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ABSTRACT
Antipodromia new genus (Diptera: Empididae:
Hemerodromiinae) is described. The genus is monotypic
with Hemerodromia radialis Collin, 1928 its type species
and is endemic to New Zealand. Systematic relationships
with Hemerodromia Meigen and other Hemerodromiinae
are discussed.
Keywords. Diptera, Empididae, Hemerodromiinae,
Antipodromia, new genus, Hemerodromia.
INTRODUCTION
Hemerodromiinae (Diptera: Empididae) is a subfamily of small
predatory ies with characteristic inated raptorial forelegs (Fig.
1). The subfamily is well represented in New Zealand (Collin
1928) where despite recent revisions (Plant 1993; 2005; 2007)
more taxa await description and many species may be incorrectly
generically assigned (Plant, 2010). Hemerodromia Meigen
is a large genus to which 140 species are currently assigned,
mostly from the Northern Hemisphere and southern Africa
(Plant 2010). Plant and Sinclair (2008) considered the genus
to be well founded despite considerable regional variation but
supported the views of Collin (1993) and Smith (1969) that the
single species from New Zealand, H. radialis Collin, 1928 had
been incorrectly assigned to Hemerodromia. A cladistic analysis
of Hemerodromiinae based on adult morphology (Plant 2010)
recovered strong support for a monophyletic Hemerodromia
based on several synapomorphies but H. radialis was retrieved
in sister group relationship to Hemerodromia in a poorly
resolved clade which also included Neoplasta, Cladodromia,
Metachela and Chelifera. H. radialis differs from ‘typical’
Hemerodromia in two important apomorphies of the wing;
a short and curved vein R2+3, and loss of cell dm (Fig. 2). H.
radialis retains a plesiomorphic free vein Sc, and presence of
cell cup unlike Hemerodromia. There can be no doubt that H.
radialis is incorrectly assigned to Hemerodromia. The present
work describes a new genus to accommodate H. radialis and
redescribes and illustrates the species.
MATERIALS AND METHODS
Specimens were borrowed from and/or deposited deposited in
the following institutions: AMS, Australian Museum, Sydney,
Australia; BMNH, Natural History Museum, London, UK;
NMWC, National Museum of Wales, Cardiff, UK; MNHN,
Muséum national d’Histoire naturelle, Paris, France; NZAC, New
Zealand Arthropod Collection. The general morphological terms
of McAlpine (1981) and antennal nomenclature of Stuckenberg
(1999) were followed. Interpretation of genitalic homology
followed Cumming et al. (1995) and Sinclair (2000). Maceration
of genitalia was performed in hot (90°C) lactic acid (85% v/v).
RESULTS
Antipodromia new genus
Type species. Hemerodromia radialis Collin, 1928; 42–43. (Figs. 1–5)
Diagnosis
A characteristic genus of the Empididae subfamily
Hemerodromiinae with raptorial forelegs widely separated from
the mid legs and fore femur bearing distinct regular rows of
setae ventrally. Antipodromia can be distinguished from other
Hemerodromiinae by the combination of (1) R2+3 very short,
curving to join C before middle of wing near apex of R1; (2) Wing
vein forks R4+5 and M1+2 present; (3) Vein Sc present, free; (4) cell
dm absent; (5) cell cup present, complete; (6) cell bm narrower
and shorter than br; (7) female abdomen apically elongate and
modied as ovipositor; (8) laterotergite bare.
Description
Head. Subspherical, slightly dorsoventrally compressed; viewed
laterally slightly narrowed anteriorly, about 1.2X as long as deep
at vertex with posterior margin of occiput broader and more
obtuse; viewed dorsally somewhat oval, slightly longer than wide.
Eyes viewed from side rather reniform, longer above than below
with hind margin smoothly concave; narrowly separated on face
and frons. Frons rather longer than face, 3–4X as long as distance
between ocelli, widening gradually behind, anterior ommatidia
enlarged. Ocellar setae small but distinct, diverging. Face narrow,
parallel below eyes, widening towards mouth. Frons & face with
small ne erect setae at eye margins, but frontal setae shorter.
Lower occiput prominent; one pair vertical setae and one pair of
upper postocular occiputals more or less distinguished from sparse
covering of ne hairs on vertex and occiput. Antennae inserted
slightly above middle of head in lateral view; scape slightly longer
than wide, almost bare; pedicel as long as wide, a subapical circlet
of setulae stronger dorsally; postpedicel lanceolate, apically
pointed, about 2X long as wide, covered with minute pile which
is longer dorsoapically; stylus about as long as postpedicel,
microscopically pubescent with a minute bristle-like apical
mechanoreceptor. Mouthparts directed downwards, no longer
than head is deep; proboscis narrow, slightly curved posteriorly,
apically pointed; labellum slightly shorter, apically obtuse with a
few longish setae posteriorly; palp elongate, 4–5X long as wide,
apically pointed, covered with ne pile and longer setae distally.
The Hemerodromiinae (Diptera: Empididae) of New Zealand III.
Antipodromia new genus
A. PLANT
Department of Biodiversity and Systematic Biology, National Museum of Wales, Cathays Park, Cardiff CF10 3NP, United Kingdom.
E-mail: adrian.plant@museumwales.ac.uk
New Zealand Entomologist | Vol 34: 52-55 | February 2011
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Thorax. Elongate, narrowing anteriorly. Scutum in prole
very evenly arched but quite at in prescutellar area (Fig. 1);
in dorsal view about 2X as long as wide between wing bases,
narrowed anteriorly, wider behind with posterior margin almost
linear and postalar calli prominent. Postpronotal lobe weakly
developed but proepisternum slightly produced dorsally, the
suture separating it from scutum contiguous posteriorly with
postpronotal depression and anteriorly with much stronger suture
behind prominent prothoracic ‘collar’. Fused anepisternum +
katepisternum broadly triangular, in prole its longer ventral side
not very strongly arched and only slightly concave on anterior
0.25 immediately behind front coxa but slightly convex on
posterior 0.75 in front of mid coxa. Scutellum irregularly rugose
dorsally. Strong setae limited to a single notopleural and a pair
of apical scutellars. Other setae small and hair-like comprising
numerous biserial acrostichals and uniserial dorsocentrals, and
usually several small hairs near margin of scutum between
notopleural and postpronotal areas; a pair of ne setulae outside
strong scutellar pair. Laterotergite bare.
Legs. Front leg widely separated from mid leg by distance similar
to length of front coxa (Fig. 1); mid and hind leg close together.
Front coxa elongate, about 0.7X length of thorax, all setulae
ne and hair-like. Front femur 1.2X length of front coxa; evenly
inated, widest 0.5 from base where 2X wide as front coxa; two
parallel rows of minute denticle-like setulae ventrally on distal 0.8
and a posteroventral and anteroventral series of stronger bristle-
like setae outside the double row. Front tibia geniculate basally,
slightly curved, 0.8X length of front femur, armed ventrally with
a single row of short sharply pointed setae which articulate with
the double row of denticles on the front femur when the limb is
reexed; a short pointed apicoventral spur with a strong spine-
like seta emerging near its base; a fringe of short closely spaced
proclinate setulae dorsally. Mid and hind coxae short, mid and
hind femora and tibiae slender, short haired; hind tibia with
posteroapical swelling at extreme apex. All tarsi with segments 1–
4 progressively shorter and apical segment about as long as third.
Abdomen. Male abdomen with tergites and sternite quite
strongly sclerotized; terminalia (Fig. 3) normally reexed
vertically but sometimes strongly reexed anteriorly with
part of hypandrium retracted under eighth sternite and tergite.
Hypandrium moderately large, approximately triangular, strongly
keel-shaped posteroventrally; membraneous, very weakly
sclerotized but apically with a strongly sclerotized blunt ended
process (postgonite ?) which is partially fused with membrane
of hypandrium. Epandrium free, rather ovate, narrowed basally,
outer face shallowly convex, more or less enclosing proximal
part of cercus. Cercus free; broad in lateral view (although largely
hidden by enclosing epandrial lobe), narrow in anterior view
(Fig. 4); apical margin with numerous internally directed strong
setae and two peg-like processes. Phallus elongate, reaching far
beyond tip of cerci when extended but more usually retracted such
that its apex does not reach beyond level of apex of epandrium.
Female abdomen with terminal segments elongate (about as long
as segments 5–7 combined), strongly sclerotized, ovipositor-like
(Fig. 5); tergite 8 and sternite 8 narrowly triangular, apically
pointed, slightly upturned, a few minute erect hairs on sternite
8; tergite 10 short, triangular, apically pointed; cercus small,
short, apically blunt, bare apart from minute erect apical hairs.
Spermatheca spherical.
Wing. (Fig. 2). Narrow, anal lobe hardly developed. Vein C
circumambient but weak along posterior margin. Sc free, closely
parallel to R1 for its entire length, fading abruptly apically before
reaching C. Crossvein h present. R1 joining C well before middle
of wing. Rs short with origin distinctly distal to crossvein h at
about 0.3 from base; weak basally where junction with R1 obtuse,
stronger distally about junction with R2+3. R2+3 very short, evenly
curving anteriorly to join C before middle of wing just beyond
apex of R1. Fork R4+5 present, obtuse, R5 2–3X length of R4 and
0.6–0.7X length of stem of R4+5. Fork M1+2 present, acute. Cell br
long, about 0.3 as long as wing, conspicuously broad apically;
cell bm narrower and slightly shorter. Cell cup present, about 0.5
length of bm; A1 and CuA2 strong, their junction at posteroapical
apex of cell cup a smooth but almost rectangular curve; A1
continued as a fold in membrane for long distance beyond apex
of cup. Membrane clear but usually darkened near base of R2+3,
R4+5 and M1+2.
Etymology. The specic epithet is a contraction of ‘antipode’
in reference to the genus’ Southern Hemisphere occurrence and
‘dromia’ (from the Greek dromos, running), a common sufx applied
to hemerodromiine genera in reference to their cursorial habits.
Distribution. Antipodromia appears to be endemic to New
Zealand, being found on North, South and Stewart Islands.
Antipodromia radialis (Collin, 1928). comb. nov.
Hemerodromia radialis Collin, 1928: 42–43. (Figs 1–5)
Description
Head. Black, subshining, ne pale dusting on margins of frons,
Figure 1. Antipodromia radialis comb. nov.: female
habitus. Scale = 1mm.
Antipodromia New Genus
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vertex and occiput; frons and lower postocular margin with
stronger silver-grey microscopic pile; mouthparts and antenna
pale yellowish; setae dark brown or blackish but ne erect setae
on frons and a few hairs around mouth pale.
Thorax. Subshining black; proepisternum and sometimes
extreme front of mesonotum paler, almost yellowish. Mesonotum
greyish dusted, viewed from in front a more polished stripe just
inside line of dorsocentral setae extending posteriorly about
as far as level of notopleural seta; viewed laterally a polished
area outside of line of dorsocentrals extending from behind
postpronotal area, above notopleura, as far as supra-alar area.
Prescutellar depression and scutellum evenly dusted. Pleura with
dusting conspicuous behind insertion of front coxa and along
anterior, dorsal and ventral margins of fused anepisternum +
katepisternum; all setae yellowish brown.
Legs. Yellow, apical tarsal segment vaguely darker; setae and
hairs yellow but front femur with double row of about 20–26
minute black denticle-like ventral setulae and front tibia with
line of pointed ventral seta and apicoventral spine black.
Male. Abdomen black including terminalia, rather evenly but
sparsely covered with ne brownish or black setae; female
abdomen similar but ovipositor contrastingly brownish yellow.
Wing. Membrane tinged greyish, obviously darker and blackish
patch near base of R2+3, R4+5 and M1+2. Veins greyish yellow, darker
about black patch on membrane. Halter white, stem yellowish.
Remarks. This species is widely distributed in New Zealand
from Auckland and the Hauraki Gulf in the North Island as far
south a Stewart Island. Capture dates were between October and
December in the North Island and during January and February
in the South Island and Stewart Island. Plant (2010) mentioned
another species referred to as Hemerodromia Morph A which
resolved in the same clade as A. radialis from which it was
distinguished by having a less strongly keel-shaped hypandrium.
Hemerodromia Morph A however appears to be conspecic with
A. radialis as no signicant differences in hypandrial structure
were seen in fully macerated specimens in this study. The
hypandium is weakly sclorotized, becoming almost transparent
even with slight maceration in lactic acid, and while usually
evident in wet-preserved material, in dry-mounted specimens it
sometimes collapses and is retracted into the eighth sternite and
tergite becoming inconspicuous.
Type material
Collin (1928) described Hemerodromia radialis from a
syntypical series of ‘four males, ve females, Ohakune,
xi., xii.1922, iii.1923’ collected by T.R. Harris (BMNH).
Discrepancies between the published and actual dates on the
labels are probably due to misreading of the label data (Pont,
1995) which read correctly as.- 1 xi.1922; 2 , 3 , xi.1923; 1
,; 1 , 7–14.xii.1922 and 1 , iii.1923.
A male labelled 1–14.xii.1922 is here designated lectotype and
a circular purple lectotype label attached. The other males and
females are here designated paralectotypes and rectangular pink
paralectotype labels attached. Ohakune is in NEW ZEALAND,
North Is., Rangitikei.
Additional material
NEW ZEALAND, North Is.: 1 , Little Barrier Is.
(Coromandel), Awaroa Str., swept on river at from leaf litter
and sparse vegetation, 12.xi.1985, A.R. Plant (NMWC); 1
Tawarau Forest (Taupo), Apple Tree Road, Waterfall Track 1,
7.xii.1986, A.R. Plant (NMWC); 1 1 , Henderson (Auckland),
Opanuku Walkway, on plants by stream between Chardon Pl
and Henderson Park, 28.x.2006, S.E. Thorpe. (NZAC); 1 1 ,
Hamilton (Waikato), Parana Park, 10–11.xi.2006 (Bioblitz), S.E.
Thorpe (NZAC): South Is., 1 , Bluff Hill (Southland), Glory
Track, 15.ii.1982, sweeping, C.F. Butcher (NZAC); 1 , Owaka
(Southland), 15.i.1959, ES Gourlay (NZAC): Stewart Is., 1
1 , Murray Beach, 46.7888S, 168.0017E, NZMG 5368450N
2128220E, 20.i.2000, R.K. Didham (NMWC).
DISCUSSION
Antipodromia is clearly differentiated from Hemerodromia by
several important characters. The apomorphic fusion of vein
Sc with C for a short distance beyond the base and absence
of crossvein h only occurs in Hemerodromia and the Central
American genus Colabris Melander while in Antipodromia
and all other Hemerodromiinae, Sc is free and h is present. In
Antipodromia cell dm is lost with cell bm closed apically by
crossvein bm–cu whereas in Hemerodromia cell dm is apparently
present but fused with bm and crossvein bm–cu is absent (the
combined cell bm+dm being obviously longer than cell br).
This interpretation is supported by cell bm being very short in
Antipodromia (as in most other Hemerodromiini) and not at all
elongated, as might be expected if it were fused with cell dm.
The apomorphic short anteriorly curved vein R2+3 usefully
separates Antipodromia from Hemerodromia although a similar
condition is also found in certain undescribed hemerodromiine
taxa from the Australasian Region. Plant (2010) noted a specimen
from New Caledonia (Nouvelle Caledonie, For. Imf. Mt Mou,
200–250m M.B. Ruiss, 16.xi.1983, L. Matile [MNHN]) as being
possibly congeneric with A. radialis and there are specimens of
a pattern winged species from several Tasmanian localities (in
AMS) which also have R2+3 very short and curved. However,
in both the Tasmanian and New Caledonian species, cells bm
and dm are fused and they can not be assigned to Antipodromia;
Figure 2. Antipodromia radialis comb. nov.: male wing showing
nomenclature of some major veins and cells. Scale = 1mm.
2
Antipodromia New Genus
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additionally, vein Sc is free and h is present precluding their
inclusion in Hemerodromia. The closed anal cell (cell cup)
found in Antipodromia is probably the plesiomorphic state in
Hemerodromiinae (Plant 2010). In contrast the anal cell has been
completely lost in most Hemerodromia although CuA2 (the vein
closing the cell apically) is more or less present in many species
from Australia and southern Africa.
Antipodromia thus appears to be monotypic and endemic to New
Zealand. Although it is abundantly distinct from Hemerodromia
its relationships to other Hemerodromiini are unclear. Plant
(2010) found a sister-group relationship between Hemerodromia
and the remaining Hemerodromiini (Antipodromia, Neoplasta,
Cladodromia, Metachela, Chelifera and several undescribed
genera) characterized by an apomorphic distal origin for Rs and
a female ovipositor more or less developed. Antipodromia is
probably most closely related to Neoplasta, Cladodromia and an
undescribed Australian genus designated GENAU[D] by Plant
(2010). The taxonomy of these forms is currently very confused
as for example, Australasian Neoplasta and Cladodromia are
probably not identical with the nominate genus in the New World.
Knowledge of the systematic position of Antipodromia awaits a
more detailed analysis of these closely related Southern taxa.
So far as is known, the larvae of all Hemerodromiini are aquatic
and adults are usually found in proximity to water bodies. Two
of the site localities for A. radialis are specically noted as being
adjacent to streams. Nearctic species of Neoplasta have a well
developed ovipositor of similar form to that in Antipodromia. The
ovipositor appears to be an adaptation for inserting ova into cracks
in the bark of submerged wood (JR. Harkrider, pers. com.) and it
seems likely that this habit will also be found in Antipodromia.
ACKNOWLEDGEMENTS
I would like to thank Dan Bickel, Trevor Crosby,
Christophe Daugeron, Theresa Howard, Erica McAlister and
Stephen Thorpe for their help in arranging loans of material used
in this study and J. Robert Harkrider for useful discussion of the
form and function of the ovipositor.
REFERENCES
Collin JE. 1928. New Zealand Empididae. Pp. 1-110 British
Museum (Natural History), London.
Collin JE. 1933. Diptera of Patagonia and South Chile,
Part IV Empididae. Pp. 1–334. British Museum (Natural
History), London.
Cumming JM, Sinclair BJ, Wood DM. 1995. Homology
and phylogenetic implications in male genitalia in Diptera
– Eremoneura. Entomologica Scandinavica 26: 121–151.
McAlpine JF. 1981. Morphology and terminology – Adults
Chapter 2. In: Manual of Nearctic Diptera, (co-ordinators.
McAlpine JF, Peterson BV, Shewell GE, Teskey HJ.,
Vockeroth, JR. & Wood DM) Volume 1. Pp. 9–63.
Agriculture Canada Monograph 27. (available free online at
http://www.esc-sec.ca/aafcmono.html)
Plant AR. 1993. Sexual dimorphism in the genus Monodromia
(Diptera: Empididae: Hemerodromiinae). New Zealand
Journal of Zoology 20: 207–210.
Plant AR. 2005. The Hemerodromiinae (Diptera, Empididae)
of New Zealand I. Phyllodromia Zetterstedt. Studia
dipterologica 12: 119–138.
Plant AR. 2007. The Hemerodromiinae (Diptera: Empididae) of
New Zealand II. Chelipoda Macquart. Zootaxa 1537: 1–88.
Plant AR. 2010. Hemerodromiinae (Diptera: Empididae):
a tentative phylogeny and biogeographical discussion.
Systematic Entomology published online 30/09/2010
(DOI:10.1111/j.1365-3113.2010.00547.x).
Plant AR, Sinclair BJ. 2008. New species of Hemerodromia
Meigen from Fiji (Diptera: Empididae). In: Evenhuis, N.L.
& Bickel, D.J. (eds.), Fiji Arthropods XI. Bishop Museum
Occasional Papers 98: 31–52.
Pont AC. 1995. The Type-Material of Diptera (Insecta)
Described by G.H. Verrall and J.E. Collin. Pp 1–223.
Oxford University Press, Oxford.
Sinclair BJ. 2000. 1.2. Morphology and terminology of
Diptera male terminalia. In: PaPp, L. & Darvas, B. (eds.),
Contributions to a Manual of Palaearctic Diptera 1: General and
APplied Dipterology 54–74.
Smith KGV. 1969. The Empididae of Southern Africa (Diptera).
Annals of Natal Museum 19: 1–347.
Stuckenberg BR. 1999. Antennal evolution in the Brachycera
(Diptera), with a reassessment of terminology relating to the
agellum. Studia dipterologica 6: 33–48.
Figures 3-5. Antipodromia radialis comb. nov.: 3, male terminalia, lateral,
scale = 0.1mm; 4, male cercus, anterodrosal, scale = 0.05mm; 5, female
cercus, lateral, scale = 0.4mm. Abbreviations: cer, cercus; epan, epandrium;
hypan, hypandrium; pg, postgonite(?); ph, phallus; S8, sternite 8; T8, tergite
8; T10, tergite 10.
New Zealand Entomologist | Vol 34: 52-55 | February 2011
