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The species of the two genera of Khoikhoiinae (Hymenoptera, Braconidae) are revised. Thirteen species are recognized, of which five are new and eight were previously described: Khoikhoia anthelion Sharkey, sp. n., K. lission Mason, 1984, K. oligospilos Sharkey, sp. n., K. semiadusta Mason, 1983, K. solata Mason, 1983, K. townesi Mason, 1983, K. turneri Mason, 1984, Sania browni Sharkey, sp. n., S. capensis Mason, 1983, S. henryi Mason, 1983, S. marjoriae Mason, 1983, S. masneri Sharkey, sp. n., and S. masoni Sharkey, sp. n.. All are from the Cape Region of South Africa, and all but one species are confined to the western Cape. A dichotomous key to species is presented; links to electronic interactive keys and to distribution maps are also included. Based on phylogenetic position and morphological characters, speculations on life history are made, and it is suggested that some species may be parasitoids of wood- or stem-boring Lepidoptera. The DELTA data matrix and images for the key are available at 10.3897/zookeys.20.108.app.1.ik; Intkey files are available at 10.3897/zookeys.20.108.app.2.ik; Lucid files in LIF and SDD format are available at doi:10.3897/zookeys.20.108.app.3.ik and doi:10.3897/zookeys.20.108.app.4.ik. Publishing of DELTA raw data will facilitate future workers to edit keys and to add newly discovered taxa.
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Revision of Khoikhoiinae (Hymenoptera, Braconidae) 299
Revision of Khoikhoiinae (Hymenoptera, Braconidae)
Michael Sharkey1,†, Simon van Noort2,‡, James Whit eld3,§
1 Department of Entomology, University of Kentucky, Lexington, Kentucky 40546-0091, USA 2 Natural
History Division, South African Museum, Iziko Museums of Cape Town, PO Box 61, Cape Town 8000, South
Africa 3 Department of Entomology, 320 Morrill Hall, University of Illinois at Urbana-Champaign, 505 S.
Goodwin Ave, Urbana, IL 61801 USA
urn:lsid:zoobank.org:author:77B8EC3A-442C-4A7A-AF85-A31C27E257F2
urn:lsid:zoobank.org:author:7CCD166F-F1FA-43DA-B582-4E84EAF59AD1
§ urn:lsid:zoobank.org:author:7A98AB5F-552D-4437-8F5D-C593CA713506
Corresponding author: James Whit eld (jwhit e@life.uiuc.edu)
Academic editor: Norman Johnson|Received21 February 2009|Accepted 17 June 2009|Published 14 September2009
urn:lsid:zoobank.org:pub:4AA0293D-DEC8-424C-9053-1CDD965CCC82
Citation: Sharkey M, van Noort S, Whit eld J (2009) Revision of Khoikhoiinae (Hymenoptera, Braconidae). In:
Johnson N (Ed) Advances in the sys tematics of Hymenoptera. Festschrift in honour of Lubomír Masner. ZooKeys 20:
299–348. doi: 10.3897/zookeys.20.108
Abstract
e species of the two genera of Khoikhoiinae (Hymenoptera, Braconidae) are revised.  irteen species
are recognized, of which  ve are new and eight were previously described: Khoikhoia anthelion Sharkey,
sp.n., K. lission Mason, 1984, K. oligospilos Sharkey, sp. n., K. semiadusta Mason, 1983, K. solata Mason,
1983, K. townesi Mason, 1983, K. turneri Mason, 1984, Sania browni Sharkey, sp. n., S. capensis Mason,
1983, S. henryi Mason, 1983, S. marjoriae Mason, 1983, S. masneri Sharkey, sp. n., and S. masoni Sharkey,
sp. n. All are from the Cape Region of South Africa, and all but one species are con ned to the western
Cape. A dichotomous key to species is presented; links to electronic interactive keys and to distribution
maps are also included. Based on phylogenetic position and morphological characters, speculations on
life history are made, and it is suggested that some species may be parasitoids of wood- or stem-boring
Lepidoptera.  e DELTA data matrix and images for the key are available at 10.3897/zookeys.20.108.
app.1.ik; Intkey  les are available at 10.3897/zookeys.20.108.app.2.ik; Lucid  les in LIF and SDD for-
mat are available at: doi:10.3897/zookeys.20.108.app.3.ik and doi:10.3897/zookeys.20.108.app.4.ik.
Publishing of DELTA raw data will facilitate future workers to edit keys and to add newly discovered taxa.
Keywords
South Africa, parasitoid wasps, Lepidoptera, Khoikhoia, Sania, taxonomy, systematics
ZooKeys 20: 299–348 (2009)
doi: 10.3897/zookeys.20.108
www.pensoftonline.net/zookeys
Copyright Michael Sharkey et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
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RESEARCH ARTICLE
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
300
Introduction
Khoikhoiinae is a small subfamily of braconids in the microgastroid complex that is
restricted to the Cape Region of South Africa. It was proposed by Mason (1983), who
included two new genera, Khoikhoia and Sania, and six species, three of each genus. He
also discussed its phylogenetic position within the microgastroid complex, and di er-
entiated it from the morphologically somewhat similar Cardiochilinae. Mason (1984)
added two species that he discovered in the British Museum and added characters that
further di erentiated the two genera. Here we describe  ve new species and re-describe
Masons species. A fully illustrated dichotomous key is presented and links to on-line
interactive and dichotomous keys are included.
Mason proposed generic limits and synapomorphies, and described eight species
based on only 11 specimens. We examined 76 specimens in the course of this study.
Naturally some of Masons (1983, 1984) conclusions now seem to be erroneous in
light of the new data. For example, in Masons (1983) key, he used several characters
to distinguish the two genera that we have discovered to be present in both. We too
are still working with a very limited set of specimens and our conclusions are tentative.
Materials and methods
All species descriptions and the dichotomous key were generated using DELTA soft-
ware (http://delta-intkey.com). Data, species names, characters and character states
were entered into Delta Editor (Dallwitz 1980; Dallwitz et al. 1999).  e tokey”
le was edited to select and weight the characters used for the dichotomous key,
and the modi ed  le was exported from DELTA to produce the key which was then
lightly edited to produce the  nal version (Dallwitz 1974; Dallwitz 1980; Dallwitz
et al. 1993).  e interactive key was produced in a similar manner using the DELTA
le “toint” and the software IntKey (Dallwitz 1980; Dallwitz et al. 1993; Dallwitz
et al. 1995). All source  les and images used in this publication are available http://
sharkeylab.org/sharkeylab/Misc/datasets/DeltaFiles/KhoikhoiinaeDeltaFiles.zip and
in Appendix 1 of the present paper (doi:10.3897/zookeys.20.108.app.1.ds).  ese
les are open to the public and future researchers are welcome to download them if
they wish to modify, correct, or add to it for publication. All new species have been
registered with Zoobank (Polaszek et al. 2005).
Online interactive matrix and dichotomous keys were also produced
using Lucid (www.lucidcentral.org), and are available on Waspweb at: http://
www.waspweb.org/Ichneumonoidea/Braconidae/Keys/index.htm. The key to
Khoikhoiinae is available in three formats. Lucid Phoenix keys are dichotomous
and a choice needs to be made at each key couplet to continue. Lucid matrix keys,
on the other hand, use a different approach where relevant states from multiple
character features can be selected independently until identification is achieved.
For more information concerning Lucid keys visit www.lucidcentral.org <http://
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 301
www.lucidcentral.org>. Files are provided as appendices in two formats enabling
conversion of the Lucid matrix key to other platforms. 1. Lucid Interchange
Format version 3 (LIF3) files are XML-based files that store all the Lucid3 key
data, allowing exchange of the key with other key developers. 2. SDD files are
XML-based files structured using the internationally agreed SDD (Structure of
Descriptive Data) Schema. SDD files may be used to exchange Lucid keys with
other SDD-compliant applications.
Measurements were taken with a Microcode II stage measurement device manu-
factured by Boeckeler Instruments.
Images were taken with three di erent instrument systems: 1. Automontage soft-
ware using a 3CCD JVC digital camera attached to a Leica MZ16 stereo microscope.
2. EntoVision micro-imaging system.  is system included a Leica M16 compound
microscope with a JVC KY-75U 3-CCD digital video camera attached that fed im-
age data to a notebook.  e program Cartograph 5.6.0 was then used to merge an
image series into a single in-focus image. Lighting was achieved using techniques
summarized in Bu ngton et al. (2005), Kerr et al. (2009) and Bu ngton and Gates
(2009). 3. SEM photographs of gold coated specimens and ESEM photographs of
uncoated type series specimens were taken with a Philips XL30 ESEM-FEG  eld-
emission environmental scanning electron microscope at the Imaging Technology
Group, Beckman Institute, University of Illinois, with digital images captured di-
rectly onto computer.
Species descriptions are for the holotype specimen. If the holotype was a male and
female specimens were known, the female specimens was used to describe female spe-
ci c characters. Within the holotype description, variation in other specimens of both
sexes is given in angled brackets, < >. Where there are clear di erences in male and
female specimens in non-sexual characters these are mentioned in a separate section
below the holotype description.
Museum depositories:
American Entomological Institute (AEI) 3005 SW 56th Avenue, Gainesville, FL
32608-5047, USA.
Canadian National Collection of Insects (CNCI) Eastern Cereal and Oilseed Research
Centre, Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling
Ave. Ottawa, Ontario K1A 0C6, CANADA
Illinois Natural History Survey (INHS) 1816 South Oak Street, MC 652, Cham-
paign, IL 61820.
Iziko South African Museum (SAMC) Natural History Division, South African
Museum, Iziko Museums of Cape Town, PO Box 61, Cape Town 8000, South
Africa.
Hymenoptera Institute Collection (HIC) Department of Entomology, University of
Kentucky, Lexington, Kentucky 40546–0091, USA.
e Natural History Museum (BMNH) Cromwell Road, London SW7 5BD, UK.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
302
Life history and evolution
Host associations are unknown for all species of Khoikhoiinae; however, morphological
and phylogenetic clues allow for some conjecture. All reliable host records for members
of the microgastroid braconids are Lepidoptera, although occasional uncorroborated
records from other insect orders have been published, and one reasonable record is
from Trichoptera (Achterberg 2002). Phylogenetic studies of the microgastroids (re-
Figure 1. Ovipositor morphology, showing variation within Khoikhoiinae A Sania marjoriae Mason
1983 B Sania masoni Sharkey, sp. n. C Sania browni Sharkey, sp. n. D Sania masneri Sharkey, sp. n. E
Khoikhoia turneri Mason, 1984 F Khoikhoia townesi Mason, 1983.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 303
Figure 2. Scanning electron micrographs A Khoikhoia anthelion Sharkey, sp. n., female, mesosoma and
rst two metasomal tergites, dorsal view B Sania browni Sharkey, sp. n., female, propodeum and  rst three
metasomal tergites, dorsal view C Sania marjoriae Mason, 1983, female, propodeum and laterotergites,
lateral view D Sania masneri Sharkey, sp. n., male, genitalia E Sania masoni Sharkey, sp. n., male, meta-
soma, lateral view, showing genitalia F Khoikhoia anthelion Sharkey, sp. n., female, ovipositor, lateral view.
viewed in Murphy et al. 2008), although far from robust in support for some clades,
clearly place Cheloninae as sister to the remaining subfamilies, so we can infer that
members of Khoikhoiinae are parasitoids of Lepidoptera. All microgastroids are larval
or egg-larval parasitoids (attacking the egg stage and developing in the larval stage).
Since egg-larval parasitism occurs in all known chelonines and is also present in many
microgastrines, it could possibly be part of the ground plan for the non-chelonine
microgastroids (Ruberson and Whit eld 1996 ).  e very thin, sharp ovipositors of
khoikhoiines (Figs. 1, 2F) are consistent with this scenario.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
304
Many ichneumonoids that emerge from the host larvae after it has spun a pupal
chamber have twisted mandibles that are thought to be e ective in cutting through
the silken threads of their host.  is is the case with many members of Microgastrinae.
e family Agathidinae consists exclusively of parasitoids of Lepidoptera larvae, and
most members have twisted scissor-like mandibles.  e one known exception to this is
in the clade composed of the genera Crassomicrodus and Agathirsia, both of which are
found primarily in semiarid regions of the southwest of the USA, Mexico and Central
America (Pucci and Sharkey 2004). Although it is necessary to extract from only one
Figure 3. Scanning electron micrographs A Sania marjoriae Mason, 1983, female, head lateral view B
Sania masoni Sharkey, sp. n., female, head, lateral view C Khoikhoia anthelion Sharkey, sp. n., female,
antennal scape (showing apical  ange) and details of antennal scrobe, lateral view D Khoikhoia anthelion
Sharkey, sp. n., female, subalar region, lateral view E Sania marjoriae Mason, 1983, female, subalar region,
lateral view F Sania masoni Sharkey, sp. n., female, subalar region, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 305
host record (Acontia cretata Noctuidae), these are parasitoids of caterpillars that do
not spin cocoons, and cutting mandibles are not required to escape the host’s silken
chamber. We suspect, therefore, that the hosts of khoikhoiines do not spin cocoons.
Another odd morphological characteristic possessed by many species of Khoikhoiinae
is a well-excavated antennal scrobe (Figs. 3A–B).  is is a feature that, in the Braconidae
at least, is restricted to taxa attacking wood-boring hosts.  e antennal scrobes function to
protect the antennae as the adult braconid emerges from woody substrate.  ese braconids,
e.g., Helcon, Capitonious, also have robust mandibles much like those of khoikhoiines.
Perhaps the most informative characteristic of Khoikhoiinae is the  ared scape.  e
scape of all species of Khoikhoia, and two species of Sania (S. marjoriae, and S. masneri),
is  ared apically and the apical surface is not a simple ridge, but rather it includes an area
with considerable surface (Figs. 3A–C).  is is another characteristic that, within the
Braconidae, is restricted to species attacking wood-boring hosts, e.g. many Braconinae
including Cyclaulax and Hemibracon (see Figs. 37 and 41 in Quicke 1997). We do not
believe that there has been any published conjecture on function, but we suggest that
these structures are used to help prevent back-slippage when the adult wasp is escaping
its host’s habitat through a wooden tunnel. While moving forward through the tunnel
the scape would be directed posteriorly and housed within the antennal scrobe.  e
anged part of the scape would be directed posterodorsally. As the wasp moved forward
in the tunnel the scape would be pushed against the scrobe but any backwards move-
ment would cause the  ange of the scape to lock into the dorsal surface of the tunnel.
In essence the movement of the wasp forward in the tunnel is analogous to a ratchet.
Four of the six species of Sania do not have a strongly  ared scape and also lack a
well-developed antennal scrobe. Also, the species of Sania have less robust mandibles
compared with those of Khoikhoia.  e four species of Sania that lack a  ared scape (S.
browni, S. capensis, S. henryi, and S. masoni) may not be attacking wood-boring hosts.
In summary, we posit that some members of Khoikhoiinae attack wood-boring (or
possibly stem-boring or woody in orescence boring) Lepidoptera. In orescences and in-
fructescences of Protea species are bored by the larvae of a range of coleopteran and lepidop-
teran species (Coetzee and Giliomee 1987, Wright and Samways 1999, 2000).  e most
likely candidate would be a basal lepidopteran clade that is restricted, or mostly con ned,
to the Cape Region of South Africa, where all species of Khoikhoiinae have been collected.
Another potential host candidate is a lepidopteran that feeds on roots, in which case the
antennal and mandibular modi cations of Khoikhoiinae might be used to tunnel through
densely packed soil. One candidate host clade is the enigmatic Prototheoridae (Lepidoptera:
Hepialoidea).  is family is con ned to southern Africa, and most diverse in the Cape
region of South Africa (Davis 1996). Unfortunately knowledge of its life history is lacking.
Khoikhoiinae are associated with the Fynbos and Succulent Karoo biomes (Ru-
therford et al. 2006), which form the major component of the Cape Floristic region.
ese biomes have been recognized as globally important biodiversity hotspots (My-
ers et al. 2000; Latimer 2005), as well as a center of distribution and endemism for
plants (Cowling et al. 1998), and numerous invertebrate taxa, including the  gitid sub-
family Pycnostigminae and the recently discovered insect order Mantophasmatodea,
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
306
both groups having their center of species richness in the Cape Region (Picker et al.
2002, Bu ngton and van Noort 2007, Damgaard et al. 2008). It is plausible that the
Khoikhoiinae may have diversi ed with the evolution and radiation of the Cape Flo-
ristic Region 3–5 million years ago (Goldblatt 1997; Linder et al. 1992; Linder 2003).
Several of the new species described here were collected in signi cant numbers in
Malaise traps. For example, 21 specimens of Sania masoni were collected over a 20 day
period from a single Malaise trap.  is suggests that khoikhoiines may be locally com-
mon, and is consistent with the idea some species or their hosts are gregarious.
Relationships among member subfamilies in the microgastroid complex are dis-
cussed by Mason (1983) and Whit eld and Mason (1994) with respect to morpho-
logical evidence, and by Murphy et al. (2008) with respect to molecular evidence from
seven genes. Despite analyses of a relatively large amount of DNA sequence data (Mur-
phy et al. 2008) and of a signi cant data set of comparative morphological characters
(Whit eld and Mason 1994), the exact placement of Khoikhoiinae among the non-
cheloninae microgastroids is still uncertain, although its inclusion within this complex
is highly supported. Our understanding is likely to improve considerably when the host
biology and larval stages of khoikhoiines become known, and when a larger sample of
the fauna can be included in molecular phylogenetic analysis (the study by Murphy
et al. (2008) featured only one Khoikhoia and three Sania species).  e recent study
of bracovirus origins in the microgastroid complex by Bezier et al. (2009) attempted
to con rm the presence of bracovirus structural genes across the complex using PCR,
but was unable to con rm the presence of these genes in either Sania or Khoikhoia. It
is nevertheless likely, due to the phylogenetic position of khoikhoiines, that they do
contain mutualistic bracoviruses, and that the negative PCR results were due to either
mismatches in primer design or marginal specimen quality for molecular study.
Key to genera and species of Khoikhoiinae
An illustrated, interactive IntKey is available at http://sharkeylab.org/sharkeylab/shar-
keyKeys.php. Interactive Lucid matrix and Lucid Phoenix keys are available at: http://
www.waspweb.org/Ichneumonoidea/Braconidae/Keys/index.htm
1. Clypeus with medial tooth. ..................................................Khoikhoia .....2
1’ Clypeus without medial tooth..................................................... Sania .....8
11'
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 307
22'
2(1). Several large punctures arranged in a longitudinal row on middle lobe of
mesoscutum posteromedially...............Khoikhoia anthelion Sharkey, sp. n.
2’ Medial lobe of mesoscutum lacking longitudinal row of large punctures pos-
teromedially ................................................................................................ 3
33'
3(2). Notauli entirely crenulate. ..........................................................................4
3’ Notauli crenulations weak and partly e aced ................................................
......................................................... Khoikhoia oligospilos Sharkey, sp. n.
44'
4(3). Mesopleuron at least partly smooth with punctures ....................................6
4’ Mesopleuron entirely rugosopunctate or rugose ..........................................5
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
308
7(6). Median tergite 1 mostly smooth with weak microsculpture ..........................
............................................................................. Khoikhoia solata Mason
7’ Median tergite 1 mostly rugose .............................Khoikhoia lission Mason
77'
6(4). Mesosoma entirely black except tegula somewhat paler .................................
.....................................................................Khoikhoia semiadusta Mason
6’ Mesosoma partly brown ..............................................................................7
66'6'
55'
5(4). Middle lobe of scutum smooth with punctures anteromedially, otherwise ru-
gosopunctate ....................................................... Khoikhoia turneri Mason
5’ Middle lobe of scutum entirely rugosopunctate ............................................
...........................................................................Khoikhoia townesi Mason
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 309
9(8). Metapleuron mostly rugose, lacking a large smooth lightly punctate area. .....
................................................................................ Sania capensis Mason
9’ Metapleuron partly rugose but with a large smooth lightly punctate area. .....
..................................................................... Sania browni Sharkey, sp. n.
99'
8(1). Discrimen smooth or with a few barely perceptible crenulae .......................9
8’ Discrimen crenulate ..................................................................................10
8'8
1010 10'
10(8). Scape ared apicoanteriorly with expanded apical surface .........................11
10’ Scape not  ared apicoanteriorly and lacking expanded apical surface ........12
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
310
Taxonomy
Khoikhoiinae Mason, 1983
Description. Head. Number of  agellomeres variable within species ranging from
about 20–40, antenna often sexually dimorphic, female  agellum compressed api-
cally with  agellomeres shorter and wider than males, sometimes  agellomeres trans-
verse in females, though usually slightly to 3× longer than wide;  agellomeres some-
times sexually dimorphic with  agellum of females tapering strongly towards apex,
males tapering slightly as in most other microgastroids; (this type of dimorphism
is also found in a number of species of Cheloninae, e.g., Ascogaster abdominator
(Dahlbom, 1833));  agellar placodes short, about 1/3 to 1/4 length of  agellom-
eres in males, not arranged into two rows; antennal scrobe present, though weak
in most species of Sania; frons with elevated, coarsely sculptured, protuberance lat-
erad antennal scrobe, e ectively increasing functional depth of scrobe, protuberance
11 11'
12'12
12(10). Median tergite 1 partly smooth posterolaterally, rugose anteriorly .................
...................................................................... Sania masoni Sharkey, sp. n.
12’ Median tergite 1 completely rugose .............................Sania henryi Mason
11(10). Median tergite 1 partly smooth posterolaterally, rugose anteriorly; posterola-
teral margin straight. .....................................Sania masneri Sharkey, sp. n.
11’ Median tergite 1 completely rugose; posterolateral margin curved ................
.............................................................................. Sania marjoriae Mason
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 311
variously developed from weak to strong depending on species and generally more
developed in members of Khoikhoia; malar suture distinct; mandible thick and long,
not greatly twisted to function as scissors; temple large, presumably to hold large
mandibular muscles; maxillary palpus 5-segmented, basal two segments fused; la-
bial palpus 4-segmented; labrum setose, semicircular, usually exposed but capable
of folding behind the clypeus, clypeus weakly concave (Sania) or with median and
sometimes lateral teeth (Khoikhoia).
Mesosoma. Propleuron lacking carina on posterolateral margin and lacking
ventral  ange overlapping ventral corner of pronotum; subalar region of meso-
pleuron with a smooth glabrous area posteriorly, more pronounced in species of
Khoikhoia; notauli impressed, crenulate and meeting posteriorly where they gener-
ally extend to the transscutal articulation; transscutal articulation complete and
depressed; postscutellar depression well developed, more so in Khoikhoia; propo-
deum mostly rugose with median longitudinal carina; epicnemial carina absent;
tarsal claws simple; hind tarsomeres lacking longitudinal ridge of setae; hind basi-
tarsus and to a lesser degree some other hind tarsomeres laterally compressed; api-
cal abscissa of Rs of forewing decurved and not tracheated; forewing 1a crossvein
long and strong; forewing 2a crossvein absent; forewing 1cu-a far apical to M;
forewing 2cu-a absent; forewing 2nd submarginal cell quadrate, tapering apically
with r-m crossvein mostly unsclerotized; hindwing crossvein r long but not tubu-
lar; hindwing r-m absent.
Metasoma. Median tergite 1 with a median longitudinal groove; laterotergite 1
membranous (or absent according to Masons 1983 interpretation) and with micro-
striae, spiracle on laterotergite 1; hypopygium size variable from more than half length
of metasoma to approximately 1/5 length of metasoma; ovipositor length variable from
barely exserted to almost as long as the metasoma; setae of ovipositor sheath sometimes
restricted to apex where they can be very long, e.g., K. anthelion.
Distribution. Restricted to the Western, Eastern and Northern Cape Provinces of
South Africa, in localities varying from sea level to 1000 meters. Distribution map is
available at http://sharkeylab.org/sharkeylab/Misc/generalmapper.php?table=khoikhoi
inae&subfamily=Khoikhoiinae.
Biodiversity. Including the new species proposed here, there are 13 species of
Khoikhoiinae. Based on morphological evidence, most of these seem quite distinct,
although K. turneri and K. townesi may constitute one species. Nonetheless, there are
undoubtedly many more species. Of the seven species of Khoikhoia, all but two are
represented by one specimen, and the two exceptions are known from two specimens.
Two of these species were collected during intensive programs of Malaise trap sam-
pling at two localities spanning a couple of years suggesting that in contrast to Sania,
Khoikhoia species may be rare. Malaise traps seem e ective in capturing specimens,
and further intensive Malaise trap sampling targeting under-collected habitats in the
Cape and perhaps other areas of southern Africa will undoubtedly result in the discov-
ery of many more species.
Hosts. Unknown, probably larval Lepidoptera (See Life History section above).
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
312
Khoikhoia Mason, 1983
Type species: Khoikhoia townesi Mason, 1983
Diagnosis: Clypeus with median and sometimes lateral teeth; mandible large, face
rugose to rugosopunctate; subalar area of mesopleuron with vertical carina weak or
absent, and with posterior convex smooth glabrous area.
Distribution: Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia
Etymology: “ e genus name is feminine, dedicated to the original inhabitants of
the South African cape region, the Khoikhoi, who were known to the Dutch settlers as
Hottentots because of their unique clicking speech.” (Mason 1983: p. 53).
Species Descriptions of Khoikhoiinae
Khoikhoia anthelion Sharkey, sp. n.
urn:lsid:zoobank.org:act:D613345A-984A-4DEF-89AA-C5A91BD6FA84
Holotype female. Body. Length. 6.8 mm. Color. Mostly black with laterotergite 1
pale and the following areas yellowish brown: gena, posterodorsal apex of pronotum,
tegula, legs except coxae, trochanters, and some tarsomeres (Fig. 4A–C). Forewing
mostly weakly infuscate but less so anterobasally and at midlength (Fig. 5E). <Male
paratype very similar to holotype, with somewhat more extensive yellowish brown
color on the pronotum.> Head. Number of  agellomeres 29 <29–30>. Scape  ared
apicoanteriorly with expanded apical surface (Fig. 3C). Antennal scrobe shallow (Fig.
3C); rugose medially, smooth laterally except with few transverse striae immediately
posterad antennal insertion (Fig. 3C). Gena with wide irregular punctures, lacking
distinct rugose striae (Fig. 5A). Face entirely rugose (Fig. 4E). Area between antennal
scrobe and inner orbit of eye with weak protuberance (Fig. 4E). Vertex with strong
coarse punctures (Fig. 4F). Posterior orbit of eye with distinct crenulate margin (Fig.
5A). Mesosoma. Middle lobe of scutum entirely smooth with punctures (Fig. 4D).
Notauli entirely crenulate (Fig. 4D). Mesopleuron entirely rugosopunctate or rugose
(Figs. 3D, 4C). Sternaulus long, occupying most of length of mesopleuron, indicated
by vertically elongated crenulae (Figs. 3D, 4C). Discrimen (median longitudinal
ventral sulcus between mesopleura) crenulate. Metapleuron mostly rugose, lacking
a large smooth lightly punctate area (Fig. 4C). Propodeum entirely rugose without
smooth area posterolaterally (Fig. 5C). Metasoma. Median tergite 1 not distinctly
narrowed posteriorly (Figs. 2A, 5C), completely rugose (Figs. 2A, 5C). Hypopygium
less than 1/3 length of metasoma (Fig. 5D); extending past apical tergum (Fig. 5D).
Ovipositor barely exserted, much shorter than metasoma (Figs. 2F, 5D). Length of
setae of ovipositor sheath as much as 4x as long as width of sheath (Figs. 2F, 5D).
Male. Antenna highly sexually dimorphic with the female antennae shorter and
tapering much more rapidly than those of male.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 313
Diagnosis. Mesopleuron entirely rugosopunctate or rugose (Fig. 4C); middle lobe
of scutum entirely smooth with punctures (Fig. 4D).
Material Examined. Holotype female: South Africa, Western Cape, Kogelberg Na-
ture Reserve, MT, KO98-M42, Mesic Mtn. Fynbos, last burnt c. 1988, 34°16.481’S,
19°01.033’E, 199m, 16.IX-16.X.1999, S. van Noort. SAM-HYM-P0024705 (SAMC)
Paratype. Male same data as holotype (HIC).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia&species=anthelion.
Figure 4. Khoikhoia anthelion Sharkey, sp. n., female, holotype A habitus, lateral view B habitus, dorsal
view C mesosoma, lateral view D mesosoma, dorsal view E head, anterior view F head, dorsal view.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
314
Etymology. Anthelion (Greek) is the diminutive of anthele, tuft or plume of a reed.
e name refers to the unique tuft of setae on the ovipositor sheaths.
Khoikhoia lission Mason, 1984
Holotype female. Body Length. 4.7 mm. Color. Mottled dark brown and yellowish
brown except laterotergite 1 whitish, otherwise metasoma mostly pale brown, head and
Figure 5. Khoikhoia anthelion Sharkey, sp. n., female, holotype A head, lateral view B head, anterior view
C propodeum, metasomal tergites, dorsal view D ovipositor, lateral view E wings F data labels.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 315
mesothorax predominantly yellowish brown (Fig. 6). Forewing entirely infuscate (Fig.
7E). Head. Number of  agellomeres 34. Scape  ared apicoanteriorly with expanded
apical surface. Antennal scrobe deep (Fig. 7B); entirely transversely rugosostriate. Gena
transversely rugosostriate anteriorly, rugosopunctate at midlength, longitudinally ru-
gosostriate posteriorly (Fig. 7B). Face entirely rugose (Fig. 7A). Area between antennal
scrobe and inner orbit of eye with protuberance (Fig. 7A). Vertex with weak punctures
(Fig. 7B). Posterior orbit of eye with distinct crenulate margin (Fig. 7B). Mesosoma.
Middle lobe of scutum entirely smooth with punctures (Fig. 7C). Notauli entirely crenu-
Figure 6. Khoikhoia lission Mason, 1984, female, holotype A habitus, dorsal view B habitus, lateral view
C head, mesosoma, lateral view.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
316
late (Fig. 7C). Mesopleuron at least partly smooth with punctures (Fig. 6C). Sternaulus
weakly indicated by a slight depression, not di erentially sculptured (Fig. 6C) <this is
likely to be variable>. Discrimen (median longitudinal ventral sulcus between mesop-
leura) crenulate. Metapleuron mostly rugose, lacking a large smooth lightly punctate area
(Fig. 6C). Propodeum entirely rugose without smooth area posterolaterally (Fig. 7C).
Metasoma. Median tergite 1 not distinctly narrowed posteriorly (Fig. 7D), completely
rugose but with rugae weak posterolaterally (Fig. 7D).
Diagnosis. Mesopleuron at least partly smooth with punctures (Fig. 6C); notauli entirely
crenulate; mesosoma partly or entirely brown; median tergite 1 completely rugose (Fig. 7C).
Figure 7. Khoikhoia lission Mason, 1984, female, holotype A head anterior view B head, lateral view C
mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 317
Male. Unknown.
Material Examined. Holotype female: South Africa, Witzenberg Vall., S.W. Cape
Prov., 914m, 19.I.1921, R.E. Turner (BMNH).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia&species=lission.
Etymology. Mason (1984) did not include the etymology but the interpreta-
tion seems rather straightforward. Lission (Greek) is the diminutive of lissos, meaning
smooth or polished and referring to the mesosoma which is smooth relative to other
members of the genus.
Figure 8. Khoikhoia oligospilos Sharkey, sp. n., male, holotype A habitus, dorsal view B habitus, lateral
view C mesosoma, lateral view.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
318
Khoikhoia oligospilos Sharkey, sp. n.
urn:lsid:zoobank.org:act:55AB2D05-C992-4B88-A7C3-B93E6BCD70A4
Holotype male. Body. Length. 6.0 mm. Color. Mottled black and brown except
laterotergite 1 whitish, otherwise metasoma mostly brown (Fig. 8). Forewing mostly
weakly infuscate but less so anterobasally and at midlength (Fig. 8A). Head. Number
of  agellomeres 31. Scape  ared apicoanteriorly with expanded apical surface (Fig.
9B). Antennal scrobe deep (Fig. 9E); entirely transversely rugosostriate, somewhat
Figure 9. Khoikhoia oligospilos Sharkey, sp. n., male, holotype A head anterior view B head, lateral view
C mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E head, dorsal view F data labels.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 319
smoother dorsomedially. Gena with wide irregular punctures, lacking distinct ru-
gose striae (Fig. 9B). Face entirely rugose (Fig. 9A). Area between antennal scrobe
and inner orbit of eye with protuberance (Fig. 9A). Vertex with weak punctures
(Fig. 9E). Posterior orbit of eye with distinct crenulate margin (Fig. 9B). Mesosoma.
Middle lobe of scutum entirely smooth with punctures (Fig. 9C). Notauli crenula-
tions weak and partly e aced (Fig. 9C). Mesopleuron at least partly smooth with
punctures (Fig. 8C). Sternaulus absent (Fig. 8C). Discrimen (median longitudinal
ventral sulcus between mesopleura) crenulate. Metapleuron mostly rugose, lacking a
large smooth lightly punctate area, or partly rugose but with a large smooth lightly
punctate area. Propodeum entirely rugose without smooth area posterolaterally (Fig.
9D). Metasoma. Median tergite 1 not distinctly narrowed posteriorly (Fig. 9D),
partly smooth with punctures especially posteriorly, with weak rugae or microsculp-
ture anteriorly (Fig. 9D).
Female. Unknown.
Diagnosis. Notaular crenulations weak and partly e aced (Fig. 9C).
Material Examined. Holotype female: South Africa, Western Cape, Lang-
berg Farm, 3 km 270° W Langebaanweg, MT, LW02-N2-M229, Sand Plain Fyn-
bos, 32°58.461’S, 18°07.344’E, 51m, 29.X-5.XI.2003, S. van Noort. SAM-HYM-
P0024985 (SAMC).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia&species=oligospilos.
Etymology. Oligo- is from the Greek, oligos meaning few, and spilos meaning spot.
e name refers to the reduction of crenulae on the notauli.
Khoikhoia semiadusta Mason, 1983
Holotype male. Body. Length. 7.0 mm. Color. Mostly black with reddish brown infu-
sions on temple, lateral frons, tegula, and parts of legs; laterotergite 1 whitish; lower
gena and malar space yellowish white (Fig. 10). Forewing clear basally, infuscate in
apical half (Fig. 11E). Head. Number of  agellomeres 37. Scape  ared apicoanteriorly
with expanded apical surface (Fig. 11B). Antennal scrobe deep (Fig. 11B); entirely
transversely rugosostriate. Gena entirely longitudinally rugosostriate, striae weaker
posteriorly (Fig. 11B). Face entirely rugose (Fig. 11A). Area between antennal scrobe
and inner orbit of eye with protuberance (Fig. 11A). Vertex with strong coarse punc-
tures (Fig. 10A). Posterior orbit of eye with distinct crenulate margin (Fig. 11B). Meso-
soma. Middle lobe of scutum entirely smooth with punctures (Fig. 11C). Notauli
entirely crenulate (Fig. 11C). Mesopleuron at least partly smooth with punctures (Fig.
10C). Sternaulus weakly indicated at midlength with slightly deeper sculpture (Fig.
10C). Discrimen (median longitudinal ventral sulcus between mesopleura) crenulate.
Metapleuron mostly rugose, lacking a large smooth lightly punctate area. Propodeum
entirely rugose without smooth area posterolaterally (Fig. 11C). Metasoma. Median
tergite 1 not distinctly narrowed posteriorly (Fig. 11D), completely rugose (Fig. 11D).
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320
Female. Unknown.
Diagnosis. Body of mesosoma entirely black except tegula brown (Fig. 10).
Material Examined. Holotype male: South Africa: Jonkershoek, near Stellen-
bosch, 33°57’33.74”S, 18°55’10.23”E, 227m, 24.XII.1970 , V. Whitehead (AEI).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia&species=semiadusta.
Etymology. “From the Latin meaning half tanned by the sun in reference to the
wings, which are brown apically and hyaline basally.” (Mason 1983: p. 55).
Figure 10. Khoikhoia semiadusta Mason, 1983, male, holotype A habitus, dorsal view B habitus, lateral
view C head, mesosoma, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 321
Khoikhoia solata Mason, 1983
Holotype male. Body. Length. 7.2 mm. Color. Mottled black and brown, except
laterotergite 1 whitish, otherwise metasoma mostly brown with some pale yellowish-
brown areas on mesosoma and ventral gena (Fig. 12). Forewing entirely infuscate
(Fig. 13E). Head. Number of  agellomeres 32 <32–33>. Scape  ared apicoanteri-
orly with expanded apical surface (Fig. 13A, B). Antennal scrobe deep (Fig. 13B);
entirely transversely rugosostriate (Fig. 13B). Gena entirely longitudinally rugosos-
Figure 11. Khoikhoia semiadusta Mason, 1983, male, holotype A head anterior view B head, lateral view.
C mesosoma, dorsal view D metasomal tergites, dorsal view E wings F data labels.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
322
triate, striae weaker posteriorly (Fig. 13A, B). Face entirely rugose (Fig. 13A). Area
between antennal scrobe and inner orbit of eye with protuberance (Fig. 13A). Vertex
with weak punctures (Fig. 12A). Posterior orbit of eye with distinct crenulate margin
(Fig. 13B). Mesosoma. Middle lobe of scutum entirely smooth with punctures (Fig.
13C). Notauli entirely crenulate (Fig. 13C). Mesopleuron at least partly smooth with
punctures (Fig. 12C). Sternaulus absent or di cult to discern due to dense sculpture
of mesopleuron (Fig. 12C). Discrimen (median longitudinal ventral sulcus between
mesopleura) crenulate. Metapleuron mostly rugose, lacking a large smooth lightly
Figure 12. Khoikhoia solata Mason, 1983, male, holotype A habitus, dorsal view B habitus, lateral view.
C head, mesosoma, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 323
punctate area (Fig. 12C). Propodeum entirely rugose without smooth area posterola-
terally (Fig. 13C, D). Metasoma. Median tergite 1 not distinctly narrowed posteriorly
(Fig. 13D), partly smooth with punctures especially posteriorly, with weak rugae or
microsculpture anteriorly (Fig. 13D).
Female. Unknown.
Diagnosis. Tergite 1 partly smooth with punctures posteriorly (Fig. 13D); notauli
entirely crenulate (Fig. 13C).
Figure 13. Khoikhoia solata Mason, 1983, male, holotype A head anterior view B head, lateral view C
mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
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324
Material Examined. Holotype male: South Africa: Jonkershoek, near Stellen-
bosch, 33°57’33.74”S, 18°55’10.23”E, 227m, 18.XII.1970, V. Whitehead (AEI).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia&species=solata.
Etymology. “From Latin meaning sunburned, an allusion to the color and to the
fully infuscated wings.” (Mason 1983: p. 54).
Figure 14. Khoikhoia townesi Mason, 1983, female, holotype A habitus, dorsal view B habitus, lateral
view C head, mesosoma, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 325
Khoikhoia townesi Mason, 1983
Holotype female. Body Length. 6.4 mm. Color. Mottled dark brown and yellowish
brown except laterotergite 1 and margins of most metasomal sclerites whitish, head
and mesothorax predominantly yellowish brown (Fig. 14). Forewing mostly weakly
infuscate but less so anterobasally and at midlength (Fig. 15E), or clear basally, in-
fuscate in apical half. Head. Number of  agellomeres 35 <31–39>. Scape  ared api-
coanteriorly with expanded apical surface (Fig. 15B). Antennal scrobe deep (Fig. 15B);
entirely transversely rugosostriate. Gena transversely rugosostriate anteriorly, longitu-
Figure 15. Khoikhoia townesi Mason, 1983, female, holotype A head anterior view B head, lateral view
C mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
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326
dinally rugosostriate posteriorly (Fig. 15A, B). Face entirely rugose (Fig. 15A). Area
between antennal scrobe and inner orbit of eye with protuberance (Fig. 15A). Vertex
with strong coarse punctures (Fig. 14A). Posterior orbit of eye with distinct crenulate
margin (Fig. 15B). Mesosoma. Middle lobe of scutum entirely rugosopunctate (Fig.
15C). Notauli entirely crenulate (Fig. 15C). Mesopleuron entirely rugosopunctate or
rugose (Fig. 14C). Sternaulus absent or di cult to discern due to dense sculpture of
mesopleuron (Fig. 14C). Discrimen (median longitudinal ventral sulcus between mes-
opleura) crenulate. Metapleuron mostly rugose, lacking a large smooth lightly punc-
tate area (Fig. 14C). Propodeum entirely rugose without smooth area posterolaterally
(Fig. 15C, D). Metasoma. Median tergite 1 not distinctly narrowed posteriorly (Fig.
15D); completely rugose (Fig. 15D). Hypopygium less than 1/3 length of metasoma
(Fig. 1F, 14B); not extending past apical tergum (Fig. 1F, 14B). Ovipositor barely
exserted, much shorter than metasoma (Fig. 1F, 14B). Setae of ovipositor sheath not
longer than 2× width of ovipositor sheath (Fig. 1F).
Male. Unknown.
Diagnosis. Middle lobe of scutum entirely rugosopunctate (Fig. 15C); mesopleuron
entirely rugosopunctate (Fig. 14C). Khoikhoia townesi is very similar to K. turneri,
and may be a senior synonym.  e only appreciable di erences are slight sculptural
di erences on the face and the middle lobe of the mesoscutum, and rather slight color
di erences (see  gure associated with couplet 5 in the key).
Material Examined. Holotype female: South Africa, Grahamstown, 33°18’37.50”S,
26°31’30.22”E, 545m, II.1972, Fred Gess (AEI).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia&species=townesi.
e sole specimen of this species, together with the single specimen of Sania henryi
(Northern Cape Province) are the only representatives of Khoikhoiinae collected out-
side of the Western Cape Province. Like its congeners, Khoikhoia townesi is probably
still associated with Fynbos, as Grahamstown lies at the extreme eastern limits of this
biome.
Etymology. Although not speci ed in Mason (1983), this is a patronym for
Henry Townes.
Khoikhoia turneri Mason, 1984
Holotype female. Body. Length. 6.8 mm. Color. Mottled black and brown except
laterotergite 1 whitish, body, particularly metasoma, predominantly dark brown to
black (Fig. 16). Forewing entirely infuscate (Fig. 17E). Head. Number of  agellomer-
es 34 <30–38>. Scape  ared apicoanteriorly with expanded apical surface (Fig. 17B).
Antennal scrobe deep (Fig. 17B); entirely transversely rugosostriate. Gena entirely
longitudinally rugosostriate, striae weaker posteriorly (Fig. 17A). Face entirely rugose
(Fig. 17A). Area between antennal scrobe and inner orbit of eye with protuberance
(Fig. 17A). Vertex with strong coarse punctures (Fig. 16A). Posterior orbit of eye
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 327
with distinct crenulate margin (Fig. 17B). Mesosoma. Middle lobe of scutum smooth
with punctures anteromedially, otherwise rugosopunctate (Fig. 17C). Notauli entire-
ly crenulate (Fig. 17C). Mesopleuron entirely rugosopunctate or rugose (Fig. 16C).
Sternaulus absent or di cult to discern due to dense sculpture of mesopleuron (Fig.
16C). Discrimen (median longitudinal ventral sulcus between mesopleura) crenulate.
Metapleuron mostly rugose, lacking a large smooth lightly punctate area (Fig. 16C).
Propodeum entirely rugose without smooth area posterolaterally (Fig. 17D). Meta-
soma. Median tergite 1 not distinctly narrowed posteriorly (Fig. 17D); completely
Figure 16. K hoikhoia turneri Mason, 1984, female, holotype A habitus, dorsal view B habitus, lateral
view. C head, mesosoma, lateral view.
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328
rugose (Fig. 17D). Hypopygium less than 1/3 length of metasoma (Fig. 1E, 16B); not
extending past apical tergum (Fig. 1E, 16B). Ovipositor barely exserted, much shorter
than metasoma (Fig. 1E, 16B). Setae of ovipositor sheath not longer than 2× width
of ovipositor sheath (Fig. 1E).
Male. Unknown.
Diagnosis. Middle lobe of scutum smooth with punctures anteromedially, oth-
erwise rugosopunctate (Fig. 17C); mesopleuron entirely rugosopunctate (Fig. 16C).
Material Examined. Holotype female: South Africa, Ceres, Cape Province,
33°22’14.08”S, 19°18”22.10”E, 457m, I.1921, R.E.Turner (BMNH).
Figure 17. Khoikhoia turneri Mason, 1984, female, holotype A head anterior view B head, lateral view.
C mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 329
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Khoikhoia&species=turneri.
Etymology. Although not speci ed in Mason (1984), this is a patronym for R.
Turner, the collector of the holotype.
Sania Mason, 1983
Type species. Sania marjoriae Mason, 1983
Diagnosis. Clypeus weakly concave, lacking teeth; face smooth to weakly
punctate; subalar area of mesopleuron with strong vertical carina, and with
posterior  at smooth to weakly convex glabrous area, not nearly as well developed
as that of Khoikhoia.
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Sania.
Etymology.A feminine generic name, dedicated to the San people, a few of whom
still follow their ancient hunting life in remote and arid parts of southern Africa.
(Mason 1983: p. 51).
Sania browni Sharkey, sp. n.
urn:lsid:zoobank.org:act:035CFE8B-7302-4940-A085-1A58E9E99762
Holotype female. Body Length. 3.5 mm <3.4 – 4.4 mm>. Color. Mostly yellowish
brown except as follows: head mostly black except ventrally; mesosoma including legs
with most sclerites with some black infusions; metasoma mostly brownish black, with
lateral areas of terga yellowish white, and pleural area of tergum 1 white; ovipositor
sheath white basally, brown apically; <quite variable, from mostly black to mostly yel-
lowish brown (Fig. 18); mesosoma always with extensive orangish brown; di ering
from most other species in that all specimens have extensive pale whitish yellow on
all lateral terga (Fig. 18A)>. Forewing entirely infuscate (Fig. 19E). Head. Number
of  agellomeres, 23 <23–25>. Scape not  ared apicoanteriorly and lacking expanded
apical surface (Fig. 18C,F). Antennal scrobe  at (Fig. 18F); entirely smooth with weak
punctures. Clypeus mostly smooth with weak punctures (Fig. 18E); without median
tooth (Fig. 18E, 19A). Gena mostly smooth with weak punctures (Fig. 18E,F). Face
mostly punctate with some weak rugae especially dorsally (Fig. 18E). Area between
antennal scrobe and inner orbit of eye without protuberance (Fig. 18E). Vertex with
weak punctures (Fig. 18D). Posterior orbit of eye lacking distinct crenulate margin
(Fig. 18C). Mesosoma. Middle lobe of scutum entirely smooth with punctures (Fig.
18D). Notauli entirely crenulate (Fig. 18D). Mesopleuron at least partly smooth with
punctures (Fig. 18C, 19C). Sternaulus only present at midlength (Figs. 18C, 19C),
or absent (Fig. 18C). Discrimen (median longitudinal ventral sulcus between mesop-
leura) smooth or with a few barely perceptible crenulae (Fig. 19B). Metapleuron partly
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
330
rugose but with a large smooth lightly punctate area (Figs. 18C, 19C). Propodeum
entirely rugose without smooth area posterolaterally (Figs. 18D, 19D). Metasoma.
Median tergite 1 distinctly narrowed posteriorly (Figs. 2B,19D); mostly smooth with
few rugae near midlength (Figs. 2B, 19D). Hypopygium less than 1/3 length of meta-
soma (Figs. 1C, 18A); not extending past apical tergum (Fig. 1C). Ovipositor barely
exserted, much shorter than metasoma (Figs. 1C, 18A). Length of setae of ovipositor
sheath none longer than 2x width of ovipositor sheath (Fig. 1C).
Male. Antenna not sexually dimorphic.
Figure 18. Sania browni Sharkey, sp. n., female, paratype A habitus, lateral view B habitus, dorsal view
C head, mesosoma, lateral view D head, mesosoma, dorsal view E head, anterior view F head, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 331
Diagnosis. Discrimen smooth or with a few barely perceptible crenulae (Fig. 19B);
metapleuron partly rugose but with a large smooth lightly punctate area (Fig. 19C).
Material Examined. Holotype Female: South Africa, Western Cape, 15km N
Citrusdal, Koedoeskop Farm, Malaise across hillside trail, 32°29’18”S, 18°57’30”E,
220m, 5–25.X.2004, ME Irwin, FD Parker. SA-34 (SAMC)
Paratypes. South Africa, Western Cape: 2 females, 3 males, same data as holo-
type. 1 female, 9 males, 10 km S. Clanwilliam, Malaise on sandy hill, 32°13’39”S,
18°50’50”E, 140m, 5–25.X.2004, ME Irwin, FD Parker, M Hauser. 2 females, 1 male,
Figure 19. Sania browni Sharkey, sp. n., female, paratype A head ventral view B discrimen C mesosoma,
lateral view D mesosoma & metasoma, dorsal view E wings F data labels.
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332
14km NW Robertson@hwy R60, Malaise in dry wash, 33°46.65’S 19°45.69’E, 270
m, 7.X.2002, ME Irwin, FD Parker, RSA-15 (SAMC, AEI, HIC, CNCI, INHS).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Sania&species=browni.
Note: is species is referred to as Sania sp. n. 3 in the molecular study by Murphy
et al. (2008).
Etymology. Named in honor of Brian Brown, a dipterist of outstanding repute
and long time collaborator of the senior author.
Figure 20. Sania capensis Mason, 1983, male, holotype A habitus, dorsal view B habitus, lateral view C
head, mesosoma, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 333
Sania capensis Mason, 1983
Holotype male. Body Length. 4.0 mm. <4.0–4.1 mm> Color. Mottled black and
brown except laterotergite 1 and margins of most metasomal sclerites whitish, left and
right sides of the holotype vary considerably with the right side much lighter (Fig. 20).
Forewing mostly weakly infuscate, but less so anterobasally and at midlength (Fig. 21E),
or clear basally, infuscate in apical half. Head. Number of  agellomeres 26 (23–29).
Scape not  ared apicoanteriorly and lacking expanded apical surface (Fig. 21B). An-
Figure 21. Sania capensis Mason, 1983, male, holotype A head anterior view B head, lateral view C
mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
334
tennal scrobe  at (Fig. 21A); smooth with weak punctures medially, weakly and ir-
regularly transverse-striate laterally. Clypeus mostly smooth with weak punctures (Fig.
21A); without median tooth (Fig. 21A). Gena mostly smooth with weak punctures
(Fig. 21A). Face weakly punctate throughout (Fig. 21A). Area between antennal scrobe
and inner orbit of eye without protuberance (Fig. 21A). Vertex with weak punctures
(Fig. 20A). Posterior orbit of eye lacking distinct crenulate margin (Figs. 20C, 21B).
Mesosoma. Middle lobe of scutum entirely smooth with punctures (Fig. 21C). Notauli
entirely crenulate (Fig. 21C). Mesopleuron at least partly smooth with punctures (Fig.
20C). Sternaulus only present at midlength (Fig. 20C). Discrimen (median longitudi-
nal ventral sulcus between mesopleura) smooth or with a few barely perceptible crenu-
lae. Metapleuron mostly rugose, lacking a large smooth lightly punctate area (Fig. 20C).
Propodeum entirely rugose without smooth area posterolaterally (Fig. 21C). Metasoma.
Median tergite 1 distinctly narrowed posteriorly (Fig. 21D); partly smooth with punc-
tures especially posteriorly, with weak rugae or microsculpture anteriorly (Fig. 21D).
Female. Unknown.
Diagnosis. Discrimen smooth or with a few barely perceptible crenulae; metap-
leuron mostly rugose, lacking a large smooth lightly punctate area (Fig. 20C).
Material Examined. Holotype male: South Africa: Jonkershoek, near Stellen-
bosch, 33°57’33.74”S, 18°55’10.23”E, 227m, 18.XII.1970, V. Whitehead (AEI)
Paratypes. 2 males, South Africa: Jonkershoek, near Stellenbosch, 33°57’33.74”S,
18°55’10.23”E, 227m, 11.I.1971, V. Whitehead (AEI).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Sania&species=capensis.
Etymology. Not mentioned in Mason (1983), but obviously a reference to the
Cape region of South Africa.
Sania henryi Mason, 1983
Holotype male. Body Length. 4.4 mm. Color. Mottled black and brown except lateroter-
gite 1 and margins of most metasomal sclerites whitish, otherwise metasoma mostly brown
(Figs. 22A, B). Forewing entirely infuscate (Fig. 22). Head. Number of  agellomeres 24
<20–28>. Scape not  ared apicoanteriorly and lacking expanded apical surface (Fig. 23B).
Antennal scrobe shallow (Fig. 23B); mostly smooth lacking punctures and microsetae,
with very weak irregular sculpture laterally. Clypeus mostly smooth with weak punctures
(Fig. 23A); without median tooth (Fig. 23A). Gena with weak rugose microsculpture (Fig.
23A, B). Face mostly punctate with some weak rugae especially dorsally (Fig. 23A). Area
between antennal scrobe and inner orbit of eye without protuberance (Fig. 23A). Vertex
with weak punctures (Fig. 22A). Posterior orbit of eye with distinct crenulate margin (Figs.
22C, 23B), or lacking distinct crenulate margin. Mesosoma. Middle lobe of scutum en-
tirely smooth with punctures (Fig. 23C). Notauli entirely crenulate (Fig. 23C). Mesopleu-
ron at least partly smooth with punctures (Fig. 22C). Sternaulus long, occupying most of
length of mesopleuron (Fig. 22C). Discrimen (median longitudinal ventral sulcus between
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 335
mesopleura) crenulate. Metapleuron mostly rugose, lacking a large smooth lightly punctate
area (Fig. 22C), or partly rugose but with a large smooth lightly punctate area (Fig. 22C).
Propodeum entirely rugose without smooth area posterolaterally (Fig. 23D). Metasoma.
Median tergite 1 distinctly narrowed posteriorly (Fig. 23D); completely rugose (Fig. 23D).
Female. Unknown.
Diagnosis. Scape not  ared apicoanteriorly and lacking expanded apical surface
(Fig. 23B); discrimen crenulate; mesosoma partly or entirely brown (Fig. 22C).
Material Examined. Holotype male: South Africa, Garies, Cape, 30°33’35.77”S,
17°59’24.91”E, 223m, 23.IX.1970, H. & M. Townes (AEI).
Figure 22. Sania henryi Mason, 1983, male, holotype A habitus, dorsal view B habitus, lateral view C
head, mesosoma, lateral view.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
336
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Sania&species=henryi.
Together with the single specimen of Khoikhoia townesi (Eastern Cape Province) these
are the only two representatives of Khoikhoiinae collected outside of the Western Cape
Province. Sania henryi is the only Khoikhoiinae so far recorded as being associated with
the Succulent Karoo biome, a biodiversity hotspot (Myers et al. 2000). We expect that
further undiscovered species will be collected in this habitat.
Etymology. Not mentioned in Mason (1983) but clearly a patronym for Henry Townes.
Figure 23. Sania henryi Mason, 1983, male, holotype A head anterior view B head, lateral view C meso-
soma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 337
Sania marjoriae Mason, 1983
Holotype female. Body Length. 4.0 mm <3.4 – 4.0 mm.>. Color. Mostly dark brown
to black; mottled black or dark brown and pale brown except laterotergite 1 and mar-
gins of most metasomal sclerites whitish, left and right sides of the holotype vary con-
siderably with the right side much lighter (Fig. 24), <the extent of pale brown color
varies considerably but dark brown to black always predominates in the examined speci-
mens>. Forewing mostly weakly infuscate but less so anterobasally and at midlength
(Fig. 25E), or clear basally, infuscate in apical half. Head. Number of  agellomeres 28
<24 – 28>. Scape  ared apicoanteriorly with expanded apical surface (Figs. 3A, 25B).
Antennal scrobe shallow (Figs. 3A, 25B), or  at; mostly smooth lacking punctures and
microsetae, with very weak irregular sculpture laterally. Clypeus mostly smooth with
weak punctures (Fig. 25A); without median tooth (Fig. 25A). Gena with weak rugose
microsculpture (Fig. 25A). Face mostly punctate with some weak rugae especially dor-
sally (Fig. 25A). Area between antennal scrobe and inner orbit of eye without protu-
berance (Fig. 25A). Vertex with weak punctures (Fig. 24A). Posterior orbit of eye with
distinct crenulate margin, or lacking distinct crenulate margin (Fig. 25B). Mesosoma.
Middle lobe of scutum entirely smooth with punctures (Fig. 25C). Notauli entirely
crenulate (Fig. 25C). Mesopleuron at least partly smooth with punctures (Figs. 3E,
24C). Sternaulus long, occupying most of length of mesopleuron (Figs. 3E, 24C). Dis-
crimen (median longitudinal ventral sulcus between mesopleura) crenulate. Metapleu-
ron mostly rugose, lacking a large smooth lightly punctate area (Fig. 24C). Propodeum
mostly rugose with smooth area posterolaterally (Fig. 25C). Metasoma. Median tergite
1 distinctly narrowed posteriorly (Figs. 2C, 25D) completely rugose (Figs. 2C, 25D).
Hypopygium subequal to half length of metasoma (Figs. 1A, 24B); extending past api-
cal tergum (Figs. 1A, 24B). Ovipositor more than half as long as metasoma (Figs. 1A,
24B). Setae of ovipositor sheath 1 to 2× as long as sheath width (Figs. 1A, 24B).
Male. Antenna slightly sexually dimorphic with the female antennae shorter and
tapering slightly more rapidly than those of male.
Diagnosis. Scape  ared apicoanteriorly with expanded apical surface; median
tergite 1 distinctly narrowed posteriorly and completely rugose (Fig. 25D).
Material Examined. Holotype female: South Africa: Jonkershoek, near Stellen-
bosch, 33°57’33.74”S, 18°55’10.23”E, 227m, 18.XII.1970, V. Whitehead (AEI)
Other specimens. 5 females, 9 males, South Africa: Western Cape, 10 km S.
Clanwilliam, Malaise on sandy hill, 32°13’39”S, 18°50’50”E, 140m, 5–25.X.2004,
ME Irwin, FD Parker, M Hauser (SAMC, HIC, INHS).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Sania&species=marjoriae.
Etymology. “Dedicated to Marjorie Townes, whose quiet labors over many years
have contributed so greatly to the production of the ‘Townes and Townes’ team.”
(Mason 1983: p. 51).
Note. is species is referred to as Sania sp. n. 4 in the molecular study by
Murphy et al. (2008).
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
338
Sania masneri Sharkey, sp. n.
urn:lsid:zoobank.org:act:49E6B48F-F40B-479C-9704-8EDFAE402EB9
Holotype female. Body Length. 5.2 mm <male 4.2. mm>. Color. Mottled black and
brown except laterotergite 1 and margins of most metasomal sclerites whitish, otherwise
metasoma mostly brown (Fig. 26). Forewing clear basally, infuscate in apical half (Fig.
27E). Head. Number of  agellomeres 28 <male also 28>. Scape  ared apicoanteriorly
with expanded apical surface (Fig. 27B). Antennal scrobe shallow (Fig. 27B); smooth
medially, transversely striate laterally. Clypeus mostly smooth with weak punctures (Fig.
Figure 24. Sania marjoriae Mason, 1983, female, holotype A habitus, dorsal view B habitus, lateral view
C head, mesosoma, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 339
27A); without median tooth (Fig. 27A). Gena mostly smooth with weak punctures
(Fig. 27A). Face mostly punctate with some weak rugae especially dorsally (Fig. 27A).
Area between antennal scrobe and inner orbit of eye with protuberance, or without pro-
tuberance. Vertex with weak punctures (Fig. 26A). Posterior orbit of eye with distinct
crenulate margin (Fig. 27B), or lacking distinct crenulate margin. Mesosoma. Middle
lobe of scutum entirely smooth with punctures (Fig. 27C). Notauli entirely crenulate
(Fig. 27C). Mesopleuron at least partly smooth with punctures (Fig. 26C). Sternaulus
long occupying most of length of mesopleuron, or only present at midlength, or absent
or di cult to discern due to dense sculpture of mesopleuron (Fig. 26C). Discrimen
Figure 25. Sania marjoriae Mason, 1983, female, holotype A head anterior view B head, lateral view C
mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
340
(median longitudinal ventral sulcus between mesopleura) crenulate. Metapleuron most-
ly rugose, lacking a large smooth lightly punctate area (Fig. 26C). Propodeum entirely
rugose without smooth area posterolaterally (Fig. 27C). Metasoma. Median tergite 1
distinctly narrowed posteriorly (Fig. 27C); partly smooth with punctures especially pos-
teriorly, with weak rugae or microsculpture anteriorly (Fig. 27C). Hypopygium sub-
equal to half length of metasoma (Figs. 1D, 26B); extending past apical tergum (Figs.
1D, 26B). Ovipositor more than half as long as metasoma (Figs. 1D, 26B). Setae of
ovipositor sheath as much as 3 or 4× as long as width of sheath (Figs. 1D, 26B).
Figure 26. Sania masneri Sharkey, sp. n., female, holotype A habitus, dorsal view B habitus, lateral view
C head, mesosoma, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 341
Figure 27. Sania masneri Sharkey, sp. n., female, holotype A head anterior view B head, lateral view C
mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
Male. Antenna highly sexually dimorphic with the female antennae shorter and
tapering much more rapidly than those of male. Genitalia large (Fig. 2D).
Diagnosis. Scape  ared apicoanteriorly with expanded apical surface (Fig. 27B);
median tergite 1 distinctly narrowed posteriorly and smooth with punctures postero-
laterally (Fig. 27D).
Material Examined. Holotype female: South Africa, Western Cape, 10 km S.
Clanwilliam, Malaise on sandy hill, 32°13’39”S, 18°5:0’50”E, 140m, 5–25.X.2004,
ME Irwin, FD Parker, M Hauser (SAMC)
Paratypes. 2 males, same data as holotype (SAMC, HIC).
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
342
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Sania&species=masneri.
Etymology. Dedicated to Lubomír Masner for his dedication to systematics and
for the inspiration that he has given to so many of us.
Note. is species is referred to as Sania sp. n. 1 in the molecular study by
Murphy et al. (2008).
Figure 28. Sania masoni Sharkey, sp. n., female, holotype A habitus, dorsal view B habitus, lateral view
C head, mesosoma, lateral view.
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 343
Sania masoni Sharkey, sp. n.
urn:lsid:zoobank.org:act:DDC57A2D-20B9-4003-9B29-468B1100B7A4
Holotype female. Body Length. 4.8 mm <4.7–5.2>. Color. All black except: mandible
and parts of all segments of fore- and midlegs reddish or yellowish brown, hind tibia red-
dish brown basally, laterotergite and sternum of metasomal tergum 1 whitish (Fig. 28)
<pale color, reddish or yellowish brown, varies from as in the holotype to completely ab-
sent, although laterotergite and sternum of metasomal tergum 1 always whitish>. Forew-
ing mostly weakly infuscate but less so anterobasally and at midlength, or clear basally,
Figure 29. Sania masoni Sharkey, sp. n., female, paratype A head anterior view B head, lateral view C
mesosoma, dorsal view D propodeum, metasomal tergites, dorsal view E wings F data labels.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
344
infuscate in apical half (Fig. 29E). Head. Number of  agellomeres 24 <22–24>. Scape
not  ared apicoanteriorly and lacking expanded apical surface (Figs. 3B, 29A, B). Anten-
nal scrobe  at (Figs. 3B, 29B); variable, from mostly transverse-striate, to mostly smooth
with punctures. Clypeus mostly smooth with weak punctures (Fig. 29A), without median
tooth (Fig. 29A). Gena mostly smooth with weak punctures (Fig. 29A). Face mostly punc-
tate with some weak rugae especially dorsally (Fig. 29A). Area between antennal scrobe
and inner orbit of eye with protuberance (Fig. 29A), or without protuberance. Vertex
with weak punctures (Fig. 28A). Posterior orbit of eye lacking distinct crenulate margin
(Fig. 29B). Mesosoma. Middle lobe of scutum entirely smooth with punctures (Fig. 29C).
Notauli entirely crenulate (Fig. 29C). Mesopleuron at least partly smooth with punctures
(Figs. 3F, 28C). Sternaulus only present at midlength, or absent (Figs. 3F, 28C). Discri-
men (median longitudinal ventral sulcus between mesopleura) crenulate. Metapleuron
mostly rugose, lacking a large smooth lightly punctate area (Fig. 28C). Propodeum en-
tirely rugose without smooth area posterolaterally (Fig. 29C). Metasoma. Median tergite
1 not distinctly narrowed posteriorly (Fig. 29D); partly smooth with punctures especially
posteriorly, with weak rugae or microsculpture anteriorly (Fig. 29D). Hypopygium less
than 1/3 length of metasoma (Figs. 1B, 28B); not extending past apical tergum (Figs. 1B,
28B). Ovipositor barely exserted, much shorter than metasoma (Figs. 1B, 28B). Setae of
ovipositor sheath not longer than 2× width of ovipositor sheath (Figs. 1B, 28B).
Male. Antenna slightly sexually dimorphic with the female antennae shorter and
tapering slightly more rapidly than those of male. Metasoma (Fig. 2E).
Diagnosis. Scape not  ared apicoanteriorly and lacking expanded apical surface
(Figs. 3B, 29A, B); discrimen crenulate; mesosoma black (Fig. 28A, C).
Material Examined. Holotype female: South Africa, Western Cape, 10 km S.
Clanwilliam, Malaise on sandy hill, 32°13’39”S, 18°50’50”E, 140m, 5–25.X.2004,
ME Irwin, FD Parker, M Hauser (SAMC)
Paratypes. 3 females, 18 males, South Africa, Western Cape, 15km N Citrus-
dal, Koedoeskop Farm, Malaise across hillside trail, 32°29’18”S, 18°57’30”E, 220m,
5–25.X.2004, ME Irwin, FD Parker, SA-34 (SAMC, AEI, HIC, CNCI, INHS).
Distribution. Distribution map is available at http://sharkeylab.org/sharkeylab/
Misc/generalmapper.php?table=khoikhoiinae&genus=Sania&species=masoni.
Etymology. Dedicated to the late W.R.M. (Bill) Mason for his many contribu-
tions to Hymenopterology including the discovery of the subfamily Khoikhoiinae.
Acknowledgements
is article is dedicated to Lubomír Masner. Each of the three authors has been touched
by his enthusiasm and each of us has bene ted from his extensive and profound knowl-
edge of Hymenoptera. We all wish him many more productive and happy years. We
hope that including his name alongside those of his mentors and friends, Henry and
Marjorie Townes, and Bill Mason in the subfamily Khoikhoiinae will please him. Sup-
port for MS was provided by NSF EF-0337220 and by a South African National Re-
search Foundation grant GUN 61497 (SvN) and an Italian/South African Scienti c
Revision of Khoikhoiinae (Hymenoptera, Braconidae) 345
and Technological Co-operation grant GUN 2068865 awarded to SvN and Prof. M.
Olmi (University of Tuscia). anks to David Wahl, Gavin Broad and Andy Bennett
for providing specimens, to David Wagner for insights into possible lepidopteran hosts,
and to Scott Robinson and Alex Wild for help and direction with the ESEM images.
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Revision of Khoikhoiinae (Hymenoptera, Braconidae) 347
Appendix 1.
DELTA data matrix, images, and other  les to the key of the Khoikhoiinae (Hymenop-
tera, Braconidae). doi:10.3897/zookeys.20.108.app.1.ik.
Copyright notice:  is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). 
e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and use
this Dataset while maintaining this same freedom for others, provided that the original
source and author(s) are credited.
Citation: Sharkey M, van Noort S, Whit eld J (2009) DELTA data matrix and images to the key of the of
Khoikhoiinae (Hymenoptera, Braconidae). doi:10.3897/zookeys.20.108.app.1.ik. Dataset published in: Zookeys 20:
299–348. doi:10.3897/zookeys.20.108.
Appendix 2.
Interactive key, in IntKey format, to the species of Khoikhoiinae (Hymenoptera, Bra-
conidae). doi:10.3897/zookeys.20.108.app.2.ik.
Note: To run the identi cation key, you will need Windows 95/NT or a later ver-
sion. You also need to download Intkey software and reboot your computer, if it is
not already installed.  e software package, Intkey, can be downloaded from http://
delta-intkey.com/www/programs.htm. Once Intkey is installed you need only click on
the .ink  le (below) and the key will open. Click on any character on the left to begin.
More details on how to use Intkey e ciently are found at http:// orabase.calm.wa.gov.
au/help/keys/intkey_tutorial.pdf.
Copyright notice:  is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). 
e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and use
this Dataset while maintaining this same freedom for others, provided that the original
source and author(s) are credited.
Citation: Sharkey M, van Noort S, Whit eld J (2009) Interactive key, in IntKey format, to the species of Khoikhoiinae
(Hymenoptera, Braconidae). doi:10.3897/zookeys.20.108.app.2.ik. Dataset published in: Zookeys 20: 299–348.
doi:10.3897/zookeys.20.108.
Michael Sharkey, Simon van Noort & James Whit eld / ZooKeys 20: 299–348 (2009)
348
Appendix 3.
Lucid Interchange Format version 3 (LIF3)  le.  is is an XML-based  le that sto-
res all the Lucid3 key data, allowing exchange of the key with other key developers.
doi:10.3897/zookeys.20.108.app.3.ik.
Copyright notice:  is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/).  e Open Database License (ODbL)
is a license agreement intended to allow users to freely share, modify, and use this Dataset
while maintaining this same freedom for others, provided that the original source and
author(s) are credited.
Citation: Sharkey M, van Noort S, Whit eld J (2009) Lucid Interchange Format version 3 (LIF3)  le to the species of
Khoikhoiinae (Hymenoptera, Braconidae). doi:10.3897/zookeys.20.108.app.3.ik. Dataset published in: Zookeys 20:
299–348. doi:10.3897/zookeys.20.108
Appendix 4.
Lucid SDD  le.  is is a XML-based  le structured using the internationally agreed SDD
(Structure of Descriptive Data) Schema.  is SDD  le may be used to exchange the Lu-
cid key with other SDD-compliant applications. doi:10.3897/zookeys.20.108.app.4.ik.
Copyright notice:  is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/).  e Open Database License (ODbL)
is a license agreement intended to allow users to freely share, modify, and use this Dataset
while maintaining this same freedom for others, provided that the original source and
author(s) are credited.
Citation: Sharkey M, van Noort S, Whit eld J (2009) Lucid SDD  le to the species of Khoikhoiinae (Hymenopte-
ra, Braconidae). doi:10.3897/zookeys.20.108.app.4.ik. Dataset published in: Zookeys 20: 299–348. doi:10.3897/zoo-
keys.20.108
... We do not consider Proteropinae as part of the microgastroid complex, as members of that subfamily utilize sawfly larvae as hosts ( van Achterberg, 1976;Sharkey et al., 2021). Rather, we regard Proteropinae as sister to the microgastroid complex, as members of the latter utilize, or in the case of Khoikhoiinae likely utilize (Sharkey et al., 2009), Lepidoptera larvae as hosts (Quicke and van Achterberg, 1990;Whitfield, 1997;Murphy et al., 2008;Whitfield et al., 2018;Fernandez-Triana et al., 2020). The relationships between Mendesellinae and Khoikhoiinae with the rest of the complex have varied slightly among authors (Mason, 1983;Whitfield and Mason, 1994;Whitfield, 1997;Belshaw et al., 1998;Banks and Whitfield, 2006;Murphy et al., 2008;Sharanowski et al., 2011) but in all cases, including our present work, Cheloninae has been recovered as sister to all other microgastroids and Microgastrinae as sister to Cardiochilinae + Miracinae. ...
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The DELTA system is a flexible data-coding format for taxonomic descriptions, and an associated set of programs for producing and typesetting natural-language descriptions and keys, for interactive identification and information retrieval, and for conversion of the data to formats required for phylogenetic and phenetic analysis. This manual is a comprehensive guide to the data format and the program directives.