Image Credit: Hectonichus, CC BY-SA 3.0, Image Cropped

Would you rather stab, or be on the receiving end of a stab? This may seem like a confronting question, but it’s the dilemma many flatworms face when a mating opportunity arises. 

Option 1: You are a flatworm and have just been stabbed by a stubby penis. You now have puncture wounds that must heal, after which you must carry fertilized eggs which you need to lay and protect upwards of 24 hours. Oh, the energy demands! 

Option 2: Flatworm victory! You have successfully stabbed your opponent with your stubby penis before they could stab you. Your sperm has now fertilized their eggs. With this win, you move on with life and wait for your next mating “opponent”.

Which option do you choose? If you still can’t choose, it’s a good thing you aren’t a simultaneous-hermaphroditic flatworm. These flatworms have both fully functional male and female reproductive capabilities that can be used interchangeably, unlike other hermaphroditic species who switch back and forth during different phases of life. One might say these individuals have the capability to “choose” what role they want to play, male or female. Although, those forced into the role of reproductive female may disagree…

It is believed that individuals fight to “remain male” (i.e., not be fertilized) because sperm is biologically cheaper to produce than eggs, and males can produce more offspring than females over a lifetime. This type of fight has been thought to be “pure evolutionary selfishness”.

It was only discovered recently, after Dr. Leslie Newman and Dr. Nicholas Michiels spent 20 hours continuously watching pairs of captured flatworms. They observed that when an individual encounters another, both assume a fighting stance, curling their bodies back to display their penises. Next, they began to fight, each attempting to stab the other, which could last from 20 to 60 minutes.

Different species fight with different strategies. For example, racing-line flatworms (Pseudocerotidae bifurcus) use their penis to repeatedly strike at one another until one succeeds, injecting sperm under the skin of the other. Once the sperm is injected, it moves through the body to find and fertilize the eggs. Persian carpet flatworms (P. bedfordi, pictured above) instead use their penis like a water gun, ejaculating anywhere on their opponent’s body. With a sperm cocktail that dissolves flesh, it burns its way through various tissues until it reaches and fertilizes the eggs. 

Penis fencing is the term scientists use to describe this behavior to “remain male”. This mating behavior isn’t seen amongst all flatworm species, only certain species within the family Pseudocerotidae. In the 1990’s there were only two species of flatworm known for this behavior, however as of 2020, the number has grown to 16. 

Evolution of Penis Fencing

Species of flatworms can use sexual reproduction (need both gametes; sperm and egg), asexual reproduction (does not require both gametes, obtain all DNA from parent), or both. Those that use both, do so depending on which strategy is favoured by the environmental conditions. For example, sexual reproduction is favored under harsher, more unpredictable conditions, since genetically variable offspring are often better able to adapt and survive these conditions. Asexual reproduction may be favored when individuals are scarce, however it tends to be avoided as there is on average a 50% loss of genetic diversity per generation, subsequently increasing the probability of inbreeding in future generations. If asexual reproduction does occur, it can occur through budding or transverse fission. Budding occurs when ‘buds’ (i.e., outgrowth) grow out of the flatworm’s body until they are large enough to break off as new individuals. Fission, on the other hand, involves an individual being cut in half, with each half becoming a new individual. 

A species may employ different hermaphroditic strategies of cross-fertilization depending on their ecological niche. These include delivery of sperm to a sperm-receiving organ of the mating partner, or hypodermic insemination of sperm into the cellular tissue by a modified penis that enables individuals to pierce the body wall of their partner. It is believed that the willingness to invest as little resources as possible into their offspring is very strong in hermaphroditic species, leading to these extreme mating behaviors such as penis fencing.  

Yet penis fencing does not always occur when individuals meet. Four possible scenarios have been observed when individuals encountered one another: 

1. Both partners were receptive to mating and penis fencing was observed, 
2. Both partners were receptive but no penis fencing was observed, 
3. Only one partner was receptive and no penis fencing was observed however insemination was successful, and 
4. Neither were receptive to mating. 

If penis fencing occurs, it typically leads to successful sperm insemination for one or both individuals. Number 3 may be the result of other mating behaviors. For example, mating Starry flatworms (P. stellae) will curl around each other, swimming in circular motions in attempts to inseminate each other. 

Outcomes of Penis Fencing

A more recent study in 2020 found that penis fencing results in three outcomes; 1) both individuals were inseminated, 2) one individual was inseminated, or 3) neither were inseminated. These researchers found penis fencing to be more of a duel or contest mating ritual, rather than an aggressive, violent behavior as was originally thought. This is because they found different scenarios where penis fencing occurred that resulted in neither individual being inseminated, or where no penis fencing occurred resulting in at least one individual being inseminated. Although we may think of penis fencing a little differently now, one thing that will forever remain constant are the words of David Attenborough, “its only solace is knowing it’s young will carry the genes of a master swordsman”.  

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Jennifer Merems is a writer and researcher focusing on behavioral and nutritional ecology. She is currently a PhD candidate in the Department of Forest and Wildlife Ecology with the University of Wisconsin-Madison. You can learn more about Jennifer by following her on Twitter at @atyourcervid.