Recently found a good summary of what makes strepsipterans so twisted!
https://academic.oup.com/bioscience/article/54/5/383/416832 Most people have never heard of strepsipterans and almost no one has knowingly seen one. Last year, when an article appeared in
Proceedings of the National Academy of Sciences (24 June 2003) describing how this parasitic insect eludes the defenses of its host by enclosing itself in a bag composed of the host's epidermal cells, the insect became a media darling. Two of the article's authors, entomologists Jeyaraney Kathirithamby (University of Oxford, United Kingdom) and Spencer Johnston (Texas A&M University, College Station), then found their “exotic little insect parasite” featured in “
Nature, Science, BBC, Discovery, and 16 other radio, TV, and journal publications.”
“Strepsipterans are the most enigmatic order of insect,” says entomologist David Grimaldi, of the American Museum of Natural History in New York. They are called the “twisted-wing” parasite, because in these insects, the hind wings develop into normal wings, but the front wings develop into halteres, or balancing organs that are modified wings shaped like knobs. “It is this character that stimulated early workers to think that Strepsiptera were related to beetles,” remarks Johnston.
Strepsipterans are a very old group. Grimaldi is studying a specimen found in Cretaceous amber—which is at least 65 million years old. In modern-day strepsipterans, males and females are parasitic. But Grimaldi explains that the Cretaceous fossil is “more primitive than the primitive living ones,” because it appears as if only the male larval forms grew within a host species.
One of the problems of strepsipteran systematics is that no one knows where they belong phylogenetically. Looking at them morphologically, at least four orders of insects have been proposed as their closest relatives, says Grimaldi. Genomic data show “they're closely related to flies,” but Grimaldi indicates that there is doubt about such a relationship. Johnston says that “more recent data (unfinished) do not support this association.”
Weird biology
Kathirithamby and colleagues noted in the
PNAS article that there are nearly 600 species of Strepsiptera, which parasitize “7 orders and 34 families of Insecta.” But unless one is specifically looking for species they parasitize or is dissecting a host species specimen for another reason, one would never know the parasite was present. In most cases, male and female strepsipterans parasitize one host species, though the male is parasitic only through early stages of development. But in members of the family Myrmecolacidae, Kathirithamby notes, the males parasitize ants and the females parasitize orthopterans. Not only do the sexes parasitize different species of insects, they also are so sexually dimorphic that of 108 species described, males and females have been recorded in only two of the species, and females alone were recorded for eight species. The sexes of these organisms are so different that even if a female conspecific of a known male can be found, the only way to be sure the two are the same species would be to use DNA analysis. Females, notes Johnston, look like worms, while the males look vaguely like
Drosophila with twisted wings (halteres in front of wings).
Their behavioral adaptations and their effects on their hosts are as strange as their morphology. For example, strepsipterans may parasitize their hosts at various times during the host's life cycle, but when the host is an adult, the parasites extrude their mouthparts and cephalothoraxes (females) or cephalothecae (males); the mouthparts are then lost. Kathirithamby explains that the body parts extruded by the female contain an opening in the head that receives sperm and functions as an exit for the first instar larvae.
Being parasitized—or stylopized, as it is called—extends the host's life span. For example, Kathirithamby explains that when the plant hopper,
Javesella dubia, is parasitized by the strepsipteran
Elenchus tenuicornis, the host fifth instar nymphs and adults live five or more times longer than nonparasitized members of the host species.
Stylopization may change the hosts' behavior, as well. When stylopized, one species of wasp,
Polistes dominulus, may leave its nest immediately before its strepsipteran parasites extrude their cephalothoraxes (in the case of females) and cephalothecae (in the case of males). These wasps then form mating aggregations—not for the wasps, but for their parasitic strepsipteran,
Xenos vesparum. In these aggregations, free-living adult male
X. vesparum emerge from their wasp hosts' bodies and mate with the females, whose cepthalothoraxes are just barely visible poking out of a host wasp's body. “This is the first example in
Polistes wasps of behavioral change induced by a parasite,” says Kathirithamby. The wasps that house the female parasites then overwinter. During this time, young develop within the viviparous female parasite. When the first instars of the parasite emerge, the host wasp dies—after an extended life span.
Why have eyes?
Free-living adult male strepsipterans live for only a few hours. During those hours, the male must find a stylopized host that contains a female with which it can mate. As the female is just visible as a small dot of a cephalothorax protruding from a host, this is not necessarily an easy task. Some researchers, such as evolutionary neurobiologist Elke Buschbeck, of the University of Cincinnati in Ohio, hypothesize that vision must be important in aiding free-living males in their search for mates. Others, such as Jochen Zeil, of the Australian National University in Canberra, suspect that strepsipteran vision has limited resolution and is more likely to be used for “flight stabilization, but not for ‘object vision.’”
Buschbeck has been studying the structure of the unique eye of
Xenos peckii, a strepsipteran parasite of the paper wasp,
Polistes fuscatus. She came to Strepsiptera by chance, working on fly vision in a lab at Cornell with someone who had been studying wasp behavior. “By looking at the eyes of Strepsiptera, it was immediately obvious: They're strange,” she says. She noticed that they did not follow the pattern of the insect compound eye. “The entire image is put together adding all of the little parts of images into one big image,” she explains. Buschbeck calls each eye section an eyelet so as not to confuse it with named structures from other insect eyes. She notes that it is “still not really resolved what they are in terms of homology.” What is known is that about 75 percent of the male strepsipteran brain is composed of visual cells.
The life cycle of
Xenos peckii is similar to that of other strepsipterans. The viviparous female is a wormlike parasite within the wasp host. Larvae develop within the host, but adult males emerge from their puparia and are free-living for several hours while they try to find a mate, whose cephalothorax is poking out of an opening in the host's abdomen. Photograph: Elke Buschbeck.
Left: The pupa of a male
Xenos peckii protruding from the body of the wasp host,
Polistes fuscatus. Right: Larvae (small) and pupae (large) of
Xenos peckii, removed from their
Polistes host. Photograph: Elke Buschbeck.
Adult male strepsipteran,
Pseudoxenos species, emerging from its puparium in the hymenopteran
Odynerus bicolour Saussure. Photograph: Jeyaraney Kathirithamby.
Free-living adult male strepsipteran. Note the reduced front wings (halteres), the larger hind wings, and the compound eyes. Photograph: Elke Buschbeck.