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A review of the occurrence and diversity of the sphragis in butterflies (Lepidoptera, Papilionoidea)

Authors: Ana Paula S. Carvalho; Albert G. Orr; Akito Y. Kawahara;

A review of the occurrence and diversity of the sphragis in butterflies (Lepidoptera, Papilionoidea)

Abstract

Males of many butterfly species secrete long-lasting mating plugs to prevent their mates from copulating with other males, thus ensuring their sperm will fertilize all future eggs laid. Certain species have further developed a greatly enlarged, often spectacular, externalized plug, termed a sphragis. This distinctive structure results from complex adaptations in both male and female genitalia and is qualitatively distinct from the amorphous, internal mating plugs of other species. Intermediate conditions between internal plug and external sphragis are rare. The term sphragis has often been misunderstood in recent years, hence we provide a formal definition based on accepted usage throughout most of the last century. Despite it being a highly apparent trait, neither the incidence nor diversity of the sphragis has been systematically documented. We record a sphragis or related structure in 273 butterfly species, representing 72 species of Papilionidae in 13 genera, and 201 species of Nymphalidae in 9 genera. These figures represent respectively, 13% of Papilionidae, 3% of Nymphalidae, and 1% of known butterfly species. A well-formed sphragis evolved independently in at least five butterfly subfamilies, with a rudimentary structure also occurring in an additional subfamily. The sphragis is probably the plesiomorphic condition in groups such as Parnassius (Papilionidae: Parnassiinae) and many Acraeini (Nymphalidae: Heliconiinae). Some butterflies, such as those belonging to theParnassius simogroup, have apparently lost the structure secondarily. The material cost of producing the sphragis is considerable. It is typically offset by production of a smaller spermatophore, thus reducing the amount of male-derived nutrients donated to the female during mating for use in oogenesis and/or somatic maintenance. The sphragis potentially represents one of the clearest examples of mate conflict known. Investigating its biology should yield testable hypotheses to further our understanding of the selective processes at play in an ‘arms race’ between the sexes. This paper provides an overview, which will inform future study.

Keywords

Cossina, Insecta, Arthropoda, sperm precedence, Nephrozoa, Protostomia, Basal, mate conflict, Review Article, Carbotriplurida, sexual competition, Circumscriptional names of the taxon under, sperm guarding, Panorpida, Animalia, Bilateria, Eumetabola, Galacticoidea, Pterygota, Nymphalidae, Papilionoidea, Papilionidae, Strashila incredibilis, Circumscriptional names, Lepidoptera, Heteroneura, Boltonocostidae, PapilionoideaCephalornis, Bombycina, mating plug, QL1-991, Notchia, LepidopteraAnimalia, Circumscriptional name, Ecdysozoa, Amphiesmenoptera, Zoology, Porina, Ditrysia, Coelenterata

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citations
This is an alternative to the "Influence" indicator, which also reflects the overall/total impact of an article in the research community at large, based on the underlying citation network (diachronically).
BIP!Citations provided by BIP!
popularity
This indicator reflects the "current" impact/attention (the "hype") of an article in the research community at large, based on the underlying citation network.
BIP!Popularity provided by BIP!
influence
This indicator reflects the overall/total impact of an article in the research community at large, based on the underlying citation network (diachronically).
BIP!Influence provided by BIP!
impulse
This indicator reflects the initial momentum of an article directly after its publication, based on the underlying citation network.
BIP!Impulse provided by BIP!
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