Carabus insulicola

Two methods that male C. insulicola use to increase their reproductive success are post-copulatory guarding and spermatophore displacement.

[1] Post-copulatory guarding is when a male remains on the female’s back after copulation,[2] which increases the likelihood of the first mate fertilizing the eggs.

[6] Larvae can be found under objects and debris on the ground, while pupae grow in cells made of soil buried a few inches underground.

They hunt smaller invertebrates for their primary food source, including snails, earthworms, and caterpillars.

[4] In laboratory experiments using C. insulicola, they can be fed minced beef and chopped apples to mimic their natural diet.

This allows for sperm competition to occur, where the female can then ensure that her eggs will be fertilized and increase the genetic quality of her brood.

[3] Adults emerge from the pupal stage in late summer to autumn, followed by hibernation throughout the winter without mating.

[1] The spermatophore, a hemispherical capsule that contains sperm, is deposited through an opening at the end of the endophallus called the gonopore.

The spermatheca is a skinny tube-like structure responsible for storing the sperm that is taken from the spermatophore as it is digested in the bursa copulatrix and fertilizing the eggs.

Sperm can only be transferred to the spermatheca if the spermatophore is located at the innermost corner of the bursa copulatrix and attached to the vaginal apophysis.

Research has found that males have evolved to have longer copulatory pieces in order to increase their reproductive success.

A longer vaginal appendix lessened this fitness cost by decreasing egg dumping and increasing fertilization.

During a weak rejection, the female will extend segments of her abdomen to block the aedeagus from reaching the opening.

For a strong rejection, she will bend her abdominal terminalia upwards to prevent deeper insertion of the aedeagus.

Acquiescence occurs when the genitals are coupled and the female stops resisting the male, allowing him to copulate.

[2] During the copulation stage, the aedeagus enters the bursa copulatrix and the male’s copulatory piece is hooked inside the female’s vaginal appendix.

Sperm displacement is a tactic used by males of many species to ensure paternity when females mate multiply.

[4] When a female mates multiply, the latest male’s spermatophore is the one that is ultimately located on the innermost part of the bursa copulatrix, making it closest to the spermatheca.

While spermatophore displacement is an effective offensive tactic for males, it is not a foolproof way to guarantee paternity.

[2] It benefits the male to stay on for as long as possible because post-copulatory guarding increases the chance of paternity.

Eggs laid earlier, or closer in time to the end of copulation, were more likely to be fertilized by the second male.

Honshu, Japan, where Carabus insulicula is found