Mozambique Diary: Not all flies fly

Tsetse fly (Glossina sp.) from Gorongosa feeding on my blood. Luckily, tsetses in this area do not carry the dreaded sleeping sickness (but it does not make it any less painful).
Tsetse fly (Glossina sp.) from Gorongosa feeding on my blood. Luckily, tsetses in Gorongosa do not carry the dreaded sleeping sickness.

After a long hike in the scorching heat of the African savanna the cool, shady patch of tall miombo forest looked like heaven to us. I was in the southern part of Gorongosa, looking with a few friends for some elusive species of arthropods. But we were having little luck finding any and after several hours of strenuous walking the morale was low. As we stepped under the dark, inviting canopy of the forest, the drop in the temperature was palpable and we all relaxed, slowed down the pace, and the mood in the group immediately improved. But then, suddenly, somebody yelped “Ouch!” and at the same moment I felt a painful pin-prick at the back of my neck. Crap, tsetse flies! We looked around – they were everywhere. Clouds of them. We could see groups of dozens clumping on vegetation, taking into the air the instant they noticed the movement of our bodies. We ran.

A painting (undoubtedly the first and only) of a bat fly (Penicillidia sp.) burrowing in the fur of a Long-winged bat (Miniopterus).
A painting (undoubtedly the first and only) of a bat fly (Penicillidia sp.) burrowing in the fur of a Long-winged bat (Miniopterus).

Tsetse flies have long had a reputation for being one of the scourges of Africa, alongside malaria, crocodiles, and the plague of locusts. And deservedly so – some species of tsetses, all members of the genus Glossina, are vectors of nasty protozoans, including Trypanosoma brucei, the cause of the deadly sleeping sickness. Luckily for us, Gorongosa tsetses carry a different Trypanosoma species, T. congolense. This protozoan does not affect humans but unfortunately causes the chronic Nagana disease in cattle and horses, which explains the nearly complete absence of these animals around the park and in almost the entire region of central Mozambique. But knowing that tsetse bites are not going to kill us did not make them any more pleasant. Tsetses are large flies, about the size of a bee, and their skin-piercing mouthparts are much thicker than those of a mosquito. In other words, it hurts like hell when one jabs you with its proboscis, and you flail your arms like a madman to shoo it away while the fly escapes unharmed.

Members of the family Streblidae, such as this Raymondia sp., collected from the Hildebrandt's horseshoe bat (Rhinolophus hildebrandtii), often exhibit interesting adaptations in their wing morphology, such as the ability to fold them longitudinally along the back. This presumably helps them move swiftly in the pelage of their hosts.
Members of the family Streblidae, such as this Raymondia sp., collected from the Hildebrandt’s horseshoe bat (Rhinolophus hildebrandtii), often exhibit interesting adaptations in their wing morphology, such as the ability to fold them longitudinally along the back. This presumably helps them move swiftly in the pelage of their hosts.

But count yourself lucky. Imagine instead that you cannot shoo them away. You try to smack one but it runs, hides in your hair or some place where you are not able to reach, and it continues to bite. It only leaves your body to give birth somewhere in your house but then immediately runs back, guided by your scent and body heat. Oh, and imagine that this fly is the size of your fist (or a small puppy). Welcome to the world that bats are forced to live in.

Tsetses are members of a large group of flies, the superfamily Hippoboscoidea, all of which are exclusively hematophagous – blood is the only food that they are interested in. The tsetse family (Glossinidae) is the most basal (unsophisticated, one might say) member of this lineage of insects – they are always looking for a blood meal but never evolved the ability to stay with their tasty host. Bats are unlucky to have been colonized by two much savvier families of flies, the Nycteribiidae and Streblidae. These insects know the value of a good host and, once they landed on the furry back of a bat, they never leave it again. Over millions of years of coevolution with their mammalian hosts the bat flies have undergone a remarkable transition. From a free-flying ancestor, most likely very similar to today’s tsetse flies, emerged several lineages of highly modified, often completely wingless, spider-like creatures. Their body became flattened and very hard, making it almost impossible to squash them against the skin. In the family Nyctiberiidae the head turned into a small appendage that can be safely tucked away in a protective groove on the back and all traces of wings completely disappeared. These flies cannot survive for long outside of their host’s body and only feel at home when scurrying at an alarming speed in its dense fur. Their feet are armed with large claws that make it almost impossible to dislodge them from the hair of their host. They really don’t look like flies and when a friend spotted one on the body of a bat she called me to collect the bat’s “pet spider.”

In the closely related family Streblidae the wings may or may not be present, but even in the winged species the body is modified for squeezing through the fur, and members of the subfamily Ascodipterinae go even further in their commitment to the host. Much further. Once a female lands on a bat she sheds her wings and legs (yes, legs) and burrows head-first into the skin. Once there, her head and thorax sink into her own abdomen, and the skin of the bat overgrows her body. She becomes one with her host.

Penicillidia bat flies (Nycteribiidae) are some of the most unusual members of the order Diptera and hardly resemble their winged relatives. This individual was collected from a Long-winged bat (Miniopterus natalensis) in Gorongosa, Mozambique.
Penicillidia bat flies (Nycteribiidae) are some of the most unusual members of the order Diptera and hardly resemble their winged relatives. This individual was collected from a Long-winged bat (Miniopterus natalensis) in Gorongosa, Mozambique.

Female bat flies, like their relatives tsetse flies, are remarkably good mothers. The great majority of insects relies on what ecologist call “r-selection” in their reproduction – they lay hundreds or thousands of eggs, betting on one or two of them making it to adulthood. Bat flies, on the other hand, rely on “K-selection” – like humans, they prefer to invest a lot in a much smaller number of offspring, hoping that they will all make it to the reproductive age. They are larviparous – instead of laying eggs the female gives birth to a single, fully developed larva, which immediately turns into a pupa. While in her mother’s body, the larva feeds on “milk glands”, analogous to the mammalian mammary glands (if they were located in the uterus), and develops safely protected from the elements and predators. When the time comes for the mother to give birth she walks off the bat’s body and attaches the larva to the wall of the bat’s roosting place, usually a cave (which explains why bats that roost in rolled-up leaves and other less permanent places have fewer ectoparasites). Then she turns back and runs towards her host, guided by the smell and the heat of its body.

The recent Ebola crisis brought back the attention of the medical community to bats as potential reservoirs of the virus. Although there is no evidence that bats are in fact harboring the virus, there seems to be some correlation between instances of the outbreak and the presence of large numbers of bats in the affected areas. While reading the literature on both Ebola and bat flies I found it rather curious that nobody has tested bat flies for the presence of the virus – these are relatively very long lived (195 days on average) insects, who always stay (as pupae) at the roosting sites of bats, even when the hosts leave to forage elsewhere. They often move from one host species to another and, this point makes me really wonder why nobody has seriously looked at these flies as potential vectors, occasionally drop on and bite people. We know that they harbor a slew of pathogens – a recent study conducted in Gorongosa National Park on bats Rhinolophus landeri and Hipposideros caffer showed that flies living on these animals are vectors of Trypanosoma species that are ancestral to those that cause Chagas disease. Add to this the fact that one of the first cases of Marburg disease in Zimbabwe (caused by a virus related to Ebola) was caused by a bite of an arthropod (by default all unidentified bites seem to be classified by the medical community as “spider bites” and spiders in the area were tested, predictably unsuccessfully, for Marburg). It is far more likely that the bite was caused by a fly that fell off a bat.

A friend of mine recently expressed her dismay at “lowly” parasites. I beg to differ – if anything, parasites, including bat flies, are incredible examples of evolution at its best, organisms capable of both adapting to life in the most hostile of environments (the very substrate you live on wants you dead!) and resisting diseases that live inside your body. I cannot promise that I will not try to smack the next tsetse fly that lands on me but at least I promise that I will do it in the most respectful, considerate way.

Louse flies (Hippoboscidae) are close, equally modified for ectoparasitic lifestyle family of flies. This Lipoptena sp. was collected from a Nyala antelope while it was fitted with a GPS collar. Louse flies are parasites of large mammals and birds, and some are considered serious pests of sheep.
Louse flies (Hippoboscidae) are closely related to bat flies and equally modified for ectoparasitic lifestyle. This Lipoptena sp. was collected from a Nyala antelope while it was being fitted with a GPS collar. Louse flies are parasites of large mammals and birds, and some are considered serious pests of sheep.

14 Comments Add yours

  1. Rose says:

    Once again… many thanks for these amazingly interesting articles. For someone who doesn’t even like insects and the sort, I have been completely wowed reading them and learning so much. Thank you!

  2. Evita says:

    Hello Piotr, I found this
    https://t.co/YtKbsmfTx0?amp=1
    and was wondering if the picture with the parasite in the face of the bat can be real.
    What’s the size of the parasite, and what’s the size if the bat?
    Thank you! All the best, Evita

  3. achei cpean seu tipo de design! qual você usa?
    tenho um site que fala sobre o mexmo raciocínio, quem queiser ddar uma olhada..

  4. Mike Huben says:

    By the way, those are the most outstanding pictures I’ve ever seen of these groups.

  5. Mike Huben says:

    Bats have wonderful assemblages of parasites. I’ve collected Streblids, bedbugs, and Spinturnicidae from live bats. The latter are large, flat mites that walk on the wings of bats with enormous claws. Bats also have an assortment of other mite families that I’ve not yet collected.

    One Spinturnicid is colloquially known as the “bat asshole mite” because that’s the usual place it has been found. I have a (donated) specimen in my collection.

    Only birds have more diverse parasites, IMHO.

  6. Sonia says:

    Very interesting read. I’ve had the privilege of working with members of the Streblidae family in the past and was able to really appreciate their morphology up close. They are indeed an extraordinary group to learn about. Glad to see them featured on social media and hope to see more on them in the future! I agree about the Ebola link. It has my attention for sure.

  7. Harm says:

    Super cool story, and intersting link with ebola. However there’s one thing I dont understand. You write that the parasitic flies on the bats shed their wings and legs, but later walk their newborns off the back of the bat to the caveside. How do they walk there without their legs?

    1. The wingless and legless females are found only in members of the subfamily Ascopteridinae. In this group the female is complete encased in the host’s skin and cannot move, and only the tip of her abdomen is sticking out of the skin. In this position she gives birth to a live, fully developed larva, which then falls to the ground and turns into a puparium.
      In some species of Nycteribiidae the larva also simply drops to the ground so that the female does not need to leave the host, and in some Hippoboscidae the larva turns into a puparium directly on the host and develops in its fur.

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