 6 undivided, with minute unmodified setae; tergite 7 undivided, with long
posteromarginal setae; tergite 8 unmodified. Sternites yellowish, with scattered minute setae, sternite 8
with long, posteromarginal setae. Gland-like structure present between tergites 4 and 5, ampoule-like.
Terminalia (Figs 2—4) rather small, with cerci, apical part of right epandrial lamella and left surstylus
brownish, otherwise yellow. Right epandrial lamella bifid (Fig. 2); dorsal lobe with setae on dorsal
margin. Tip with a brush of densely set, short setae. Ventral lobe with strong brown setae on apex. Right
and left cerci equally long. Right cercus with a single long subapical seta, other setae short. Feft cercus
(Fig. 3) bifid; outward lobe with 3 strong brown setae on apex. Tip of left epandrial lamella with 4 setae:
1 weak and 3 stronger. Feft surstylus also bifid, with short setae (Fig. 4).
Female
Similar to male; abdomen without ampoule-like gland-like structure; intersegmental area between
tergites 4 and 5 set with short hairs, anteriorly with 2 openings; segment 8 moderately long, tergite
and sternite 8 separated, yellowish brown; tergite 8 entire, with several short to moderately long setae
posteriorly; sternite 8 entire, covered with moderately long setae; cercus small, yellowish.
Distribution
Bali, Indonesia.
Remarks
Elaphropeza triseta sp. nov. is actually the only yellow Oriental species with three anterodorsal setae
on the hind tibiae. In the key of Grootaert & Shamshev (2012), E. triseta sp. nov. resolves closest to E.
acrodactyla (Melander, 1928), described from a female from the Philippines, but the latter has only two
anterodorsal bristles on the hind tibiae and has the postpedicel about three times as long as wide. In E.
triseta sp. nov. the postpedicel is nearly 1.5 times as long as wide.
Elaphropeza balinensis sp. nov.
urn:lsid:zoobank.org:act:D744E850-5F65-4C29-A671-9F9D0872A6A9
Figs 5-10
Diagnosis
A species of the E. biuncinata group; thorax yellow, scutum with narrow, short, brownish patch above
wing; antenna with short postpedicel, about 2.0 times as long as wide; legs yellow, but all tibiae and tarsi
faintly brownish tinged, including apical tarsomere of all legs; hind tibia with 1 short, black anterodorsal
bristle at about middle and 1 similar subapical bristle anteriorly; halter brownish. Scutum entirely
covered with setae (acrostichals and dorsocentrals not differentiated), reaching base of scutellum.
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European Journal of Taxonomy 112: 1-10 (2015)
Etymology
The specific epithet refers to the provenance from Bali.
Type material
Holotype
INDONESIA: Bali, Amed, Bunutan, beach, Arya Amed, 8°20’38.89” S, 115°40’21.81” E, 2 Jan.
2014, leg. P. Grootaert & I. Van de Velde (MZB).
Paratypes
INDONESIA: 3 $$ (1 $ dissected), 2 $ $, Bali, Amed, Bunutan, beach, 8°20’19.44” S, 115°39’53.39”
E, 26 Dec. 2013, leg. P. Grootaert & I. Van de Velde (RBINS).
Description
Male (Fig. 5)
Length. Body 1.8 mm, wing 1.6 mm
Head. Occiput entirely black, largely shining, with yellowish setae; inner verticals long inclinate, outer
verticals short. Anterior ocellars long proclinate, posterior ocellars minute. Frons subshining, very narrow,
above antennae about as broad as anterior ocellus. Antenna with scape and pedicel yellow, postpedicel
and stylus brownish; pedicel with circlet of subequally short setulae; postpedicel short, nearly 2 times as
long as wide; stylus with short pubescence, very long, 2.5-3.0 times as long as postpedicel. Proboscis
brownish yellow. Palpus yellow, small, rounded, bearing scattered setulae, subapical seta short.
Thorax. Almost entirely yellow, scutum with narrow, short, brownish yellow patch above wing on
each side (indistinct in paler specimens), metanotum slightly shadowed on middle part (sometimes only
posteriorly or entirely yellow), mesopleuron posteriorly and sternopleuron on lower part usually with
small, brownish yellow patch; shining, with yellowish setation; spiracles brownish yellow. Prothoracic
episterna with long, upturned seta just above fore coxa. Postpronotal seta not prominent. Mesonotum
with 2 long notopleurals, 1 short postsutural supra-alar, 1 moderately long postalar and 4 scutellars
(apical pair long, cruciate; lateral pair short); scutum covered with uniform, minute setae, extending to
base of scutellum. Acrostichals and dorsocentrals not differentiated, except for 1 pair of long, prescutellar
dorsocentrals.
Legs. Robust, yellow, but all tibiae and tarsi yellowish-brownish tinged. Coxae and trochanters with
unmodified setation. Fore femur slightly thickened, with rows of minute anteroventral and postero ventral
setae, long seta near base and 1 subapical bristle anteriorly. Fore tibia lacking prominent bristles (except
subapicals). Mid femur with rows of minute anteroventral and somewhat longer posteroventral setae, 1
long seta near base and 1 moderately long subapical bristle anteriorly. Mid tibia with row of black, short,
anteroventral spines on apical 34, lacking prominent setae (except subapicals). Hind femur with minute
anteroventrals and 3 erect, dorsal setae near base. Hind tibia with 1 short, black anterodorsal bristle on
about middle and 1 similar subapical bristle anteriorly. Tarsi of all legs unmodified, with unmodified
setation.
Wings. Normally developed, finely infuscate, covered with uniform microtrichia; veins mostly yellowish
brown; basal section of M 1+2 , crossveins bm-cu and r-m pale. Costal vein with moderately long, uniform
setulae along anterior margin. Costal bristle moderately long, brown. Costal index: 29/22/26. Vein
Rs long, nearly as long as crossvein bm-cu. Vein R 0+3 evenly bowed. Veins R 4+5 and M 1+2 somewhat
divergent near wing apex, both slightly bowed. Vein CuA { reaching wing margin. Vein A l lacking.
Crossvein bm-cu oblique. Crossvein r-m near middle of cell bm. Squama yellowish brown, with long,
brownish setae. Halter brownish.
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GROOTAERT R et al ., New coastal Elaphropeza (Hybotidae) from Bali
Abdomen. Tergite 1 largely pale yellow, very narrow, divided medially; tergite 2 broadly concave
medially, undivided; tergite 3 broadest, subrectangular, with squamiform setae; tergites 4-5 narrow,
both with squamiform setae; tergites 6-7 broader than tergite 5, the latter with moderately long,
posteromarginal setae; stemites 3-4 broadly divided, sternite 5 entire, concave anteriorly, stemites 6-7
entire, subrectangular, with scattered minute setae. Gland-like structure present between tergites 3—4
and 4-5, plate-like. Terminalia (Figs 6-10) rather small; epandrium brown, otherwise yellow. Right
epandrial lamella (Fig. 6) with base brown, apex (surstylus) yellowish; basal half of dorsal margin with 4
long, strong setae. Feft epandrial lamella without the usual set of setae dorsally. Feft surstylus composed
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European Journal of Taxonomy 112: 1-10 (2015)
of 3 pieces (Fig. 10), hidden by the large left cercus: lsl with apex indented (Fig. 9), on inner side with
at least 6 setae; ls2 with apex saw-toothed; ls3 with short setae (Fig. 10). Phallus long, with partly coiled
tip (Fig. 8). Right cercus short, with rather long setae. Left cercus very wide, covering the left surstylus;
with only minute setae.
Female
Similar to male but mid tibia without anteroventral spines; abdominal tergites 3-7 of subequal width;
segment 8 short, tergite 8 separated from sternite 8, deeply cleft posteriorly; cercus short, yellowish.
Distribution
Bali, Indonesia.
Remarks
In the key of Grootaert & Shamshev (2012), E. balinensis sp. nov. resolves closest to E. monospina
Shamshev & Grootaert, 2007, known only from Singapore. The latter also has a somewhat maculated
scutum, but it has the hind tarsomere 1 with a spine-like ventral seta near the base. The structure of the
male genitalia is also very different.
Figs 6-10. Elaphropeza balinensis sp. nov. Male terminalia. 6. Right epandrial lamella. 7. Detail of right
surstylus. 8. Epandrium with cerci. 9. Left surstylus, lsl. 10. Left surstylus. lc = left cercus; lsl, ls2, ls3
= 3 sclerites of the left surstylus; ph = phallus; re = right cercus. Scale bar = 0.1 mm.
8
GROOTAERT R et al ., New coastal Elaphropeza (Hybotidae) from Bali
Discussion
Shamshev & Grootaert (2007) proposed two species groups in the Oriental Elaphropeza based on
morphological characters. In species of the E. biuncinata group a long upturned bristle is present just
above the fore coxa and the abdominal tergites 3, 4 and 5 bear squamiform setae (Fig. 5). In species of
the E. ephippiata group the proepisternum is bare just above the fore coxa and the abdominal tergite
3 is covered with ordinary setulae, lacking squamiform setae, which are present on tergites 4 and/or 5
only. In addition, there are several characters in the male genitalia, such as fusion or not of the right
surstylus with the right epandrial lamella, fusion or not of both cerci, short or very long phallus and
more (Shamshev & Grootaert 2007). This preliminary and informal classification is confirmed here by
the COI gene (Fig. 11), with a bootstrap value of 86 for the biuncinata- group. However, it would not
be practical to elevate the two species groups to genus level since the morphological characters are not
always easy to see.
As can be seen further on Fig. 11, the genetic distance between all species is large and always more than
5%, showing that at least for the selected species from the Oriental realm, this separation is already of
considerable age.
20
23
37
56
34
26
23
42
37
66
76
74
991
- AB491266 57_Elaphrapeza_malayensis_Singapore_S ungei_B uloh _ma le
-- AB42570227_Elaphnopez3_malayensis_5irg apone_P ulau_Ubin_m ale
- AB49126639_Elapftropeza_fowi_Singap(we_Semakau_feinale
- A B42570217_E taph na pez a_c han ae_Singapore_S em akaii_ma]e
■ A B4 2570251_E laphropez a_fem inata_S i rcga pore_P u lau_Ubin_fema le
-AB42570229_Elaphropeza_goUae_Singap*re_Semakau_mal'e
39 i- AB42570260_ElaphJopeza_temasek_SingapQne_$ema)cau_male
AB42570214_ElaplvopezaJemase k_S i ngapore_Sungei_Buloh_male
- AB42570245_Ela phropez a_pal pala_$ingapofe_Pu lau_Ubm_fem ale
-A B4 2570220_E I apfiropez a_asi optii la_S ingapore_Pulau_U bi njnale
93 r- AB49l2662G_Elaphropeza_5prcata_SingapDre_Semakau_1iemale
A B42570255_E I aphropeza_spicata_S i ngapore_S ungei_Butoh_male
AB42570219 Elaphropeza trlseta Bali = Buniitan = female
33 AB42570213 Elaphropeza trlseta Bali_Bunutan_male
99 I AB4912fi656_Elaphropeza_kranjiersis_Singapore_Serrakau_male
' AB49126644_Ela phropez a_k ra ojieri si s_Singapore_Serrakau_female
-A B425 7G259_E laphropeza_s[ , .a solhii_S ingaporeJ3emakau_male
- AB42570224_E la phropez a_asexa_Si nga pore_Sema kau_male
39
, A B42570236_E laphropeza^asex ajSinga.pare_Semaka u_fern ale
781 A84257023Q_E laphrapeza_asex a_Singapofe_Semakau_fem ale
■ A B42570226_E laph no peza = ubineo si S_Smgapore_Pulau_Ubin jnale
-- AB42570249_E la phropez a_bezzii_S*ngapore_S unge i_B uloh_ma le
-AB42570221 _EI aphropez a_bezz i l_Singapone_P ul au_U bi n_m ale
- A B42570239_E laphropezajnonos pina_Singapore_P ulauJJ tsn_m al e
99
C
AB42570233_E laphropeza_mondspma_SmgapDre_P ulau_Ubin_male
AB49126616_E la phropez aJurea_Singapore_Sungei_Biiloh_noale
99 r AB42570231 Elaphropeza balmensis Bali^BuhutarJemale
AB42570225 Elaphropeza balmensis Bali_Bumjtan_male
-AB42570243_E laphropeza_biu ncmata_Sirgapore_Sungei_Bulbb_rrale
-A B42 570215_Elaphropeza_acantha_S ingapore_Pulau_IJbin_female
iAB42570244_E.laph ropeza_crassicenc us_Singapof e_P ulau JJ bi n_m ale
39 L AB42570236_E laphropeza_crass icercus_S ingaponeJ 3 ulau_Ubi rwnale
0.02
ephippiata
group
biuncinata
group
Fig. 11. Neighbour-Joining tree of the COI barcodes of a number of selected Southeast Asian Elaphro¬
peza species showing the position of E. triseta sp. nov. and E. balinensis sp. nov. (both underlined) in
the ephippiata- group and biuncinata- group, respectively. The bootstrap value is indicated at the nodes.
The scale of the genetic distance is 2%.
9
European Journal of Taxonomy 112: 1-10 (2015)
Acknowledgements
The authors thank Mr Johan De Grayter for the barcoding of the species. Mr Jonathan Brecko helped
with the photography.
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Manuscript received: 22 October 2014
Manuscript accepted: 10 December 2014
Published on: 13 February 2015
Topic editor: Koen Martens
Desk editor: Danny Eibye-Jacobsen
Printed versions of all papers are also deposited in the libraries of the institutes that are members of
the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National Botanic Garden
of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; National History
Museum, Fondon, United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium;
Natural History Museum of Denmark, Copenhagen, De nm ark
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