Years ago I met Richard Fortey at the inaugural meeting of Spain’s new evolution society, and found him an affable and lovely guy. He’s a paleontologist and writer, and I had the pleasure of reading and giving a positive review to his first book, Life: A Natural History of the first Four Billion Years on Earth, which is well worth reading (he’s written several other books, including Trilobite: Eyewitness to Evolution (also a good read).
And it’s four trilobite species that are the subject of Fortey’s new paper coauthored with Alan D. Gishlick, a geophysical sciences professor at Bloomsburg University, in PNAS, a paper you can read for free by clicking the title below (it’s free with the legal Unpaywall app., the pdf is here, the reference is at bottom, and judicious inquiry might yield a pdf if you can’t see the paper). Trilobites are common fossils, and were marine arthropods that went extinct without leaving descendants.
The upshot is that Gishlick and Fortey analyzed fossils of one species of trilobite found in Morocco, deriving from the Devonian (400 million years ago). This species, Walliserops trifurcatus, had a long trident attached to the front of their bodies, and tried to figure out what it was for. They also found one adult individual whose trident was a bit deformed (see below). Their conclusion is that these were weapons used by males to fight with other males, almost surely to compete for females. They are, posit the authors, the arthropod equivalent of reindeer horns. The other possible functions (feeding, digging, etc.) were largely ruled out.
Here are four species of Walliserops, shown below. All specimens bear a rigid cephalic trident. W. trifurcatus has a slightly recurved trident that bends upwards, while the other species have tridents more flush with the surface of the sediment (all captions come from the paper):
The obvious question is: what is this damn thing for? And there are several hypotheses, all assuming that the structure was molded by natural selection (which includes sexual selection). The authors find evidence against all but one possible function. Here are the alternatives (of course, it could have been used for several things, but it’s likely that selection was wholly or largely on one function). Indented bits are quotes from the paper. The rest of the discussion concerns W. trifurcatus:
A.) Defense. Perhaps the structure could have been used to ward off predators, like the spines found on other trilobites. Here’s how the authors rule this out:
However, such a function would have been difficult given the overall anatomy of the trident and the trilobite. The trident is rigidly attached and cannot be moved independently from the cephalon; it could only be flexed in a dorsal-ventral plane by the trilobite raising and lowering its cephalon. This would create further difficulties since the long genal spines limit how high the head could be angled without lifting the entire body. The trident, therefore, could not be employed in a versatile way, nor be presented as to defend from a predator attacking from above or behind. This morphology is not consistent with a defensive structure.
B.) A feeding structure. Doesn’t seem likely:
A second possible function for the trident would be as an aid to feeding. Like all members of the Phacopida, Walliserops was probably a scavenger/predator, and it might be considered as a possibility that the trident was a comparatively sophisticated sensory device concerned with early detection of prey species—such as buried annelid worms—which could then be grasped by the endopods of the ventral limbs.
C.) Sensory detection of the environment. This is also deemed unlikely from inspection of the structure:
However, examination of the trident in optical and scanning electron microscopy failed to find the arrays of cuticular pits or tubercles usually indicative of the presence of sensilla in fossil arthropods. Most groups of trilobites include species with exterior exoskeletal pitting that is preserved even if the intracuticular canals have been removed by calcite reorganization—and there is no evidence of such exterior pitting on the trident of Walliserops. The absence of evidence for specialized organs on the tines makes it unlikely that it was primarily a sensory apparatus.
D.) A spear to pierce prey: Unlikely because the structure was inflexible, so the animal would have no way of accessing speared prey.
E.) An apparatus to dig, perhaps for prey. The way it’s shaped and angled seems to preclude this (remember, it’s slightly recurved upward; see below):
Another possibility is that the trident may have been used to agitate sediment to disturb prey items, which could then be trapped by the limbs. It is difficult to conceive of W. trifurcatus digging into sediment because to engage sufficiently with the substrate the cephalon would have to tilt at an angle greater than would be allowed by movement on the posterior occipital margin. Equally, if the thorax was arched, the pygidial spines themselves would dig into the sediment.
F.) A combat device on males molded by sexual selection mediated by male-male competition for mates. The authors consider this most likely, especially because the tridents resemble the structure of male dynastine (rhinoceros) beetles, which use them to fight for females.
Here’s a picture of three of those beetles which have similar projections as do the Walliserops trilobites (the one at the extreme right).
The authors did a complex morphometric analysis of body and horn shape of W. trifurcatus, comparing it with living rhinoceros beetles to see if the trident could have been used for shoveling/prying, grasping, or fencing—the three types of male-male combat seen in living beetles. The analysis puts the trilobite in the group of living rhinoceros beetles whose males fight by fencing/shoveling: jousting with the structure in front and then trying to shovel the opponent over onto its back. I won’t go into the gory statistical details, which involve principal-components analysis, but the recurved structure of the trilobite’s “trident” is similar to that of shoveling, prying, and fencing beetles (left column: observed means of fighting of living beetles; center: the cephalic structures used; right: the species name [trilobite at the bottom]).
As you see, and as the statistical groupings show, W. trifurcatus is similar to the structures used in rhinoceros beetles for fencing, prying and shoveling. Here is Gishlick and Fortey’s scenario of how the males battled it out in the competition to pass on their genes:
We would hypothesize a fighting scenario in Walliserops similar to that of Trypoxylus. The trilobites would meet and at first spar with their forks, pushing and poking. At some point, they would shift to trying to slide the fork under the other, in an attempt to flip them over. Given the morphology of Walliserops, flipping would be a very effective combat technique. Although the appendages of Walliserops are unknown, it is likely that they were like those of other phacopids in not extending beyond the carapace. This is seen in the Devonian Chotecops, asteropygines Asteropyge, and Rhenops, and recently described in three-dimensional material from the Silurian Dalmanites. Once the trilobite was inverted, righting would not be a simple matter, especially if the dorsally directed spines had snagged in the sediment. An upended trilobite would probably be even more helpless than a beetle in this position and thus excluded from sexual competition.
It might also be dead!
Now the first thing that struck me when I saw this paper was the question that would have occurred to many of you: WHERE ARE THE BLOODY FEMALES?? One of the signs of male-male competition is that the structures used to compete are present in males but almost never in females, as they’re of no use in that sex—and detrimental to fitness if you don’t use them. Male deer have antlers, females do not. Body size, used for combat in elephant seals, is huge in the males, and much, much smaller in females. So if these trilobite horns really were tools used for the “combat” form of sexual selection (the other form, as pointed out by Darwin, is female preference), the females should be around but lack the ornaments. Where are they?
Gislick and Fortey suggest that the females were indeed around, but because they lack the tridents they have not been identified as females of Walliserops trifurcata:
Since the diagnostic synapomorphy [JAC: shared derived trait] for Walliserops is the anterior trident, it would be likely that the female of the species has been classified in a different genus. That leaves two possibilities: either the females of the relevant species are at present unknown, or they are known but placed in another trilobite genus within Asteropyginae.
That mandates a search for trilobites that resemble the males but lack the horns. The authors raise another possibility: the females weren’t preserved or were offstage, living elsewhere, but this seems less likely:
If we extend the beetle analogy further, it is possible that the females are not preserved if some trilobites, like many dynastines, engaged in sex-specific aggregations; in this case, the females were not always present in the same locations as the males, although it is difficult to explain why the latter were selectively caught up in obrution events. [JAC: “Obrution” is rapid burial in the sediments, the way these creatures must have died and been preserved.]
I favor the “females not yet found” hypothesis. There’s one more hypothesis, which is mine: both males and females have tridents. I don’t know why this would be the case, although you could think that it’s used to take other individuals out of action in conspecific competition for food. But that makes little sense.
Finally, the authors found one example of W. trifurcatus with a deformed trident, having an extra spike (a “quadent”?). Here it is on the right. Note that the branching pattern can be asymmetrical in the normal three-pronged structure).
Because the individual on the right was an adult, Gishlick and Fortey suggest that the deformed structure did not prevent the bearer from growing up and thriving, and thus was unlikely to be used for some vital function like feeding. This adds a little more weight to the sexual-selection hypothesis.
The Upshot: The authors’ analyses and explanations seem plausible to me, though they’d be even stronger if they could find the females. That might be tough: in living species you could find them by looking at mating pairs or even seeing that the DNA was nearly identical, but this isn’t possible with fossilized trilobites, especially because in some living and sexually dimorphic species the females look very different from males. If the authors are right, and I think they are, then this quote from the paper is correct:
Walliserops provides the earliest example in the fossil record of combat behavior, very likely ritualized in competition for mates. Although fossil life habits are difficult to prove, the consilience of morphology, teratology, and biometric data all point to the same interpretation, making it one of the more robust examples of paleoecological speculation.
Gishlick, A. D. and R. A. Fortey. 2023. Trilobite tridents demonstrate sexual combat 400 Mya. Proc. Nat. Acad. Sci. USA 120 (4) e2119970120 (in press).