Mating marks on museum specimens reveal breeding patterns in species of Pterostichus Bonelli (Carabidae, Pterostichini)

We found distinct and consistently placed, species- and sex-specific abrasions of the cuticle on museum specimens of 14 species of the Pterostichus Bonelli, 1810 (Carabidae, Pterostichini) subgenusHypherpes Chaudoir, 1838. We deduced that these marks are generated during mating and, therefore, can be used to distinguish between preserved specimens of beetles that had previously mated at the time of capture and those that had not mated. In addition to describing and detailing the occurrence of the marks and providing evidence that they are the result of mating, we demonstrate their utility for inferring life history using a museum voucher collection. By scoring these indications of mating from pinned specimens, we describe life cycle patterns in two similar, relatively closely related and sympatric species of the subgenus Hypherpes, P.vicinus Mannerheim, 1843 and P.californicus (Dejean, 1828). Both were sampled during a pitfall trap study in Contra Costa, California, USA from 2014–2019 and deposited in the Essig Museum of Entomology, UC Berkeley. Both species had very low adult activity through the drought and end of drought period prior to the spring of 2017 and are significantly more abundant in the post-drought period. Based on mating marks, both species responded to accumulated precipitation ending the drought by the emergence of an active, mostly unmated cohort of adults. The spring activity peak, following the end of the drought, was dominated by unmarked and presumably unmated beetles, but samples from subsequent springs included a nearly equal mix of beetles showing mating marks and apparently unmated beetles. The beetle activity appears to correspond more with the accumulated rainfall of the preceding rainy season than with the rains of the sample year. Beetles sampled in autumn and winter (rainy season) predominantly show mating marks. The occurrence throughout the year of beetles that are marked as having mated is consistent with iteroparous beetles with a lifespan of more than one year and also consistent with dynamic phenotypic polyvariance in which the adult activity period is synchronised by adjusting development time. The dominant pattern fits with a life cycle that is typically annual univoltine, or possibly biennial semivoltine in dry years, rainy season breeding (autumn-winter) iteroparous, with adult summer aestivation and possibly facultative larval hibernation. However, unmarked and so apparently unmated individuals and teneral adults were captured during peak activity periods regardless of the season, suggesting that either the beetles diapause as teneral adults that then complete development and become active at various points during the year and/or there are multiple periods of breeding and oviposition each year in at least some portion of the population.