endozoochory

by

Camel Crickets and the Dust Seeds of Parasitic Plants

A common way for plants to disperse their seeds is to entice animals to eat their seed-bearing fruits – a strategy known as endozoochory. Undigested seeds have the potential to travel long distances in the belly of an animal, and when they are finally deposited, a bit of fertilizer joins them. Discussions surrounding this method of seed dispersal usually have birds and mammals playing the starring roles – vertebrates, in other words. But what about invertebrates like insects? Do they have a role to play in transporting seeds within themselves?

Certain insects are absolutely important in the dispersal of seeds, particularly ants. But ants aren’t known to eat fruits and then poop out seeds. Instead they carry seeds to new locations, and some of these seeds go on to grow into new plants. In certain cases there is an elaisome attached to the seed, which is a nutritious treat that ants are particularly interested in eating. Elaisomes or arils have also been known to attract other insects like wasps and crickets, which may then become agents of seed dispersal. But endozoochory in insects, at first, seems unlikely. How would seeds survive not being crushed by an insect’s mandibles or otherwise destroyed in the digestion process?

camel crickets eating fruits of parasitic plants (via New Phytologist)

While observing parasitic plants in Japan, Kenji Suetsugu wanted to know how their seeds were dispersed. Many parasitic plants rely on wind dispersal, thus their seeds are minuscule, dust-like, and often winged. However, the seeds of the plants Suetsugu was observing, while tiny, were housed in fleshy fruits that don’t split open when ripe (i.e. indehiscent). This isn’t particularly unusual as other species of parasitic plants are known to have similar fruits, and Suetsugu was aware of studies that found rodents to be potential seed disperers for one species, birds to be dispersers of another, and even one instance of beetle endozoochory in a parasitic plant with fleshy, indehiscent fruit. With this in mind, he set out to identify the seed dispersers in his study.

Suetsugu observed three achlorophyllous, holoparisitic plants – Yoania amagiensis, Monotropastrum humile, and Phacellanthus tubiflorus. While their lifestyles are similar, they are not at all closely related and represent three different families (Orchidaceae,  Ericaceae, and Orobanchaceae respectively). All of these plants grow very low to the ground in deep shade below the canopy of trees. Air movement is at a minimum at their level, so seed dispersal by wind is not likely to be very effective. Using remote cameras, Suetsugu captured dozens of hours of footage and found camel crickets and ground beetles to be the main consumers of the fruits, with camel crickets being “the most voracious of the invertebrates.” This lead to the next question – did the feces of the fruit-eating camel crickets and ground beetles contain viable seeds?

Monotropastrum humile via wikimedia commons

After collecting a number of fecal pellets from the insects, Suetsugu determined that the seeds of all three species were “not robust enough to withstand mastication by the mandibles of the ground beetles.” On the other hand, the seeds passed through the camel crickets unscathed. A seed viability test confirmed that they were viable. Camel crickets were dispersing intact seeds of all three parasitic plants via their poop. The minuscule size of the seeds as well as their tough seed coat (compared to wind dispersed seeds of similar species) allowed for safe passage through the digestive system of this common ground insect.

In a later study, Suetsugu observed another mycoheterotrophic orchid, Yoania japonica, and also found camel crickets to be a common consumer of its fleshy, indehiscent fruits. Viable seeds were again found in the insect’s frass and were observed germinating in their natural habitat. Seutsugu noted that all of the fruits in his studies consumed by camel crickets are white or translucent, easily accessible to ground dwelling insects, and give off a fermented scent to which insects like camel crickets are known to be attracted. Camel crickets also spend their time foraging in areas suitable for the growth of these plants. All of this suggests co-evolutionary adaptations that have led to camel cricket-mediated seed dispersal.

Yoania japonica via wikimedia commons

Insect endozoochory may be an uncommon phenomenon, but perhaps it’s not as rare as we once presumed. As mentioned above, an instance of endozoochory by a beetle has been reported, as has one by a species of cockroach. Certainly the most well known example involves the wetas of New Zealand, which are large, flightless insects in the same order as grasshoppers and crickets and sometimes referred to as “invertebrate mice.” New Zealand lacks native ground-dwelling mammals, and wetas appear to have taken on the seed dispersal role that such mammals often play.

Where seeds are small enough and seed coats tough enough, insects have the potential to be agents of seed dispersal via ingestion. Further investigation will reveal additional instances where this is the case. Of course, effective seed dispersal means seeds must ultimately find themselves in locations suitable for germination in numbers that maintain healthy populations, which for the dust seeds of parasitic plants is quite specific since they require a host organism to root into. Thus, effective seed dispersal in these scenarios is also worth a more detailed look.

Further Reading:

———————

For more stories of seed dispersal check out the first issue of my new zine, Dispersal Stories.

Advertisement
by

Seed Dispersal by Way of Tree Climbing Goats

Goats are surprisingly good climbers. Given the opportunity, they’ll climb just about anything, including each other. So what’s stopping them from climbing a tree, especially if there is something up there they can eat? And so they do. Tree climbing goats are such a fascinating sight, they even have their own calendar. But the story doesn’t end there. The goats find food in the trees, entertaining humans as they go; meanwhile, the trees have a reliable partner in the goats, who inadvertently help disperse the tree’s seeds.

In general, goats don’t need to climb trees to find food. Goats aren’t known to be picky eaters, and there is usually plenty for them to eat at ground level. However, in arid climates where food can become limited, ascending trees to eat foliage and fruits is a matter of survival. This is the case in southwestern Morocco, where goats can be found in the tops of argan trees every autumn gorging on the fruits of this desert tree.

goats in Argania spinosa via wikimedia commons

Argan (Argania spinosa) is a relatively short tree with a sprawling canopy and thorny branches. It is the only species in its genus and is endemic to parts of Morocco and neighboring Algeria. The tree is economically important to the area due to the oil-rich seeds found within its bitter fruits. Argan oil has a variety of culinary uses and is also used medicinally and in cosmetics. To get to the oil, goats are often employed in harvesting the fruits. The goats retrieve the fruits from the tops of the trees and consume their fleshy outer layer. The hard, seed-containing pits are expelled, collected, and cracked open to get to the seeds.

This is where a team of researchers from Europe come in. There has been some confusion as to how the pits are expelled, with some reports claiming that they pass through the goats digestive track and are deposited in their manure. This is a common way for the seeds of many other plant species to be dispersed, and is carried out not only by goats and other ruminants, but also by a wide variety of mammals, as well as birds and even reptiles. However, considering the average size of the pits (22 mm long x 15 mm wide), the researchers thought this to be unlikely.

fruits of Argania spinosa via wikimedia commons

Others reported that the seeds were spat out in the goats’ cud while they ruminated. Goats, like other ruminants, have stomachs composed of multiple compartments, the first of which being the rumen. Partially digested food, known as cud, is sent back into the mouth from the rumen for further chewing and may be spat out or swallowed again. Goats are known to ruminate in the same location that they defecate, which results in confusion as to when and how certain seeds, like those of the argan tree, are deposited.

By feeding various fruits to a group of goats, the researchers were able to test the hypothesis that seeds could be regurgitated and spat from the cud and that this is a viable method of seed dispersal. The researchers reported that larger seeds were more commonly spat out than smaller seeds, but that “almost any seed could be ejected during, mastication, spat from the cud, digested, or defecated.” The viability of spat out seeds was tested, and over 70% of them were found to be viable.

pits and seeds of Argania spinosa via wikimedia commons

This discovery suggests that seed dispersal via spitting by ruminants could be a common occurrence – possibly far more common than previously considered. The researchers postulate that studies that have only considered seeds dispersed in manure “may have underestimated an important fraction of the total number of dispersed seeds” and that the seeds spat from the cud likely represent different species from those commonly dispersed in dung. In addition, the seeds of some species don’t survive the digestive tract of ruminants, so “spitting from the cud may represent their only, or at least their main, dispersal mechanism.”

This study surrounding the argan trees was followed up by the same group of researchers with a literature review that was published last month. The review looked into all available studies that mentioned seed dispersal via regurgitation by ruminants. While they considered over 1000 papers, only 40 published studies were found to be relevant for the review. From these studies, they determined that the seeds of 48 plant species (representing 21 different families) are dispersed by being spat from a ruminant’s cud, and that most of these plant species are trees and shrubs whose fruits contain large seeds. Also of note is that ruminants across the globe are doing this – representatives from 18 different genera were mentioned in the studies.

ruminating goat via wikimedia commons

The researchers conclude that this is a “neglected” mechanism of seed dispersal. It’s difficult to observe, and in many cases it hasn’t even been considered. Like so many other animals, ruminants can disperse seeds in a variety of ways. Seeds can attach to their fur and be transported wherever they go. They can pass through their digestive track and end up in their dung, potentially far from where they were first consumed. And, as presented here, they can be spat out during rumination. Investigations involving all of these mechanisms and the different plant species involved will allow us to see, in a much clearer way, the role that ruminants play in the dispersal of seeds.