taxonID	type	description	language	source
03C36B2DB831521C78835DE8FECE2F34.taxon	description	The sterigma of female Hypermnestra is not obviously specialised to reject plugging by the male but is highly modified from the ancestral condition found in most non-sphragisbearing butterflies and the ostium is still rather exposed. Moreover, the fleshy projection seems to support the sphragis. The projection itself is relatively soft (at least when macerated), but after a male smears it with a large amount of secretory material, it provides a core, supporting a sphragis at the right angle to the abdomen to create an obstacle to remating with another male. If so, the female genitalia of Hypermnestra may be secondarily adapted to retain a sphragis. Possibly the females actively grip a small sphragis that will inhibit remating; in this way they might receive a larger spermatophore. Probable sphragis-retaining female genitalia among sphragis-bearing Papilionidae are otherwise only known in Parides proneus (Hübner) (Orr 1988, 1995). The female genitalia in this species are exposed but equipped with a sclerotised bar in front of the ostium, which anchors a small sphragis. It is possible that this is a female adaptation to retain the sphragis meaning that a small sphragis is still effective, thus enabling males to commit more secretory material to a large spermatophore. In general, it is possible that in certain circumstances females evolve the capacity to hold a sphragis when additional matings are no longer advantageous. This may be because the male has already provided a significant nuptial gift, together with the small sphragis, or, in cases where male ejaculates are small and spermatophores have little nutritional value, bearing a sphragis may free a female from the risks and costs of unwanted copulation (i. e. waste of time and energy, risk of predation or physical damage). Orr and Rutowski (1991) demonstrated that females of Cressida cressida, which receive no nutrients from the male (Orr 1988), display the sphragis to avoid unwanted second matings, which in this species may prove fatal (Orr 1999). Wedell et al. (2002) demonstrated that there is a genetic variation in female mating frequency in Pieris napi (Linnaeus) in Sweden and suggested that the variation can be related to a trade-off between the nutritional benefit and time available for oviposition in the unpredictable Swedish weather. Mated female H. helios may benefit from bearing a sphragis while ovipositing, because the species lives in a dry open habitat and females lay eggs singly on a small hostplant (Igarashi 1979), hence ovipositing females probably conspicuously visit many hostplants and they may frequently encounter and be harassed by searching males, a situation where the female may benefit by retaining an effective sphragis.	en	Matsumoto, Kazuma, Orr, Albert G. (2021): The occurrence of a sphragis and genital modification in Hypermnestra, Archon and Sericinus (Lepidoptera: Papilionoidea: Papilionidae). Journal of Natural History 55 (15 - 16): 1033-1057, DOI: 10.1080/00222933.2021.1930227, URL: http://dx.doi.org/10.1080/00222933.2021.1930227
03C36B2DB834521B78835FC4FBBA2FC4.taxon	description	The invaginated genital plate of Sericinus is probably difficult to plug, because it requires a relatively large investment of the plug material, if the hollow space must be blocked by completely filling it. It may have evolved as an anti-plugging device. The lidlike structure of the sphragis is a male counter-adaptation to overcome it while saving sphragidal material. The secretion lining the inner wall, including the bottom where the ostium opens, may not be easily penetrated and extracted. The male accessory glands which produce the sphragis are only moderately hypertrophied (Orr 1988, 1995) and their capacity to produce sphragidal material is probably quite limited, presumably the reason why the sphragis seldom fills the invagination and why many mated females lack a sphragis. The sharply hooked tips of the robust valvae may be used for removing the sphragis, if the opportunity arises. The short ductus bursae may be a plug-rejecting structure effective in past, when the genital plate was flat. This evidence suggests an arms race between the two sexes in the past, but we suspect the intersexual conflict in this species is now relaxing, especially because of the somewhat low frequency of the sphragis and high incidence of monoandry in wild females as indicated by the spermatophore counts. In Sericinus, the sphragis may be becoming redundant. A relatively semelparous oviposition schedule and short adult life span in female Sericinus may reduce the need for the sphragis as a sperm guarding device. Monastyrskiy and Kotlobay (1995) reported that female S. montela from Maritime Territory of Russia began to oviposit 10 min – 15 min after copulation and oviposited every other day for 5 days with 65 % of the eggs being laid on the first day. Li et al. (2016) reported that female S. montela of Gansu Province, China mated on the day of eclosion and laid eggs on the same day or the next day and then died. The reason for these somewhat different results is unknown, but both studies indicate that females live rather short lives and lay most of their eggs soon after the mating. In such a situation they would gain little from male nutrients in oogenesis. Our sample also indicates old females are rather sparse in the field, suggesting a short female adult life (Tables 4 and 5). The female lays eggs in large clusters, probably only a few times during their adult life. Accordingly, the period during which a mated female has high reproductive value (Fisher 1930) should be short. In this case the sperm guarding value of the sphragis quickly declines with time after mating. Moreover, the females are cryptic and inactive (Monastyrskiy and Kotlobay 1995; Li et al. 2016), hence not easy to locate by searching males. In this situation, the males may not be selected to invest much in a sphragis nor to produce many sphragides. It is now confirmed that a sphragis, in some form, occurs in almost all species of all genera in Parnassiinae. The subfamily includes genera having a protosphragis (Hypermnestra), a vestigial sphragis (Bhutanitis and Zerynthia), various types of large elaborate sphragides (Parnassius, Luehdorfia, Archon, Sericinus and Allancastria) as well as those that have probably secondarily lost the sphragis (Parnassius simo group). Archon and Sericinus may be close to an evolutionary tipping point in terms of losing the sphragis. Parnassiinae is thus an important model group for investigating the process of sphragis evolution. As well as the functional morphological analysis presented here, investigations coordinating ecological, behavioural and physiological factors and relating these to the presence or absence of a sphragis are needed. Ecological factors include habitat type, hostplant dispersion, population structure and life tables with oviposition and survivorship schedules; behavioural traits include mate location and mating behaviour and encounter rates; physiological traits chiefly relate to the material benefits females may gain from mating and material costs to males of sphragis production. With such information it would be possible to construct a more holistic model of sphragis evolution and hopefully also gain insight into the circumstances leading to several anomalous forms.	en	Matsumoto, Kazuma, Orr, Albert G. (2021): The occurrence of a sphragis and genital modification in Hypermnestra, Archon and Sericinus (Lepidoptera: Papilionoidea: Papilionidae). Journal of Natural History 55 (15 - 16): 1033-1057, DOI: 10.1080/00222933.2021.1930227, URL: http://dx.doi.org/10.1080/00222933.2021.1930227
