The Benefits of Being Ugly

...include siring more offspring- so long as you are a firefly..

MALE COURTSHIP ATTRACTIVENESS AND PATERNITY SUCCESS IN FIREFLIES

So this is a way cool study in fire flies.. Briefly, the researchers do an experiment to determine if mating success correlates with reproductive success (actual parentage) They allow females to choose and mate with their preferred male- then they allow a second copulation (to have a 2nd copulation is typical for this promiscuous species), but now they impose the choice- in some cases, they allow an attractive male to mate, in others, an unattractive.. To add more mystery to the story, attractive and unattractive males offer spermatophores of similar size and quality...Below are the results: less attractive males ultimately sire more offspring and have greater reproductive success than do attractive males.

The authors offer a few possible explanations:

1. Trade-off. Here males may choose one strategy.. either be attractive and have high mating success, of be unattractive, and win in the end... Seems like this is a no brainer... there is not way I can see for the possibility of a mixed ESS... We would not expect to see the attractive male strategy in the population.
2. Post-copulatory female choice makes up for error in pre-copulatory choice... Basically, that the attractive male signal is not honest, but instead is manipulative. Again, I'm pretty sure that natural selection would remove the honest signal pretty quickly.. Could all females be mistaken? I don't think this is a very good explanation either..
   

So this leads me to ask.. Why are there males investing a lot of time and energy, and risk in a mating strategy that does not pay off? Why do females mate with these "attractive males", then bias paternity towards less attractive.. The usual thing is to say that the the female gains direct benefits, but that does not seem to be the case here.. If attractive males have lower relative fitness that the ugly guys- how do they maintain in the population.. I wonder if there is some pleiotropic effect.. Maybe ugly fireflies are more susceptible to predation, etc... Maybe attractive fireflies are more resistant to infectious diseases, live longer, ect...I'm perplexed.

We always think:

1. that mating success is positively correlated with actual reproductive success... I wonder how often these 2 are in opposition?