Annales de la Société Entomologique de France

International Journal of Entomology

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Kauriphanes n. gen., a new genus of braconid

parasitoid wasp (Hymenoptera: Braconidae:
Doryctinae) from New Zealand

Sergey A. Belokobylskij , Sara Fadia Ceccarelli & Alejandro Zaldívar-Riverón

To cite this article: Sergey A. Belokobylskij , Sara Fadia Ceccarelli & Alejandro Zaldívar-Riverón
(2011) Kauriphanes n. gen., a new genus of braconid parasitoid wasp (Hymenoptera: Braconidae:
Doryctinae) from New Zealand, Annales de la Société Entomologique de France, 47:3-4, 394-401,
DOI: 10.1080/00379271.2011.10697733

To link to this article: https://doi.org/10.1080/00379271.2011.10697733

Published online: 31 May 2013.

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 ARTICLE Ann. soc. entomol. Fr. (n.s.), 2011, 47 (3–4) : 394-401

Kauriphanes n. gen., a new genus of braconid parasitoid wasp

(Hymenoptera: Braconidae: Doryctinae) from New Zealand

Sergey A. Belokobylskij (1), Sara Fadia Ceccarelli (2) & Alejandro Zaldívar-Riverón (2)
(1)
Museum and Institute of Zoology PAN, Wilcza 64, 00–679 Warsaw, Poland
(2)
Colección Nacional de Insectos, Instituto de Biología, Universidad Nacional Autónoma de México, 3er. circuito exterior s/n,
Cd. Universitaria, Copilco, Coyoacán, A. P. 70–233, C. P. 04510., D. F., México

Abstract. A new genus belonging to the braconid wasp subfamily Doryctinae, Kauriphanes n. gen.
(type species K. khalaimi n. sp.), is described from New Zealand. This genus is placed within the
doryctine subtribe Caenophanina. The extent of this subtribe is discussed and the phylogenetic
relationships of three of its genera were investigated using one mitochondrial and one nuclear DNA
sequence markers. Similar to previous studies, the Bayesian analyses performed significantly support
a clade with the included members of Caenophanina as a sister group of a clade with the examined
species of Spathiini sensu stricto. The placement of the Caenophanini within Doryctini, however, is
left pendant to further exhaustive phylogenetic studies. A key to genera and subgenera belonging to
Caenophanina is given.
Résumé. Kauriphanes n. gen., un nouveau genre de guêpe parasitoïde de la famille des
braconides (Hymenoptera : Braconidae : Doryctinae) de Nouvelle-Zélande. Un nouveau genre de
la famille des braconides, sous-famille des Doryctinae, est décrit de Nouvelle-Zélande : Kauriphanes
n. gen. (espèce-type: K. khalaimi n. sp.). Le genre est placé parmi la sous-famille des doryctines
et la sous-tribu Caenophanina. L’interprétation de cette sous-tribu est discutée et les relations
phylogénétiques de trois des genres sont l’objet de recherches grâce à un marqueur mitochondrial et
un marqueur d’ADN nucléaire. Comme dans des études précédentes, les analyses bayésiennes ont
supporté de manière significative un clade qui inclut les membres des Caenophanina comme groupe-
frère d’un clade avec les espèces de Spathiini sensu stricto examinées. Toutefois, le placement des
Caenophanini parmi les Doryctini reste pendante en attendant une étude phylogénétique complète.
Une clé des genres et sous-genres des Caenophanina est fournie.
Keywords: Caenophanina, Spathiini, phylogeny; mitochondrial DNA, nuclear DNA.

T he subfamily Doryctinae represents a large group

of braconid parasitoid wasps distributed in almost
all terrestrial ecosystems, and is one of the most speciose
groups of parasitoid wasps in the tropics. Doryctines
are commonly assumed to be especially diverse in
the Neotropics (Marsh 1997, 2002; Belokobylskij et
al. 2004b; Zaldívar-Riverón et al. 2008); however,
recent surveys are showing the vast diversity of this
group in other biogeographic regions. For instance, a
recent revision of the Australasian (including Australia,
New Guinea, New Zealand, New Caledonia and Fiji)
doryctine fauna revealed the existence of 49 genera,
12 of which were new to science (Belokobylskij et
al. 2004a). Moreover, 12 of these genera (24%) are
endemic to the Australian continent, and 21 (43%)
have only been recorded in Australasia. The generic
doryctine diversity in the Australasian region was also
E-mail: sb@zin.ru
Accepté le 16 juin 2011
found in the above study to have a strong Oriental-East
Palaearctic component, with 15 genera (31%) shared
between these regions, whereas practically no Australian
genera are shared with the Neotropics, suggesting that
the latter two faunas originated independently.
Despite the above revision, the current knowledge
of the New Zealand doryctine fauna is still poor, with
only six genera recorded for this country. One of these
genera, Caenophanes Foerster 1862, does not have
any identified species (Belokobylskij et al. 2004a).
On the other hand, species of two other genera are
endemic to New Zealand, the monotypic Doryctopsis
Belokobylskij, Iqbal et Austin 2004 (with type species
D. neozealandicus Belokobylskij, Iqbal et Austin
2004) and Pseudosyngaster pallidus (Gourlay 1928)
(Belokobylskij et al. 2004a). The members of the
remaining three doryctine genera recorded from New
Zealand were undoubtedly introduced perhaps from
Europe: the originally Palaearctic Spathius exarator
(Linnaeus 1758) and Ontsira antica (Wollaston 1858),
and the widely distributed Monolexis fuscicornis Foerster
1862 (Belokobylskij et al. 2004a).

394

Published online 31 May 2013

New genus of doryctine wasp from New Zealand

Figures 1–12
Kauriphanes khalaimi n. gen. et sp. (female, paratype). 1, head, front view. 2, head, dorsal view. 3, head, lateral view. 4, six basal segments of antenna. 5,
hind femur. 6, hind coxa. 7, mesosoma, lateral view. 8, mesonotum, dorsal view. 9, propodeum, dorsal view. 10, fore wing. 11, hind wing. 12, metasoma
and ovipositor, dorsal view.

395

Here we describe a new doryctine genus from
New Zealand, Kauriphanes n. gen., which is placed
within the subtribe Caenophanina based both on
morphological and DNA sequence data. We also
discuss the composition of the Caenophanina and
provide a key to genera and subgenera belonging to
this subtribe.
Material and methods
Specimens and terminology
Specimens of the new genus were collected in 2009 by A. I.
Khalaim and D. Ward using sweep nets and Malaise traps in the
Kauri forest, located in the Auckland Province of New Zealand.
The Kauri forest is dominated by the tree species Agathis
australis (D. Don) Loudon (Araucariaceae). All specimens
were examined using a MC-2 ZOOM stereomicroscope.
Photographs were taken with a Leica IC 3D digital camera that
was mounted on a Leica MZ16 microscope and using the Leica
Application Suite® imaging system. All photographs and figures
were edited with the program Adobe Photoshop®.
The terminology employed for morphological features, sculpture
and measurements follows Belokobylskij & Maeto (2009). The
wing venation nomenclature follows Belokobylskij & Tobias
(1998), Belokobylskij & Maeto (2009) and Sharkey & Wharton
(1997) (in parentheses). The holotype of the new species is kept
in the New Zealand Arthropod Collection (Auckland, New
Zealand – NZAC), the paratype in the Zoological Institute,
Russian Academy of Sciences (St. Petersburg, Russia - ZISP).
Phylogenetic relationships
We assessed the phylogenetic placement of the new genus
using two of the most commonly examined genetic markers
for braconid wasps: the barcoding region, consisting of a 658
bp fragment of the cytochrome oxidase I (COI) mt DNA
gene, and a ~650 bp fragment belonging to the second and
third domain regions of the nuclear ribosomal 28S gene. We
generated DNA sequence data for the holotype specimen
assigned to the new genus (GenBank accession nos JN710472,
JN710480) and for the following taxa: Caenophanes sp. -
Mexico, Jalisco, Estación de Biología Chamela (GenBank
accession no. HQ201002); Doryctes gylak Telenga (GenBank
accession nos JN710477, JN710485), Hypodoryctes bilobus
(Shestakov) (GenBank accession nos JN710474, JN710482),
and H. fuga Belokobylskij & Chen (two specimens) (GenBank
accession nos JN710475-6, JN710483-4) - Russia,
Primorskiy kray, Anisimovka; Dendrosotinus sp. (GenBank
accession nos JN710473, JN710481) and Rhyssaloides
antipoda Belokobylskij (two specimens) (GenBank accession
nos JN710478-9, JN710486-7) - New Zealand, North Island,
Auckland Province. All amplification and DNA sequencing
procedures were those employed in Zaldívar-Riverón et al.
(2006), except for the COI primers (LepF1, LepR1; Hebert et
al. 2004) and their amplification program, which are described
in Zaldívar-Riverón et al. (2010). We also included in the
analyses a total of 66 previously generated sequences belonging
to 47 doryctine genera and four outgroups (see Appendix),
rooting the reconstructed trees with the mesostoine genus
Mesostoa. These sequences were already published in three of
our previous studies (see Zaldívar-Riverón et al. 2006, 2007,
2008 for GenBank accession numbers).
396
S. A. Belokobylskij, S. F. Ceccarelli & A. Zaldívar-Riverón
Genetic distances and number of nucleotidic differences among
the examined specimens were assessed for the two examined
markers with the p uncorrected distances using PAUP
version 4.0b10 (Swofford 2002). We carried out separate and
simultaneous Bayesian analyses with MrBayes version 3.1.2
(Ronquist & Huelsenbeck 2003). All the analyses were run
in Bioportal (University of Oslo; https://www.bioportal.uio.
no). For each data set, analyses consisted of two independent
runs of 10 million generations each, sampling trees every
1000 generations and using four chains and uniform priors.
The GTR + I + Γ (Lanave et al. 1984) was selected as the
most appropriate model of sequence evolution for the two
gene markers examined using the program jMODELTEST
(Posada 2008) with the AIC criterion. Stationarity occurred
before 500,000 generations in all analyses, but we followed a
conservative approach and discarded the trees sampled during
the first five million generations. The remaining trees were
employed to build a majority rule, fully resolved consensus tree
with posterior probabilities (PP), considering those ≥ 0.95 as
significantly supported (Huelsenbeck & Ronquist 2001).
Results and discussion
Systematic part
Kauriphanes Belokobylskij & Zaldívar n. gen.
Type-species. Kauriphanes khalaimi Belokobylskij & Zaldívar
n. sp.
Diagnosis. Kauriphanes n. gen. is similar to Caenophanes Foerster
and differs from the latter in the loss of second radiomedial vein
(r-m), the presence of four abscissae of radial vein (r, 3RSa,
3RSb, 3RSc) of fore wing, and the fused submedially on long
distance radial (3RSb) and medial (2M) veins.
Description. Head weakly transverse (fig. 2), its width 1.4 times
median length (dorsal view). Ocelli small, in triangle with base
1.2–1.3 times its sides (figs 2, 17). Frons flat, without median
keel. Eyes glabrous. Occipital carina present, complete dorsally,
shortly obliterated below and not fused with hypostomal carina.
Malar suture absent or very fine (fig. 1). Clypeal suture deep
laterally and more shallow upper. Hypoclypeal depression small
and round (figs 1, 15). Face with fine depressions above clypeal
suture. Postgenal bridge present, but narrow. Maxillary palpi
long and 6-segmented, labial palpi short and 4-segmented.
Scape wide and rather long, without apical lobe and basal
constriction. First flagellar segment 0.85–0.9 times as long as
second segment (figs 4, 15).
Mesosoma not depressed (figs 7, 16). Neck of promesosoma
rather short, weakly convex dorsally (lateral view). Pronotal keel
fine, situated in anterior 0.3 of pronotum. Propleural posterior
lobe distinct and rather narrow. Mesonotum distinctly and
gently-roundly elevated above pronotum (figs 7, 16). Median
lobe of mesonotum without anterolateral corners. Notauli deep,
wide, complete, crenulate with granulation, fused to each others
behind middle of mesoscutum, then following shallow and
wide single depression (figs 8, 17). Prescutellar depression long,
deep and sculptured. Scuto-scutellar suture distinct. Scutellum
convex, without lateral carinae, its length equal to maximum
width. Metanotum with short and rounded median tooth.
Subalar depression rather deep and wide. Mesopleural pit deep
and weakly elongated. Sternaulus distinct, rather narrow, long,
almost straight, distinctly crenulate with granulation (figs 7,
16, 19). Prepectal carina distinct. Postpectal carina absent.
New genus of doryctine wasp from New Zealand

Figures 13–21
Kauriphanes khalaimi n. gen. et sp. (female, paratype, photos). 13, body, lateral view. 14, body, dorsal view. 15, head and basal segments of antenna, front
view. 16, basal segments of antenna, head and mesosoma, lateral view. 17, head and mesosoma, dorsal view. 18, posterior part of mesosoma and metasoma,
dorsal view. 19, posterior part of mesosoma, hind leg and metasoma, lateral view. 20, metasoma and ovipositor, lateral view. 21, fore and hind wings.

397

Metapleural flange short, narrow, subpointed apically.
Propodeum with areas distinctly delineated by carinae; lateral
tubercles and propodeal bridge absent (fig. 9).
Pterostigma of fore wing (figs 10, 21) narrow; radial vein (r)
arising distinctly behind middle of pterostigma. Parastigma
almost indistinct. Radial (marginal) cell not shortened. First
radiomedial vein (2RS) mostly absent, present only its more
or less short basal abscissa. Second radial (3RSa) and second
medial (2M) abscissae fused distally on rather long distance
(3RSa+2M), then again divergent. Recurrent vein (1m-cu)
short. Mediocubital vein (M+CU) almost not curved to lon-
gitudinal anal vein (1A). Nervulus (1cu-a) short and weakly or
distinctly postfurcal. Discoidal (first discal) cell petiolate. Paral-
lel vein (2CU) interstitial. Brachial (first subdiscal) cell distally
closed distinctly before recurrent vein. Both transverse anal
veins (1a, 2a) absent. Hind wing (figs 11, 21) with 3 hamuli.
Nervellus (cu-a) present. Submedial (subbasal) cell present and
small. First abscissa of mediocubital vein (M+CU) about half
as long as second abscissa (1M). Recurrent vein (m-cu) present,
but short, strongly antefurcal. Medial (subbasal) cell narrow,
0.4 times as long as hind wing. Radial vein (RS) very strongly
unsclerotised; radial (marginal) cell without additional trans-
verse vein (r). First abscissa of costal vein (C+Sc+R) 0.45–0.5
times as long as second abscissa (SC+R).
Legs. Fore and middle tibiae with dense and slender spines
arranged in almost single row. Hind coxa medium-sized, with
distinct basoventral tooth (fig. 6). All femora with fine dorsal
protuberances. Hind femur 3.4–3.6 times longer than wide (figs
5, 19). Hind tibia with raw of dense white setae on inner distal
margin. Hind tibial spurs rather short. Basitarsus of hind tarsus
0.7–0.8 times as long as second-fifth segments combined.
Metasoma (figs 12, 18–20). First tergite not petiolate, wide
(fig. 12). Acrosternite about 0.2 times as long as first tergite,
its apical margin situated distinctly before spiracles. Dorsope
of first tergite rather small; basolateral lobes absent. Spiracular
tubercles small, situated in basal third of tergite; dorsal carinae
present only basally. Suture between second and third tergites
absent. Second-fifth tergites without separate laterotergites.
Ovipositor about as long as metasoma (fig. 20).
Etymology. From “kauri”, the indigenous name of the tree, in
the forest of which this taxon was collected, and “Caenophanes”,
part of the name of the most related doryctine genus. The
gender is masculine.
Biology. Unknown. Kauriphanes n. gen. is closely
related to Caenophanes Foerster, and thus it might
parasitize larvae of the beetles from the families
Bostrichidae, Buprestidae, Cerambycidae, or
Curculionidae (including Scolytinae) (Shenefelt &
Marsh 1976; Yu et al. 2005; Belokobylskij & Maeto
2009).
Distribution. Australasian region (New Zealand).
Kauriphanes khalaimi Belokobylskij & Zaldívar
n. sp.
(figs 1–21)
Type material. Holotype, ♀ “New Zealand, North Island,
Auckland reg., Huia [37°0’ S, 174°35’ E], coastal forest,
CNIN649, 19.xi.2009, D. Ward” (NZAC). Paratype: 1 ♀,
“New Zealand, North Island, Auckland reg., Huapai [36°46’ S,
398
S. A. Belokobylskij, S. F. Ceccarelli & A. Zaldívar-Riverón
174°33’ E], kauri forest, 25.xi.2009, A.I. Khalaim” (ZISP).
Description. Female. Body length 1.5 (paratype) – 2.7
(holotype) mm; fore wing length 1.6 (paratype) – 2.5 (holotype)
mm.
Head behind eyes (dorsal view) roundly narrowed; transverse
diameter of eye 1.1 times longer than temple (dorsal view).
POL 1.5–2.0 times Od, 0.35–0.40 times OOL; Od 0.15–0.25
times OOL. Eye 1.3 times as high as broad. Malar space 0.6–
0.7 times as high as eye, 1.2–1.3 times higher than basal width
of mandible. Face width 1.2–1.3 times eye height and about
1.3 times height of face and clypeus combined. Clypeus with
distinct flange along lower margin. Hypoclypeal depression oval,
its width 0.6–0.7 times distance from depression to eye, 0.35–
0.40 times width of face. Tentorial pits small. Head strongly
and weakly-roundly narrowed below eyes (front view).
Antenna slender, filiform, more than 22-segmented. Scape 1.8–
2.0 times longer than pedicel. First flagellar segment weakly
curved, 5.0–5.2 times longer than its apical width. Subapical
segments 3.0–3.5 times longer than their width.
Mesosoma. Length 1.8–1.9 times its height. Prescutellar
depression about 0.3 times as long as scutellum. Subalar
depression densely granulate and with short striation. Furrow
before meso-metapleural suture distinctly, densely and shortly
crenulate.
Wing. Length of fore wing 3.6–3.7 times its maximum width.
Pterostigma 7.0–8.0 times longer than width, 1.0–1.1 times
as long as metacarp (R1a). Length of first radial abscissa (r)
1.0–1.2 times maximum width of pterostigma. Radial vein
(r) arising from posterior 0.60–0.65 of pterostigma. All
radial abscissae (r, 3RSa, 3RSb, 3RSc) straight. Second radial
abscissa (3RSa) 2.8–3.0 times longer than first radial abscissa
(r), 2.2–2.6 times longer than third (fused with medial vein)
abscissa (3RSb+2M), 0.5–0.6 times as long as fourth abscissa
(3RSc). First medial abscissa ((RS+M)a) weakly or very weakly
S-shaped, about 0.5 times as long as second abscissa (2M), 3.7–
4.0 times longer than recurrent vein. Basal (1M) and recurrent
(1m-cu) veins distinctly divergent posteriorly. Discoidal (first
discal) cell 1.5–1.6 times longer than width, 1.5–1.6 times
longer than brachial (first subdiscal) cell. Distance between
tip of brachial (first subdiscal) cell and parallel vein (2CU)
1.3–1.6 times longer than recurrent vein. Distance between
nervulus (1cu-a) and basal vein (1M) 0.3–0.8 times nervulus
(1cu-a) length. Hind wing 5.4–5.8 times longer than width.
First mediocubital abscissa (M+CU) 0.5–0.55 times as long as
second abscissa (1M).
Legs. Hind tarsus 0.8–0.9 times as long as hind tibia. Second
tarsal segment short, 0.4 times as long as basitarsus, about 0.9
times as long as fifth segment (without pretarsus).
Metasoma 1.0–1.1 times as long as mesosoma. Length of first
tergite 1.1–1.15 times its apical width; apical width 2.3–2.5
times its basal width. Length of second and third tergites
combined 0.85–0.90 times basal width of second tergite,
0.6–0.7 times their maximum width. Following tergites weakly
protruding behind third tergite. Ovipositor sheath 0.9–1.1
times as long as metasoma, 1.0–1.5 times as long as mesosoma,
0.4–0.6 times as long as fore wing.
Sculpture and pubescence. Head densely granulate; face
granulate or finely granulate-coriaceous, sometimes medially
almost smooth; temple below finely coriaceous to smooth.
Mesoscutum and scutellum very densely and coarsely granulate,
mesoscutum in posterior half with fine or very fine rugulosity.
New genus of doryctine wasp from New Zealand
Mesopleuron entirely densely granulate. Propodeum densely
granulate, sometimes partly with fine striation, with rugosity
along posterior and lateral margins, with areas distinctly
delineated by carinae, areola short and wide, almost as wide
as long, basal carina long, 1.0–1.3 times as long as anterior
fork of areola. Hind coxa densely granulate, finely granulate
or granulate-coriaceous below. Hind femur small reticulate-
granulate or reticulate-coriaceous, finely coriaceous to smooth
below. First tergite entirely densely striate, with additional fine
or distinct reticulation. Remaining tergites entirely smooth.
Vertex almost glabrous or with very short and sparse setae.
Mesoscutum widely glabrous, with very short, semi-erect, and
sparse setae along notauli and marginally, sometimes also with
setae in anterior third. Hind tibia on dorsal side with very
short, semi-erect and rather dense setae; length of these setae
much shorter than maximum width of hind tibia.
Colour. Head brownish yellow or light brown with infuscate
anterior part of frons and face. Mesosoma reddish-brown or
light reddish brown, sometimes (paratype) almost black,
mesoscutum yellowish brown or light reddish brown; metasoma
reddish brown or dark reddish brown. Palpi pale yellow. Antenna
yellow to yellowish brown in basal half, reddish brown to dark
reddish brown or almost black in apical half. Legs yellow or
Figure 22
Bayesian 28S+COI phylogram showing the placement of Kariophanes n. gen. within the subtribe Caenophanina (two independent runs of 10 million
generations each, burning after 5 million generations in both runs). The black circles above nodes indicate posterior probabilities of clades ≥ 0.95.
yellowish brown, hind coxa and femur brown or reddish brown,
sometimes dark reddish brown. Ovipositor sheath black, brown
in basal half. Fore wing distinctly maculate, partly infuscate,
with some hyaline spots. Pterostigma yellowish brown, pale
yellow in basal 0.3–0.4.
Male. Unknown.
Etymology. In honour of Dr. Andrey I. Khalaim, Russian
hymenopterist, one of the collectors of the type material of this
new genus.
Genetic distances and phylogenetic relationships
Our 28S and 28S+COI analyses recovered a
significantly supported clade with the two specimens
of Caenophanes (Caenophanes), the specimen of
Dendrosotinus (Gildoria) and the holotype specimen of
Kauriphanes (only the 28S+COI bayesian tree is shown;
fig. 22). Within this clade, the Mexican specimen
of Caenophanes, which represents the first record
for the genus in the New World, was not recovered
as sister taxon to the Old World Caenophanes in the
simultaneous analysis, though this could be due to the
399

lack of the 28S sequence for the former specimen. The
separate and simultaneous analyses recovered the clade
with the above three genera as the sister group of a
clade with the included species of Spathius.
The COI p uncorrected distances observed
between the specimens of Caenophanes from Old
and New Worlds and among the specimens assigned
to Kauriphanes, Caenophanes and Dendrosotinus were
considerably high, ranging from 86 (14.4%) and 78
to 97 bp (13.7–15.5%), respectively. The COI genetic
distances varied from 77 to 101 bp (13.1–17.4%)
between the members of the above three genera and
the species of Spathius. The p uncorrected distances for
28S among the specimens of the former three genera
and among them and the species of Spathius varied
from 32 to 36 bp (7.9–8.8%) and from 46 to 82 bp
(11.3–17%), respectively.
The subtribe Caenophanina (Belokobylskij 1992)
was erected for several doryctine genera that are
distinguished by having the following diagnostic
features: parallel vein (2CU) of fore wing always
interstitial to cubital vein (1CU); brachial (first
subdiscal) cell closed apically before or at the level
of recurrent vein (1m-cu); subcubital (subbasal) cell
of hind wing short and first mediocubital abscissa
(M+CU) distinctly shorter than second abscissa
(1M); recurrent vein (1m-cu) postfurcal (when first
radiomedial vein (2RS) is present); first radiomedial
vein (2RS) sometimes widely reduced; mesoscutum
and scutellum densely granulate; propodeum usually
with areas delineated by carinae; frons often without
lateral protuberances; ocellar triangle equilateral or
with base larger than its sides; hind wing of male
without stigma-like enlargement; metasomal tergites
of male without any transformation; and basal ring
of male genitalia not elongated. This subtribe was
previously represented by three genera: Caenophanes
Foerster (with two subgenera: nominative and
Pacificophanes Belokobylskij 2010), Dendrosotinus
Telenga 1941 (with two subgenera: nominative and
Gildoria Hedqvist 1974) and Eodendrus Belokobylskij
1998 (Belokobylskij 1992; Belokobylskij et al. 2005;
Belokobylskij 2010). Aforementioned character
combination is also present in Kauriphanes n. gen.
The morphological and molecular evidences therefore
indicate that this new doryctine genus should be also
placed within the subtribe Caenophanina (Doryctini).
Similar to some of our previous molecular
phylogenetic studies (Zaldívar-Riverón et al. 2007,
2008) a clade with the members of Caenophanes and
its related genera consistently appears as the sister
group of a clade with specimens assigned to Spathius
Nees 1819 (Spathiini sensu stricto), which indicates
400
S. A. Belokobylskij, S. F. Ceccarelli & A. Zaldívar-Riverón
the polyphyletic condition of the tribe Doryctini,
from which the subtribe Caenophanina belongs
(Belokobylskij 1992). However, we have decided to
maintain the validity of this subtribe despite the above
molecular evidence until more taxa and genetic markers
are analysed together with morphological information
in order to define the placement and composition of
the different taxa currently assigned to Doryctini.
Key to the World genera and subgenera of the
subtribe Caenophanina
1. First radiomedial vein (2RS) of fore wing always and
almost completely present. Hind coxa usually without
basoventral tooth ........................................................ 2
– First radiomedial vein (2RS) of fore wing always
completely or mainly absent. Hind coxa usually with
basoventral tooth ......................................................... 4
2. Second metasomal tergite with rather distinct and
more or less convergent posteriorly longitudinal
furrows. Acrosternite of the first tergite of female long,
longer than 0.4 of the tergite length. – Palaearctic,
Oriental, Australasian and Afrotropical regions
.................................................. Eodendrus Belokobylskij
– Second metasomal tergite without furrows. Acrosternite
of the first tergite of female always short, shorter than
0.3 of the tergite length (Dendrosotinus Telenga) ...... 3
3. First flagellar segment of antenna more or less
widened, weakly depressed, and at least partly
sculptured on its outer margin. Second metasomal
tergite curved. Brachial (first subdiscal) cell of fore
wing rather wide. – Palaearctic and Oriental Regions
........................................... Dendrosotinus Telenga s. str.
– First flagellar segment of antenna not widened,
subcylindrical, and smooth on its outer margin. Second
metasomal tergite absent or more or less straight.
Brachial cell (first subdiscal) of fore wing usually
narrow. – Palaearctic, Oriental and Afrotropical regions
............................................................ Gildoria Hedqvist
4(1) Second radial (3RSb) and medial (2M) veins widely
fused submedially. Radial vein with four abscissae (r,
3RSa, 3RSb, 3RSc). Second radiomedial vein (2RS)
absent. – Australasian region .......... Kauriphanes n. gen.
– Second radial (3RSb) and medial (2M) veins widely
separated submedially. Radial vein with three abscissae
(r, 3RSa, 3RSb). Second radiomedial vein (2RS) present
and long (Caenophanes Foerster) ................................ 5
5. Propodeum with areas distinctly delineated by carinae,
at least with basolateral areas and areola. Dorsope of the
first metasomal tergite large. Occipital carina dorsally
complete. Sides of pronotum mainly not granulate,
with submedian furrow, which more or less distinctly
or at least partly delineated by carinae; pronotal carina
present. Mesoscutum almost entirely or at least along
notauli setose. – Palaearctic, Oriental and Australasian
regions ................................ . Caenophanes Foerster s. str.
New genus of doryctine wasp from New Zealand
– Propodeum without delineated areas. Dorsope of
the first metasomal tergite small. Occipital carina
dorsally interrupted or obliterated. Sides of pronotum
entirely or almost entirely densely granulate, without
submedian furrow; pronotal carina absent. Mesoscutum
almost entirely glabrous. – Australasian region
........................................... Pacificophanes Belokobylskij
Acknowledgements . The authors are very thankful to Dr
Donald Quicke (Ascot, U.K.) and Dr Kees van Achterberg
(Leiden, the Netherlands) for the review of manuscript. This
work was supported by the Russian Foundation for Basic
Research (grant No. 10–04–00265) and by the Presidium RAS
Program “Origin of biosphere and evolution of geobiological
system” to S.A.B., and by grants given by the “Ministerio de
Ciencia e Innovación” (CGL2010-15786; Spain) and the
Consejo Nacional de Ciencia y Tecnología (CONACyT-511;
Mexico) to A.Z.R.
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Appendix
List with the terminal taxa (with their voucher numbers)
included in the phylogenetic analyses. GenBank accession
numbers and localities for these specimens are found in
Zaldívar-Riverón et al. (2006, 2007, 2008), except for
the specimens sequenced for this work
INGROUP: Acrophasmus IB2, Aivalykus Jo725SB, Aivalykus
Jo741SB, Aivalykus Jo840VA, Allorhogas IB18, Aphelopsia Jo838VA,
Barbalhoa Jo723_739SB, Caenopachys Jo793, Caenophanes
Jo781SB, Caenophanes Mex CNIN-ASDOR613, cf. Ontsira
Jo577, Curtisella IB3, Curtiselloides Jo640SB, Dendrosoter 670938,
Dendrosoter Jo573VA, Dendrosoter Jo920, Dendrosotinus CNIN
648, Doryctes gylak CNIN653, Doryctophasmus Jo922, Euscelinus
Jo871, Hansonorum Jo728SB, Hecabolus Jo650, Heerz Jo715,
Heterospilus IB01, Heterospilus IB04, Heterospilus prosopidis Jo601,
Histeromeroides Jo866, Hypodoryctes bilobus CNIN650, Hypodoryctes
fuga CNIN651, Hypodoryctes fuga CNIN652, Hypodoryctes sibiricus
Jo981, Ipodoryctes Jo929, Johnosonius Jo719, Kauriphanes khalaimi
CNIN649, Labania, Leptorhaconotus Jo665, Liobracon Jo714,
Masonius Jo578SB, Monitoriella Jo733, Neodoryctes AL235_
233VA, Neurocrassus haconensis Jo970IB33, Notiospathius IB05,
Notiospathius Jo594VA, Notiospathius Jo596VA, Odontobracon
Jo712SB, Ondigus Jo834VA, Ontsira imperator Jo971, Pedinotus
Jo742SB, Percnobracon IB19, Platyspathius AL183SB, Plyctes
JO928, Plyctes JO936, Polystenus Jo710SB, Rhaconotus AL0185,
Rhaconotus Jo720SB, Rinamba Jo588SB, Schlettereriella Jo673,
Schlettereriella Jo677, Semirhytus Jo628, Spathiomorpha Jo711SB,
Spathius (Stenophasmus) 671005, Spathius (Stenophasmus) Jo675SB,
Spathius agrili, Spathius IB067, Spathius Jo591VA, Stenocorse Jo750,
Stephanospathius Jo674, Syngaster lepidus, Tarasco Jo926_587.
OUTGROUPS: Andesipolis AY935485, Aspilodemon AY935487,
Ephedrus californicus AJ302938, Mesostoa kerri AJ302930,
Rhyssaloides antipoda CNIN645, Rhyssaloides antipoda CNIN647.
401