Research Article |
Corresponding author: Simon van Noort ( svannoort@iziko.org.za ) Academic editor: Denis Brothers
© 2021 Simon van Noort, Rosali Smith, Julie A. Coetzee.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
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
|
The Brazilian waterweed, Egeria densa Planchon, 1849 (Hydrocharitaceae), is an invasive species in South Africa where it is a host plant for the aquatic leaf-miner Hydrellia egeriae 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 Hydrellia egeriae. By comparison with the type specimens, these species have been determined to be Ademon lagarosiphonae van Achterberg, 2012 (Braconidae: Opiinae), Chaenusa anervata van Achterberg, 2012 and Chaenusa seminervata van Achterberg, 2012 (Braconidae: Alysiinae: Dacnusini), all previously recorded as parasitoids of an ephydrid dipterous aquatic leaf-miner, Hydrellia lagarosiphon Deeming, 2012, on Lagarosiphon major (Ridley, 1886) Moss ex Wager (Hydrocharitaceae) in South Africa. The chalcidoid, Janicharis africanus Gumovsky & Delvare, 2006 (Eulophidae), was also reared from Hydrellia egeriae 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).
Ademon, Alysiinae, aquatic weeds, biocontrol, Chaenusa, Entedoninae, Hydrellia egeriae, Hydrocharitaceae, Janicharis, Opiinae
Egeria densa 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 (
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, Hydrellia egeriae Rodrigues-Júnior, 2015, a phytophagous biological control agent attacking Egeria densa. The Ephydridae contain a number of important naturally-occurring biological control agents of submerged invasive waterweeds (
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.
SANC National Collection of Insects, ARC, Pretoria (Curator: Vivienne Uys).
Four parasitoid wasp species belonging to two families (Braconidae and Eulophidae) were reared from puparia of Hydrellia egeriae on Egeria densa. The three braconids, through comparison with the type specimens, were determined to be Ademon lagarosiphonae van Achterberg, 2012 (Opiinae), Chaenusa anervata van Achterberg, 2012 and Chaenusa seminervata van Achterberg, 2012 (Alysiinae), described from specimens reared from an aquatic ephydrid leafminer, Hydrellia lagarosiphon Deeming, 2012 on Lagarosiphon major, (Ridley, 1886) Moss ex Wager also of the family Hydrocharitaceae. The chalcidoid, Janicharis africanus 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) (
http://www.waspweb.org/Ichneumonoidea/Braconidae/Alysiinae/index.htm
http://www.waspweb.org/Ichneumonoidea/Braconidae/Opiinae/index.htm
http://www.waspweb.org/Ichneumonoidea/Braconidae/Keys/index.htm
http://www.waspweb.org/Chalcidoidea/Eulophidae/Entedoninae/Janicharis/index.htm
Tribe Dacnusini Foerster, 1862
Alysia (Chaenusa) Haliday, 1839: 19. Type species: Bracon conjungens Nees von Esenbeck, 1812 (1811), by monotypy; type destroyed.
Chorebidea
Viereck, 1914: 32. Type species: Alysia (Chorebus) nereidum Haliday, 1839, by original designation and monotypy. Treated as Chaenusa (Chorebidea) Viereck, 1914 (
Chorebidea Nixon, 1943: 28. Preoccupied. Type species: Alysia (Chorebus) naiadum Haliday, 1839, by original designation and monotypy.
Chorebidella Riegel, 1950: 125. Type species: Chorebidella bergi Riegel, 1950, by original designation and monotypy.
See
Afrotropical, Australian, Nearctic, Neotropical, Oriental and Palaearctic Regions (
Modified after van
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 Lagarosiphon major Hydrocharitaceae (SANC examined).
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 Lagarosiphon major Hydrocharitaceae (
South Africa • 1 male; KwaZulu-Natal, Midmar Dam, 29°32'21.35"S, 30°11'40.15"E, June 2019, R. Smith, Collected from Egeria densa infestation with Hydrellia egeriae pupae, reared in lab, MDMR-FOR-R05, SAM-HYM-P092803 (
Parasitoid of aquatic leaf-mining flies (Ephydridae): Hydrellia lagarosiphon on Lagarosiphon major (Hydrocharitaceae) and Hydrellia egeriae on Egeria densa (Hydrocharitaceae).
South Africa.
According to van
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 Lagarosiphon major Hydrocharitaceae (SANC examined).
South Africa • 1 female; Mpumalanga Prov., Belfast, Lakenvlei Wetland; 25°60'S, 30°05'E (
South Africa • 1 male; KwaZulu-Natal, Midmar Dam, 29°32'21.35"S, 30°11'40.15"E, June 2019, R. Smith, collected from Egeria densa infestation with Hydrellia egeriae pupae, reared in lab, MDMR-FOR-R01, SAM-HYM-P092800 (
Parasitoid of aquatic leaf-mining flies (Ephydridae): Hydrellia lagarosiphon on Lagarosiphon major (Hydrocharitaceae) and Hydrellia egeriae on Egeria densa (Hydrocharitaceae).
South Africa.
The Neotropical species Chaenusa aurantium Kula & Martinez, 2009 was reared from an undescribed species of Hydrellia in Argentina under evaluation for control of Egeria densa in the United States (
Chorebidea testacea Granger, 1949.
Madagascar • 2 females, 1 male; (
Unknown.
Madagascar.
Tribe Opiini Blanchard, 1845
Subtribe Ademonina Fischer, 1964
Ademon Haliday, 1833: 266. Type species: Bracon decrescens Nees von Esenbeck, 1812 (1811), by monotypy; type destroyed.
Lytacra Foerster, 1863: 266. Type species: Lytacra stygia Foerster, 1863, by monotypy and original designation.
Giardinaia de Stefani-Perez, 1902. Type species: Giardinaia urinator de Stefani, 1902, by monotypy.
Analostania Viereck, 1916. Type species: Analostania tenuipes Viereck, 1916, by monotypy and original designation.
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 (
Afrotropical, Nearctic, Oriental and Palaearctic Regions (van
Line drawings modified from van
Angola • female; CAS-TYPE-10370 (
Unknown.
Angola.
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 Lagarosiphon major (Hydrocharitaceae) (SANC examined).
South Africa • 1 male; Eastern Cape, East London, Nahoon River; 32°57'45.41"S, 27°54'41.59"E; 12 December 2018; R. Smith; Collected from Egeria densa infestation with Hydrellia egeriae pupae; SAM-HYM-P095098 (
Parasitoid of aquatic leaf-mining flies (Ephydridae): Hydrellia lagarosiphon on Lagarosiphon major (Hydrocharitaceae) and Hydrellia egeriae on Egeria densa (Hydrocharitaceae).
South Africa.
This species exhibits a disparate range of colour forms, from being completely black (Figs
Ademon lagarosiphonae females SAM-HYM-P092756 (A–D), SAM-HYM-P092796 (E), SAM-HYM-P092744 (F) (
Janicharis Gumovsky & Delvare, 2006
Janicharis africanus Gumovsky & Delvare, 2006, by monotypy and original designation.
Cameroon • female; Maroua, Djarengol; Malaise trap; 26.ix.1984; G. Delvare (
Cameroon • 3 females, idem (
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 Hydrellia egeriae pupa collected from host plant Egeria densa; SAM-HYM-P092807 (
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 (
Previously unknown. Here, we record the species as a parasitoid associated with immature stages of the aquatic ephydrid fly Hydrellia egeriae, having been reared from puparia along with specimens of Ademon lagarosiphonae and Chaenusa seminervata 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. Janicharis africanus is morphologically similar to the genera Hakuna Gumovsky & Delvare, 2006 and Pediocharis Bouček, 1988 (
Cameroon, Nigeria, Madagascar and South Africa (new country record).
The reared specimens were identified by corroboration of morphological character attributes with the original description and type photographs of J. africanus (
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 (
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.