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Ephemeroptera: Heptageniidae) Oriental Insects ISSN: 0030-5316 (Print) 2157-8745 (Online) Journal homepage: http://www.tandfonline.com/loi/toin20 The last two moulting processes of Parafronurus youi and possible emergence evolution of Mayflies (Ephemeroptera: Heptageniidae) Juan-Yan Luo, Ze Hu, Zhen-Xing Ma & Chang-Fa Zhou To cite this article: Juan-Yan Luo, Ze Hu, Zhen-Xing Ma & Chang-Fa Zhou (2017): The last two moulting processes of Parafronurus youi and possible emergence evolution of Mayflies (Ephemeroptera: Heptageniidae), Oriental Insects, DOI: 10.1080/00305316.2017.1415172 To link to this article: https://doi.org/10.1080/00305316.2017.1415172 Published online: 21 Dec 2017. Submit your article to this journal Article views: 2 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=toin20 ORIENTAL INSECTS, 2017 https://doi.org/10.1080/00305316.2017.1415172 The last two moulting processes of Parafronurus youi and possible emergence evolution of Mayflies (Ephemeroptera: Heptageniidae) Juan-Yan Luo , Ze Hu, Zhen-Xing Ma and Chang-Fa Zhou The Key Laboratory of Jiangsu Biodiversity and Biotechnology, College of Life Sciences, Nanjing Normal University, Nanjing, China ABSTRACT ARTICLE HISTORY Moulting processes of last instar nymph and subimago, Received 10 February 2017 longevity of subimago and imago of Parafronurus youi is Accepted 6 December 2017 observed and filmed in laboratory for the first time in China. KEYWORDS The videos show clearly this species emerges in the afternoon Mayfly; emergence; moult; (ca. 5:30 PM to 9:30 PM) on the object surface in water. The subimago; Heptageniidae subimagos moult 14–19 h later before noon. Imago can survive for 3 to 4 days in lab. The data in the present paper and others as well as phylogeny do not provide clear evolutionary pattern of mayfly emergence location or time but a possibility maybe present: both the primitive mayfly nymph and adult can live and emerge on moist land, then adapt to aquatic habitats. Introduction Mayflies are unique winged insects because of their two adult instars (subimago and imago). However, the ecdysis of aquatic mayfly nymphs are difficult to observe in the field and their subimaginal stage is very brief. So there are few detailed reports on metamorphic pattern of mayflies although some researchers sum- marised this process. Kimmins (1941) reported the Heptagenia lateralis nymph (now in the genus Electrogena) can emerge underwater or on the surface of water. Minshall (1967) and Hall et al. (1975) observed, respectively, the Epeorus pleralis and Tricorythodes atratus can moult in water too. Peters and Peters (1977) show the species Dolania americana life history. Russev (1987) took some pictures about moulting behaviour and process of Palingenia longicauda. The latter two species emerge on the water surface. Generally, Edmunds and McCafferty (1988) pro- posed some mayflies (like Isonychia and Coloburiscus) can emerge on the surface of water or on the land but most mayflies emerged at surface of the water, such as CONTACT Chang-Fa Zhou [email protected] © 2017 Informa UK Limited, trading as Taylor & Francis Group Published online 21 Dec 2017 .J.-Y. LUO ET AL 2 some species of Baetidae and Ephemerellidae. Brittain (1982) and Bauernfeind and Soldán (2012) indicated mayflies can emerge at surface of water, under water or above water. For instance, Siphlonurus, Isonychia and Baetisca mayflies will climb out of water to moult. But the evolutionary trends of them have not been hypothesised so far. According to Kukalová-Peck (1978, 1991), the original mayflies probably lived in the moist environment and then became aquatic. That means living in the water of mayfly larvae is secondary. So in our own opinion, if more species emergence processes are detailed, they can help us solve some phylogenetic and evolutionary puzzles of Ephemeroptera. In China, only few mayfly species biology was mentioned and discussed briefly (Zhou and Peters 2003, 2004; Wang et al. 2011), but no specific research on this issue. In 2015, the Parafronurus youi Zhou and Braasch (2003) were reared, observed and recorded in lab. Therefore, the biology of this species, such as emer- gence of nymph, moulting of subimago, longevities of subimago and imago are clear to us and described here. Their possible phylogeny and evolutionary value are discussed. Methods Mayfly rearing (Fig. 1) Nymphs are collected with a hand net (40 um) or a hand nylon mesh (35 um) in the ponds and creeks of Zijin Mountain, Nanjing, Eastern China. Mature nymphs (with black wingpads) were taken back to laboratory and reared in the white plastic trays (size 40 × 25 × 8 cm) which were filled with stones (some parts above water), withered leaves and water gathered in the creek where the insects live. The rearing water depth is about 5 cm under the temperature 18–25 °C. Those trays were covered with mosquito nets to catch the adults (Fig. 1). Video taking In order to observe and see clearly the moulting process, we took videos with smartphones. When we find mature nymphs show some abnormal behaviour like moving fast and irregularly, we focus on them and record. The videos will replay on laptop and key information will be written down. Totally 22 emergence videos of nymphs and 3 moulting episodes of subimagos are filmed. They show exactly the last two moulting process of the mayfly P. y oui and some information of its life history. ORIENTAL INSECTS 3 Figure 1. The rearing equipment. Results Moulting of the nymph (Fig. 2A–C) The last instar nymphs of Parafronurus youi moult at about 5:30 PM to 9:30 PM (Beijing time). Just before that, the nymphs will climb to or grasp the stone surface in the water (Fig. 2A). This phenomenon can happen several times. At the same time, the colour of ongoing moulting nymphs becomes conspicuously darker and the vibrations of gills are quicker than normal. Before moulting, the gas bubbles can be seen in head, thorax and wingpads first. This process lasts several minutes. During this period, they will move forth and back if they are touched by other larvae or be shaken. Also in this process, the nymphs can climb to stone tips above the water one to several times, expose their dorsal head or half body into the air 1–2 s and move back down to the water. Just before the emergence, three caudal filaments of nymphs will put together quickly. After the gas bubbles appeared in the body about 6 min, pro- and mesonotum will split in the middle, and then extend to vertex. Soon the head and thorax of subimago come out of nymphal exuviae and pull forward slowly, and then legs, wings, abdomen and caudal fila- ments appeared progressively. This emergence process can last 11s–126s (Table 1). During this process, the packed wings show silver white appearances. After they take off their exuviae, the subimagos will move to the water surface immediately. As soon as they touch the air, the wings of them stretched out fully and became brown to black immediately. .J.-Y. LUO ET AL 4 Figure 2 A–H. Different stages of Parafronurus youi; A, nymph; B–C, nymph moulting to subimago; D, female subimago; E–G, moulting subimago; H, male imago. Longevity of the subimago Under laboratory conditions (20–25 °C), the observed living time of three subima- gos (Fig. 2D) (from coming out of nymphal skin to next moult) is about 14–19 h. Moulting of the subimago (Fig. 2E–G) Before subimagos begin to moult, their vertically positioning wings will flap down and hold laterally step by step. When wings are at horizontal position, they bend backwards slightly. The whole body, meanwhile, is shaking quickly. Then the head and thorax of imago come out from their shells and move forward slowly or roll over to pull abdomen and caudal filaments out of cuticle. The average lasting time ORIENTAL INSECTS 5 Table 1. The moulting time of the Parafronurus youi nymph. Video The start time of moulting The time of metamorphosis 1 19:22:57 112s 2 21:12:26 11s 3 19:33:25 119s 4 17:47:11 111s 5 18:05:58 126s 6 18:01:12 95s 7 18:39:41 88s 8 18:47:21 94s 9 18:59:09 81s 10 19:01:01 87s 11 19:16:08 98s 12 19:24:21 86s 13 19:40:31 107s 14 18:37:14 110s 15 18:57:31 93s 16 19:10:23 108s 17 19:17:51 113s 18 18:07:32 111s 19 18:20:11 121s 20 19:19:41 109s 21 20:32:20 99s of this moulting is 233 s. The time of subimaginal moulting is from about 10:00 to 12:00 AM (Beijing Time) (Table 2). Longevity of the imago The imagos (Fig. 2H) in mosquito net can live 3–4 days under laboratory condi- tions. Mating, swarming and oviposition were not seen. Discussion In order to compare the emergence patterns of mayflies, we list the related infor- mation in the literatures we can find (Table 3). From it, besides most mayflies emerge at the water surface, no other obvious and congruent evolutionary trend can be summarised generally, either the emergence location or time. For exam- ple, the heptageniid and leptophlebiid nymphs have three moulting types: at the surface of water, under water or above water. The ephemerellid and leptohyphid species have two types. Furthermore, the species in same genus can have more than one moulting type, like Ecdyonurus, Habroleptoides and Neoephemera. Even Table 2. The moulting time of the Parafronurus youi subimago. The time of the The start time of The preparation The time of meta- whole moulting Video moulting time of moulting morphosis process A 10:15:28 81s 203s 284s B 11:30:57 – 153s 153s C 11:47:15 81s 180s 261s Average time 81s ≈179s ≈233s 6 Table 3.
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