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Pteronymia sao(Hübner, [1813]) Figure 8 Females examined: 56. 15 with complete sphragides, 12 bearing incomplete or amorphous sphragides. Description: Coloration—golden-brown. Shape—circular shape when viewed ventrally, irregular in shape with several lacunae when viewed laterally. Structure—mostly hollow. Scales—few scales present, not ordered., Published as part of Carvalho, Ana Paula S., Mota, Luísa L. & Kawahara, Akito Y., 2019, Intersexual ' Arms Race' and the Evolution of the Sphragis in Pteronymia Butterflies, pp. 1-13 in Insect Systematics and Diversity 3 (2019) on page 7, DOI: 10.1093/isd/ixy021, http://zenodo.org/record/4551929

Pteronymia obscuratus(Fabricius, 1793) Figure 4 Females examined: 15. 11 with complete sphragides; 3 bearing incomplete or amorphous sphragides. Description: Coloration—golden-brown. Shape—rounded anteriorly with two symmetrical lateral cavities, an elevated projection medially, and the sphragis curves upward and over the sterigma posteriorly, overhanging the sterigma. The tip of the sphragis is pointed and nearly touches the papillae anales. Structure—mostly solid. Scales—present, ordered, hair-like. Attached to leading edge of the overhanging posterior half of sphragis. Comments: We observed one female with a spermatophore attached to the sphragis outside of the female body (Fig. 4D). It is likely that there was already a sphragis in place on the female's sterigma, and that the second male was unsuccessful in depositing his spermatophore inside the female's reproductive tract, but still produced a complete sphragis on top of the first one., Published as part of Carvalho, Ana Paula S., Mota, Luísa L. & Kawahara, Akito Y., 2019, Intersexual ' Arms Race' and the Evolution of the Sphragis in Pteronymia Butterflies, pp. 1-13 in Insect Systematics and Diversity 3 (2019) on pages 5-6, DOI: 10.1093/isd/ixy021, http://zenodo.org/record/4551929

Pteronymia ozia(Hewitson, 1870) Figure 6 Females examined: 12. 1 with a complete sphragis; 2 bearing incomplete or amorphous sphragides. Description: Coloration—dark chocolate-brown. Shape—platelike. Structure—mostly hollow. Scales—absent. Complexity category—likely 2, but more specimens are needed to properly assign a complexity category. Comments: The sphragis figured here is the only one that was complete among the sphragides examined. It is placed laterally in the sterigma, and we suspect it was not properly deposited by the male or that it was displaced after attachment., Published as part of Carvalho, Ana Paula S., Mota, Luísa L. & Kawahara, Akito Y., 2019, Intersexual ' Arms Race' and the Evolution of the Sphragis in Pteronymia Butterflies, pp. 1-13 in Insect Systematics and Diversity 3 (2019) on page 6, DOI: 10.1093/isd/ixy021, http://zenodo.org/record/4551929

Pteronymia andreas(Weeks, 1901) Figure 2 Females examined: 22. 1 with a complete sphragis; 12 bearing incomplete or amorphous sphragides. Description: Coloration—golden-brown to yellowish off-white. Shape—plate-like. Structure—mostly hollow. Scales—found in some specimens, but not ordered, probably accidentally attached. Complexity category—2. Comments: The complete sphragis that we observed was likely deformed due to the way it was attached to the sterigma. Nearly all (12/13, 92%) of the sphragides observed for this species were incomplete. Incomplete sphragides were in the form of a plate, and always completely internalized inside the bowl-like sterigma. The amount of sphragidal material was variable, but often it only covered the ostium bursae and immediate surroundings., Published as part of Carvalho, Ana Paula S., Mota, Luísa L. & Kawahara, Akito Y., 2019, Intersexual ' Arms Race' and the Evolution of the Sphragis in Pteronymia Butterflies, pp. 1-13 in Insect Systematics and Diversity 3 (2019) on page 5, DOI: 10.1093/isd/ixy021, http://zenodo.org/record/4551929

Pteronymia simplex(Salvin, 1869) Figure 9 Females examined: 94. 39 with complete sphragides; 41 bearing incomplete or amorphous sphragides. Description: Coloration—white to golden yellowish brown. Shape—bowl-like. Margin of the sphragis curled. Structure—mostly hollow. Scales—few, not ordered. Comments: We found five females bearing two sphragides each (e.g., Fig. 16), but we could not locate the spermatophore of the second male outside of the body of any of these females, and they were likely broken. Similar to P. alissa and P. picta, the level of sphragis completeness of P. simplex was highly variable. The shape of these incomplete sphragides was comparable to those that had complete sphragides, differing only in its smaller size or placement, which was entirely within the bowl-like sterigma. We also observed some structures that resembled a complete sphragis, but which were barely externalized beyond the margins of the bowl-like sterigma. It was occasionally difficult to determine whether the sphragis was complete, and this is due to the fact that the amount of sphragidal material produced by males in SB Pteronymia is extremely inconsistent., Published as part of Carvalho, Ana Paula S., Mota, Luísa L. & Kawahara, Akito Y., 2019, Intersexual ' Arms Race' and the Evolution of the Sphragis in Pteronymia Butterflies, pp. 1-13 in Insect Systematics and Diversity 3 (2019) on page 7, DOI: 10.1093/isd/ixy021, http://zenodo.org/record/4551929

Pteronymia calgiriaSchaus, 1902 Figure 3 Females examined: 1 bearing complete sphragis. Description: Coloration—golden-brown. Shape—round, cupped, extremities curled inward. Structure—mostly solid. Scales—absent. Complexity category—likely 2, but more specimens are needed in order to accurately assign a category., Published as part of Carvalho, Ana Paula S., Mota, Luísa L. & Kawahara, Akito Y., 2019, Intersexual ' Arms Race' and the Evolution of the Sphragis in Pteronymia Butterflies, pp. 1-13 in Insect Systematics and Diversity 3 (2019) on page 5, DOI: 10.1093/isd/ixy021, http://zenodo.org/record/4551929

Carvalho, Ana Paula S., Mota, Luísa L., Kawahara, Akito Y. (2019): Intersexual 'Arms Race' and the Evolution of the Sphragis in Pteronymia Butterflies. Insect Systematics and Diversity 3 (2019), No. 3: 1-13, DOI: 10.1093/isd/ixy021, URL: http://dx.doi.org/10.1093/isd/ixy021

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 the Parnassius simo group, 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.