53urn:lsid:arphahub.com:pub:AF883919-64C9-54D5-9D76-F85E1978BFEEurn:lsid:zoobank.org:pub:B44ABE57-762D-42BD-A81A-2AD84D9E8B5CAfrican InvertebratesAI1681-55562305-2562Pensoft Publishers10.3897/afrinvertebr.62.6284262842Research ArticleBraconidaeEulophidaeIdentification keyTaxonomyAfricaIdentity of parasitoid wasps (Hymenoptera, Braconidae and Eulophidae) reared from aquatic leaf-mining flies (Diptera, Ephydridae) on invasive Brazilian waterweed Egeriadensa in South AfricaNoortSimon van12svannoort@iziko.org.zahttps://orcid.org/0000-0001-6930-9741SmithRosali3CoetzeeJulie A.3Research and Exhibitions Department, South African Museum, Iziko Museums of South Africa, P.O. Box 61, Cape Town, 8000, South AfricaIziko Museums of South AfricaCape TownSouth AfricaDepartment of Biological Sciences, University of Cape Town, Private Bag, Rondebosch, 7701, South AfricaUniversity of Cape TownCape TownSouth AfricaDepartment of Zoology and Entomology, Centre for Biological Control, Rhodes University, Grahamstown, South AfricaRhodes UniversityGrahamstownSouth Africa
Corresponding author: Simon van Noort (svannoort@iziko.org.za)
Academic editor: D. Brothers
2021090320216212873140321137F-E48A-5A16-9DC4-D9B172218B54DD1C3651-0E8A-4F50-8D66-32474969C0F346095170601202101022021Simon van Noort, Rosali Smith, Julie A. CoetzeeThis is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.http://zoobank.org/DD1C3651-0E8A-4F50-8D66-32474969C0F3
The Brazilian waterweed, Egeriadensa Planchon, 1849 (Hydrocharitaceae), is an invasive species in South Africa where it is a host plant for the aquatic leaf-miner Hydrelliaegeriae Rodrigues-Júnior, 2015 (Ephydridae, Diptera). Efficacy of the biocontrol agent can potentially be affected by parasitoids. Three species of braconid parasitoid wasps were reared from puparia of Hydrelliaegeriae. By comparison with the type specimens, these species have been determined to be Ademonlagarosiphonae van Achterberg, 2012 (Braconidae: Opiinae), Chaenusaanervata van Achterberg, 2012 and Chaenusaseminervata van Achterberg, 2012 (Braconidae: Alysiinae: Dacnusini), all previously recorded as parasitoids of an ephydrid dipterous aquatic leaf-miner, Hydrellialagarosiphon Deeming, 2012, on Lagarosiphonmajor (Ridley, 1886) Moss ex Wager (Hydrocharitaceae) in South Africa. The chalcidoid, Janicharisafricanus Gumovsky & Delvare, 2006 (Eulophidae), was also reared from Hydrelliaegeriae and is possibly a hyperparasitoid of the braconids. South Africa is a new country record for J.africanus. We provide comprehensive images of all species including the braconid types and illustrated identification keys to the Afrotropical species of the two braconid genera are also provided. All images and online keys are available on WaspWeb (http://www.waspweb.org).
AdemonAlysiinaeaquatic weedsbiocontrolChaenusaEntedoninaeHydrelliaegeriaeHydrocharitaceaeJanicharisOpiinaeDepartment of Environmental Affairs501100007646http://doi.org/10.13039/501100007646Citation
van Noort S, Smith R, Coetzee JA (2021) Identity of parasitoid wasps (Hymenoptera, Braconidae and Eulophidae) reared from aquatic leaf-mining flies (Diptera, Ephydridae) on invasive Brazilian waterweed Egeria densa in South Africa. African Invertebrates 62(1): 287–314. https://doi.org/10.3897/AfrInvertebr.62.62842
Introduction
Egeriadensa Planchon, 1849, Hydrocharitaceae (Brazilian waterweed; also known as leafy elodea or dense waterweed) is a rooted submerged aquatic macrophyte native to Brazil, Argentina and Uruguay (Cook and Urmi-König 1984; Cabrera Walsh et al. 2013). Stems are elongate, with a diameter of between 1 and 3 mm and irregular side branching (Cook and Urmi-König 1984). It prefers still to slow-moving water and can grow until it reaches the water surface (Yarrow et al. 2009). Leaves are small (10 to 44 mm long, 1.5 to 4.5 mm wide) and are characteristically grouped as four leaves per whorl (Cook and Urmi-König 1984). Vegetative growth occurs at double nodes on stems, which can either produce flowers, lateral branches or root buds (Yarrow et al. 2009). Egeriadensa has been distributed around the world because it is an “oxygenator” and has a simple anatomy, making it an ideal aquarium and study plant (Yarrow et al. 2009; Coetzee et al. 2011a; June-Wells et al. 2012). Due to its ability to grow from fragments, it has established and become invasive in many countries (Hussner et al. 2017). In South Africa, it has taken advantage of open, eutrophic water following the successful management of floating aquatic weeds (Coetzee et al. 2011a). Egeriadensa grows vigorously in eutrophic systems, which allows it to form dense uniform stands, with negative ecological, economic and societal impacts on the invaded system (Vundla et al. 2017; Smith et al. 2019).
A biocontrol investigation, conducted by the Centre for Biological Control at Rhodes University (South Africa) on the invasive Brazilian waterweed, produced four species of parasitoid wasps belonging to two families. These were reared from puparia of the ephydrid fly, Hydrelliaegeriae Rodrigues-Júnior, 2015, a phytophagous biological control agent attacking Egeriadensa. The Ephydridae contain a number of important naturally-occurring biological control agents of submerged invasive waterweeds (Wheeler and Center 2001; Baars et al. 2010; Coetzee et al. 2011a, b; Cabrera Walsh et al. 2013; Bownes 2014). These flies are attacked by braconid parasitoid wasps mainly belonging to the genera Ademon Haliday, 1833 and Opius Wesmael, 1835 (subfamily Opiinae); and Chaenusa Haliday, 1839 and Chorebus Haliday, 1833 (tribe Dacnusini, subfamily Alysiinae) (Yu et al. 2016). Both Ademon and Chaenusa contain aquatic parasitoid species of Hydrellia Robineau-Desvoidy, 1830 flies (Diptera, Ephydridae) (Kula and Zolnerowich 2008; van Achterberg and Prinsloo 2012; Kula and Harms 2016). We here provide determinations and photographs of the four species of parasitoid wasps recorded from puparia of the aquatic leaf-miner, Hydrelliaegeriae developing on Brazilian waterweed Egeriadensa to facilitate ongoing investigation into the efficacy of Hydrelliaegeriae as a biocontrol agent in South Africa. We also provide illustrated identification keys to the species of the two braconid genera occurring in Africa.
Materials and methodsPhotography
Images were acquired at SAMC with a Leica LAS 4.9 imaging system, comprising a Leica Z16 microscope with a Leica DFC450 Camera and 0.63× video objective attached. The imaging process, using an automated Z-stepper, was managed using the Leica Application Suite V. 4.9 software installed on a desktop computer. Diffused lighting was achieved using a Leica LED 5000 Dome.
Depositories
CASCCalifornia Academy of Sciences, San Francisco, USA (Curator: Brian Fisher);
CIRADCentre de Coopération Internationale en Recherche Agronomique pour le Développement (Curator: Philippe Lachenaud);
MNHNMuséum national d’Histoire naturelle, Paris (Curator: Bernardo Santos);
NHMUKNatural History Museum, London (Curator: Gavin Broad);
RMNHNaturalis Biodiversity Center, Leiden (Curator: Luc Willemse);
SAMCIziko South African Museum, Cape Town, South Africa (Curator: Simon van Noort);
SANC National Collection of Insects, ARC, Pretoria (Curator: Vivienne Uys).
Results
Four parasitoid wasp species belonging to two families (Braconidae and Eulophidae) were reared from puparia of Hydrelliaegeriae on Egeriadensa. The three braconids, through comparison with the type specimens, were determined to be Ademonlagarosiphonae van Achterberg, 2012 (Opiinae), Chaenusaanervata van Achterberg, 2012 and Chaenusaseminervata van Achterberg, 2012 (Alysiinae), described from specimens reared from an aquatic ephydrid leafminer, Hydrellialagarosiphon Deeming, 2012 on Lagarosiphonmajor, (Ridley, 1886) Moss ex Wager also of the family Hydrocharitaceae. The chalcidoid, Janicharisafricanus Gumovsky & Delvare, 2006 (Eulophidae) was also reared from this ephydrid fly and is likely a hyper-parasitoid of the braconids.
All images included in this paper, as well as additional images and online interactive keys to the braconid species are available on WaspWeb (http://www.waspweb.org) (van Noort 2021):
AnimaliaHymenopteraBraconidaeAB9570BC-590B-5755-BA41-067D5AACB864ChaenusaHaliday, 1839Alysia (Chaenusa)Haliday, 1839: 19. Type species: Braconconjungens Nees von Esenbeck, 1812 (1811), by monotypy; type destroyed.ChorebideaViereck, 1914: 32. Type species: Alysia (Chorebus) nereidum Haliday, 1839, by original designation and monotypy. Treated as Chaenusa (Chorebidea) Viereck, 1914 (Zaykov 1986).ChorebideaNixon, 1943: 28. Preoccupied. Type species: Alysia (Chorebus) naiadum Haliday, 1839, by original designation and monotypy.ChorebidellaRiegel, 1950: 125. Type species: Chorebidellabergi Riegel, 1950, by original designation and monotypy.Diagnosis.
See Kula and Zolnerowich 2008.
Distribution.
Afrotropical, Australian, Nearctic, Neotropical, Oriental and Palaearctic Regions (Kula and Zolnerowich 2008; Yu et al. 2016).
Key to the Afrotropical species of the genus Chaenusa Haliday
Modified after van Achterberg and Prinsloo 2012.
1
Vein 2–1A of forewing completely developed, resulting in a closed first subdiscal cell (A); vein 1-SR+M of forewing absent or, when present, largely unsclerotised (A); apical half of metasoma of female largely depressed (B); body black; face strongly narrowed in females, ventrally narrower than high (C) (subgenus Chaenusa Haliday, 1839)
2
–
Vein 2–1A of forewing largely absent, resulting in an open first subdiscal cell (a); vein 1-SR+M of forewing sclerotised, complete (a); apical half of metasoma of female strongly compressed; body largely reddish-brown; female face slightly wider than high (subgenus Chorebidea Viereck, 1914)
Chaenusatestacea
2
Frons (fr) and face polished (A); face and eyes (ep) sparsely pubescent (A); each scutellar fovea (sf) with 2 or 3 longitudinal septa (A); first tergite (1t) longer than posteriorly wide, laterally strigous, medially rugulose (B); vein 1-SR+M of forewing pigmented, but largely unsclerotised (C); distal end of pterostigma (pt) comparatively elongate (C)
Chaenusaseminervata
–
Frons (fr) and face granulate (a); face and eyes (ep) densely pubescent (a); each scutellar fovea (sf) with no obvious longitudinal septa (a); first tergite (1t) as long as posteriorly wide, overall rugulose without longitudinal strigae (b); vein 1-SR+M of forewing absent (c); distal end of pterostigma (pt) comparatively shorter and more robust (c)
South Africa • female; Mpumalanga Prov., Lydenburg Fisheries, Lydenburg; 25°11'S, 30°48'E; 21.xi.2008; J.R. Baars, J.A. Coetzee, G.D. Martin; ex leaf-miner on LagarosiphonmajorHydrocharitaceae (SANC examined).
Chaenusaanervata Holotype female TYPH01927 (SANC) A habitus, lateral view B habitus, dorsal view C head and mesosoma, lateral view D head and mesosoma, dorsal view E head, anterior view F head, ventrolateral view.
https://binary.pensoft.net/fig/518154Paratypes.
South Africa • 1 female; KwaZulu-Natal Prov., Mooi River, Mearns Dam; 29°25'S, 29°97'E; 18.xi.2008; J.R. Baars, J.A. Coetzee, G.D. Martin; ex leaf-miner on LagarosiphonmajorHydrocharitaceae (RMNH not seen) • 1 male: Underberg, Farm Dam on Bushman’s Nek road; 30°27'S, 29°14'E; 16.xi.2008 (SANC examined) • 1 male; Mooi River, Mearns Weir, 29°15'S, 29°57'E, 17.xi.2008 (RMNH not seen).
Chaenusaanervata Holotype female TYPH01927 (SANC) A propodeum and metasoma, dorsal view B propodeum and metasoma, lateral view C scutellum, metanotum and propodeum, dorsal view D first tergite, dorsal view E wings F data labels.
South Africa • 1 male; KwaZulu-Natal, Midmar Dam, 29°32'21.35"S, 30°11'40.15"E, June 2019, R. Smith, Collected from Egeriadensa infestation with Hydrelliaegeriae pupae, reared in lab, MDMR-FOR-R05, SAM-HYM-P092803 (SAMC).
Biology.
Parasitoid of aquatic leaf-mining flies (Ephydridae): Hydrellialagarosiphon on Lagarosiphonmajor (Hydrocharitaceae) and Hydrelliaegeriae on Egeriadensa (Hydrocharitaceae).
According to van Achterberg and Prinsloo (2012), this species is similar to Chaenusaglabra Kula, 2009 (India, Pakistan), which has a two-segmented labial palp (three-segmented in C.anervata and C.seminervata), females with 12–14 antennal segments (at least 15 in C.anervata and 16–18 in C.seminervata) and males with 14–17 segments (22 in C.anervata and 17–24 in C.seminervata). The body is brown or brownish-yellow as opposed to black in C.anervata and C.seminervata (van Achterberg and Prinsloo 2012). Other sexually dimorphic characters in C.anervata encompass the forewing pterostigma, which is light brown in females and dark brown in males; the male first metasomal tergite is generally longer (1.1–1.3 times its apical width) than in the female (1.1 times apical width) (van Achterberg and Prinsloo 2012); in addition, the compound eyes in females converge far more strongly (narrowest ventral width of face 0.55× broadest dorsal width) towards the clypeus than in the males (narrowest ventral width of face 0.90× broadest dorsal width); however, the overall relative dimensions of the face relative to the broadest width just below the toruli are equivalent in the two sexes, being slightly wider than high (females H:W = 9:10; males H:W = 10:11).
South Africa • female; Mpumalanga Prov., Lydenburg Fisheries, Lydenburg; 25°11'S, 30°48'E; 21.xi.2008; J.R. Baars, J.A. Coetzee, G.D. Martin; ex leaf-miner on LagarosiphonmajorHydrocharitaceae (SANC examined).
Chaenusaseminervata Holotype female TYPH01928 (SANC) A habitus, lateral view B habitus, dorsal view C head and mesosoma, lateral view D head and mesosoma, dorsal view E head, anterior view F head, ventrolateral view.
https://binary.pensoft.net/fig/518158Paratypes.
South Africa • 1 female; Mpumalanga Prov., Belfast, Lakenvlei Wetland; 25°60'S, 30°05'E (RMNH not seen) • 1 male; KwaZulu-Natal Prov., Stillwater Dam, Rosetta; 29°30'S, 29°97'E; 17.xi.2008; J.R. Baars, J.A. Coetzee, G.D. Martin; ex leaf-miner on LagarosiphonmajorHydrocharitaceae (RMNH not seen) • 4 males; Dieu Donne Farm Dam, Sani Pass; 29°68'407"S, 29°48'949"E; 16.xi.2008; J.R. Baars, J.A. Coetzee, G.D. Martin; reared from leaf-miner on Lagarosiphonmajor (Hydrocharitaceae) (SANC examined, RMNH not seen).
Chaenusaseminervata Holotype female TYPH01928 (SANC) A propodeum and metasoma, dorsal view B propodeum and metasoma, lateral view C mesosoma and first tergite, dorsal view D propodeum and first tergite, dorsal view E wings F data labels.
Chaenusaseminervata female SAM-HYM-P092801 (SAMC) A habitus, lateral view B habitus, dorsal view C head and mesosoma, lateral view D head and mesosoma, dorsal view E head, anterior view F head, ventrolateral view.
https://binary.pensoft.net/fig/518160Biology.
Parasitoid of aquatic leaf-mining flies (Ephydridae): Hydrellialagarosiphon on Lagarosiphonmajor (Hydrocharitaceae) and Hydrelliaegeriae on Egeriadensa (Hydrocharitaceae).
Chaenusaseminervata female SAM-HYM-P092801 (SAMC) A metasoma, dorsal view B metasoma, lateral view C scutellum, metanotum, propodeum and first tergite, dorsal view D first tergite, dorsal view E wings, inset: data labels F vertex of head.
The Neotropical species Chaenusaaurantium Kula & Martinez, 2009 was reared from an undescribed species of Hydrellia in Argentina under evaluation for control of Egeriadensa in the United States (Kula et al. 2009). Chaenusaseminervata keys to C.aurantium (terminal maxillary palpomere is brown) in the key to New World species (Kula and Zolnerowich 2008; Kula et al. 2009), but it is distinct from C.aurantium in head colour, sculpture and setation of the frons, as well as mesoscutal, mesopleural and propodeal sculpture. Sexual dimorphism of Chaenusaseminervata is exhibited in the following characters: females with 16–18 antennal segments, males with 17–24 segments; the forewing pterostigma is light brown in females and dark brown in males; in addition, the compound eyes in females converge less strongly (narrowest ventral width of the face 0.75× broadest dorsal width) towards the clypeus than in C.anervata, but still more strongly than in males (narrowest ventral width of the face 0.81× broadest dorsal width); however, the overall relative dimensions of the face relative to the broadest width just below the toruli are equivalent in the two sexes, being slightly wider than high (females H:W = 10:12; males H:W = 9:11).
Chaenusaseminervata male SAM-HYM-P092802 (SAMC) A habitus, lateral view, inset: data labels B habitus, dorsal view C head and mesosoma, lateral view D head, anterior view.
Madagascar • 2 females, 1 male; (MNHN). Types not seen.
Biology.
Unknown.
Distribution.
Madagascar.
Subfamily Opiinae Blanchard, 1845
Tribe Opiini Blanchard, 1845
Subtribe Ademonina Fischer, 1964
AnimaliaHymenopteraBraconidae00C3BA6D-8267-58DB-BF03-3AE518AEF5A1AdemonHaliday, 1833AdemonHaliday, 1833: 266. Type species: Bracondecrescens Nees von Esenbeck, 1812 (1811), by monotypy; type destroyed.LytacraFoerster, 1863: 266. Type species: Lytacrastygia Foerster, 1863, by monotypy and original designation.Giardinaiade Stefani-Perez, 1902. Type species: Giardinaiaurinator de Stefani, 1902, by monotypy.AnalostaniaViereck, 1916. Type species: Analostaniatenuipes Viereck, 1916, by monotypy and original designation.Diagnosis.
Heavily-sculptured body with presence of an epicnemial carina (= prepectal carina), more or less developed ventrally; occipital carina complete dorsally; malar suture absent; crenulate depression above eye present; vein SR1 of forewing incomplete, not reaching the wing margin and resulting in an open marginal cell; medio-posteriorly scutellum with continuation of lateral elevated area; hind tibia, tarsus and tarsal claws very slender; second metasomal tergite distinctly longer than third tergite and second metasomal suture distinct (Wu et al. 2014; Wharton 2021).
Distribution.
Afrotropical, Nearctic, Oriental and Palaearctic Regions (van Achterberg and Prinsloo 2012).
Key to the Afrotropical species of the genus Ademon Haliday
Line drawings modified from van Achterberg and Prinsloo 2012.
1
Vein 3-SR of forewing 5.0 times as long as vein r; vein 1-SR+M slightly sinuate; vein r-m inclivous (A)
Ademonangolanus
–
Vein 3-SR 2.1–3.5 times as long as vein r; vein 1-SR+M usually strongly sinuate; vein r-m vertical or nearly so (a)
South Africa • female Mpumalanga, Lydenburg Fisheries, Lydenburg, 25°11'S, 30°48'E, 21.xi.2008, J.R. Baars, J.A. Coetzee, G.D. Martin, reared from leaf-miner on Lagarosiphonmajor (Hydrocharitaceae) (SANC examined).
Ademonlagarosiphonae Holotype female TYPH01926 (SANC) A habitus, lateral view B habitus, dorsal view C head and mesosoma, lateral view D head and mesosoma, dorsal view E head, anterior view F head, ventroanterior view.
https://binary.pensoft.net/fig/518163Paratypes.
South Africa • 1 female, 1 male; same data as for holotype (SANC examined).
Ademonlagarosiphonae Holotype female TYPH01926 (SANC) A head and mesosoma, laterodorsal view B metasoma, lateral view C metasoma, dorsal view D metasomal terminal tergites, dorsal view.
Ademonlagarosiphonae Holotype female TYPH01926 (SANC) A scutellum, metanotum and propodeum, dorsal view B metasomal tergites 1–2, lateral view C forewing D data labels.
https://binary.pensoft.net/fig/518165Biology.
Parasitoid of aquatic leaf-mining flies (Ephydridae): Hydrellialagarosiphon on Lagarosiphonmajor (Hydrocharitaceae) and Hydrelliaegeriae on Egeriadensa (Hydrocharitaceae).
Distribution.
South Africa.
Comments.
This species exhibits a disparate range of colour forms, from being completely black (Figs 14 A–F; 15 A–F; 16A, B) through to being completely yellowish-orange, as in the holotype (Figs 10 A–F; 11 A–D; 12 A, B) with intermediate grades present (Figs 13A, B; 16E, F). This extent of intra-specific colour difference is also present in the Palaearctic type species of the genus, A.descrescens (Nees, 1812 [1811]) (van Achterberg and Prinsloo 2012).
Ademonlagarosiphonae Paratype female (A, B) and paratype male (C, D) (SANC) A habitus, lateral view B habitus, dorsal view, inset: data labels C habitus, lateral view, inset: data labels D habitus, dorsal view.
Ademonlagarosiphonae female SAM-HYM-P092756 (SAMC) A habitus, lateral view B habitus, dorsal view C head and mesosoma, lateral view D head and mesosoma, dorsal view E head, anterior view F head, ventroanterior view.
Ademonlagarosiphonae female SAM-HYM-P092756 (SAMC) A head and mesosoma, laterodorsal view B metasoma, lateral view C metasoma, dorsal view D metasomal terminal tergites, dorsal view.
Ademonlagarosiphonae females SAM-HYM-P092756 (A–D), SAM-HYM-P092796 (E), SAM-HYM-P092744 (F) (SAMC) A scutellum, metanotum and propodeum, dorsal view B metasomal tergites 1–2, lateral view C forewing D data labels E habitus, lateral view, inset: data labels F habitus, lateral view, inset: data labels. Figures E and F illustrate intermediate colour forms.
Cameroon • female; Maroua, Djarengol; Malaise trap; 26.ix.1984; G. Delvare (MNHN not seen).
Paratypes.
Cameroon • 3 females, idem (CIRAD) • 3 females, idem (RMNH) • Nigeria • Oyo, Ibadan, Iita Compound; x.1987; J. Noyes (NHMUK) • Madagascar • Lac Alaotra; on Oryzasativa, 17.x.1991; P. Bousses (CIRAD). Paratypes not seen.
Other material.
South Africa • 1 female: Eastern Cape, East London, Nahoon river; 32°57'45.41"S, 27°54'41.59"E; NHN-WINTER-FOR-R02; July 2019; R. Smith; reared in laboratory; ex Hydrelliaegeriae pupa collected from host plant Egeriadensa; SAM-HYM-P092807 (SAMC) • 1 female: idem; except for NHN-WINTER-FOR-R29 and SAM-HYM-P092808 (SAMC) • 1 female: idem; except for: February 2019; R. Smith; Collected from Egeriadensa infestation with Hydrelliaegeriae pupae; SAM-HYM-P095099 (SAMC) • 1 male: idem; except for SAM-HYM-P095100 (SAMC).
Diagnosis.
Uniquely defined by two large foveae situated anterio-medially on propodeum; anterolateral propodeal strip wide, somewhat angulate above spiracle. Characters shared with morphologically-similar congeners: pronotum dorsally reduced, placed significantly below the level of mesoscutum; propodeum with anterolateral propodeal strips; anterior propodeum with basal cup and foveae on sides; metanotum with anteriorly-delimited foveae at sides of dorsellum; long postmarginal vein (Gumovsky et al. 2006).
Biology.
Previously unknown. Here, we record the species as a parasitoid associated with immature stages of the aquatic ephydrid fly Hydrelliaegeriae, having been reared from puparia along with specimens of Ademonlagarosiphonae and Chaenusaseminervata and hence is potentially a hyperparasitoid attacking the braconids rather than the fly. The eulophid subfamily Entedoninae harbours a wide range of life style strategies including species that are usually solitary or gregarious endoparasitoids (more rarely ectoparasitoids or hyper-parasitoids) of concealed dipteran, lepidopteran, coleopteran, hymenopteran or hemipteran larvae or rarely of eggs or pupae. Janicharisafricanus is morphologically similar to the genera Hakuna Gumovsky & Delvare, 2006 and Pediocharis Bouček, 1988 (Gumovsky et al. 2006) and may, therefore, exhibit similar lifestyle strategies. The monotypic species Hakunamatata Gumovsky & Bouček, 2006 was reared from conical insect galls on a forest plant and each gall contained several pupae in separate cells; adult wasps emerged through a single hole at the apex of the gall. Based on these notes by the collector, Gumovsky et al. (2006) hypothesised H.matata to be a parasitoid of the gall former. Biology of Pediocharis is unknown and, together with the lack of detail concerning the biology of Hakuna, makes it difficult to predict whether J.africanus is, indeed, a hyper-parasitoid or not. A more remote possibility of morphological congeneric similarity for Janicharis is to Chrysocharis Foerster, 1856 (Gumovsky et al. 2006), but this genus contains a very broad range of lifestyle strategies, including endo- and ectoparasitoids and facultative hyperparasitoids (Yu et al. 2016), which does not provide any further enlightenment as to potential possibilities for Janicharis. The only recourse is to pursue direct investigation of the biology for J.africanus in the field.
Distribution.
Cameroon, Nigeria, Madagascar and South Africa (new country record).
Comments.
The reared specimens were identified by corroboration of morphological character attributes with the original description and type photographs of J.africanus (Gumovsky et al. 2006). The genus is monotypic.
Janicharisafricanus female SAM-HYM-P092808 (SAMC) A habitus, lateral view B habitus, dorsal view C head, anterior view D data labels.
https://binary.pensoft.net/fig/518170Discussion
This paper contributes towards an understanding of the impact of parasitoid wasps on potential biocontrol agents of invasive waterweeds, specifically that of Brazilian waterweed, which is targeted by biocontrol agents in the genus Hydrellia. Numerous studies have been conducted in the New World and Oriental Region assessing, in particular, species delimitation and host relationships of Chaenusa species (Kula et al. 2006; Kula and Zolnerowich 2008; Kula 2009; Kula et al. 2009; Kula and Harms 2016). Efficacy of the biocontrol agent can potentially be affected by parasitoids and the extent to which these Chaenusa parasitoid wasps, as well as species of Ademon and Janicharis, affect these waterweed biocontrol programmes in various parts of the world is under ongoing investigation, including in South Africa (Rosali Smith, in prep.). Janicharisafricanus may be a hyper-parasitoid of the braconids, further complicating the impact of the biocontrol efficacy of Hydrelliaegeriae, although if it is a hyper-parasitoid, this species should potentially have a positive impact on the biocontrol programme. Elucidation of the precise lifestyle strategy of J.africanus is under ongoing investigation by the Centre for Biological Control at Rhodes University (South Africa). Ramifications of adaptation by the indigenous parasitoids, documented in this paper, to introduced hosts released as potential biocontrol agents against targeted invasive species in South Africa, will be detailed in an accompanying publication (Smith, Coetzee, van Noort, in prep). This paper provides a taxonomic contribution towards resources that will facilitate identification of parasitoid wasps reared in the process of establishing effective control programmes for invasive waterweeds in Africa.
Acknowledgements
We thank Denis Brothers, Mark Shaw, Cees van Achterberg and an anonymous reviewer for their critical reviews that improved the manuscript. This research was funded through the Department of Environmental Affairs, Natural Resource Management Programme’s Working for Water programme. Further funding for this work was provided by the South African Research Chairs Initiative of the Department of Science and Technology and the National Research Foundation of South Africa (grant nos. 89967, 109244 and 109683 to Martin Hill, SARCHi Chair). Simon van Noort was funded by a South African NRF FBIP grant no. 98115.
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