"So, naturalists observe, a flea has smaller fleas that on him prey; and these have smaller still to bite ’em; and so proceed ad infinitum."
- Jonathan Swift

July 11, 2024

Parvatrema spp.

Parasites are known for their complex life cycles, especially among parasitic flatworms such as flukes. And the flukes that are being featured in today's post have life cycles that make them a fluke among flukes. This blog post is about an extensive study that culminated from 25 years of work, where a group of researchers were able to identify and describe five different fluke species with a peculiar life cycle adaptation.

Parvatrema sp. “quadriramis” cercaria stage (left), young metacercaria (centre), parthenogenetic metacercaria containing fully-formed metacercariae (right). Insert: Parvtrema parthenogenetic metacercariae in the hepatopancreas of a limpet.
Photos from Fig. 4 and Fig. 11 of the paper

In the cold waters of the northern European seas and the Sea of Okhotsk, there is a group of parasites with life cycles that defy the conventions of its class. They are five closely related species of flukes in the genus Parvatrema, and they spend parts of their lives lurking quietly in the bodies of clams and snails. In many ways, they're just like other digenean flukes, with multi-host life cycles that involve turning their first mollusc host into a clone factory, producing clonal larvae which go off to infect a second host, and culminating in sexually mature adults living in the gut of vertebrate animals. But these Parvatrema flukes have evolved to do some things differently once they reach their second host.

These flukes infect sea snails such as limpets and periwinkles as their second host. Some species sit in the extrapallial space - the fluid-filled gap between the snail's fleshy mantle and the shell, others get into the gonads and digestive organs. Usually, this is a relatively dormant stage of the fluke life cycle, where they literally sit and wait to be swallowed by the appropriate final host. But with these Parvatrema flukes, instead of simply sitting around waiting to be eaten by a bird like other flukes would, they have another round of asexual reproduction, as a treat.

Each of those immature flukes become filled with numerous miniature clones of itself until it is stuffed to the point of exploding. Some of them take it further, with unlimited consecutive generations of parthenogenetic clones, each fluke exploding into multiple clones and then each of those clones explodes into even more clones and so on, like a never ending series of Matryoshka dolls. On top of that, some of those Parvatrema doing unlimited fluke works are also able to produce cercariae - the free-swimming larval stages - which then go off to infect more sea snails to start the asexual cycle again. 

So why did they evolve this unique developmental stage? Parvatrema are tiny flukes, the adult stage only live for a few days in the gut of a bird, and they produce less than a hundred eggs - a relatively low number compared with other flukes which may produce thousands or even millions of eggs over their lifetime. To make things worse, the likelihood of any one fluke successfully infecting the right hosts at each consecutive stages of its life cycle is astronomically low, so they need to multiply their numbers at every chance they get.

Furthermore, the final hosts for these flukes are migratory birds which only come once a year - so they need to make the most of their brief stay by making sure that if they only eat one infected snail, instead of just getting a single or a dozen flukes in each snail, they're getting the whole gaggle of fluke clones arriving en masse into the bird's gut in their hundreds or thousands, ready to get on with the business of producing the next generation.

Essentially, these Parvatrema flukes recapitulate the process that most other digenean flukes only undergo in the first host. Asexual reproduction in the first host is arguably one of the key evolutionary innovation of digenean flukes, allowing them to offset the losses associated with the process of being transmitted from one host to the next. Since Parvatrema seems to do asexual reproduction at every possible opportunity, they can provide us with insights into how flukes evolved their one weird asexual trick that gave them an edge in the transmission game.

Galaktionov, K. V., Gonchar, A., Postanogova, D., Miroliubov, A., & Bodrov, S. Y. (2024). Parvatrema spp.(Digenea, Gymnophallidae) with parthenogenetic metacercariae: diversity, distribution and host specificity in the Palaearctic. International Journal for Parasitology. 54: 333-355

June 10, 2024

Forficuloecus pezopori

Parasites are a major part of biodiversity, but they spend most of their time hidden in plain sight. Even with some animals that have been known to science for centuries, their parasite fauna remains completely unknown. This can either be due to the lack of research interest into their parasites, or the host animal is just really rare, so there has been very little to no opportunities to study the parasites that live on or in them. In some cases, those rare animals are at risk of going extinct, which means their parasites and symbionts may also disappear before we even know they exist. This post is a story about a ground parrot and its hidden louse.

Left: Forficuloecus pezopori louse viewed under light microscope, Right: a western ground parrot (Pezoporus flaviventris).
Both photos from graphical abstract of the paper

The parasite being featured in this post is Forficuloecus pezopori, and it is the first known parasite from the western ground parrot (Pezoporus flaviventris), also known as Kyloring. Kyloring is one of the rarest parrots in the world and is considered critically endangered, which is bad news for F. pezopori, because we have barely gotten to know this little insect, and it may already be at risk of disappearing along with its feathered host. Lice are particularly vulnerable to co-extinction as they are completely helpless off the host's body and are often specifically adapted to living on just one particular host species.

Forficuloecus pezopori was found on some captive ground parrots at Perth Zoo, the lice were hanging around the feathers at the back of the birds' head and nape. As far we know, the western ground parrot is the only host for this parasite. There is a slim chance that it might also be found on the western ground parrot's closest relative, the eastern ground parrot (Pezoporus wallicus), but we can't verify that at this point, because we know so little about the parasites of ground parrots in Australia.

For example, a subspecies of eastern ground parrot in Tasmania was found to support a type of feather mite called Dubininia pezopori but it is uncertain whether that mite is unique to just that particular subspecies, or if it is also found on the mainland eastern ground parrots, since nothing is actually known about the parasites and symbionts of those birds. Given the eastern and western ground parrot have been separated since the Pleistocene about 2 million years ago, there would have been enough separation in time and space for the two species to develop their own distinct collection of parasites. So as far as we can tell, the Kyloring is the only host for F. pezopori.

Parasites on endangered hosts such as the Kyloring are in a precarious position, because not only are they at risk of dying out alongside their hosts, historically, there have been cases of parasites being wiped out in the process of people trying to conserve their hosts. For example, during the California condor breeding program, a unique species of condor louse was wiped out due to the pesticide-based delousing that the birds received when they were taken into captivity. And the California condor louse is not the only victim of extinction via conservation efforts. The Iberian lynx louse also suffered the same fate

Even more tragic is the case of Rallicola extinctus - a louse of the Huia, which was a species of New Zealand bird that became extinct early in the 20th century, with its last confirmed sighting in 1907. But the louse that it hosted was not even formally described until 1990 - many decades after the host had already gone extinct, hence the species name R. extinctus. Forficuloecus pezopori and many other lice species are at risk of such entangled fates, or become victims of well-meaning conservation efforts.

While a lot of people may not mourn the loss of parasites, it might be their hosts that end up missing them the most. The presence of parasites may help them develop a properly functioning immune system, and their absence could leave the host with a range of physiological disorders. And these parasites might be better off together with their hosts as they can tell us a lot about how the host animals live, and the ecosystem they exist in.

To rectify the mistakes of the past, the researchers suggested that any future studies on wild populations of ground parrots should incorporate a routine louse check to see how common F. pezopori are among those birds, but not to remove any of the lice that are found, and just let those little insects be. Especially since they don't seem to harm healthy, wild parrots. At the same time, lice infection on captive birds can serve as an opportunity to learn more about F. pezopori - saving the host along with its parasites at the same time

Living organisms are intertwined within a network of ecological interactions, if you pull on one loose thread you might trigger a series of co-extinctions and unravel the entire tapestry. Because of those connections, we could be losing more species than we realise.  Though they are often hidden out of sight, and thus out of minds, losing parasites and symbionts would leave us with emptier ecosystems and a lesser world.


May 10, 2024

Dracunculus insignis

The Guinea Worm, Dracunculus medinensis, is an agonising parasite for those who have to endure its wrath. The female worm can grow up to 80 cm long and when it comes time for it to release its offspring, it does so by poking its body partially out of the host's arms or legs, all while causing a fiery pain that forces the host to immerse their limbs into the water, allowing the worm to release its larvae. This parasite has afflicted humans since antiquity, with description of pathologies and treatment associated with the worm dating from ancient Egypt, and depiction of the parasite in a 15th century painting

In the modern era, the Guinea worm has been the subject of an eradication effort by the World Health Organization (WHO) since the 1980s. An obituary was even written about this parasite in 2013. But while this campaign has been largely successful, the effort to completely eradicate the Guinea worm has hit an obstacle in some regions as the worm has taken to using dogs as alternative hosts in place of humans.

Left: Large bundle of Dracunculus insignis in the paws of a river otter (Lontra canadensis), Right: A Dracunculus worm being removed from a river otter (Lontra canadensis)

But aside from the infamous Guinea worm, there are many other species of Dracunculus out there which are found in a wide range of animals, many of which are actually reptiles. Of those, Dracunculus insignis is considered the most important because in addition to parasitising many species of wildlife,
it can also parasitise cats and dogs. The female worm can grow to 30 cm long, and about 300 days after the initial infection, the mature worm - now loaded with larvae - will migrate to the extremities and exit through a lesion, to explosively release a load of baby worms to begin the cycle anew.

This study looked at Dracunculus worms in river otters from North America. The worms the researchers examined came from various sources, including wildlife parasite surveys, as well as dead otters which were obtained from trappers. In addition, they also collected some worms from an otter in Florida that was recovering in a rehabilitation centre after being struck by a car. During its stay in rehab, worms started emerging on their own out of the otter's body. It was just one thing after another for that unlucky otter.

The worms dwelled in swollen abscesses under the skin on the otter's back, and examination of dead otters obtained from trappers revealed that some of the worms were also located in swellings deep in the limb joints or in the otter's paws, particularly D. insignis. In total, the researchers found four different Dracunculus species in the otters - alongside D. insignis, there was also D. lutrae, as well as two other unique lineages of Dracunculus, one of which was first discovered in a Virginia opossum. It seems that otters are just a cornucopia of different Dracunculus species, some of which are currently undiscovered. Just last year, another newly found species of Dracunculus - D. jaguape - was described from neotropical otters (Lontra longicaudis).

Like other Dracunculus, those worms have larvae that stowaway in tiny crustaceans called copepods where they moult and grow. But unlike the Guinea worm which usually infect people when they drink from stagnant water that contains parasitised copepods, in order to get inside otters, the larval Dracunculus would need to take a detour up the food chain into larger aquatic animals such as a fish or amphibians which are on these otters' menu. Incidentally, that is also the suspected route through which the Guinea worm is infecting dogs in places like Chad, because in contrast to how humans drink water, the way that dogs lap water with their tongue means they are unlikely to end up swallowing infected copepods.

While most of the research on Dracunculus have focused specifically on the Guinea worm and its "classical" route of transmission through infected copepods, this has blinded us to the other potential ways that these parasites can circulate in the environment. Understanding how D. insignis and other wildlife-borne Dracunculus complete their life cycles can provide insight into the different ways that these parasites reach their hosts, which in turn can help us better understand how to control the Guinea worm in affected communities.


April 11, 2024

Anoplocephala gorillae

Tapeworms are found in all kinds of vertebrate animals, and while their life cycles and transmission usually rely upon parasitised prey being eaten by predatory final hosts, some tapeworms have evolved ways to infect herbivorous animals as well. Anoplocephala is a genus of tapeworms that parasitise a wide range of herbivorous mammals including elephants, rhinos, hyrax, zebras, and more. The most well-studied species is Anoplocephala perfoliata because it happens to be a parasite of horses, and heavy infection with that tapeworm can cause gastrointestinal diseases. But the species featured in this post are found in a close relative of humans, specifically the Mountain Gorilla (Gorilla beringei beringei), and its name is Anoplocephala gorillae.

Left: Anterior of four Anoplocephala gorillae with their scolices (attachment organ) visible. Right: Proglottids (reproductive segments) of Anoplocephala gorillae collected from faecal samples.
Photos of the parasite from Figure 2 and Figure 4 of the paper

This post is about a study which took place at the Volcanoes National Park (VoNP), in the Rwandan part of the Virunga Massif - a complex of protected areas spanning the borders of Rwanda, Uganda and the Democratic Republic of the Congo (DRC). The aim of the study was to examine the epidemiology of tapeworms in mountain gorillas, and to improve the diagnostic tools for detecting such parasites. To do so, researchers examined faecal samples which were collected by park personnel and Gorilla Doctors veterinarians from groups of habituated gorillas in the VoNP. Whenever possible, each of those samples were identified to specific gorilla individuals, allowing veterinarians to keep track of each gorilla's health and parasite status.

Researchers estimated the abundance of tapeworms in the gorillas by counting the number of tapeworm eggs in each gram of faeces. Generally speaking, more eggs means more worms, but egg production varies between individual worms at different times, so multiple samples needed to be taken to ensure a more accurate count. Out of the 1500 samples they examined, about seven percent had egg counts of over a thousand eggs per gram of faeces, though the average was much lower at 384 eggs per gram. While A. gorillae seems to dominate the tapeworm fauna of these gorillas, the faeces of one gorilla, an infant male named Inkingi, also had another tapeworm species in the genus Bertiella. It is relatively easy to distinguish the eggs from those two different tapeworms - Anoplocephala has quadrangular or triangular-shaped eggs with flat sides and thick shells, whereas Bertiella has spherical eggs with thin shells

In addition to those faecal samples, any gorillas that had died were retrieved from the wild and necropsied as a part of the local veterinary surveillance program. For the purpose of this study, five deceased gorillas that were recovered between 2015–2018 were necropsied and examined for tapeworms. In total, 53 A. gorillae tapeworms were collected, and they varied in size from 1.5 to 13 cm long. Most of them were found in the small intestine, but there were also some in the caecum and colon.

So how do the gorillas end up with all those tapeworms in the first place? While the eggs are released into the environment packaged in the gorilla's faeces, they cannot infect the gorillas directly. Like other tapeworms, they have to go through an intermediate host, which as mentioned earlier, is usually a prey animal. But since gorillas are herbivores, how can tapeworms gain entry into their guts? 

Based on what is known for other Anoplocephalidae tapeworms, gorillas become infected by swallowing mites that are parasitised by the tapeworm's larvae. These mites are tiny, barely pinhead-size, thus can be easily swallowed among a mouthful of foliage. While the prevalence of Anoplocephala among mites might be extremely low, like other herbivorous mammals, gorillas go through a lot of plant matter, eating 18-45 kilograms of vegetation a day. So just a few infected mite sprinkled in would be enough to ensure that the gorillas get infected,

While the deceased gorillas that were necropsied in this study had large numbers of tapeworms dwelling in their gut, they were all in good condition, and had died from other sources of trauma rather than disease. So in contrast to A. perfoliata which can cause major pathologies in horses, A. gorillae is content with a more peaceful existence, just living quietly as a part of the gorilla's regular gut symbiont fauna.


March 9, 2024

Veneriserva pygoclava

There are many ways to become a parasite, and there are parasites with vastly different ancestries that end up joining the same path on the road of parasitism. In some cases, sharing the same path can also mean adopting a certain shape. This post is about Veneriserva pygoclava, a worm that lives inside a worm, more specifically it is a polychaete worm that has evolved to parasitise another type of polychaete worm which are commonly called "sea mice".

Top left: Ventral view of an infected Aphrodita longipalpa with a Veneriserva pygoclava parasite inside. Bottom Left: MicroCT scan image of an Aphrodita longipalpa with Veneriserva pygoclava female highlighted in yellow and juvenile highlighted in blue. Right: A female Veneriserva pygoclava (top) and a male (bottom).
Photos from Fig. 1 and Fig. 3 of the paper

The genus name of this parasitic polychaete translates into "Venus' servant" though this worm is certainly a servant for nobody but itself. You'd think that living inside the body of another animal would restrict how big it can get, but the female Veneriserva grows to about seven centimetres long, which is twice as long as its host. Surprisingly enough, being longer than the host is not unusual among these kinds of parasitic polychaete worms. Despite its size and the amount of space it occupies within the host, it does not seem to cause any injuries or damage to the host's internal organs.

Living this endoparasitic lifestyle requires some specialised adaptations, and over the course of its evolution, Veneriserva has ended up with a body plan which is very similar to that of tapeworms. Despite both being called "worms", tapeworm and polychaete worms are from entirely separate animal phyla and their path to this "tapeworm body plan" (for the lack of a better term) were very different.

Tapeworms evolved from free-living flatworms, which are fairly simple animals, at least in terms of their body plan. A flatworm has no body cavity, its gut is more or less a blind-end sac (with some branches in larger flatworms), and it doesn't even have a circulatory system. If anything, in order to adapt to a parasitic lifestyle, tapeworms have evolved to become more complex than their free-living ancestors. Over the course of the tapeworm's evolution, they have gained a new attachment organ - the scolex - which is a heavily modified head, while the rest of the body has become an efficient conveyor belt of reproductive organs. These parasitic flatworms have even evolved a brand new type of "skin" called a tegument which allows it to absorb nutrients as well as protect itself against the host's enzymes, and some tapeworms even have the most complex central nervous system among all the flatworms, enabling them to navigate and maneuver in the dark, fleshy tunnels that are their host's intestinal tract.

In contrast, polychaetes are segmented worms, and are actually more similar to us in their body plan, equipped with a full body cavity, muscular gastrointestinal tract, and a closed circulatory system with blood vessels. But Veneriserva has abandoned much of that, because when you're living inside another animal, being built like a tapeworm seems to be the way to go.

Veneriserva does have a mouth of sorts, but it is not connected to any digestive tract to speak of. In fact, the digestive tract has been reduced down to a throat with a blind-end. Instead, the mouth of Veneriserva serves as a grabber to hold the parasite in place, functioning much like a tapeworm's scolex. Additionally, Veneriserva has also evolved its own version of the tapeworm's tegument, which is covered in fine microscopic finger-like projects (rather like the lining of your small intestine, just inside out) allowing it to absorb nutrients through its skin. There are also patches of cilia on the skin which may serve to stir the host's bodily fluids in order to bring more nutrients into contact with the parasite's skin.

However, when it comes to sex, there is one key difference between Veneriserva and tapeworms. Tapeworms are hermaphroditic - any tapeworm can mate with any other individual of the same species, or even with itself if it is desperate and alone. In contrast Veneriserva have separate female and male sexes which are clearly distinguishable - male worms are tiny compared to their much larger partners (see accompanying photo).

This "attachment organ + loads of gonads" type of body plan that tapeworms and Veneriserva have both independently evolved is also found in other internal parasites. For example, acanthocephalans - thorny-headed worms - are parasitic worms which live in the gastrointestinal tract of vertebrate animals, and are somewhat related to rotifers. Despite being in a different phylum, they share some key anatomical similarities to tapeworms, with their own version of the tegument, a body dominated by gonads, and a prickly anchor at its "head" to stay attached to the host's intestinal wall. Another example is Thyonicola, the parasitic snail which uses a thin stalk to attach itself to the intestines of its sea cucumber host, while the rest of the body is simply a long tube of reproductive organs and developing eggs. There are even some parasitic dinoflagellates that have evolved to resemble tapeworms. 

Judging from how common this "tapeworm-style" anatomy is across different parasite groups, it seems that when you are an internal parasite, you have to get into shape - and that shape happens to be that of a tapeworm.


February 12, 2024

Ascarophis globuligera

To land-dwelling humans, deep sea hydrothermal vents would seem like a vision of hell, amidst the crushing darkness you have plumes of superheated water, mixed with noxious sulfides, erupting from fissures on the seafloor. But for many deep sea animals, this "hell" is in fact a vibrant oasis in the middle of the abyss. This lively habitat is made possible thanks to bacteria that are able to extract energy from the sulphurous waters billowing from those vents. In the absence of sunlight, these chemoautotrophs form the foundation of the food chain. Some tube worms have been able to co-opt the power of these bacteria by housing the microbes in their gills, enabling them to grow to enormous sizes. Their tubes form dense, forest-like habitats for many other animals including other polychaete worms, fishes, crustaceans, and molluscs. This sets the stage for all kinds of complex ecological interactions, and that includes parasitism.

Left: Anterior of Ascarophis globuligera from Fig. 6 of the paper, Right: Photo of Thermarces cerberus (pink vent fish) by Dr Lauren Dykman, used with permission 

This post is about a paper reporting on three newly described species of Ascarophis nematodes that have been found in the guts of some deep sea hydrothermal vent fishes. Some of those worms were collected as a part of a larger study which focused on looking for parasites from hydrothermal vent animals, and along with freshly caught specimens, the researchers also examined preserved fishes collected by past expeditions. While they only managed to recover a few specimens of Ascarophis nematodes, some of which were fragmentary, those were enough to provide a scientific description for three different species - A. justinei, A. globuligera, and A. monofilamentosa.

The three species differed slightly in which fish species they infect - A. justinei is found in both the pink vent fish and a species of viviparous brotula, whereas A. globuligera has only been found in the pink vent fish, and A. monofilamentosa lives in a species of zoarchid fish named Pyrolycus manusanus. While it is not possible to conduct experimental infections to work out exactly how these nematodes transmit between hosts, their life cycles can be inferred based on what is known about other Ascarophis species which are found in shallower waters. This usually involves a crustacean, often amphipods, serving as the intermediate host for the parasite's larvae. Amphipods are plentiful around hydrothermal vents, and these crustaceans are eaten by a range of animals including deep sea fishes such as the pink vent fish, making them the ideal vehicle for Ascarophis to complete its life cycle.

The need for Ascarophis to reach an amphipod host may explain why each of those Ascarophis species has differently shaped eggs. For example, A. justinei has eggs which are regular, ovoid shape rather similar to other known species of Ascarophis, but the eggs of A. globuligera have a bulge on their side (which gave the species its name), and A. monofilamentosa eggs have a long filament dangling from them which is about fifteen times the length of the egg itself. These differently shaped eggs could mean slightly different transmission strategies. The extra ornament on the eggs of A. globuligera might serve to entice a hungry amphipod by resembling something edible (as with some tapeworm eggs that infect crustaceans by mimicking diatoms), or in the case of A. monofilamentosa, its long filament may prevent the eggs from drifting away into the empty abyss by wrapping them around a structure, or entangle them around something which might get eaten by an amphipod.

Some Ascarophis species are actually known to take a shortcut with their life-cycles -  instead of waiting for a fish host to come along, they become sexually mature and start laying eggs inside the amphipod, bypassing the need to enter a fish host to complete their life cycles. It is unknown whether any of the newly described deep sea species are capable of doing this, but in an ephemeral habitat like hydrothermal vents, it would be useful to have such an option as insurance.

There are many biomes on this planet which are completely inhospitable to humans. But that does not stop them from being as rich and vibrant as those that we are more familiar with, and wherever there is a thriving ecosystem, you will find parasites taking part in its web of interactions.

Moravec, F., Dykman, L. N., & Davis, D. B. (2024). Three new species of Ascarophis van Beneden, 1871 (Nematoda: Cystidicolidae) from deep-sea hydrothermal vent fishes of the Pacific Ocean. Systematic Parasitology 101: 2.

January 7, 2024

Prosthogonimus cuneatus

Prosthogonimus is a genus of flukes that live in a special part of a bird's anatomy. It is usually found either in the Bursa of Fabricius, an organ that only birds have, or in the oviduct, and it's this latter location which lend this parasite its common name, the oviduct fluke. This fluke is found all over the world in many different species of birds, and while it doesn't seem to cause much issues for wild birds, it presents a major problem for the poultry industry.

Left: Dragonflies Sympetrum vulgatum (top) and Sympetrum depressiusculum (bottom), Right: A metacercaria cyst of Prosthogonimus cuneatus. Photos from Fig. 2 and Fig. 3 of the paper

Since this fluke lives by clinging to and feeding on the surface of mucosal membranes, its activities can leave lesions and cause inflammations, and heavy infection of Prosthogonimus can lead to all kinds of oviduct disorders in chickens. This includes leaking milky discharges from the cloaca, laying soft-shelled or malformed eggs, or even egg peritonitis, where egg yolk material gets displaced into the hen's body cavity, leading to secondary infections and death. In some cases, the fluke can even end up getting bundled into the egg itself, which seems pretty mild compared with what I have mentioned above, but it would nevertheless be a nasty surprise for anyone looking to make an omelette. To make matters worse, there are currently no effective treatments available for getting rid of this fluke once a bird is infected.

So how do birds end up with this peculiar parasite? Prosthogonimus has a multi-host life cycle that takes it across three very different animals - freshwater snails, dragonflies, and birds. In the dragonfly, the larval Prosthogonimus lies in wait as a dormant cyst called a metacercaria, waiting for its host to get eaten by a bird. That is why this parasite is usually associated with free-range chickens, as they have more opportunity to feed on a variety of things. Most studies on Prosthogonimus have focused on the effects it has on the bird hosts, but surprisingly fewer studies have investigated the source of the infection - parasitised dragonflies.

To rectify this oversight, a group of researchers undertook a truly herculean effort to investigate the presence of Prosthogonimus in dragonflies from the Heilongjiang province, China. The researchers collected over TEN THOUSAND dragonflies, composed of 12 different species from 41 locations. They identified each of the dragonflies before dissecting them for Prosthogonimus metacercariae, which are usually located in the abdominal muscles. The researchers noticed that infected dragonflies tend to have softer abdominal muscles, possibly due to injuries caused by the presence of the Prosthogonimus cysts.

They found three different species of Prosthogonimus in those dragonflies, of which Prosthogonimus cuneatus was the most common. Overall, about 20% of the dragonflies they examined were infected by Prosthogonimus, but it was more common in some species than others. The spotted darter (Sympetrum depressiusculum) was most frequently infected (28.53% prevalence), followed closely by the vagrant darter (Sympetrum vulgatum) (27.86% prevalence) and the autumn darter (Sympetrum frequens) (20.99% prevalence). The highest number of fluke larvae in a single dragonfly goes to an unlucky Sympetrum kunckeli which was packed with 157 Prosthogonimus metacercariae in its abdomen.

But dragonflies are aerial predators - how do they end up being infected with fluke larvae which are shed from freshwater snails? Well, before becoming acrobatic flying hunters, dragonflies spend their early life as underwater predators. But this aquatic life also expose them to Prosthogonimus' waterborne larvae, which are drawn into the dragonfly nymph's body through its respiratory current - in other words, they get sucked through the dragonfly nymph's butt whenever it takes a breath. Even as the dragonflies metamorphose into airborne adults, they carry the legacy from their youth in the form of Prosthogonimus cysts

Overall, the study found that Prosthogonimus was most common in Heihe, which might be due to the presence of large wetlands in the area. Those wetlands are home to high levels of biodiversity which help support the life cycle of this parasite - they provide habitats for numerous snails that can host the asexual stage of Prosthogonimus, along with wild birds that would usually act as the final host for this parasite. Just add dragonflies, which are always common around water bodies, and the circle of life is complete for Prosthogonimus.

Studying and elucidating the life cycles and ecological role of parasites in their natural context is an important part of disease ecology research. Understanding what these parasites actually do in nature can help us prevent them from causing problems in the animals that we raise.

Li, B., Lan, Z., Guo, X. R., Zhang, A. H., Wei, W., Li, Y., Jin, Z. H., Gao, Z. Y., Zhang, X. G., Li, B., Gao, J. F., & Wang, C. R. (2023). Survey of the Prosthogonimus spp. metacercariae infection in the second intermediate host dragonfly in Heilongjiang Province, China. Parasitology Research 122: 2859-2870.

December 14, 2023

Euglenaformis parasitica

This parasite is invisible to the naked eye, can kill its host in 3 days, and it can be found lurking in the waters of rice fields. What I have just described may sound like a nightmare pathogen from a b-horror movie, but it is actually a microscopic flagellated protozoan that has given up a solar-powered life for one fuelled by the blood of its victims. The name of this microscopic monster is Euglenaformis parasitica, and it belongs to a group of otherwise innocuous single-celled critters called Euglenids.

Left: Scanning electron micrograph of Euglenaformis parasitica. Top Right: E. parasitica extracted from an ostracod host. Bottom Right: E. parasitica visible in the appendage of an infected ostracod.
Photos from Fig. 1 and Fig. 7 of the paper

Euglenids are single-celled flagellated organisms often found in freshwater. The most well-known and well-studied genus is Euglena, which is literally the textbook example of the group, appearing in many biology books as an example of a single-celled eukaryote organism. Many euglenids are photosynthetic, and historically they have been treated as sharing affinity with plants (due to their photosynthetic capabilities) or with animals (due to their active flagellum and being able to take in nutrient via heterotrophy), before getting shunted into a group called the "protist" which is just a jumble of different organisms that scientists couldn't classify into plants, animals, or fungi.

Euglena and its kin are mostly free-living, photosynthesizing when the sun's out, absorbing organic matter from the environment when it's dark. But the ancestor of E. parasitica was not content with this mostly peaceful lifestyle, and has evolved to live inside the body of animals. These euglenids were found parasitising ostracods (also known as seed shrimps) and flatworms in a rice field in Ibaraki, Japan.  Ostracods and flatworms belong to two entirely different phyla of animals, and most parasites that infect different phyla of host animals do so at different stages of their complex, multi-stage life cycles. But E. parasitica has just a simple life cycle, which makes it quite remarkable that it is able to adapt to the very different internal environments presented by ostracods and flatworms.

When E. parasitica is in an ostracod, it dwells in the body cavity, swimming in the hemolymph and bathing in its nutrients. Whereas in flatworms, since they don't have any body cavities to speak of, E. parasitica lives in the space between the spongy tissue that forms the bulk of a flatworm's body, burrowing between the cells of the parenchymal tissue. But whether it is in an ostracod or a flatworm, once E. parasitica establishes itself in the host's body, it starts absorbing the literal lifeblood of their host, using it to fuel its exponential growth as it divides and conquers from within. 

What started with just a single or a few E. parasitica soon turns into a swarm. This is particularly noticeable in flatworms - uninfected flatworms are semi-translucent, but infected flatworms darken in colour as their body becomes filled with brownish to blackish granules which are actually rapidly dividing E. parasitica. The same goes for ostracods as their blood becomes saturated with the parasite's progenies. After three days, the insides of the host are completely consumed by the swarm of E. parasitica, which proceeds to exit into the surrounding water, leaving behind an empty husk.  

There are still a lot of mysteries surrounding this flagellated organism, such as how it is able to make use of such radically different hosts as flatworms and seed shrimps, or how it enters the host's body in the first place. Does it somehow bore through the body wall, or perhaps it tricks the host into eating it, and then burrows through the digestive tract to other parts of the body? If so, it won't be the only parasite to use that trick. There are also questions about its evolutionary origin. Euglenaformis parasitica's close relatives are photosynthetic euglenids, so what made it abandon a solar-powered life in favour of living and reproducing in the bodies of small aquatic animals? Understanding that process would provide us with another clue as to how various different organisms ending up following the path of parasitism.

Kato, K., Yahata, K., & Nakayama, T. (2023). Taxonomy of a New Parasitic Euglenid, Euglenaformis parasitica sp. nov.(Euglenales, Euglenaceae) in Ostracods and Rhabdocoels. Protist 174: 125967.

November 14, 2023

Stylops ater

Strepisptera is an order of parasitic insects with some very unique characteristics.They are also known as twisted wing parasites, based on the twisted hindwings on the male parasite. They infect many different orders of insects, but mostly target wasps and bees where they up take up residency in the host's abdomen. If you know what to look for, you can immediately spot their presence. In fact, there's even a special term for describing bees and wasps that are parasitised - they get "stylopized".

Top: A male Stylops ater (indicated by red arrow) attempting to mate with a female in a bee's abdomen.
Bottom left: Female Stylops ater adult (indicated by red arrow) in a bee's abdomen,
Bottom right: Male Stylops ater pupa casing (indicated by red arrow) in a bee's abdomen. 
From Fig. 1 of the paper

And it's not just the hindwings of stresipterans that are a bit twisted, these insects have extreme sexual dimorphism, so much so that if you didn't know any better, you'd think the females and males are completely different types of animals. The female stresipteran looks like a grub, and she spends her entire life inside the abdomen of the host, with just her head partially poking out from between the segments of the host's abdomen.

In contrast, the males have a pair of giant compound eyes, prominent branched antennae and the "twisted wings" that give this group of insects its name. They have a short and frantic adulthood - after emerging from the host, he only lives for a few hours and his sole mission in life is to find and mate with an elusive female strepisteran, hidden away in the abdomen of a host insect. And he does the deed with an appendage that entomologist Tom Houslay once vividly called a "stabby cock dagger". The technical term for this form of mating is "hypodermic insemination" - where the male basically stabs and inject his sperm into the female, and the sperm somehow find their way to the eggs. Strespiterans are not alone in having this type of appendage, male bed bugs also have a stabby cock dagger - but that's another story.

The study being featured in this post focuses on Stylops ater, a species which parasitises Andrena vaga, the grey-backed mining bee. Unlike the honeybees that most people are familiar with, these are solitary bees, with no castes. And while they do gather into an aggregation to nest, each bee just builds and looks after their own nest. The researchers examined a population of these bees in Lower Saxony, Germany. They sampled over two periods, during late winter, when all 508 bees they looked at were stylopized, and late spring, when they only managed to find two stylopized bees out of a total of 150.

Almost two-third of the stylopized bees were female, but these parasites seem to prefer hosts that are of the same sex as themselves. Since female bees live longer and can provide more nutrients than male bees, this works out well for the life history of female Stylops as it gives them more time and nutrients to grow her brood. After mating, the female Stylops can release up to 7000 offspring, which crawl off to find other bees. While each larva is merely 0.2 millimetre long, they can traverse long distances by hitching a ride on the hair, pollen sacks, or even the crop of bees, to end up in a new bee nest, filled with fresh hosts.

While most bees only hosted a single parasite, some had two or three, and the researchers did find one very unlucky bee that was harbouring four Stylops in its abdomen. But even a single Stylops can take a severe toll on its host. In fact, this parasite is so demanding that it wouldn't grow as big if it had to share its host with another Stylops. As a result, bees infected with Stylops are unable to develop eggs or only produce poorly developed eggs.

But aside from effectively sterilising the bee, Stylops also tinkers its host's biological clock, making it emerge out of hibernation a few weeks earlier than uninfected bees - hence why the researchers found so many stylopized bees in late winter. Making the bees such early risers ensures that there will be plenty of female Stylops around for the male Stylops to find, which will be emerging at that time to live out their extremely short lives. It also gives the female Stylops' larvae more time to develop, so they will be able to crawl off in time to find new hosts in the bee's brood cells. This type of behaviour manipulation is comparable to what's found in Sphaerularia vespae, a nematode that alters the seasonal biological clock of hornet queens.

In order to make these changes to the bee's internal clock, Stylops would have to manipulate the host's hormones, but this also results in some side effects on the bee's body. Female bees that get stylopized tend to have a hairier back, skinnier legs, and the hairs on said legs become shorter and more sparse. In short, they take on characteristics that are more similar to that of regular male bees.

So next time you are out and about, keep an eye out for a bee that looks a bit different from the rest. It might be flying under the influence of a parasite tucked away in its abdomen, looking to make a rendezvous with her short-lived partner.


October 10, 2023

Atriophallophorus winterbourni

In Lake Alexandrina of New Zealand lives a species of tiny freshwater snail called Potamopyrgus antipodarum. These snails are capable of alternating between sexual and asexual reproduction and can be extremely abundant. So much so that they have become invasive in many other parts of the world. Outside of their original home, they are free to proliferate to their heart's content. But back in New Zealand, these snails don't always have things go their way. They are held back by a whole menagerie of flukes which parasitise them - at least 20 different species in fact.

Top: Photo of the snail Potamopyrgus antipodarum by Michal Maňas, used under Creative Commons (CC BY-SA 4.0) license. Bottom: The metacercariae cysts of Atriophallophorus winterbourni, from Figure 1 of the paper.

These flukes have a range of different life cycles, but all of them use P. antipodarum as a site of asexual reproduction - converting the snail's insides into a clone factory and rendering it sterile in the process. These flukes might be the reason why these snails continue to engage in sex every now and then, despite asexual reproduction being much more efficient. Sex is necessary to maintain genetic variations - the key ingredient in the evolutionary arms race against all those flukes.

Researchers who have been studying these snails and their flukes noticed that while all 20 species of those parasite are essentially body snatchers that take over the insides of their unwitting host, one species - Atriophallophorus winterbourni - goes beyond simply messing with their host's physiology and seems to be influencing the snail's behaviour too. Snails infected with A. winterbourni tend to be found in the shallow areas of the lake. Among snails collected from the shallow water margin of the lake, they represent 95% of the infections. Is this because those areas just happen to be hot spots for snails to get infected with A. winterbourni? Or are these flukes actually coaxing the snails into hanging out in the shallows?

To figure out if there's something special about A. winterbourni, researchers compared snails infected with A. winterbourni with those that were infected with a different species of fluke - Notocotylus - to see if such behavioural change is simply a side-effect of fluke infection, or if it is something specific to A. winterbourni. The researchers did this by setting up a series of ten 5 metres long tubular mesh cages that stretched across different depth clines in the lake, from less than 0.8 metres at the shallow end to 2.8 metres at the deep end, with different sections of the cage corresponding to different levels of water depth. Using snails collected from two high infection prevalence sites at the lake, they added about 800 snails to the deepest section of each cage, and the snails were allowed to freely roam between the different sections. After eleven days, samples of snails were randomly collected from each depth level and examined for parasites.

There are some key differences in the life cycles of A. winterbourni and Notocotylus that makes them good for comparisons. Just like other flukes, A. winterbourni undergoes asexual reproduction inside the snail host, producing a whole load of clonal larvae. But unlike many other flukes, these clonal larvae stay in the snail and transform into cysts, where they wait to be eaten by a duck hungry for snails. In contrast, snails that are infected with Notocotylus release those clonal larvae into the surrounding waters, and they do so continuously over the course of about 8 months. These larvae attach themselves to vegetation or the shells of other snails, and are transmitted to grazing ducks that accidentally ingest them. Therefore, unlike A. winterbourni, their transmission is largely decoupled from the snail's own movement and behaviour.

So after those eleven days of allowing infected snails to roam in the cages, what did the researchers find? Well, snails infected with A. winterbourni were heavily distributed towards the shallow end, with over a third of the snails in that section being infected, which is over three times higher than the expected background infection level (11%). In the deepest section of the cage, A. winterbourni-infected snails were rare, representing only 3-5% of the snails in that section, and some of them were immature infections. In contrast, those infected with Notocotylus were found to have distributed themselves fairly evenly across the entire depth cline. It is unclear what exactly A. winterbourni is doing to the snails that makes them favour shallow water, but more importantly, why would they do this? What's in it for the fluke? Well, the final hosts for A. winterbourni are dabbling ducks that only feed in the shallow parts of the lake. So in order for A. winterbourni to make a successful rendezvous with its final host and complete its life cycle, it will have to prod its snail host into the shallows.

Atriophallophorus winterbourni belong to a family of flukes called Microphallidae, and there are a few other species in this family which are also known host manipulators. For example, Gynaecotyla adunca is a species that infects marine mudsnails, and it coaxes its mudsnail host into stranding itself onto beaches, which brings them closer to the crustaceans that serve as the next host in the parasite's life cycle. There's also Microphallus papillorobustus, which infects little sand shrimps (amphipods), and it alters their behaviour in a number of different ways that makes them more visible to hungry birds. Even though not all members of Microphallidae are host manipulators, it's a trait that does seem pretty common in this fluke family. Sometimes, in order to complete a life cycle, you just have to drag that snail to where you need it to be.