"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

January 24, 2016

Artystone trysibia

The tongue-biter Cymothoa exigua is arguable one of the most (in)famous fish parasite in the world. It was famous enough to get a mention on the Colbert Report, and while the world recoil in collective horror at the sight of a fish which had its tongue replaced by a parasite, among its fellow parasitic crustaceans, tongue-biter's modus operandi is actually rather quaint. It can easily be upstaged by other parasitic isopods in the horror department, and today's post is about one those species.

The parasite we are featuring today is Artystone trysibia - it is in the same taxonomic group as the tongue-biter (Cymothoidae), and it parasitises a number of freshwater fish in the Amazonian basin. But unlike its more famous cousin which is content with merely living in the host's mouth, A. trysibia cranks the nightmare fuel up to eleven and lives inside a fleshy capsule in the host's body cavity.
Photo from Figure 2 of this paper
This parasite and others like it are actually relatively common. This parasite has been documented from a range of freshwater fish from South America, and has also been reported from aquarium ornamental fish. They're so common that A. trysibia has gained a common name - "ghili" - among the Kichwa people.

Female A. trysibia (right) and Female A. trysibia with larvae
Photo from Figure 3 of this paper
This study documented its presence in the Bristlemouth Armoured Catfish (Chaetostoma dermorhynchum). Despite its armouring, this fish has no protection against A. trysibia. Usually, the only sign of the parasite's presence is a small, gaping hole on their belly or their flank. But that hole serves as a window for the parasite within. For this study, these catfish were sampled from three pristine sites at the Tena River in the Amazonian region of Ecuador.

These catfish are fairly small fish, and most of them are about 15 - 20 cm long (6-8 inches), but A. trysibia can grow to 1.5 - 3 cm (0.6-1.2 inches) long and takes up quite a lot of space within the catfish. For comparison, it would be the equivalent of having something the size of a pet rabbit living in your torso. As mentioned above, the only contact the parasite has with the outside world is with a tiny hole through which they breath and release their offspring, and they can reproduce in prodigious numbers - one female isopod was recorded to be carrying 828 larvae. Each catfish was found to (thankfully?) only ever have a single A. trysibia, and it seems that the bigger the host, the bigger the parasite, possibly because a larger host would give the parasite more room to grow.

Artystone trysibia is not alone in its style of parasitism, there are other similar species which are found in both freshwater and marine fish around the world (for example, see here). So the next fish which you come across might not just have a tongue-biter in its mouth, it might also have a ghili in its belly.

Junoy, J. (2016). Parasitism of the isopod Artystone trysibia in the fish Chaetostoma dermorhynchum from the Tena River (Amazonian region, Ecuador). Acta Tropica 153, 36-45.

January 10, 2016

Cardicola orientalis

Tunas are one of the most graceful animals of the sea. These sleek and powerful predators spend their lives in motion, cruising the open seas for prey. But despite being such formidable fast movers, this does not make them immune to parasitic burdens, indeed the parasite we are featuring today are found in the heart these pelagic predators.

Scientists examining Pacific bluefin tuna (Thunnus orientalis) at a tuna ranch at the Wakayama prefecture, Japan came across some unfamiliar-looking flukes living in the heart of the tunas, which they described and named Cardicola orientalis in this study. Aside from feeding on the tuna's blood, the eggs that these parasites produce can become lodged in various tissues, obstructing blood vessels and causing harmful lesions and inflammations. In fact, these wayward eggs are more debilitating to the host than the adult parasite itself.
Photos of Cardicola orientalis from this paper
If you think the fish host have it bad with these parasites, the invertebrate host have it much worse. These parasites have a complex lifecycle that alternates between a vertebrate and an invertebrate host. The adult parasite reproduce sexually in the fish host but the eggs that they produce (if it they don't get lodged in somewherein the host) are released into the environment and hatch into a larva call a miracidium that infects mollusc or polychaete worm (depending on the fluke species). Inside the invertebrate host, the fluke larva transforms into a sausage-shape stage call a sporocyst, which then multiply via cloning inside their body, turning them into a parasite factory (see photo above).

This newly-commissioned biological factory then churn out another larval form of the parasite, called cercariae, which are shed into the surrounding waters where the fish hosts are found. So if you want to stop the tuna from getting infected with blood flukes like C. orientalis, you have to figure out which invertebrate is acting as the parasite factory in the fluke's lifecycle.

Of the 136 known species of fish blood flukes, the full lifecycle is known for a handful of them. Because they are one of the few fluke species that can severely impair or even kill their fish host, fish blood flukes are a major concern to the aquaculture industry. Considering the number of marine invertebrates that can serve as potential host for C. orientalis, it would seem that these scientists had a pretty difficult task at hand. However, based on previously documented lifecycles for tuna blood flukes, they are somewhat different from those other fish blood flukes in that instead of using snail or a bivalve for their clonal stage, they use polychaete worms. Specifically they use a family of worm call the terebellids - also known as spaghetti worms - which live in burrows and crevices.

The research team found many such worms encrusted on the structure of the tuna cage, alongside other invertebrates such as sponge, seashells, and sea squirts. The most abundant species was a marine worm call Nicolea gracilibranchis. They took monthly sample of these worms from the tuna cages from January to May, dissecting 4729 worms in total and finding 349 to be infected with the clonal stage of C. orientalis. Even though the researchers found that most of those worms were living on the floats that surround the tuna cages, it was the worms encrusted on the ropes which held the cage in place that are more likely to be infected with the parasite's clonal stages.

They also noted that infected hosts became more common over course of the sampling period, and while the worms they dissected in January were mostly filled with developing parasite embryos, those sampled after February were ripe with cercariae ready to pop. These pattern seems to indicate that the worms become infected through eggs that were expelled from tunas during winter and the parasite larvae developed over spring.

Since tuna has a reputation for being a fast swimming fish, you'd think their parasites would be equally well-equipped for swimming. But instead, C. orientalis has a tiny stub of a tail which doesn't appear to be good for swimming (or much else for that matter). But somehow, they must be getting to the tuna just fine; either the infected spaghetti worms churn out so much cercariae that at least some manage to encounter their host, or they have some other adaptation that facilitates their rendezvous with a tuna. Or both.

The research team also came across one case of a different tuna blood fluke species - Cardicola fosteri - which has previously been found in Australia and was featured in a post on this blog from 2011. It is worth noting that while in Australia, that parasite infects the worm Longicarpus modestus and the southern bluefin tuna (Thunnus maccoyii), in Japanese water they were infecting a different species of terebellid worm (Amphitrites sp.) and tuna for their lifecycle. So is this ability to switch host common to all fish blood flukes, or is it just this particular group of tuna blood flukes?

This flexibility in host use would be an extremely useful adaptation, especially for a parasite like C. orientalis since its host is an open water animal which is widely distributed across the world's oceans. But this can also be a concern for fish farmers as fish species introduced for aquaculture may exchange parasites with wild fish native to a particular region. As the aquaculture industry incorporate more species to their stock, novel and/or poorly described species will emerge as new problems. The lesson here is that if you are going to farm fish, you better be prepared to come across some flukes.

Shirakashi, S., Tani, K., Ishimaru, K., Shin, S. P., Honryo, T., & Ogawa, K. (2016). Discovery of intermediate hosts for two species of blood flukes Cardicola orientalis and Cardicola forsteri (Trematoda: Aporocotylidae) infecting Pacific bluefin tuna in Japan. Parasitology international 65: 128-136.

December 27, 2015

The Worms of Beasts, Tragic Romances, and Body-Snatchers

It has been yet another year of parasitology, and this year has been my fifth year writing on a regular basis for Parasite of the Day! So what had been on the parasite menu for 2015?  First of all, some of the parasites that made their way onto the blog this year have been various worms that cause misery for everyone's favourite large mammals like dolphins, pandawhales, and baboons. But it is not just large mammals that become unwitting host for parasites, for example, the giant ocean sunfish is also host to a fluke that surrounds itself in a bag of the host's flesh

If that all sounds very snug and cosy, then one might see that some of the posts can be described as love stories, though most of them with a tragic or unsavoury twists. There was a post about treacherous journey undertaken by male pea crabs to answer a booty call, a guest post by Katie O'Dwyer about sexually transmitted infection in ladybirds, and a story of how cicadas' love songs can end in tragedy (chest-burster style).

On the subject of body-snatchers, nature certainly has no shortage of them, and insects are usually the victims - in one case, tapeworms for ants which also seem to affect the behaviour of the host's uninfected nest mates. These body-snatchers also seem to get around as well, one of these well-worn travellers is a species of roundworm that was introduced to New Zealand from Europe via earwigs. That worm is a mermithid nematodes, but its lifecycle is remarkably similar to that of another phylum of worms - the nematomorphs. More commonly known as hairworms, they which share a similar life cycle to the mermithids. This year featured a post about a species which infects and ultimately kills praying mantis, but in male mantis before this parasite takes its life, it take away its junk.

On the subject of that part of the body, there was also a post about frog bladder worms which do not always end up becoming parasitic, and whether they do so depend on its circumstances during the earliest part of its life. But even if some of those worms do no always end up as parasitising frogs, there are other worms that do, for example, the kangaroo leech. It drinks frog blood, hitches ride on crabs, and takes good care of its babies. There were also other blood suckers which were featured on the blog this year, and a rather unlikely one is the vampire snail.

As for guest posts, aside from the one contributed by Katie O'Dwyer, as usual, the students from my parasitology class also wrote stories on parasitoid wasps that force their host to weave a tangled web, tailor-made for their own purpose, but it seems that different wasps also coerce their spider hosts into weaving different webs. There was also a post about a parasite that causes rabbits to end up with a severe case of Shaft Studio head-tilt, a post about how parasites affect Monarch butterfly migration, another about how these butterflies fight back, and finally to top it off, a steaming pile of hyena poop sprinkled with tapeworm eggs.

In addition to writing about new papers about parasites, I also wrote about my experience attending the joint annual meeting for the New Zealand Society of Parasitology (NZSP) and Australian Society for Parasitology (ASP), which was held in Auckland, New Zealand this year. Among other things, in the first report I wrote about the fascinating story of giant squid parasites and its link to sharks, and in the second, I mused about the near-mythical status that Toxoplasma gondii has attained in the public consciousness.

I also wrote a post about parasite in prehistory to accompany my review paper on fossil parasites which has recently been published in the journal Biological Reviews. As a companion pieces, I also wrote an article for The Conversation which focus more specifically on dinosaur parasites (because everyone loves dinosaurs). So that about wraps it up for 2015. See you all in 2016 for another year of posts about more fascinating research into the world of parasites!

P.S. If you can't wait until next year for your parasite fix, as well as writing this blog, I have also been doing a regular radio segment call "Creepy but Curious" where I talk about parasitic and non-parasitic organisms such vegetarian spiderselectric eelshipwormsPompeii wormssirensvampire squid, brood parasitic cuckoo bees and cuckoo birds, carnivorous caterpillarsgreen sea slugs, the macabre bonehouse wasp, and a pair of unlikely parasites in the form of mussels and bitterlings. You can find links to all these and more on this page here.

P.P.S. Some of you might also know that I also do illustrations (and provide cartoons to accompany those Creepy but Curious segments), some of my drawings are about parasites, but I seem to have gone on a somewhat odd direction with those towards the end of the year...

December 10, 2015

Anomotaenia brevis

There are many examples of parasites altering the behaviour of their hosts, and some of them turn their hosts into functionally different animals compared with their uninfected counterparts. When this occurs in highly social animals, this effects can cascade onto other members of the group. Anomotaenia brevis is a tapeworm which happens to be one of many parasite species which have been documented to modify their host's appearance and/or behaviour in some way.
Photo by Sara Beros, used with permission

While the adult tapeworm lives a pretty ordinary life in the gut of a woodpecker, the larva uses a worker ant as a place to grow and a vehicle to reach the bird host. Specifically, they infect Temnothorax nylanderi - a species of ant found in oak forests of western Europe. These ants nest in naturally occurring cavities in trees such as sticks or acorns and the colony consists of a single ant queen surrounded by several dozen worker ants. These ants are a regular part of the woodpecker's diet so there's a fairly reasonable chance that the tapeworm will reach its final destination if it waited around for long enough. But A. brevis is not content with just leaving it to chance.

Worker ants can become infected through eating bird faeces which are contaminated with the parasite's eggs. As the tapeworm larvae grow inside the ant's body, these infected worker ants become noticeably different from their uninfected counterpart; they smell different (determined by the layer of hydrocarbon chemicals on their cuticle), they're smaller, they have yellow (instead of brown) cuticles, spend most of their time sitting around in the nest, and for some reason their uninfected nestmates are more willing to dote on these tapeworm-infected ants rather than healthy ones. They essentially become a different animal to the healthy workers, and other ant parasites have been known to alter their host to such a degree that parasitised individuals were initially mistaken as belonging to an entirely different species.

When scientists investigated the prevalence of A. brevis in nature, they found that about thirty percent of the ant colonies they came across have at least some infected workers. While in some nests only a few of the workers are infected, in other cases over half the workers are carrying tapeworms. Furthermore, they also found a few of the workers (2%) were infected but had yet to manifest the symptoms associated A. brevis. When over half the work force of a colony is under the spell of a body-snatching parasite, that must affect the colony in some way. So how does this affect the ant colony as a whole?

During their development, infected ants have higher survival rate and far more of them (97.2%) reach adulthood compared with uninfected (56.3-69.5%) ants. This make sense from the perspective of the parasite's transmission as it needs its host to stay alive for as long as possible to get inside a woodpecker. But it seems to also affected their uninfected sisters because uninfected worker ants in a colony which has parasitised workers also have lower survival rates than those from colonies free of any tapeworm-infected ants. But A. brevis also affects the colony's functioning in other ways as well.

The scientists behind the paper being featured today conducted a series of experiments where they manipulated the composition (and in doing so, parasite prevalence) of experimental ant colonies. Since T. nylanderi colonies regularly experience take-over and/or merging with other colonies, introducing or remove new ants into the experimental colonies would not cause them to exhibit unnatural behaviours as it is not too different what would usually occur in nature anyway. They set up colonies with different proportion of A. brevis-infected workers and tested how they responded to different types of disturbances.

They simulated a woodpecker attack by cracking open the experimental ant nests and seeing how long it took for them to evacuate. Under a simulated attack, about half of the healthy worker escaped (48-58.9%) but very few of the tapeworm-infected workers escaped (3.2%), which is exactly what the tapeworm wants - remember, the parasite needs to be eaten by a woodpecker to complete its lifecycle - so when one comes knocking, the tapeworm gets it host to sit tight and prepared to be sacrificed.

They also simulated intrusion from ants of a different colony or species by pitting individual invading ants against their experimental colonies. These invaders consisted of a mix of infected and uninfected individuals from nests which contained some or no infected nestmates. When confronting ants from other colonies, they were the most aggressive against the intruder if it was of a different species (in this case, T. affinis), but when it comes to other T. nylanderi ants, they responded more aggressively if the intruder from a different colony was harbouring tapeworm larvae.

In contrast, they were pretty chill about the presence of tapeworm-infected ants if it was one of their own nestmate. But the tapeworm also affected colony aggression in another way - the research team noted that colonies with many infected workers were also less aggressive overall towards any invaders. Not only does A. brevis alter its host's appearance and behaviour, it also seem to cause the host's nestmates to be more chilled out.

Parasites can manipulate their host in some astonishing ways, and the host's altered behaviour and/or appearance has been described as the parasite's "extended phenotype". But when the host is a social animal that is surrounded by many other group members, the parasite's influences can extend well beyond the body of its immediate host, and manifest in the surrounding kins and cohorts as well.

Beros, S., Jongepier, E., Hagemeier, F., & Foitzik, S. (2015). The parasite's long arm: a tapeworm parasite induces behavioural changes in uninfected group members of its social host. Proceedings of the Royal Society B 282: 20151473

November 23, 2015

Accacoelium contortum

Today, we are featuring a parasite that lives on the ocean sunfish (Mola mola) which happens to be the heaviest known living bony fish in the world. One can say that it is a truth universally acknowledged (by myself at least) that any sufficiently large animal must be in want of some parasite, and the sunfish is no exception. Its massive body is a prized piece of real estate for a wide variety of parasitic organisms.

Sunfish gill with arrows indicating the location of A. contortum
From Fig. 1 of the paper
Accacoelium contortum is the most commonly reported species of sunfish parasite, but even though they are numerous, not much is known about their biology. It is a digenean trematode - or parasitic fluke - and while most trematode flukes are parasites that live in the intestinal tract of their final host, A. contortum is a bit different. Occasionally you might find some of them in the gastrointestinal tract - which is where you'd expect to find most adult flukes - but more often found in the sunfish's mouth, gills, and pharynx (the part which roughly corresponds to back of the throat). Such a location is highly unusual for a trematode, it is akin to finding a seal living up a tree.

Scientists in Spain examined the parasites of 106 sunfish which were caught as bycatch and found that almost half (47.2%) were infected with A. contortum. Most of the flukes were found on the gills, some in the back of the throat near the pharyngeal teeth (which are a set of teeth that ray-finned fishes have at the back of their throat) and only a few were in the stomach. They noticed that usually the gills on the fish's right side are more heavily infected - this asymmetrical distribution is similar to what has been observed for other parasites, where they tend to congregated towards one side of the host, though in this case it's not entirely clear why they do this. In addition to preferentially hanging out on one side of the host, they also tend to congregate in clusters comprised of dozens of individuals, with those in the pharynx forming larger groups than those on the gills.

While A. contortum seems to do quite well in its rather unusual (for a trematode) habitat, the basic body plan for trematode is that of an internal parasite. So how does one modify a body plan for living inside the cosy confine of a host's body to a life clinging on to the more exposed parts of the host? Fortunately for A. contortum there are some functional overlaps between living in a fish's intestine versus living on its gills. While it lacks the hooks and sucker clamps of monogenean flatworms which are specialised for ectoparasitism, A. contortum has co-opted its large and muscular ventral sucker for hanging on to the sunfish's gills. Other trematode species use their ventral sucker to attach to the intestinal wall. In A. contortum it also function as the main attachment organ, but on a different part of the host's body. Additionally, this fluke's hind body appears to be long and prehensile, which it might to able to use to grip like a chameleon's tail (to a limited degree).
Left: Anatomical drawing of A. contortum, Right: Scanning electron microscope photo of the parasite's front
Image from Fig. 2. of the paper
The parasite's attachment also cause the surrounding host tissue to grow around them. This is most likely part of the sunfish's immune response, sealing the parasites off from the rest of the body. But this might actually work to the parasite's advantage because now it sits in a cosy little flesh bag which is tightly secured to the host body. The scientists who conducted the study also noticed that A. contortum has a series of tiny bumps and protrusions around the front of its body. They suggested that the fluke might actually be secreting growth factors which encourage host tissue growth through those bumps. Other parasitic flukes have been known to secrete proteins which manipulate host tissue growth, so it is possible that A. contortum is also capable of doing so. This is also supported by the observation that those protrusions are not found on immature flukes or those that live in the digestive tract which is more sheltered than the sunfish's gills.

The way that A. contortum apparently manipulates the sunfish's tissue is rather reminisce of how gall wasp induces their host plants to form a protective gall. While those galls protect their inhabitant against predators, in this case A. contortum, the flesh bag that it induces provide the parasite with a shelter on an otherwise exposed and turbulent location.

Ahuir-Baraja, A. E., Padrós, F., Palacios-Abella, J. F., Raga, J. A., & Montero, F. E. (2015). Accacoelium contortum (Trematoda: Accacoeliidae) a trematode living as a monogenean: morphological and pathological implications. Parasites & Vectors 8: 540.

P.S. I recently wrote a post about prehistoric/fossil parasites (which you can read here). On a related note I also wrote an article for The Conversation which focuses specifically on the fossil evidence for (non-avian) dinosaur parasites - you can check it out here.

November 12, 2015

Colubraria reticulata

Vampires have undergone a lot of image change over the centuries and they are a common part of many culture's mythology. But vampires are also a common part of nature. Blood sucking is a life style found in over 14000 known living species. Even those vampires themselves have blood suckers that feed on them. But living as a blood-sucker require special adaptations, and one particularly unlikely vampire is Colubraria (formerly known as Cumia) reticulata, the vampire snail. It is a marine snail that feed on fish blood and it belongs to a family of vampire snails called the Colubrariidae - at least six species are known to feed on blood and it is quite likely that it is a trait shared by the entire family.
Image modified from Figure 2 of the paper

So just how does a snail feed on a comparatively agile animal like a fish? First of all, they feed at night when fish are asleep, a survival tactic shared by other blood-feeders like vampire bats. They also have modified mouthpart can can slice flesh like a tiny scalpel, which is mounted at the end of a long proboscis that can stretch to three times its body length. This enables it to bypass even a parrotfish's mucus sleeping bag which normally protects it against other nocturnal blood-suckers.

But those behavioural and anatomical adaptations are just the start, most of the tools C. reticulata brings to this blood feast exist on a molecular level. The vampire snail is able to secrete a range of specialised proteins, most of which have multiple effects on the host and overlap in their functions.

First of all when the snail is about to cut into the fish's flesh, it spits out an anaesthetic similar to compounds secreted by other blood suckers like mosquitoes, to numb the area of incision. Once C. reticulata gets access under the fish's skin, other types of compounds come into play. A major problem for any would-be vampire is the natural tendency for blood to clot. Imagine drinking a smoothie and suddenly it turns into a big block of solid curd. So during feeding, C. reticulata secretes a chemical cocktail that disrupts the process of blood clotting and wound healing. Furthermore, the anti-coagulant action needs to be active until the blood is fully digested, so the snail also have secondary glands in its oesophagus that secrete other types of proteins to keep the blood liquefied as it sits in the snail's gut.

In addition to anti-coagulants, C. reticulata also spits out vasopressive compounds that increases the fish's blood pressure. This is very important to the vampire snail's feeding style because its long proboscis is actually not very muscular - so it is not that good at sucking blood. Instead, the snail injects compounds that increase the fish's blood pressure so that it will actually be pumping blood into the snail's gut. When scientists looked into the vampire snail's molecular arsenal in more details, they found that many of the proteins secrete by the vampire snail can be considered as pretty standard fare for a vampire and are similar to those found in terrestrial blood-feeders like ticks and mosquitoes.

However, C. reticulata also has a few tricks up its shell which are unique compared with other vampires, in particular the complex of protein which it secretes to temporarily suppress the fish's coagulation and healing mechanism. This is actually quite a feat because comparing with other vertebrate animals, fish are very good at repairing vascular injuries, especially in delicate blood-rich organs like the gills which are exposed to the external environment.

Another substance unique to the vampire snail is turritoxin - which is also produced by the coneshell. At this point, scientists are unsure how vampire snail (or the cone shell) uses turritoxin in their hunting behaviour, though it is possible they release it as a way of lulling the fish into a compliant state. Scientists have observed that fish which are approached by the coneshell enters a kind of "hypnotic" state before they get stung with the coneshell's highly lethal neurotoxin. Perhaps the vampire snail also release turritoxin to coax its victim into a deeper state of sleep.

By investigating the molecular arsenal of the vampire snail, scientists can gain insight into how the vampire snail evolved to be a blood-feeder. In addition, some of compounds secreted by C. reticulata can finely manipulate the physiology of their host, and examining them in detail may lead to the development of compounds with useful medical and pharmaceutical applications.

Modica, M. V., Lombardo, F., Franchini, P., & Oliverio, M. (2015). The venomous cocktail of the vampire snail Colubraria reticulata (Mollusca, Gastropoda). BMC Genomics,16: 441.

October 23, 2015

Goussia ameliae

The fate of parasites are often inextricably linked to that of their hosts, and when there are changes in the host population, the effects cascade onto their parasites. The study featured today is focused on Goussia amelia - it is a newly described single-cell protozoan parasite which infects alewives and is known to cause erosion in the intestinal wall of their fish host.
Image modified from Figure 2 and 3 of the paper
Alewife is a species of herring native to the east coast of North America. They are anadromous fish that live in the coastal marine environments as adults, but enter freshwater streams to breed, much like salmon. Sometimes populations of alewives become trapped in lakes for one reason or the other during their migratory journey. These isolated fish eventually become adapted to the freshwater environment and evolved on divergent paths to their anadromous relatives. This is a relatively common occurrence which has happened multiple time in the last few thousand years, and it is also the origin for the population of alewives found in Lake Hopatcong. This lake was originally connected via a canal to the Delaware River and alewives from the coast of New Jersey used to migrate to Lake Hopatcong to spawn. But during the start of the 1900s the canal was blocked off, and the alewives that were in the lake at the time became isolated from their relatives on the New Jersey coast.

So how did this affect parasites like G. ameliae? A pair of scientists compared G. ameliae found in alewives from Lake Hopatcong to those found in the anadromous alewives from Maurice River and noted some key differences in the two forms. For example, G. ameliae from anadromous alewives have oocysts (the infective stage of the parasite) which are comparatively shorter and wider than those from landlocked hosts.

They also have different trends in their prevalence and distribution; adult anadromous alewives are more commonly and heavily infected with G. ameliae than young fish, possibly because adult fish become stressed while migrating upstream and dealing with changing salinity levels as they move from the marine environment to a freshwater one, making them more susceptible to parasitic infections. In contrast, G. amelia was very common in younger landlocked alewives, infecting over ninety percent of young fish, but it was only found in about a third of the adult fish, which may indicate that the landlocked alewives can acquire resistance to the parasite as they mature.

Given those differences, are the anadromous and landlocked G. amelia actually different species? The scientists compared the DNA of G. ameliae from the anadromous and landlocked hosts, focusing on the 18S RNA gene which can function like a barcode for distinguish different species of parasites. They found that despite the two form having slightly different morphology and ecology, it was not enough to make them separate species - their 18S RNA gene sequences were identical. But given their differences, much like their hosts, those separate populations might be in the process of diverging into two different species - it is just a matter of time.

Lovy, J., & Friend, S. E. (2015). Intestinal coccidiosis of anadromous and landlocked alewives, Alosa pseudoharengus, caused by Goussia ameliae n. sp. and G. alosii n. sp.(Apicomplexa: Eimeriidae). International Journal for Parasitology: Parasites and Wildlife, 4: 159-170.

October 7, 2015

Marsupiobdella africana

Leeches are not endearing animals and many are literal blood-suckers. As a result they often evoke a sense of disgust in most people, and the term "leech" is usually used in a derogatory way. But most people might not realise that leeches also has a warm, maternal side too, one which is amply demonstrated in the kangaroo leech, Marsupiobdella africana. But this leech does not parasitise the kangaroo - indeed, in southern Africa where M. africanus is found there are no kangaroos - the reason it has that name actually has more to do with how it reproduces

Left: A pair of mating leech.                                   Right: Leeches riding on the legs of  a crab      (from Fig. 1 of the paper)
Marsupiobdella africana makes a living sucking blood from Xenopus laevis - the African clawed frog which is commonly used as a laboratory model for developmental biology research. When it reach sexual maturity, the leech detaches from its frog host to find a suitable mate. Some do so by simply crawling around in the environment, but they are also known to hitch-hike on the legs on crabs as if they some kind of crustacean-based Uber, admittedly an armoured, multi-legged one.

Top: Leech with spermatophore attached
Middle: Leech with filled brood pouch
Bottom: Young leeches emerging
from brood pouch
From Fig. 2 of the paper
These leech are hermaphrodites, and each individual take turns being the sperm depositor and the recipient. Mating between kangaroo leech is a very different affair to how you might imagine it, and from our perspective it is not very romantic. Instead of bringing their respective genitalia together, the leech playing the sperm depositor role actually pulls out a spermatophore - which is something like a biological hypodermic syringe filled with sperm - and stabs it into the recipient, which may end up being tagged with one to three of those sperm packets.

If the prospect of being harpooned with a sperm-filled syringe is not daunting enough, the recipient also make a habit of collecting a bunch of spermatophores from a number of different depositors, probably to ensure they can have the cream of the crop (so to speak). Once the spermatophore has made its mark, the sperm it carries are able to make their own way to the egg, no matter where the spermatophore may have initially landed on the leech's body. At this point it is not entirely clear how they accomplish this.

Once the eggs are fertilised, the sperm recipient, now playing the role of mother leech, transfer the eggs (which can be as many as 50) to a brood pouch in her belly (which is where the name kangaroo leech came from). There they will be protected and nurtured. Once the eggs hatch, the baby leeches continue to receive nutrient from their mother through her body wall and into their posterior suckers. Those developing leeches will stay in the pouch for four weeks. As a final send-off, the mother leech will find an unsuspecting clawed frog, and the young leeches are "released explosively" over the surface of the frog, thus ensuring that those blood-suckers will get the best possible start to their own lives.

Marsupiobdella africana - a loving and nurturing blood-sucker which wants nothing but the best for its babies (see also another blood sucker which goes to great lengths to care for its brood here).

Kruger, N., & Du Preez, L. (2015). Reproductive strategies of the kangaroo leech, Marsupiobdella africana (Glossiphoniidae). International Journal for Parasitology: Parasites and Wildlife 4: 142-147.

September 25, 2015

Allodero hylae

Half of all known segmented worms are oligochaetes, and the most well-known example is an earthworm. But aside from the earthworms that might be crawling under your garden, there are a wide variety of oligochaete species living in all kinds of environments, including freshwater habitats, seashore, sewage, and even glacial ice. Considering the range of environments they inhabit, it is a bit surprising that so few oligochaetes have evolved to be internal parasites.

From Figure 2 of the paper
There are only two known genera of endoparasitic oligochaetes - Chaetogaster which lives as internal parasite/symbionts of freshwater molluscs such as snails and mussels, and Dero which lives in frogs and toads, and they both belong to the family Naididae. The species featured today is Allodero (a subgenus of Dero) hylae - it lives out its life in the ureter of the Cuban tree frog Osteropilus septentionalis. Wild frogs can have more than 40 worms in the ureters, which can become dilated due to the parasite load.

The study we are featuring today investigated how these worms get from one frog to another. The researchers knew that the larval worms are pass (or pissed) into the environment via the frog's urine, but they wanted to test whether A. hylae which had been freshly expelled with frog pee can actively infect another frog, and what happens to the worms that don't end up in a frog.

First, they exposed five different species of frog and toads to some A. hylae larvae from a "donor" frog. They observed that A. hylae infect their hosts by swimming up their cloaca, but they are rather picky about whose cloaca they went up. Out of the five species of potential hosts, only the tree frogs ended up being infected. But this is not an entirely one-sided interaction - the researchers also noted that potential hosts can turn the tables on the worms by eating them before they have a chance to swim up their cloaca. If A. hylae enters a frog through its mouth instead of its cloaca, they will simply get digested.

Next they test if an uninfected frog can become infected in the presence of an infected one, and they did so by placing an uninfected frog with a frog carrying A. hylae in either a plastic container or a water-filled bromeliad (for a more naturalistic setting). For good measure, they simulate a predation event on the infected frog to ensure that some worms are expelled. In less technical terms, they scared the piss (and worms) out of an infected frog.

Photo of Allodero lutzi, a related species from southern Brazil
Photo from from Figure 1 of this paper
Sharing a room with a infected room mate is one thing, but to share a room with one that had just pissed themselves and there are parasites in their pee that wants to crawl up your cloaca is probably a bit much (even for reality TV these days). Between 60-73 percent of the tree frogs sharing a container or bromeliad with an infected room mate did end up getting worms in their ureters, showing that fresh pee from an infected frog can be a source of new infection.

Since A. hylae needs to actively seek out a host in the environment, when these worms are born, they start out well-equipped for a life swimming in the water. They have bristles (setae) on their back, well-developed gills, and a fully functional digestive tract - all necessary for making it as a free-living organisms. But once they get in a frog, within 72 hours they undergo a transformation whereby they lose all the those features and become more equipped for a life as a parasite inside a frog's ureters.

But what happens to the worms that do not end up in a frog? For most parasites, not finding a host means death. But it seems that once a larval A. hylae has been away from a frog for long enough, they don't look back. The researchers found that while worms that have been out of a frog for less than a week are attracted to frog BO, those that have been out over two weeks lose their attraction. In addition to being disinterested in frog BO, these older worms retain their bristles, gills, and fully functional digestive tract for good. Unlike their parasitic cousins who have lost all such features once they found a nice frog to settle into, these worms have become used to the outside world and are content to spend their life swimming in the water and foraging for microbes.

Animals like A. hylae, which have not evolved to be fully commit to a parasitic lifestyle, can give insight into how internal parasites have evolved from ancestors that were initially free-living organisms. Depending on its circumstances, A. hylae will end up either living in the ureters of a frog, or out hunting microbes in the water. Allodero hylae doesn't always chose the outside life, sometimes the outside life choses it

Andrews, J. M., Childress, J. N., Iakovidis, T. J., & Langford, G. J. (2015). Elucidating the Life History and Ecological Aspects of Allodero hylae (Annelida: Clitellata: Naididae), A Parasitic Oligochaete of Invasive Cuban Tree Frogs in Florida. Journal of Parasitology 101: 275-281.

September 6, 2015

Chordodes formosanus

For most insects (and other small animals), the praying mantis is a creature out of their worst nightmare; a deadly predator with giant compound eyes, a nasty set of high-speed spiky grasping limbs, and an appetite to boot. But Chordodes formosanus is a parasite that would give mantis nightmares - it is a hairworm - and regular readers of this blog will know immediately why that is justified.
From Fig. 2 of this paper
The worm starts out as a microscopic larva hidden inside the body of small insects - the mantis' usual prey - but once it is ingested by a mantis, it can then grow to several centimetres long inside its abdomen. By the time it is ready to bid farewell to its reluctant host, which comes when it reaches sexual maturity, the worm has already taken up most of the space within the mantis, leaving it a half-empty husk. The modus operandi of a horsehair worm is to then get into the water, which involves the host taking a dunk - whether it wants to or not. Apart from commandeering the mantis to go for a terminal end to their relationship, during the worm's development, it takes a massive toll. After all, one does not simply host a giant worm inside one's abdomen without any consequences.

But the said consequences is not equally distributed within the mantis population - this hairworm seems to affect male mantis more severely - especially in regards to their reproductive capacity. In a nutshell - C. formosanus shrink their testes and in some cases, they disappear altogether. However, this parasite seems more forgiving when it comes to female mantis; infected female mantis can harbour the worm and still retain intact reproductive organs. Not to say it doesn't exact a toll, just that the female mantis can still have some babies before her end comes. So why this sex bias? The reason lies in how this parasite alters the host's physiology.

When researchers looked at various aspects of the infected mantis' physical appearance, they also noticed some external changes in both sexes - they had comparatively shorter walking legs, smaller wings, and altered antennae - but it was more pronounced in the infected male mantis. Overall, the infected individuals have an appearance which bears closer resemblance to that of late-stage juvenile rather than adults. This suggest that C. formosanus might be tempering with the mantis' so-called "juvenile hormones" which control development in insects. But why is it that only the male mantis lose their reproductive organs? At this point, it is not entirely clear, but it might have something to do with the different role played said hormones in the development of each sex.

So why has C. formosanus evolved to castrate their male mantis host? From the parasite's perspective, host castration is a very effective strategy - the host does not need its gonad to survive, only to reproduce. So by tapping into this energy source, the parasite can keep the host alive while maximising the amount of resources it draws from the host.

For that, the host pays a double cost in terms of evolutionary fitness. Usually with such parasite infection which inevitably results in the host's death, the best thing for the host to do to make the best of a bad situation and reproduce as much as possible before they are eventually killed by the parasite. But in this case, the male mantis doesn't even get to do that - thanks to C. formosanus, long before it bid farewell to life, it has to bid farewell to its junk as well

Chiu, M. C., Huang, C. G., Wu, W. J., & Shiao, S. F. (2015). Morphological allometry and intersexuality in horsehair-worm-infected mantids, Hierodula formosana (Mantodea: Mantidae). Parasitology 142: 1130-1142