Sex roles in the ghost moth Hepialus humuli (L.) and a review of mating in the Hepialidae (Lepidoptera)

Recently it has been found that female Lepidoptera belonging to diverse families actively court their males, rather than play a merely passive role. Male and female Hepialus humuli have been reported to come together in three different ways: (1) females are attracted to groups or ‘leks’ of white, hovering males by visual stimuli; (2) females are attracted to the males by olfactory-substances produced on the hind-tibial brushes of the males; (3) males are attracted to sedentary females by olfactory stimuli. During my study I observed H. humili males flying on a total of 21 nights in two different parts of England. The males hovered in groups for about 20 min each evening, starting and stopping their flights in synchrony. Timing depended on light intensity, northern moths flying later in the summer evenings than southern moths. I observed a total of 18 matings. Normally, a female from outside a male lek flew into the group and up to one of the males. This male then usually followed her to a settling position, where mating took place. In a few cases females touched males; in one case a female struck a male in the air so that both fell to the ground and were copulating when examined. Photographs of hovering males show that their hind tibial brushes are fully everted in flight. The organs are folded against the body when the moth is mating, at rest or dead. Whilst hovering, the males are apparently emitting pheromones which function as primary attractants, rather than as the aphrodisiacs of many other lepidopteran males. The mating behaviour of hepialids is reviewed. It is concluded that all studied hepialids which have male brush organs (some Hepialus and Oncopera, Sthenopis, Zenophassus) exhibit similar flight and mating behaviour: males hover, sit or loop back and forth on the spot in leks; females fly into male aggregations and mate there (although some published observations suggest otherwise). In contrast, hepialids such as Fraus, Oxycanus and other Hepialus that lack male brush organs have mating behaviour in which the males are the active partner, a more standard lepidopteran method. In view of the controversies surrounding mating in hepialids, future systematic and behavioural work on Hepialidae throughout the world will be worthwhile.     

Bat defence in lekking ghost swifts (Hepialus humuli), a moth without ultrasonic hearing.

The Hepialidae represents an early branch of the Lepidoptera, whose members lack the ultrasonic hearing and other obvious predator defence systems present in other extant moths. I observed lekking male ghost swifts, Hepialus humuli, being exploited by northern bats, Eptesicus nilssonii, over a hayfield in southern Sweden. Because the moth''s display flight was restricted to a brief (30 min) period at dusk, they avoided most predators temporally but were exposed to early emerging aerial-hawking bats. Against these, they apparently employed ''acoustic crypsis'', achieved by flying close (< 0.5 m) to the vegetation, thereby hiding from the bats among clutter (echoes returning from the background). Nevertheless, the predation risk for the displaying moth males was very high (20% per night), mainly because they sometimes left the safety of the vegetation. The lack of ''advanced'' predator defence mechanisms in H. humuli requires alternative defence strategies, which, however, restrict the behavioural repertoire and still carry a high predation risk.     

The dawn flight of the gold swift Hepialus hecta: predator avoidance and the integration of complex lek behaviour (Lepidoptera,Hepialidae)

Predator pressure is a considerable evolutionary force. The evening twilight flight of species in the family Hepialidae Swift moths has been attributed to the moths (which have no hearing and so cannot detect bats) flying in a bat/bird free window. Several species deploy elaborate lek behaviour in this period. The expected flight in the dawn twilight is little reported, except in the non‐leking Hepialus (Korscheltellus) gracilis. A detailed study of the dawn flight in the leking species Hepialus (Phymatopus) hecta shows that it is less extensive than the evening lek flight (both in duration and in the number of moths participating), is confined to a much narrower window around sunrise, involves no reproductive behaviour, and functions only to re‐locate the members of copulating pairs and sessile displaying males from the emergent ground vegetation to less conspicuous roosting sites. Compilation of individual biographies over the 24‐h cycle permits a full construction of the diel activity in time and space. It consists of two twilight flight periods, separated by prolonged roosting during the hours of darkness and daylight. The moths use almost the whole available range of roosting sites, from the base of the ground vegetation to the tree canopy, and cyclically leave and re‐enter the lek site from these positions. The copulating posture and position avoid mammalian predators, and facilitate escape from spiders and wasps by the efficient use of a dead drop. The narrowness of the dawn flight is attributed to the need for this prolonged but conspicuous copulation, which precludes a morning mating, and to a demonstrated asymmetry of twilight activity in birds. The whole integrated spatio‐temporal cycle is attributable to evolutionary pressure to minimize the impact of predators, demonstrated to include birds, bats, wasps, and spiders, and probably also dragonflies, mice and shrews; the pre‐existing dawn flight is permissive to the evolution of sessile male displays during the evening lek. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 305–319.     

西藏虫草蝠蛾属一新种 (鳞翅目：蝙蝠蛾科)

 报道了采自西藏那曲地区比如基地的鳞翅目蝙蝠蛾科一新种 ,比如蝠蛾Hepialus biruensis Fu, sp.nov。该种幼虫是我国特产药材冬虫夏草的寄 主昆虫之一。     

西藏虫草蝠蛾属一新种(鳞翅目：蝙蝠蛾科)

报道了采自西藏那曲地区比如基地的鳞翅目蝙蝠蛾科一新种,比如蝠蛾Hepialus biruensis Fu, sp.nov。该种幼虫是我国特产药材冬虫夏草的寄主昆虫之一。     

阿尔泰蝠蛾幼虫血细胞形态学研究

本研究采用Wright-Giemsa染色方法,利用光学显微镜观察阿尔泰蝠蛾Hepialus altaicola Wang幼虫血细胞.根据鳞翅目昆虫血细胞形态特征对阿尔泰蝠蛾幼虫血细胞进行鉴定,计数并计算各类血细胞所占的比例.发现阿尔泰蝠蛾幼虫血淋巴中存在原血胞、浆血胞、粒血胞、类绛色细胞、珠血胞和囊血胞6种细胞类型,所...     

蝠蛾属Hepialus一新种记述(鳞翅目:蝙蝠蛾科)

记述蝙蝠蛾科Hepialidae蝠蛾属Hepialus 1新种:小金蝠蛾Hepialus xiaojinensis, sp. nov.,模式标本分别保存在西南大学和重庆市中药研究院.     

阿尔泰蝠蛾(鳞翅目:蝙蝠蛾科)幼虫低温胁迫转录组分析

[目的]本研究旨在通过转录组测序技术分析低温胁迫引起的阿尔泰蝠蛾Hepialus altaicola Wang幼虫基因转录差异及上调表达基因的主要功能类群和参与的主要代谢通路.[方法]采用Illumina HiSeq TM 2500测序平台对阿尔泰蝠蛾幼虫进行转录组测序、组装,利用Blast软件进行数据库比对和基因功能注释,用DEGSeq R软件包分析4℃低温处理与室内适温饲养试虫的差异表达基因,并对上调表达基因进行GO和KEGG代谢途径富集分析.[结果]经序列拼接后共获得100300个unigenes,总长度81600309 bp,平均长度813 bp,N50长度1719 bp.与7大数据库同源比对,共获得34691(34.59％)条unigenes.低温胁迫转录组分析得到11569个差异表达基因(DEGs),7158条基因上调,4411条基因下调.富集到47个GO类群,217个KEGG途径.其中代谢过程、催化、结合活性类群占有重要比例,核糖体、碳代谢、剪接体等途径显著富集.另外,热激蛋白、昆虫表皮蛋白、海藻糖酶、超氧化物歧化酶等非生物胁迫相关基因显著上调表达.[结论]阿尔泰蝠蛾幼虫低温胁迫转录组分析揭示,代谢过程、细胞过程、生物调节、对刺激的反应等生物学过程相关基因和部分非生物胁迫响应基因显著上调表达,提示蝠蛾幼虫可能从抗氧化防御、分子伴侣、体温调节和维持细胞的渗透平衡等多方面应对低温胁迫.     

阿尔泰蝠蛾(鳞翅目:蝙蝠蛾科)幼虫食性的观察

对阿尔泰山上的阿尔泰蝠蛾幼虫进行了为期6年的食性观察,发现阿尔泰蝠蛾幼虫在野外采食的植物是新疆芍药、块根芍药、白喉乌头、阿尔泰牡丹草、新疆藜芦、新疆猪牙花、阿尔泰羽衣草、柳兰、新疆白鲜和药用蒲公英,与蝠蛾属的其他昆虫不同。其中,大部分植物为新疆阿尔泰山上的特有植物和中药,甚至有些还有毒性。在人工饲养的情况下,阿尔泰蝠蛾喜食胡萝卜和南瓜。该观察结果将为以后阿尔泰蝠蛾幼虫的野外采集和人工饲养等研究提供一定的帮助。     

阿尔泰蝠蛾(鳞翅目:蝙蝠蛾科)幼虫低温胁迫转录组分析

[目的]本研究旨在通过转录组测序技术分析低温胁迫引起的阿尔泰蝠蛾Hepialus altaicola Wang幼虫基因转录差异及上调表达基因的主要功能类群和参与的主要代谢通路.[方法]采用Illumina HiSeq TM 2500测序平台对阿尔泰蝠蛾幼虫进行转录组测序、组装,利用Blast软件进行数据库比对和基因功能注释,用DEGSeq R软件包分析4℃低温处理与室内适温饲养试虫的差异表达基因,并对上调表达基因进行GO和KEGG代谢途径富集分析.[结果]经序列拼接后共获得100300个unigenes,总长度81600309 bp,平均长度813 bp,N50长度1719 bp.与7大数据库同源比对,共获得34691(34.59％)条unigenes.低温胁迫转录组分析得到11569个差异表达基因(DEGs),7158条基因上调,4411条基因下调.富集到47个GO类群,217个KEGG途径.其中代谢过程、催化、结合活性类群占有重要比例,核糖体、碳代谢、剪接体等途径显著富集.另外,热激蛋白、昆虫表皮蛋白、海藻糖酶、超氧化物歧化酶等非生物胁迫相关基因显著上调表达.[结论]阿尔泰蝠蛾幼虫低温胁迫转录组分析揭示,代谢过程、细胞过程、生物调节、对刺激的反应等生物学过程相关基因和部分非生物胁迫响应基因显著上调表达,提示蝠蛾幼虫可能从抗氧化防御、分子伴侣、体温调节和维持细胞的渗透平衡等多方面应对低温胁迫.     

云南西藏蝠蛾属四新种（鳞翅目：蝙蝠蛾科）

本文报道了云南西北部和西藏东南部的高寒草甸中分布的蝠蛾属４新种：剑川蝠蛾Ｈｅｐｉａｌｕｓｊｉａｎｃｈｕａｎｅｎｓｉｓｓｐ．ｎｏｖ．,甲郎蝠蛾Ｈ．ｊｉａｌａｎｇｅｎｓｉｓｓｐ．Ｎｏｖ．,察里蝠蛾Ｈ．ｚａｒｉｅｎｓｉｓｓｐ．ｎｏｖ．,异翅蝠蛾Ｈｅｐｉａｌｕｓａｎｏｍｏｐｔｅｒｕｓｓｐ．ｎｏｖ．４个新种都是冬虫夏草真菌的寄主昆虫。     

基于高通量测序的阿尔泰蝠蛾(鳞翅目:蝙蝠蛾科)幼虫转录组分析

本研究采用Illumina HiSeq TM 2500测序平台对阿尔泰蝠蛾Hepialus altaicola Wang幼虫进行转录组测序及生物信息学分析.经序列拼接后共获得100133个Unigenes,总长度86319112 bp,平均长度862 bp,N50长度1628 bp.将Unigenes与NR、COG/KOG、Pfam、Swiss-Prot、GO、KEGG数据库比对,共获得38198条Unigenes,其中Nr数据库注释的Unigenes最多,为32381条,占32.34％.通过GO功能分类,共有13216个Unigenes在GO数据库中细胞组分、分子功能和生物学过程等3大类57个分支中找到注释;KEGG通路分析,共有15058条Unigenes被注释,归属于305条代谢通路.CDS预测发现54002条序列可被编码,占全部基因的53.93％.基因注释进一步获得311个与冷适应相关的代谢调节基因,并用FPKM值对基因表达量进行评估.本研究获得的转录组信息及分析结果,为进一步研究阿尔泰蝠蛾的基因功能及低温生态适应性奠定分子基础.     

温度胁迫对小金蝠蛾幼虫抗氧化酶活性的影响

小金蝠蛾Hepialus xiaojinensis Tu.是我国名贵中药材冬虫夏草的寄主昆虫之一,适宜在高海拔低温地区(8℃-12℃)生存。本文研究了小金蝠蛾在不同温度(4℃、12℃、16℃、20℃、24℃、28℃)条件下处理4 h和12 h后4龄幼虫体内超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)、谷胱甘肽硫转移酶(GSTs)的活力变化,以及总抗氧化能力(T-AOC)和丙二醛(MDA)含量的变化。结果表明,小金蝠蛾在正常生长发育情况下SOD具有较高活性;经温度胁迫后,SOD活力持续维持在较高活性,CAT、POD活性以及T-AOC活性显著性升高(P0.05);GSTs在24℃以上活性有降低趋势,MDA含量有上升趋势。     

滇川蝠蛾四新种(鳞翅目:蝙蝠蛾科)

本文报道中支蝠蛾、叶日蝠蛾、美丽蝠蛾和里塘蝠蛾4个新种,这些新种分别采于云南和四川。     

Mating systems in primitive Lepidoptera, with emphasis on the reproductive behaviour of Korscheltellus gracilis (Hepialidae)

Adults of Korscheltellus gracilis (Grote) are crepuscular. Flight is restricted to two 20–40 minute periods each day: the first flight follows sunset during evening twilight, the other precedes dawn. Four to five times as many adults are active during the evening when courtship, mating and oviposition occur. Of these behaviours, only oviposition occurs during the pre-dawn period of activity. Field observations indicate that females release a sex pheromone while fanning their wings. Females usually fly before pairing. There is no obvious female choice or stereotyped courtship behaviour prior to pairing.
In some hepialid genera females attract males, while in others ( Hepialus Fabricius, Phymatopus Wallengren, Sthenopis Packard and Zenophassus Tindale) the calling system is reversed, with males attracting females. Cladistic analysis of the Hepialidae suggests that the plesiomorphic condition for the family is for females to release the long-distance attractant, and that the reversed calling system represents a derived condition restricted to lineages within a single clade.
Available information on the mating systems of other basal lepidopteran lineages and Trichoptera support the hypothesis that ancestral Lepidoptera possessed a female-released, long-range attractant. Support for this position comes from (1) published literature; (2) field observations made by DLW; (3) the apparent absence of male scent structures and recognizable male calling behaviour among the pre-Neolepidoptera; and (4) cases of male antennal elaboration in several basal lepidopteran lineages.     

Ultrastructural analysis of spermatozoa of Korscheltell us lupulinus L. (Lepidoptera : Hepialidae) and Micropterix calthella L. (Lepidoptera : Micropterigidae)

In Korscheltellus lupulinus (Lepidoptera : Hepialidae), both eusperm and parasperm can be found, whereas Micropterix calthella (Lepidoptera : Micropterigidae) produces only eusperm. The eupyrene sperm in K. lupulinus is 250 μm long. Spermatozoa found in K. lupulinus are very similar to those found in higher Lepidoptera, but the eusperm of M. calthella do not possess similar features. As in other Lepidoptera, the axoneme pattern of K. lupulinus is 9 + 9 + 2. Anteriorly, the axoneme is replaced by a fibrous stem similar to an acrosomal structure; posteriorly it becomes progressively uneven. The sperm of K. lupulinus has other original features. The anterior region of the sperm is rigid and contains the nucleus, which has a longitudinal groove adjacent to the axoneme. The 2 mitochondrial derivatives extend from the posterior end of the nucleus. There are no accessory bodies. Many peripheral cytoplasmic vesicles are present. A thin sheath surrounds the cell. It is thick in the parasperm and made of concentric or spiralling lamellae; it does not have any lacinate appendages.The axoneme of the sperm of M. calthella extends over the entire length of the cell and spirals in a helix around the nucleus. The nucleus is also almost as long as the cell. The 2 mitochondrial derivatives, flanked by accessory bodies, follow this spiralling. The axoneme has a 9 + 2 pattern. The plane, which contains the 2 central microtubules, is at an angle with the median plane of the nucleus.     

Visual mate-searching behaviour in the evening brown butterfly, Melanitis leda (L.) (Lepidoptera: Nymphalidae)

Abstract As in most animals, male butterflies are generally the more proactive sex with respect to seeking out mating opportunities. In most cases, males conduct their search sometime between mid-morning and mid-afternoon, but a few species are active only before dawn or after sunset. These crepuscular species offer a good opportunity to study how males deal with markedly different visual and thermal conditions. Here, I present data from a 5-month behavioural study of male Melanitis leda (Nymphalidae) at a mate-encounter site in tropical Australia. Males of this species defended perching locations along a forest edge in a similar manner to other diurnally active territorial nymphalids. They generally arrived at these sites after sunset and arrived earlier on evenings that darkened earlier. Actively mate-locating males were only seen at the site during a 25−35 min evening period, during which ambient light levels ranged between 50 and 2600 lux. Only 27% of marked territory residents were recorded again at their location of capture, but fidelity in this 'resighted' group ranged up to 23 days. A sample of males, captured under ambient temperatures of 24.0−27.2°C, maintained a mean thoracic excess of 8.25 ± 0.73°C, but did not appear to 'shiver' in the manner of other crepuscular species. Males courted conspecific females and one mating was observed. This species is an excellent candidate for further research into the evolution of mating tactics in crepuscular butterflies.