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Morphological Comparison of Proboscis Sensilla Between Plusiinae and Noctuinae (Lepidoptera: Noctuidae)

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Morphological Comparison of Proboscis Sensilla Between Plusiinae and Noctuinae (Lepidoptera: Noctuidae) See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/303325575 Morphological comparison of Proboscis Sensilla between Plusiinae and Noctuinae (Lepidoptera: Noctuidae) Article in Zoologischer Anzeiger - A Journal of Comparative Zoology · May 2016 DOI: 10.1016/j.jcz.2016.05.002 CITATIONS READS 0 44 3 authors, including: Shuang Xue Bao-Zhen Hua Northwest A & F University Northwest A & F University 3 PUBLICATIONS 5 CITATIONS 78 PUBLICATIONS 449 CITATIONS SEE PROFILE SEE PROFILE All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Bao-Zhen Hua letting you access and read them immediately. Retrieved on: 06 September 2016 Zoologischer Anzeiger - A Journal of Comparative Zoology xxx (2016) xxx-xxx Contents lists available at ScienceDirect Zoologischer Anzeiger - A Journal of Comparative Zoology journal homepage: www.elsevier.com Research paper Morphological comparison of proboscis sensilla between Plusiinae and Noctuinae (Lepidoptera: Noctuidae) Shuang Xue, Yan-Qing Hu, Bao-Zhen Hua⁎ State Key Laboratory of Crop Stress Biology for Arid Areas, Key Laboratory of Plant Protection Resources and Pest Management of the Education Ministry, Entomological Museum, Northwest A&F University, Yangling, Shaanxi 712100, China ARTICLE INFO ABSTRACT PROOF Article history: In higher Lepidoptera the proboscis and its sensilla vary from species to species in different feeding guilds. However, Received 31 March 2016 whether such morphological differences have a phylogenetic significance remains unclear. Here we compared the mor- Received in revised form 15 May 2016 phology of proboscis sensilla in 19 representative species of Plusiinae and four of Noctuinae using scanning electron Accepted 15 May 2016 microscopy. In Plusiinae four types of sensilla (sensilla chaetica, sensilla basiconica, sensilla styloconica, and sensilla Available online xxx coeloconica) were present on the proboscis, with cuticular processes only in the proximal region. In Noctuinae three types of sensilla (sensilla chaetica, sensilla basiconica, and sensilla styloconica) were found on the proboscis, with cuticular Keywords: processes on its whole length. Morphological differences of proboscis sensilla between Plusiinae and Noctuinae mainly Morphology involve the shape, number, and length of sensilla styloconica and sensilla chaetica. The sensilla styloconica of Plusiinae Mouthparts are significantly shorter and fewer than those of Noctuinae. The styli of sensilla styloconica are glabrous in Plusiinae and Proboscis Sensilla are longitudinally ribbed in Noctuinae. These significant morphological differences are useful in distinguishing the two Systematics subfamilies, suggesting that proboscis sensilla may provide useful characters in the systematic and phylogenetic analyses Phylogenetics at the subfamily level within Noctuidae. © 2016 Published by Elsevier Ltd. 1. Introduction The classification of Noctuidae has recently undergone major shifts and rearrangements, especially at the subfamily level (Speidel et al., Higher Lepidoptera typically have siphoning mouthparts, with 1996; Lafontaine and Fibiger 2006; Mitchell et al., 2006). Plusiinae, their proboscis consisting of a complex fluidic system derived from a small subfamily of Noctuidae, is regarded as monophyletic based two extremely elongated galeae. The proboscis functions like a drink- on considerable morphological, bionomical and biogeographical evi- ing straw through which floral nectar and other liquid substances are dence (Kitching, 1987; Ronkay et al., 2008), and may eventually be imbibed (Scoble, 1992; Kingsolver and Daniel, 1995; Krenn, 2010; raised to a family status (Weller et al., 1994). Monaenkova et al., 2012). Each galea bears various types of sen- In this paper, we compared the morphology of the proboscis and its silla, which play important roles in feeding activities of Lepidoptera sensilla in 19 representative species of Plusiinae and four of Noctuinae (Salama et al., 1984; Krenn et al., 2005; Krenn, 2010; Faucheux, 2013; using scanning electron microscopy in order to prove the suitability Lehnert et al., 2013; Zaspel et al., 2013). Proboscis sensilla exhibit of characters regarding proboscis sensilla for phylogenetic differenti- remarkable differences with regard to their type, morphology, loca- ation of Plusiinae and Noctuinae. tion and number, and are associated mainly with the feeding habits of their owners (Krenn et al., 2001; Krenn, 2010; Krenn and Aspöck, 2. Materials and methods 2012; Zenker et al., 2011; Faucheux, 2013; Lehnert et al., 2016). Pre- vious investigations of Lepidoptera proboscis sensilla are mainly con- 2.1. Insect collection and preparation centrated on Nymphalidae, Riodinidae, Arctiidae, Erebidae, Pyralidae, Geometridae, Saturniidae, and Noctuidae (Faucheux, 1991; Büttiker Male and female individuals of 23 species from two subfamilies et al., 1996; Paulus and Krenn, 1996; Krenn, 1998; Krenn and Penz, of Noctuidae (19 species in 17 genera of Plusiinae and four species in 1998; Walters et al., 1998; Krenn et al., 2001; Zaspel et al., 2011; four genera of Noctuinae) were examined in this study (Table 1). The Bauder et al., 2013). However, whether these morphological differ- moths were obtained from collections in the Entomological Museum, ences of sensilla have a phylogenetic significance remains unclear. Northwest A&F University, China. Dried proboscises were removed Noctuidae is the largest family in LepidopteraUNCORRECTEDwith numerous agri- at the base of the head from pinned specimens using fine-tip forceps. cultural pests of great economic significance (Zahiri et al., 2010). 2.2. Scanning electron microscopy ⁎ Corresponding author. For scanning electron microscopy (SEM), proboscises were Email address: [email protected] (B-Z Hua) cleaned with an ultrasonic cleaner in 75% ethanol for 30 s. After de- hydration in a graded ethanol series, the samples were freeze-dried in http://dx.doi.org/10.1016/j.jcz.2016.05.002 0044-5231/© 2016 Published by Elsevier Ltd. 2 Zoologischer Anzeiger - A Journal of Comparative Zoology xxx (2016) xxx-xxx Table 1 3. Results Information for specimens examined. Subfamilies Species Localities 3.1. General morphology of the noctuid proboscis Plusiinae Anadevidia peponis (Fabricius, 1775) Henan The mouthparts of adult Plusiinae and Noctuinae are of the typical Antoculeora locuples (Obethür, 1880) Shaanxi siphoning type, consisting of two extremely elongated galeae, which Autographa excelsa (Kretschmar, 1862) Shaanxi Cornutiplusia circumflexa (Linnaeus, 1767) Xinjiang are interlocked to form a sucking tube. In the resting position, the pro- Diachrysia chrysitis (Linnaeus, 1758) Xinjiang boscis is coiled in a tight spiral beneath the head between the setose Diachrysia leonine (Obethür, 1884) Jilin three-segmented labial palps. The outer surface of proboscis is cov- Diachrysia stenochrysis (Warren, 1913) Hebei ered with cuticular processes and various types of sensilla. Sexual di- Diachrysia oberthueri (Ronkay, Ronkay and Gansu morphism was not detected in the proboscis of any species. Behounek, 2008) Erythroplusia rutilifrons (Walker, 1858) Sichuan Euchalcia inconspicua (Graeser,1892) Xinjiang 3.2. Proboscis sensilla of Plusiinae Macdunnoughia confusa (Stephens, 1850) Xinjiang Panchrysia ornate (Bremer, 1864) Gansu The sensilla equipment of the 19 species of Plusiinae examined is Panchrysia tibetensis (Chou and Lu, 1982) Yunnan Plusidia cheiranthi abrostoloides (Butler, 1879) Liaoning the same in all species, all of which bear four morphological types of Plusia festucae (Linnaeus, 1758) Jiangsu sensilla on the galeae: bristle-shaped sensilla chaetica, uniporous sen- Plusia putnami festata (Graeser, 1890) Heilongjiang silla basiconica, glabrous sensilla styloconica, and particular sensilla Polychrysia esmeralda (Oberthür, 1880) Qinghai coeloconica (Fig. 1). Shapes, lengths, numbers as well as distributions Polychrysia imperatrix (Draudt, 1950) Yunnan of these sensilla do not differ significantly among species of Plusiinae Stigmoctenoplusia aeneofusa (Hampson, 1894) Yunnan Syngrapha ain persibirica (Ronkay, Ronkay and Jilin (Figs. 5 and 6), therefore we use Anadevidia peponis as the represen- Behounek, 2008) tative of this subfamily. PROOF Thysanoplusia intermixta (Warren, 1913) Shaanxi The proboscis length of A. peponis is 11.87 ± 2.25 mm Thysanoplusia orichalcea (Fabricius, 1775) Shaanxi (10.73–12.84 mm, n = 30). The proboscis can be divided into the Trichoplusia ni (Hübner, 1803) Shaanxi Zonoplusia ochreata (Walker, 1865) Yunnan proximal, bend, distal, and tip regions. When uncoiled, the proximal Noctuinae Anaplectoides virens (Butler, 1878) Shaanxi region makes up 35% of the total length; the bend region comprises Axylia putris (Linnaens, 1761) Shaanxi 12% of the total length; the distal region constitutes 41%; and the tip Euxoa sibirica (Boisduval, 1837) Shaanxi region, where fluids are taken up into the food canal, comprises 12% Xestia agalma (Pungeler, 1899) Shaanxi of the total length. The proximal region of proboscis is covered with hair-like cutic- ular processes (Fig. 1A and B). The bend, distal, and tip regions are devoid of cuticular processes. All the four types of sensilla are present tertiary butanol. They were then mounted onto SEM stubs using a dou- on the tip region, of which the sensilla chaetica and sensilla basiconica ble graphite adhesive tape, coated with gold in a sputter coater, and are rare and scattered, and the sensilla chaetica
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