First Report of the Lace Bug Neoplerochila Paliatseasi (Rodrigues, 1981) (Hemiptera: Tingidae) Infesting Cultivated Olive Trees
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Zootaxa 4722 (5): 443–462 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4722.5.3 http://zoobank.org/urn:lsid:zoobank.org:pub:0183A47A-AA1E-4AAF-8802-54CB9CCDE58C First report of the lace bug Neoplerochila paliatseasi (Rodrigues, 1981) (Hemiptera: Tingidae) infesting cultivated olive trees in South Africa, and its complete mitochondrial sequence JETHRO LANGLEY1, MORGAN CORNWALL1, CHANTÉ POWELL1, CARLO COSTA2, ELLEUNORAH ALLSOPP3, SIMON VAN NOORT4,5, ERIC GUILBERT6 & BARBARA VAN ASCH1 1Department of Genetics, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa. 2Crop Development Division, Infruitec Campus, Agricultural Research Council, Private Bag X5013, Stellenbosch 7600, South Africa. 3Agricultural Research Council, Infruitec-Nietvoorbij, Private Bag X5026, Stellenbosch 7599, South Africa. 4Research and Exhibitions Department, Iziko South African Museum, P.O. Box 61, Cape Town 8000, South Africa. 5Department of Biological Sciences, University of Cape Town, Private Bag, Rondebosch 7701, South Africa. 6Département Adaptation du Vivant, Muséum National d’Histoire Naturelle, UMR 7179, CP50, 45 Rue Buffon, 75005 Paris, France. Barbara van Asch - [email protected] ABSTRACT Olive lace bugs are small phytophagous Hemipteran insects known to cause agricultural losses in olive production in South Africa. Plerochila australis (Distant, 1904) has been reported as the species responsible for damage to olive trees; however, the diversity of olive lace bug species in the region has lacked attention. Adult olive lace bugs were collected incidentally from wild and cultivated olive trees in the Western Cape Province, and identified as P. australis and Neoplerochila paliatseasi (Rodrigues, 1981). The complete mitochondrial genome of a representative specimen of N. paliatseasi was sequenced, and used for comparative mitogenomics and phylogenetic reconstruction within the family. Furthermore, the value of DNA barcodes for species identification in Tingidae was assessed using genetic clustering and estimates of genetic divergence. The patterns of genetic clustering and genetic divergence of COI sequences supported the morphological identification of N. paliatseasi, and the utility of DNA barcoding methods in Tingidae. The complete mitogenome sequence had the typical Metazoan gene content and order, including 13 PCGs, 22 tRNAs, two rRNAs, and an AT-rich non-coding region. A+T content was high, as commonly found in Tingidae. The phylogenetic reconstruction recovered Agramma hupehanum (Drake & Maa 1954) as basal to Tingini, and as a sister species to N. paliatseasi. Stephanitis Stål 1873 and Corythucha Stål 1873 were monophyletic, but Metasalis populi (Takeya 1932) was not recovered as sister to Tingis cardui (Linnaeus 1746), as expected. The mitochondrial phylogeny of the family Tingidae has been recovered inconsistently across different studies, possibly due to sequence heterogeneity and high mutation rates. Species diversity of olive lace bugs in South Africa was previously underestimated. The presence of P. australis was confirmed in both wild and cultivated olives, and N. paliatseasi is reported in cultivated olives for the first time. These results warrant further investigation on the diversity and distribution of olive lace bugs in the Western Cape to inform pest control strategies. KEYWORDS: DNA barcoding, Olea europaea subsp. europaea, insect pest, lace bug, mitogenome, phylogeny INTRODUCTION The entomological fauna associated with wild and cultivated olive trees in South Africa is rich and specialized, in- cluding two olive fruit fly species, a variety of parasitoid and seed wasps, olive beetles, and olive lace bugs (Mkize, Hoelmer and Villet, 2008; Powell et al., 2019; Teixeira da Costa et al., 2019). This particular assemblage has most probably evolved in sub-Saharan Africa over time, along with the native African wild olive tree [Olea europaea subsp. cuspidata (Wall. ex G. Don) Cif.], as cultivated olive trees of European origin (Olea europaea subsp. euro- paea L.) were absent from the region until the colonial period. Due to its Mediterranean-like climate, South African commercial olive production is mostly concentrated in the Western Cape province. Olive orchards occupy approx- imately 3,000 ha of agricultural land interspersed with other crops and residual native Afromontane forest where Accepted by K. Menard: 5 Dec. 2019; published: 16 Jan. 2020 443 the wild olive tree is common. Due to the similarity between the African wild olive and the cultivated olive, some insects associated with olives have been found on both plant species (Powell et al., 2019). Lace bugs (Hemiptera: Tingidae) are generally phytophagous, sap-sucking insects of small size (2 to 10 mm) distributed worldwide. The family Tingidae is classified in the infraorder Cimicomorpha, and currently comprises approximately 2,500 species across 300 genera (ITIS, 2019). Lace bugs are usually host-specific insects that feed on the underside of leaves, which leads to the development of chlorotic pinprick spots. Progressive damage with detrimental consequences to plant vitality occurs as the spots necrotize. Heavy lace bug infestations cause early mortality of young shoots, which results in bushy growth (Addison, Addison and Barnes, 2015). As severe lace bug infestation may lead to total defoliation of the host, control measures are necessary when heavier infestations occur (Costa, 1998). Economically important Tingidae attacking agricultural and ornamental plants include the avocado lace bug (Pseudacysta perseae Haidemann 1908), the tea lace bug (Stephanitis chinensis Drake 1948), and the black lace bug (Amblystira machalana Drake 1948) which feeds on cassava, an important South American crop (Arias and Bellotti, 2003). Corythucha gossypii (Frabicius 1794) is a serious pest of beans and cotton (Miller and Nagamine, 2005), and Corythucha ciliata (Say 1832), commonly known as the sycamore bug, is a pest of ornamental trees of the genus Platanus spp. (Oszi, Ladányi and Hufnagel, 2006). Many other lace bug species feed on ornamental plants, such as Stephanitis pyrioides (Scott 1874) on Rhododendron spp. (Nair and Braman, 2012), Corythauma ayyari (Drake 1933) on Jasminium spp. (Haouas, Guilbert and Halima-Kamel, 2015), and Teleonemia scrupulosa Stål 1873 on Lantana camara (Guidoti, Montemayor and Guilbert, 2015). Lace bugs affecting cultivated olives in South Africa, more commonly referred to as “olive tingids” by local growers, have been reported to be Plerochila australis (Costa, 1998; Addison, Addison and Barnes, 2015). The spe- cies is endemic to sub-Saharan Africa, and its distribution most likely overlaps the natural distribution of wild olive trees, as it has been reported from South Africa to northern Ethiopia (Deckert and Gollner-Scheiding, 2006; Yirgu, Getachew and Belay, 2012). Presently, P. australis is the only species reported as using both wild and cultivated olive trees as hosts. However, olive lace bugs have been poorly studied in South Africa, and the presence of species other than P. australis was suspected, as morphologically distinct specimens were incidentally collected during a re- cent survey of Hymenoptera associated with wild and cultivated olives in the Western Cape (Powell et al., 2019). Species identification in insects has increasingly made use of integrated approaches that include morphological and molecular analyses (Foottit and Adler, 2009; Pires and Marinoni, 2010). Analyses of DNA sequences, particularly the standard COI barcoding region, using genetic clustering methods and measures of genetic distances have be- come popular in the assessment of the genetic homogeneity of insect groups suggestive of biological species (Kress et al., 2015). Mitochondrial gene regions have been employed for the reconstruction of phylogenetic relationships at all taxonomic scales, due to the maternal pattern of inheritance and effectively haploid nature of the organelle genome. Complete mitochondrial sequences represent rich sources of data that have increasingly been used for a wide range of population genetic analyses, comparative genomics, and phylogenetics across a wide range of insect groups (Cameron, 2013). In this regard, the family Tingidae is poorly represented, with only 14 mitogenomes available on Genbank, as of August 2019. The objectives of this study were 1) to identify an unusual olive bug species found feeding on cultivated olives in South Africa, using morphology and DNA-barcoding methods, 2) to generate the complete mitochondrial ge- nome for that species, and 3) to investigate its phylogenetic position within the family Tingidae using the new and publicly available mitogenome sequences. MATERIALS AND METHODS Specimen collection, morphological identification, and DNA extraction Lace bugs feeding on wild and cultivated olive trees in the Western Cape province of South Africa were inciden- tally collected between November 2015 and March 2018 during field surveys that focused on recovering species of olive-associated wasps and flies (Powell et al., 2019) (Table 1). Adult specimens were killed in 100% ethanol, and stored at -20°C until morphological identification, imaging and/or DNA extraction were performed. Adult speci- mens preserved in ethanol were identified as Plerochila australis and Neoplerochila paliatseasi, according to the current taxonomic keys (Göllner-Scheiding, 2007). Representative adult