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Description and natural history of the irst micropterous Meteorus species...
doi: 10.3897/JHR.@@.7403
REsEARCH ARtiClE
1
www.pensoft.net/journals/jhr
Description and natural history of the first micropterous
Meteorus species: M. orocrambivorus sp. n. (Hymenoptera,
Braconidae, Euphorinae), endemic to New Zealand
Helmuth Aguirre1,†, Scott R. Shaw1,‡, Jocelyn A. Berry2,§, Claudio de Sassi3,|
1 University of Wyoming Insect Museum, Department of Ecosystem Science and Management (3354), 1000
East University Avenue, Laramie, Wyoming 82071 USA 2 Heaton House, Brooklyn, Wellington, New Zealand
3 Centre for International Forestry Research, PO Box 0113 BOCBD, Bogor 16000 Indonesia
† urn:lsid:zoobank.org:author:
‡ urn:lsid:zoobank.org:author:
§ urn:lsid:zoobank.org:author:
| urn:lsid:zoobank.org:author:
Corresponding author: Helmuth Aguirre (haguirre@uwyo.edu)
Academic editor: Gavin Broad | Received 1 March 2014 | Accepted 14 May 2014 | Published @@ @@@@ 2014
urn:lsid:zoobank.org:pub:
Citation: Aguirre H, Shaw SR, Berry JA, de Sassi C (2014) Description and natural history of the irst micropterous
Meteorus species: M. orocrambivorus sp. n. (Hymenoptera, Braconidae, Euphorinae), endemic to New Zealand. Title.
Journal of Hymenoptera Research @@: @–@. doi: 10.3897/JHR.@@.7403
Abstract
Wing reduction is well known in the cyclostome lineage of Braconidae, but very unusual in non-cyclostome groups. A new species from New Zealand, Meteorus orocrambivorus, the irst micropterous species
of the non-cyclostome and cosmopolitan genus Meteorus, is described. Phylogenetic analysis places it
close to M. versicolor, a macropterous parasitoid of macrolepidoptera. Details about its host relationships, plant associations and habitat suggest that the necessity of succeeding in cryptic environments may
explain the wing modiication. A possible case of Batesian mimicry with ants could explain the extreme
sexual dimorphism.
Keywords
Wing reduction, sexual dimorphism, alpine habitat, parasitoid, Orocrambus
Copyright Helmuth Aguirre et al. 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.
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introduction
Wing reduction is relatively common within the New Zealand braconid fauna. Iqbal
et al. (2003) pointed out that 75% of doryctine species displaying wing reductions occur in the Australasian Region, and this proportion is steadily increasing (Austin and
Jennings 2009, Belokobylskij and Kula 2012, Belokobylskij and Austin 2013). Several
hypotheses, some of them overlapping, have been proposed to explain the origins of
wing reduction in groups where it is considered an exception to the general rule, i.e.
taxa mainly composed of winged species, such as the Braconidae. Such taxa rely on
light for dispersal to accomplish mating, foraging and colonization of new habitats.
Rof (1990) reviewed the prevalent knowledge about the evolution of wing reduction,
and sorted the possible causes into four groups: 1) potential tradeofs between light
capacity and fecundity, whereby the development of wings, wing muscles, and the
energetic budget to keep them are negatively correlated with the egg load; 2) relatively
stable and cold environments may boost the selection of lightless forms because migration is not required. Stable environments are characterized by a small variability in
resource supply in time and space (Rof 1990). Under such conditions, females do not
need to travel long distances, and the light apparatus is reduced in favor of increasing
reproduction; 3) increases in latitude and altitude are positively correlated with stable
environments (for example, alpine habitats), and as a consequence, they lead to wing
reduction; 4) a tight association with concealed, protected, and narrow niches drives
the loss of wings because such structures can become a handicap to moving into small
and cryptic habitats.
Wing reduction is displayed in varying degrees, from the total loss of structures
associated with wings, including the tegula, to wings being structurally well-developed
but too short to perform light. To describe this variability, Iqbal et al. (2003) used
the following terminology for parasitoid wasps: 1) macropterous, for specimens having the fore wings fully developed and reaching, or almost so, the abdominal apex; 2)
brachypterous, for specimens with the fore wing tips reaching beyond the posterior
propodeum but not the second metasomal tergite; 3) micropterous, for specimens
whose fore wing tips do not reach the posterior propodeum; and 4) apterous, for specimens with a total absence of wings or, at most, manifesting as small scales no longer
than the tegula.
he family Braconidae (Hymenoptera) is mainly comprised of winged species.
However, wing reduction is a well-known phenomenon among the cyclostome lineage of Braconidae, since 90 species in 44 genera (1.2%) have been reported showing
it (Belokobylskij and Kula 2012). In contrast, only 19 non-cyclostome species in six
genera (0.2%) are known with this character (Belokobylskij and Kula 2012). he noncyclostome euphorine clade (Euphorinae+Meteorinae+Neoneurinae) (Belshaw and
Quicke 2002) is represented by three species in the genus Cosmophorus Ratzeburg
(Belokobylskij and Kula 2012). In Cosmophorus only the male is apterous, and this
extreme sexual dimorphism is particularly remarkable in C. laricio Shaw, a parasitoid
of the bark beetle Pityogenes bistridentatus (Eichof) (Shaw 2009).
Description and natural history of the irst micropterous Meteorus species...
3
Meteorus Haliday (Euphorinae: Meteorini) is a cosmopolitan genus of koinobiont
parasitoids of Coleoptera and Lepidoptera larvae. Its most remarkable characteristic
is the distinctive pendant (meteor-like) cocoon constructed by the last larval instar
(Shaw 1997). Around 326 species have been described worldwide (Yu et al. 2012).
Huddleston (1986) studied the New Zealand fauna and reported seven species. Berry
and Walker (2004) added M. pulchricornis (Wesmael) to the list, an exotic species irst
detected in 1996. All the New Zealand Meteorus species currently known are macropterous. his paper describes the irst micropterous Meteorus species and provides information about its biology and habitat.
Methods
he sampling location was Glynn Wye station at Lewis Pass (42°22.78'S, 172°24'E),
North Canterbury Region, New Zealand (Fig. 14). he Lewis Pass traverses the Southern Alps, which run north-south along much of the South Island of New Zealand. It
is the most northern and lowest (907m) of the three main alpine passes which allow
access between the west and east coasts.
All the Meteorus specimens were reared as solitary parasitoids of caterpillars of
Orocrambus ramosellus Doubleday, O. simplex Butler (Lepidoptera: Crambidae) and
Merophyas leucaniana (Walker) (Tortricidae). he caterpillars were collected by Claudio de Sassi from four locations (42°36.73'S, 172°27.78'E; 42°36.88'S, 172°27.62'E;
42°36.72'S, 172°26.58'E; 42°38.83'S, 172°22.17'E) at three elevations (650 m, 800
m, 1000 m) from November 2008 to January 2009. he sample sites comprise alpine
and subalpine habitats dominated by a mixture of tussock grasses (Poa and Festuca),
representing the native lora component, and exotic pastures accounting for the nonnative component (Barratt et al. 2005).
he caterpillars were hand-picked from the host plants Poa cita Edgar (silver tussock) and Festuca novae-zelandiae (Hack.) (Cockayne, fescue tussock), and subsequently reared to fate in the laboratory (i.e death of caterpillar or emergence of either
adult moth or parasitoid). he collected parasitoids were preserved in vials with 95%
ethanol, and sent to the University of Wyoming Insect Museum (UWIM).
Eleven specimens were pin-mounted for taxonomic description, 21 remained in
alcohol from which 2 legs were sent to Julia Stigenberg at Stockholm University for
DNA analyses as a part of her project about the phylogeny of Euphorinae. Morphological terminology follows Sharkey and Wharton (1997) and Zitani et al. (1998).
Explanatory illustrations are provided in Aguirre et al. (2011). Sculpture terminology
is based on Harris (1979). Specimens were measured using a Leica M80 stereomicroscope with micrometer on a 10x ocular. Digital images were captured with a Leica
M205 C stereomicroscope with digital Leica DFC295 camera kit and processed with
Leica Application Suite Version 3.8.0 auto-montage software. Scanning Electron Microscopy (SEM) images were produced at the University of Wyoming, Robert A, Jenkins Microscopy Facility. Descriptions were made with the DELTA software (Dallwitz
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1974, 1980). Holotypes and paratypes are deposited at the Museum of New Zealand
Te Papa Tongarewa (MONZ). Voucher material is deposited at the University of
Wyoming Insect Museum (UWIM).
Results
Meteorus orocrambivorus Aguirre & Shaw sp. n.
http://zoobank.org/
http://species-id.net/wiki/Meteorus_orocrambivorus
Diagnosis. Occipital carina complete; ocelli small (ocelli-ocular distance 2.0–2.3
× ocellar diameter in females, 1.7 × in males); mandible stout and twisted; notauli
smooth and not distinct in females, but deeply impressed, narrow, distinct and rugose
in males; female micropterous, male macropterous; propodeum smooth in females,
but rugulose-lacunose in males; tarsal claw without lobe; dorsope and laterope absent;
ventral borders of irst tergite almost touching distally; ovipositor 1.9–2.3 × longer
than irst tergite).
Description of holotype female. Body color. Dark brown-ferruginous
Body length. 3.5 mm.
Head. (Fig. 3). Antenna with 16 lagellomeres; lagellar length/width ratios as
follows: F1 = 2.6, F2 = 2.2, F3 = 1.9, F 14 = 1, F 15 = 0.9, F 16 = 2.0; head 1.1 × wider
than high; occipital carina complete; ocelli ocular distance 2.0 × ocellar diameter; head
height 1.4 × eye height; temple length 0.7 × eyes length in dorsal view; vertex in dorsal
view not descending vertically behind the lateral ocelli; frons smooth and polished;
maximum face width 1.4 × minimum face width; face inely rugulose; minimum face
width 0.7 × clypeus width; clypeus punctate; malar space length 0.4 × mandible width
basally; mandible stout and twisted.
Mesosoma. (Figs 2, 4, 7 and 9). Pronotum in lateral view dorsally rugose; propleuron
smooth and polished; notauli smooth and not distinct; mesonotal lobes not deined;
mesoscutum smooth and polished; scutellar furrow with one carina; mesopleuron
smooth but rugulose close to tegula; sternaulus long, wide and rugose; metapleuron
mostly smooth, rugose close to the coxa; suture between propodeum and metapleuron
foveate; propodeum smooth; absence of longitudinal and transversal carinae on
propodeum; median depression on propodeum weakly present.
Wings. Very reduced, at most reaching the scutellum apex (Fig. 2).
Legs. Hind coxa slightly strigose dorsally; the remaining surface irregular and
punticulate; hind femur 4.8 × longer than it is wide; tarsal claw without lobe.
Metasoma. (Figs 1 and 4). Dorsope and laterope absent; ventral borders of irst
tergite almost touching distally; irst tergite smooth and polished except the apical
border with short and convergent costae; ovipositor 2.0 × longer than irst tergite;
ovipositor both not thickened basally and straight.
Description and natural history of the irst micropterous Meteorus species...
5
Figures 1–6. M. orocrambivorus sp. n. 1 Female lateral habitus 2 Female head and mesonotum dorsal
view. he arrow indicates the reduced wing 3 Female head frontal view 4 Female propodeum and irst
metasomal tergite dorsal view 5 Male lateral habitus 6 Male frontal view.
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Figures 7–10. Contrasting diferences between males and females. 7 female head and mesonotum dorsal view 8 male head and mesonotum dorsal view 9 female propodeum dorsal view 10 male propodeum
dorsal view.
Female variation based on paratypes. Body length 3.0–3.1 mm; head with long
and scattered setae; head 1.2 × wider than high; ocelli ocular distance 2.3 × ocellar
diameter; head height 1.3 × eye height; temples length 0.6 × eyes length in dorsal
view; maximum face width 1.3 × minimum face width; face strigulate; minimum
Description and natural history of the irst micropterous Meteorus species...
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face width 0.8 × clypeus width; malar space length 0.3–0.6 × mandible width basally;
pronotum in lateral view dorsally lacunose-foveate, faintly costate ventrally; sternaulus
carinate-foveate; propodeum smooth except a small, punctate patch dorsally; hind
coxa either strigate-punctate, strigate, or with very small and disperse punctures; hind
femur 4.6–5.2 × longer than it is wide; ventral borders of irst tergite touching for a
short distance; irst tergite with costae convergent, faintly demarcated; T2 and T3
slightly coriaceous; ovipositor 1.9–2.3 × longer than irst tergite; ovipositor neither
thickened basally nor curved.
Male variation based on paratypes. (Figs 5, 6, 8 and 10). Body black except the head
with a small testaceous patch on the temple behind the eye; wings hyaline; body length
3.7 mm; antenna with 27 lagellomeres; ocelli ocular distance 1.7 × ocellar diameter;
head height 1.6 × eye height; temples length 0.8–0.9 × eyes length in dorsal view; vertex
in dorsal view not descending vertically behind the lateral ocelli; maximum face width
1.2 × minimum face width; minimum face width equal to clypeus width; malar space
length 0.7–0.9 × mandible width basally; propleuron puncticulate and shiny; notauli
deeply impressed, narrow, distinctive and rugose, with pronounced longitudinal carina;
mesoscutal lobes well deined; central lobe of mesoscutum punctate; scutellar furrow
with seven carinae; mesopleuron smooth and polished; sternaulus rugose; propodeum
rugulose-lacunose; longitudinal and transversal carinae on propodeum absent; median
depression on propodeum absent; hind coxa with very small and dispersed punctures;
wing length 3.6–3.7 mm; second submarginal cell of fore wing not strongly narrowed
anteriorly; vein r 0.5 × length of 3Rsa (fore wing); vein 3RSa 0.8 × length of r-m (fore
wing); vein m-cu of fore wing postfurcal; vein 1M 0.8–1 × length of cu-a (HW); vein
1M 0.8 × length of 1r-m (hind wing); dorsope and laterope absent; apparent dorsopes
as deep grooves in the common dorsopes location; irst tergite with faintly demarcated
and parallel costae, which become more obvious on the apical border.
Comments. he micropterous condition of M. orocrambivorus females is unique
among all known Meteorus. However if the males are compared with the rest of the New
Zealand fauna, M. orocrambivorus seems closest to M. cobbus Huddleston (Huddleston
1986; p. 256, numeral 6 in the key). Males of both species share the following character
states: body mostly black except a small, lighter patch (yellow or testaceous) on the
temple behind the eyes; small ocelli (ocelli ocular distance ≥ 1.5 × ocellar diameter); eyes
almost parallel (maximum face width ≤ 1.2 × minimum face width); mandibles stout
and twisted; propodeum rugulose; dorsopes and lateropes absent, and ventral borders
of irst tergite touching for a short distance. M. orocrambivorus can be separated from
M. cobbus by having antennae with 27 lagellomeres (30–33 in M. cobbus), notauli
narrow, carinate and distinct (broad and reticulated in M. cobbus), and irst tergite
costate (strigose in M. cobbus).
Holotype. Female (point-mounted). NEW ZEALAND, South Island, Lewis Pass,
Hope River Valley, Glynn Wye Station, 42°36.73'S, 172°27.78'E, 650 m; host plant
Poa cita Edgar (silver tussock), host caterpillar Orocrambus ramosellus Doubleday
(Crambidae); collected as solitary parasitoid during the period November 2008 to
January 2009; Claudio de Sassi, collector.
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Figures 11–14. 11 M. orocrambivorus sp. n. cocoon 12 O. ramosellus caterpillar 13 O. ramosellus adult
14 Lewis Pass, the type-locality.
Paratypes. Two females and seven males same data as the holotype; one female same
data as holotype except collected at 42°36.88'S, 172°27.62'E, 800 m; two females and two
males same data as holotype except collected at 42°36.72'S, 172°26.58'E; three females
and two males same data as holotype except the host caterpillar collected feeding on Festuca
novae-zealandiae (Hack.) Cockayne at 1000 m; one female same data as holotype except
the host caterpillar collected feeding on F. novae-zealandiae at 42°36.88'S, 172°26.58'E;
one female same data as holotype except the host caterpillar collected feeding on F. novaezealandiae at 42°36.88'S, 172°26.58'E, 650 m; six females and four males same data as
holotype except the host caterpillar collected feeding on F. novae-zealandiae at 42°36.88'S,
172°26.58'E, 1000 m; one female and one male same data as holotype except the host
caterpillar collected feeding on F. novae- zealandiae at 42°38.83'S, 172°22.17'E, 650 m.
Distribution. NEW ZEALAND, South Island, Lewis Pass, Hope River Valley,
Glynn Wye Station.
Cocoon. (Fig. 11). Length 4.4 mm; width 1.6 mm; honey-brown translucent
except apex cap golden, posteriorly bordered by a dark ring; oval-shaped, densely
wrapped by silk, irregular cap border, anterior end (cap) nipple-like. he cocoon was
found unattached inside a structure built by the caterpillar using grass leaves and silk
(Fig. 12). No trace of a suspending thread was detected.
Biology. (Figs 12 and 13). he information gathered from the type series and
additional rearings indicates that M. orocrambivorus is a solitary parasitoid of larval O.
ramosellus, O. simplex and M. leucaniana. Parasitized Orocrambus have been collected
feeding on F. novae-zealandiae, P. cita. Holcus lanatum L., Anthoxantum odoratum L.,
Description and natural history of the irst micropterous Meteorus species...
9
Agrotis capillaris L., Festuca rubra L. and Rytidosperma setifolium (Hook. f.) Connor &
Edgar. M. leucaniana was collected on F. novae-zealandiae.
Etymology. he species name orocrambivorus is a reference to its feeding habit.
he stem of the speciic epithet refers to the genus name of the host caterpillar, Orocrambus, and the suix comes from the Latin -vorare meaning “devour.”
Discussion
he results of a molecular-based phylogenetic analysis carried out by Julia Stigenberg
(unpublished data) placed M. orocrambivorus close to Meteorus versicolor Wesmael. Meteorus versicolor belongs to clade IIB proposed by Stigenberg and Ronquist (2011);
their phylogenetic analysis partly agrees with Maeto’s work (1990), which was based
on morphology. Clade IIB corresponds with Maeto’s pulchricornis (excluding the colon subgroup) and rubens groups (Stigenberg and Ronquist 2011). heir members are
characterized by having a narrow face, strongly twisted mandibles, absence of a tubercle
on the frons, and short ovipositor (length less than 2 × the length of the irst tergite; Stigenberg and Ronquist 2011). Meteorus orocrambivorus and M. versicolor share a complete occipital carina, slender and twisted mandibles, and no dorsope However, M. orocrambivorus has the tarsal claw without a lobe (tarsal claw with a distinct basal lobe in
M. versicolor) and ventral borders of irst tergite almost touching distally (ventral borders of irst tergite completely joined along its basal half in M. versicolor, Figs 15–16).
Meteorus versicolor is a widely distributed species known from the Eastern and Western
Palaeartic Region and introduced to North America for biological control of Euproctis
chrysorrhoea L. (Lymantriidae) (Muesebeck 1923); its host range comprises about 80
lepidopteran species in ifteen families, mostly macrolepidoptera (Yu et al. 2012, Stigenberg and Shaw 2013). Despite the phylogenetic position of M. orocrambivorus in
the aforementioned analysis, it is diicult to track the origin of the wingless condition
in Meteorus since M. orocrambivorus was the only Australasian species included.
Wing reduction as an adaptation to live in concealed, small and close-itting niches
is a compelling hypothesis to explain wing reduction in several species of Doryctinae
(Seltman and Sharkey 2007, Belokobylskij and Kula 2012, Belokobylskij and Austin
2013). he frequent sampling of wasps with remarkable wing-reduction under the
leaf-litter of forested habitats suggests that the necessity of chasing hosts in cryptic
habitats has shaped the reduction (Iqbal et al. 2003, Seltman and Sharkey 2007, Belokobylskij and Kula 2012, Belokobylskij and Austin 2013). he Costa Rican species
Oroceguera andersoni Seltmann & Sharkey (Braconidae: Doryctinae) is a good example
of a wingless parasitoid associated with forest leaf-litter (Seltmann and Sharkey 2007).
Since Orocrambus caterpillars spin their cocoons at the base of tussock grasses and M.
leucaniana constructs tunnels in the detritus layer surrounding tussocks, M. orocrambivorus female wasps are pressed to succeed in tight spaces.
But a possible adaptation to live in cryptic environments does not fully explain the
remarkable sexual dimorphism. he common pattern of sexual dimorphism in Meteorus
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Figures 15–16. 15 M. orocrambivorus, ventral borders of irst tergite almost touching distally 16 M.
versicolor, ventral borders of irst tergite completely joined along its basal half. White arrows on both
pictures indicate the apical section of the structure. Black arrows indicate the most apical point where the
ventral borders converge.
species is: body size and relative eyes size smaller in males than females, relative ocelli size
larger in males than females, antennae longer in males than females, in some cases darker
body color in males than females, and the obvious absence of ovipositor in males. he
morphological deviation in M. orocrambivorus females is so extreme that the initial assignation of females to the genus Meteorus was dependent on the examination of males. A
leeting glimpse of a M. orocrambivorus female in the ield could lead to confusion because
of its ant-like appearance. he close morphological and behavioral resemblance to ants is
called myrmecomorphy, and it is outlined by a set of departures from the common bauplan in those arthropods having it (McIver and Stonedahl 1993): abdominal constriction,
well developed mandibles, elbowed or clubbed antennae, color change, loss or reduction
of wings, head enlargement and microstructural modiications (changes in surface sculpture and pubescence). Compared with males, M. orocrambivorus females display notable
diferences in color body (Figs 1–6), wing reduction (Fig 2), reduction in number of lagellomeres, relative head size (Fig 7–8) and texture of body surface (Figs 2, 4 and 7–10).
Smooth surfaces on the mesoscutum and propodeum are extremely rare in Meteorus, and
such surfaces displayed by M. orocrambivorus may be unique in the genus. Patterns of
myrmecomorphy relected by modiications on body shape and surface texture may be
explained by the Wasmannian mimicry, a special case of Batesian mimicry: when ants antennate each other, one feature they are looking for to recognize conspeciics is the texture
of the body surface (Rettenmeyer 1970). his behavior matches with changes in sculpture
present in M. orocrambivorus females, but additional ield observations are necessary to
Description and natural history of the irst micropterous Meteorus species...
11
corroborate an ant-mimicry model: 1) possible model ants sharing the same habitat with
M. orocrambivorus, 2) model ants showing a denser distribution than M. orocrambivorus,
and 3) model ants displaying an aggressive behavior or unpalatable to predators (Mappes
and Alatalo 1997).
Acknowledgements
We thank Julia Stigenberg for providing results of the DNA analysis. Dr. Zhaojie
Zhang of the University of Wyoming Microscopy Core Facility is thanked for his kind
assistance and advice with scanning electron microscopy. We are also grateful to the
Museum of New Zealand Te Papa Tongarewa and especially to Mr Ricardo Palma for
his kindness in allowing access to the museum’s Hymenoptera collection. he automontage imaging system was provided by National Science Foundation grant DEB10-20751. Any opinions, indings, and conclusions expressed are those of the author
and do not necessarily relect the views of the National Science Foundation. CdS was
supported by a University of Canterbury Doctoral Scholarship and a Hellaby Trust
Fellowship. he research was funded by the Marsden Fund (UOC-0705, grantholder
Jason M. Tylianakis) and the Miss E. L. Hellaby Indigenous Grassland Research Trust.
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