Tag Archives: wasps

Planes, trains and automobiles – insect killers?

I couldn’t not use this – it is (sadly) one of my favourite films 😊

Anyone who has driven (or walked) along a road will have come across roadkill, be it squirrels, pheasants, badgers, deer or even something more exotic, perhaps it us only us entomologists who notice the squashed invertebrates ☹

Dead carabids and mayflies Shay Lane, Staffordshire, 8th June 2021

But, lets leave the roadkill for a moment, and in the spirit of the title of the film, start in the air. The first thing I discovered when I started to search for the effects of aircraft on insects is the paucity of literature on the subject – it turns out that people are much more interested in stopping disease carrying insects being transported by air or, and coming as a bit of a surprise to me, stopping insects causing plane crashes (House et al., 2020; Grout & Russell, 2021). The aircraft industry is so concerned about the physical dangers posed to ‘planes by insects that NASA actually have a Bug Team dedicated to developing insect proof aircraft.

I am, however, more concerned about how dangerous aircraft are to insects. First, we need to know how many insects are up there and what the probability of them being struck and killed by aircraft is. I’m guessing that bug strike is pretty common, otherwise NASA wouldn’t have a Bug Team. The majority of insects in the air are found at 300-600 m, although this does vary in relation to time of day (Reynolds et al., 2005). Getting a figure for the actual number of insects in the air is as you might expect, actually quite difficult.  The first attempt to trap and collect insects using an aircraft was in 1926 in Louisiana (USA) using a specially designed trap (Glick, 1939).  These do not seem to have been particularly effective as 5 years of trapping, involving 1528 hours of flying, caught just under 30 000 insects (Glick, 1939).  Those of us who have operated pitfall traps for any length of time would consider this a very modest haul 😊

Glick (1939) The aircraft insect trap

That said, the exercise was obviously more hazardous than even collecting insects from roundabouts as this very laconic extract highlights:

 “The skill of the pilots who flew the collecting airplanes is evidenced by the fact that no fatalities occurred.  Only one major accident occurred, when a forced landing resulted in the destruction of the craft and injury to both the pilot (McGinley) and the writer. Such mishaps must be expected in a more or less hazardous undertaking.”

The distribution of catch number was very similar to that reported from the more recent UK study using radar (Reynolds et al., 2005) and is reinforced by this statement from the NASA Bug Team; “The reason we do these tests at low altitudes or do a lot of takeoffs and landings is because bug accumulation occurs at anywhere from the ground to less than 1,000 feet,” said Mia Siochi, a materials researcher at NASA Langley”.

Given the number of flights made globally and the investment being made into protecting aircraft from bug strike, I would assume that the number of insects being killed by aircraft worldwide is probably very high. I am sure that someone with the skill, time and inclination, can probably come up with a fairly realistic figure.  Over to you Dear Readers.

Next up, if we keep to the film title, are trains.  There has been a bit more work looking at the damage that trains do to insects, not a lot, but something is better than nothing.  Work collecting train kill from railway lines showed that snails were particularly vulnerable to being run over, similar to the effects on trail-following ermine moth caterpillars that I observed in Finland in 1981, with Ephemeroptera (Mayflies) in second place (Pop et al., 2020). This, as the authors suggest, was almost certainly due to the time of year and the presence of a lake nearby. Unfortunately no one has done the equivalent of a train splatometer which might be rewarding as these observations from correspondence in British Birds magazine suggest that locomotive engines are causing some mortality to flying insects.  Over to you Bug Life. How about getting the train companies to fit splatometers?

Finally, cars and their effect on insect life. There is anecdotal evidence out there, after all as drivers we have all seen moths in our headlights at night and used our windscreen washers and wipers to try and remove dried on insect corpses and their haemolymph from our front windscreens.

An observation by Ian Bedford

My front bumper – sadly (or perhaps not) much less insect spattered than in the past

Yes, anecdotally we know that insects are being hit by cars (see above) and on my front number plate, a couple of weeks ago (beginning of June) I counted 73 insects, mainly aphids after a 245 km trip. The problem as I see it, is quantifying the numbers killed and calculating the effect that this has on insect abundance. I have mentioned the splatometer in an earlier post which attempts to standardise the number plate counts and I am pleased to see that this has now been revived by Bug Life, and will hopefully carry on for many years. The idea behind this is that over the years we will be able to see if insect numbers as reflected by the change in numbers of splats are increasing, decreasing of remaining the same.  This will not, certainly as described, tell us how many insects are being killed by road using vehicles, although it would be possible if the data were collected over delineated stretches of road (Baxter-Gilbert et al., 2015).  It is not just flying insects that are killed by cars; not all flying insects fly across roads, many seem happy to walk to the other side, reckless as that may seem.

A brave, or possibly fool-hardy carabid beetle crossing the road – Guild Lane, Sutton, Staffordshire, 9th June 2021.

There have been enough studies done looking at the interactions between roads and insects for a review article to have been published fairly recently, although not all the papers deal directly with mortality effects (Munõz et al., 2015). Many studies have recorded the species affected and the number of dead individuals found but few have attempted to calculate what this means in total. Most studies, as we might expect, have been on large, easily identifiable charismatic species (Munõz et al., 2015) and it from these that we do have some idea of the magnitude of the mayhem caused by road traffic. Some of the figures are incredibly high. A survey of Odonata road kill, albeit near a wetland, of two 500 m stretches of dual carriageway in the Great Lakes region of the USA revealed that at least 88/km/day were being hit and killed by vehicles (Riffell, 1969).  Another study in the USA, this time on Lepidoptera, calculated that about 20 000 000 butterflies (mainly Pieridae) were killed in one week in September (McKenna et al., 2001). The most dramatic figures however, are those from a study in Canada which estimated that 187 billion pollinators (mainly Hymenoptera) are killed over the summer in North America (Baxter-Gilbert et al., 2015).  An unpublished study by Roger Morris (thank you Richard Wilson @ecology_digest for bringing this to my attention) also highlights the dangerous effects of cars on Hymenoptera). Despite the mounting evidence of the harm that road traffic does to insects there is remarkably little information about how this can be reduced, although I did find a paper that noted that if insects are struck by cars driving at speeds of 30-40 km/h they survive the crash whereas speeds greater than this prove fatal (Rao & Girish, 2007).  It might be possible to impose insect safe speed limits along stretches of road that go through sites of special insect interest (perhaps I should try and coin that acronym, SSII, as an additional/alternative term to SSSI (Sites of Special Scientific Interest), but I am not sure how amenable drivers would be to signs telling them to slow down because of insects😊, considering how few drivers slow down in response to the signs warning them about deer and other vertebrate hazards. Another option would be to design road vehicles so that the air flow across them pushes insects away rather than into them; this may already be fortuitously happening as Manu Saunders points in her interesting post about the ‘windscreen anecdote’.  That said, even if cars are more aerodynamic and less likely to splatter insects, the levels of road kill reported in the papers I have cited earlier, still imply that insects are being killed by traffic in huge numbers.

This one didn’t get stuck on a car, but died just the same – A519 outside Forton, Staffordshire, 15th June 2021

Even if we do accept that deaths down to direct impact with vehicles is lower than in the past, the roads on which we drive our cars are also having a negative effect on insect numbers. Roads, particularly those surfaced with tarmacadam, present an inhospitable surface to some insects which may make them reluctant to fly or walk across. It has been shown that bee and was communities can be different on different sides of a road (Andersson et al., 2017) as the road act as barriers, particularly for smaller species of bees (Fitch & Vaidya, 2021).

Despite the mortality that vehicles impose on insects, roads are not necessarily a totally bad thing for invertebrates; road verges, when sympathetically managed, can provide overwintering sites for a range of arthropod species (Saarinen et al., 2005; Schaffers et al., 2012) and some insect species seem to enjoy feeding on roadside vegetation because of the increased nitrogen content of the plants living alongside traffic (Jones & Leather, 2012).

Overall however, given the very high mortality rates directly associated with cars and other road traffic and the very real indirect effects caused by habitat fragmentation, it would seem that we have much to do to make roads safer for insects and other animals.


Andersson, P., Koffman, A., Sjödin, N.E. & Johansson, V. (2017) Roads may act as barriers to flying insects: species composition if bees and wasps differs on two sides of a large highway.  Nature Conservation, 18, 41-59.

Baxter-Gilbert, J.H., Riley, J.L., Neufeld, C.J.H., Litzgus, J.D., & Lesbarreres, D. (2015) Road mortality potentially responsible for billions of pollinating insect deaths annually. Journal of Insect Conservation, 19, 1029-1035.

Fitch, G. & Vaidya, C. (2021) Roads pose a significant barrier to bee movement, mediated by road size, traffic and bee identity. Journal of Applied Ecology, 58,1177–1186.

Glick, P.A. (1939) The Distribution of Insects, Spiders, and Mites in the air.  Technical Bulletin no. 673, USDA. https://naldc.nal.usda.gov/download/CAT86200667/PDF

Grout, A. & Russell, R.C. (2021)H Aircraft disinsection: what is the usefulness as a public health measure? Journal of Travel Medicine, 28, taaa124.

House, A.P.N., Ring, J.G., Hill, M.J. & Shaw, P.P. (2020) Insects and aviation safety: The case of the keyhole wasp Pachodynerus nasidens (Hymenoptera: Vespidae) in Australia. Transportation Research Interdisciplinary Perspectives, 4, 100096.

Jones, E.L. & Leather, S.R. (2012) Invertebrates in urban areas: a review. European Journal of Entomology, 109, 463-478.

McKenna, D.D., McKenna, K., Malcolm, S.B. & Berenbaum, M.R. (2001) Mortality of lepidoptera along roadways in Central Illinois. Journal of the Lepidopterist’s Society, 55, 63-68.

Melis, C., Olsen, C.B., Hyllvang, M., Gobbi, M., Stokke, B.G., & Røskaft, E. (2010) The effect of traffic intensity on ground beetle (Coleoptera: Carabidae) assemblages in central Sweden. Journal of Insect Conservation, 14, 159-168.

Munõz, P.T., Torres, F.P. & Megias, A.G. (2015) Effect of roads on insects: a review. Biodiversity & Conservation, 24, 659-682.

Pop, D.R., Maier, A.R.M., Cadar, A.M., Cicort-Lucaciu, A.S., Ferenți, S. & Cupșa, D. (2020) Slower than the trains! Railway mortality impacts especially snails on a railway in the Apuseni Mountains, Romania. Annales Zoologici Fennici, 57, 225-235.

Rao, R.S.P & Girish, M.K.S. (2007) Road kills: assessing insect casualties using flagship taxon. Current Science, 92, 830-837.

Reynolds, D.R., Chapman, J.W., Edwards, A.S., Smith, A.D., Wood, C. R., Barlow, J. F. and Woiwod, I.P. (2005) Radar studies of the vertical distribution of insects migrating over southern Britain: the influence of temperature inversions on nocturnal layer concentrations. Bulletin of Entomological Research, 95, 259-274.

Riffell, S.K. (1999) Road mortality of dragonflies (Odonata) in a Great Lakes coastal wetland. Great Lakes Entomologist, 32, 63-74.

Saarinen, K., Valtonen, A., Jantunen, J. & Saarnio, J. (2005) Butterflies and diurnal moths along road verges: does road type affect diversity and abundance? Biological Conservation, 123, 403-412.

Schaffers, A.P., Raemakers, I.P., & Sýkora, K.V. (2012) Successful overwintering of arthropods in roadside verges. Journal of Insect Conservation, 16, 511-522.


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Pick & Mix 61 – Terry Pratchett, chickens, trees, species conservation, wasps, bees, eating weeds and much more

Google’s new timelapse shows 37 years of climate change anywhere on earth, including your neighbourhood

Bay bees love carbs – By considering the nuances of bees’ dietary needs, we can design nutritionally balanced seed mixes that help pollinators shore up our ecosystems and food supplies.

Seirian Sumner writes about her love of wasps and why you and I should too

Why I only buy organic or truly free-range chicken and eggs – Revealed: true cost of Britain’s addiction to factory-farmed chicken – it comes at a price though. If you buy from a supermarket, a small mass reared chicken costs approximately £3, free-range’ same size, £9, and the same size organic, £18.

Jeremy Fox on which Terry Pratchett books to read and in which order – I don’t necessarily agree with him but always happy to spread the word about the late great Terry Pratchett. Fun fact, I once had his email address and used to correspond with him, Then one day, having just read the Carpet People and the Bromeliad trilogy and really enjoyed them I emailed him and said so, giving as my reason that it was like an updated and funny version of the Borrowers. Unfortunately he thought I was accusing him of plagiarism and that was the end of our relationship L

There aren’t enough trees in the world to offset society’s carbon emissions – and there never will be, but that doesn’t mean we should stop planting them

Mountain Avens – the Scottish sunflowers?

Sobering read from Charley Krebs – “There are times when we either act or give up, so if you think that the Covid epidemic, the conservation of endangered species, and the protection of old growth forests are irrelevant problems to your way of life, stop reading here. These three major problems are here and now and have come to a head as a crunch: do something or quit.”

Got a problem with Japanese Knotweed?  Try eating it 🙂

Terry McGlynn asks “Should reviewers of journal papers be paid?” 

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Pick & Mix 51 – permafrost, shifting baselines, insect pests, carrion beetles, a history of chocolate, double flowers, wasps and the most beautiful springtail you have ever seen

Very worrying news from the frozen North

You will have seen Extinction Rebellion in the news a lot complaining about lack of progress on slowing down climate change, but do they care about Nature as a whole?

Help prevent shifting baseline syndrome by talking to your grandchildren

Social media isn’t just funny cat videos – The next invasion of insect pests will be discovered via social media

Nice article on carrion beetles – and incidentally emphasising the importance of citizen science projects and social media

If you like chocolate and want an excuse to eat more, then this article might be of interest J

Did you know that flowers can grow out of flowers? Unravelling the mystery of double flowers

Professor Sierian Sunmer of University College London, explains why wasps like to join you on your picnic as summer comes to an end

Will we add a new butterfly to the British list – is the Camberwell Beauty on the way to join us?

Springtails – ubiquitous, beautiful, incredibly useful and totally overlooked by almost everyone




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Not all aphids grow up to be aphids – the enemy within

It has been said that if aphids had their own way and unlimited resources the world as we know it would be 149 km deep in the cute little beasts (Harrington, 1994 ). Last year I wrote about how predators that feed on aphids, although useful, don’t really cut the mustard when it comes to keeping them in check and suggested that their host plants played a major role in keeping aphids from taking over the World.  While they do play an important part in keeping aphid populations under control, and are aided and abetted by aphid specific predators, there are, however, some much more efficient aphid-specific natural enemies out there.  They may be less conspicuous than the brightly coloured ladybirds that we often see munching their way through aphid colonies; public perception of their name may make people wince, but these beautiful and graceful creatures make sure that our appetite for salads and exotic vegetables out of season is satisfied safely and efficiently.  Their life cycles rival that of their prey, or should that be hosts, and entomologists fondly imagine that the film Alien was inspired by them 😊

I am, of course, talking about parasitic wasps, or parasitoids as they are more commonly known.  They are called parasitoids because unlike true parasites which generally speaking keep their hosts alive, insect victims of these wasps will, if successfully parasitized, die well before their non-parasitized relatives. In case you were wondering, the term parasitoid was coined by the Finnish Hemipterist, Odo Reuter (1913).  Aphids are not the only insects that are attacked by parasitoid wasps. The action of insect parasites has been known about for over two hundred years.  Erasmus Darwin, grandfather of the more famous Charles, noted that Ichneumonid wasps parasitised cabbage white butterfly caterpillars and so should be encouraged by gardeners (Darwin, 1800).  This is not the only early mention of parasitic insects in this context; Wheeler (1928), points out that back in the 1850s, two Italian entomologists, Camillo Rondani and Vittore Ghiliana also suggested the use of parasitic insects as biological control agents.  Aphid pests of glasshouse crops originally controlled mainly by predators (van Lenteren & Woets, 1988) are now routinely controlled by the application of commercially produced Braconid and Chalcid wasps (Boivin et al., 2012; van Lenteren, 2012).

Three commonly used aphid parasitoid biological control agents in action. Images from http://biologicalservices.com.au/products/aphelinus-2.html and https://www6.inra.fr/encyclopedie-pucerons/Especes/Parasitoides/Braconidae-Aphidiinae/Praon-volucre

When people think of Hymenoptera, they tend to think of bees, Vespid wasps and ants as being the most important and abundant.  They are very much mistaken.  The Parastica, or parasitoid wasps, are, by a huge margin, the most speciose and abundant section of

Parasitoids clearly dominate the Hymenopteran fauna of the British Isles (Many thanks to Natalie Dale-Skey of the NHM for permission to use this).

the Hymenoptera both in the UK and elsewhere

In the tropics the parasitoids are even more dominant. Data from Gaston et al., (1996).

Once parasitized, the egg(s), unless they are encapsulated by the aphid ‘immune’ system, hatch and begin to feed on the internal tissues of their, presumably, unsuspecting aphid host.  The parasitoid larvae avoid feeding on vital parts of the aphid, so that it can continue to grow and develop and provide food for the parasitoid, until the parasitoid is ready to pupate. Once the parasitoid is ready to pupate it delivers the coup de grace putting the aphid out of its misery and allowing the formation of the ‘mummy’ in

The three most common types of aphid mummies.  Images from http://resources.rothamsted.ac.uk/science-stories/aphids-mummies-and-cadavershttp://biologicalservices.com.au/products/aphelinus-2.html and https://farm1.static.flickr.com/327/18532751584_becc0e56e9_b.jpg respectively.

which the parasitoid completes its development before sawing its way out to emerge as a winged adult ready to seek out new hosts, leaving a characteristic neat circular hole in mummy case. In case you were wondering why the mummy of Praon volucre looks like it is sitting on a plate, this because, unlike the other aphid parasitoids, the final instar cuts its way out of the bottom of the aphid and spins its cocoon externally underneath the remnants of the aphid, hence the ‘plate’ (Beirne, 1942).

And out she comes; emerging parasitoid – http://resources.rothamsted.ac.uk/science-stories/aphids-mummies-and-cadavers


Lysiphelbus testaceipes  Photo by J.K.Clark, University of California Statewide IPM Project

Once an aphid, now a hollow mummy; note the neat emergence holes.  Aphid parasitoids are very much tidier than the parasitic lifeform in the classic film Alien 🙂

Another aspect of their life style that makes parasitoids a breed apart from true parasites, is that as well as using aphids as egg laying sites for their larvae, the adults like to snack on them every now and then to help mature more eggs and to keep up their energy levels; sometimes quaintly described as predatism (Flanders, 1953).  Although the parasitoids can make feeding attacks at any time, they appear to feed first and then start laying their eggs (e.g. Collins et al., 1981).

Parasitoids are widely used as biological control agents in glasshouses and other protected environments as they are generally regarded as being more effective than predators (Debach & Rosen, 1991), although there is some support that generalist predators can play a significant part in biological control in the wider environment (Symondson et al., 2002; Gontijo et al., 2015).  That said, aphid parasitoids seem to be fairly host specific in that commercial companies offer specific parasitoid mixtures to control different aphid pest species e.g.  https://www.koppert.com/pests/aphids/product-against/aphipar/ [Note this is NOT an endorsement]. In fact it has been suggested that the relationship between aphids and their parasitoids can be used to clarify aphid taxonomic relationships (Mackauer, 1965). On the other hand, there are very few examples of monophagous aphid parasitoids, most being described as oligophagous (Stary & Rejmanek, 1981).   So given that there is a fair bit of evidence that the parasitoids attacking aphids do show some discrimination in their choice of hosts, how do they find them?

Parasitoids in general were originally thought to be “possessed of an unerring instinct that guided them in their search for hosts” but Cushman (1926) rebutted this idea pointing out that actually the parasitoids first home in on the habitat or food plant that their host lives in and then search for their host (Laing, 1937).   The parasitoids referred to by Cushman and Laing, are however, not parasitoids of aphids, attacking lepidopteran leaf miners and carrion feeding flies respectively, so you might perhaps think that aphid parasitoids could have a different strategy. Although habitat selection by parasitoids of lepidopteran larvae (Thorpe & Caudle, 1938) and sawfly larvae (Monteith, 1955), using olfactory cues of their host’s food plant was confirmed readily easily and early on, the situation with aphids was less clear cut. Manfred Mackauer for example, suggested that aphid parasitoids might be using visual cues, such as leaf deformities or damage to find their aphids hosts (Mackauer, 1965).  The breakthrough came when three cabbage loving entomologists from the USA used an olfactometer to first show that the Braconid parasitoid Diaeretiella rapae, responded positively to the odour of collards (what we in the UK call spring greens) and second to show a very strong preference for them to lay their eggs in the aphid Myzus persicae when it was feeding on crucifers rather than other host plants.  They attributed this to the presence of mustard oil, the chemical that gives cabbages their distinctive taste and suggested that once the aphid host plant was found then the parasitoids used visual cues to find their aphid victims (Read et al., 1970).  Six years later it was firmly established that parasitoids in general used olfactory cues both to locate the habitat of their host (long-range) and then a short-range to find and confirm the identity (contact chemicals) their insect hosts (Vinson, 1976).

It was thought that the aphid parasitoids were chemically ‘conditioned’ during their larval life within the aphid feeding on a host plant and that this influenced their adult host preferences (e.g. Sheehan & Shelton, 1989; Wickremasinghe & Van Emden, 1992).  These, and other similar results, seemed to support the Hopkins host selection principle (Hopkins, 1917) which states that adult preferences are learnt as larvae.  A very neat experiment by van Emden et al., (1996) proved this hypothesis wrong. They transferred aphid mummies from the plant on which they had been parasitized on to another host plant and this changed the preference of the emerging adult, seeming to suggest that this was how aphid parasitoids developed their host preferences.  Now comes the neat, and very tricky part; if however, the parasitoid pupae were removed (very carefully) from the mummy case and reared to adulthood in the absence of a host plant or mummy and kept in a glass tube, the emerging adults showed no preference for particular host plants, clearly showing that adult preferences were  not determined during larval development but ‘conditioned’ by exposure to the external skin of the aphid mummy on emergence (van Emden et al., 1996).  Using aphids reared on an artificial diet (Douloumpaka & van Emden, 2003) showed that the it was very likely that the mother parasitoid leaves a chemical cue in or around the egg(s) she lays and that this is later incorporated into the silk of the parasitoid pupa, thus inducing the host preference seen as an adult.

An additional twist to the story is that male and female parasitoids differ in their responses to odours.  Both sexes of Aphidius uzbekistanicus and A. ervi, parasitoids of cereal aphids in the UK, respond to plant odours, but only females respond to aphids (Powell & Zhi-Li, 1983).  Males of both species are, however, attracted to the odours of their respective females, suggesting the existence of a sex pheromone. The existence of a sex pheromone in aphid parasitoids had been suggested a few years earlier when it was shown that male D. rapae attempted to copulate with filter paper that had had female abdomens crushed on them (Askari & Alisha, 1979).  The existence of sex pheromones in aphid parasitoids has now been shown in several species (e.g. Decker et al., 1993; McNeil & Broduer, 1995).  Strangely, female parasitoids also respond to sex pheromones, but in their case, the sex pheromones of aphids.  It turns out that they ‘parasitise’ aphids in more than one way, they home in on their prey using the aphid sex pheromone and this enables them to find a suitable overwintering host (Hardie et al., 1991).  At other times of the year they also use other aphid indicators; several studies have shown that parasitoids use the presence of aphid honeydew to help them find their hosts (Budenberg, 1990; Bouchard & Cloutier, 1984; Gardner & Dixon, 1985).

Predators of aphids such as ladybirds use chemical markers to warn other ladybirds that they have laid eggs near aphid colonies, thus reducing the chances of cannibalism and competition (e.g. Oliver et al., 2006). Given that the eggs of aphid parasitoids are laid internally, they are in effect invisible, it would make sense if the parasitoids ‘marked’ their hosts in some way to avoid other parasitoids laying their eggs in an already parasitized aphid, superparasitism.  Sure enough, there is some evidence that some adult parasitoids can recognise aphids that already have larval parasitoids developing inside them although they don’t seem to be able to consistently recognise already parasitized aphids until some hours afterward (e.g. Cloutier et al., 1984).  In some cases, it seems that it is the aphid herself that prevents superparasitism by reacting more aggressively towards parasitoids after being attacked once (Gardner & Dixon, 1984) and also by the presence of dried siphuncular secretions on the aphid’s skin (Outreman et al., 2001).  The waxy secretion had an effect for up to a day or so after which the internal changes caused by the developing parasitoid larvae were enough to deter further oviposition attempts.

It is a good thing for the poor aphids that they have such a high reproductive rate, or they would truly be in dire straits.  On the other hand, as exemplified by the words of Jonathan Swift (1733),

“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

there are parasites of parasitoids, the hyperparasites, that help keep the numbers of parasitoids under control, and thus, indirectly, help aphids remain relatively abundant.



Askari, A. & Alisha, A. (1979) Courtship behavior and evidence for a sex pheromone in Diaeretiella rapae (Hymenoptera: Braconidae), the cabbage aphid primary parasitoid. Annals of the Entomological Society of America, 72, 79-750.

Beirne, B.P. (1942) Observations on the life-history of Praon volucre Haliday (Hym.: Braconidae), a parasite of the mealy plum aphis (Hyalopterus arundinis Fab.). Proceedings of the Royal Entomological Society of London, Series A, General Entomology, 17, 42-47.

Boivin, G., Hance, T. & Brodeur, J. (2012) Aphid parasitoids in biological control.  Canadian Journal of Plant Science, 92, 1-12.

Bouchard, Y. & Cloutier, C. (1984) Honeydew as a source of host-searching kairomones for the aphid parasitoid, Aphidius nigripes (Hymenoptera: Aphidiidae).  Canadian Journal of Zoology, 62, 1513-1520.

Budenberg, W.J. (1990) Honeydew as a contact kairomone for aphid parasitoidsEntomologia experimentalis et applicata, 55, 139-148.

Cloutier, C., Dohse, L.A. & Bauduin, F. (1984) Host discrimination in the aphid parasitoid Aphidius nigripes. Canadian Journal of Zoology, 62, 1367-1372.

Collins, M.D., Ward, S.A., & Dixon, A.F.G. (1981) Handling time and the functional response of Aphelinus thomsoni, a predator and parasite of the. Journal of Animal Ecology, 50, 479-487.

Cushman, R.A. (1926) Location of individual hosts versus systematic relation of hots species as a determining factor in parasitic attack. Proceedings of the Entomological Society of Washington, 28, 5-6.

Darwin, E. (1800) Phytologia: or The Philosophy of Agriculture and Gardening. P. Byrne, Grafton Street, London.

Debach, P. & Rosen, D. (1991) Biological Control by Natural Enemies, Cambridge University Press, New York.

Decker, U.M., Powell, W. & Clark, S.J. (1993) Sex pheromone in the cereal aphid parasitoids Praon volucre and Aphidius rhopalosiphiEntomologia experimentalis et applicata, 69, 33-39.

Douloumpaka, S. & van Emden, H.F. (2003) A maternal influence on the conditioning to plant cues of Aphidius colemani Viereck, parasitizing the aphid Mysuze persicae SulzerPhysiological Entomology, 28, 108-113.

Flanders, S.E. (1953) Predation by the adult Hymenopteran parasite and its role in biological control. Journal of Economic Entomology, 46, 541-544.

Gardner, S.M. & Dixon, A.F.G. (1984) Limitation of superparasitism by Aphidius rhopalosiphi: a consequence of aphid defensive behaviour. Ecological Entomology, 9, 149-155.

Gardner, S.M & Dixon, A.F.G. (1985) Plant structure and foraging success of Aphidius rhopalosiphi (Hymenoptera: Aphidiidae).  Ecological Entomology, 10, 171-179.

Gaston, K.J., Gauld, I.D. & Hanson, P. (1996) The size and composition of the hymenopteran fauna of Costa Rica.  Journal of Biogeography, 23, 105-113.

Griffiths, D.C. (1960) The behaviour and specificity of Monoctonus paldum Marshall (Hym., Braconidae), a parasite of Nasonovia ribis-nigbi (Mosley) on lettuce. Bulletin of Entomological Research, 51, 303-319.

Hardie, J., Nottingham, S.F., Powell, W. & Wadhams, L.J. (1991) Synthetic aphid sex pheromone lures female parasitoids.  Entomologia experimentalis et applciata, 61, 97-99.

Harrington, R. (1994) Aphid layer. Antenna18, 50-51.

Hopkins, A.D. (1917) Contribution to discussion.  Journal of Economic Entomology, 10, 92-93.

Holler, C. (1991) Evidence for the existence of a species closely related to the cereal aphid parasitoid Aphidius rhopalosiphi De Stefani-Perez based on host ranges, morphological characters, isoelectric focusing banding patterns, cross-breeding experiments and sex pheromone specificities (Hymenoptera, Braconidae, Aphidiinae. Systematic Entomology, 16, 15-28.


Laing, J. (1937) Host-finding byinsect parasites 1. Observations on the finding of hosts by Alysia manducator, Mormoniella vitripennis and Trichogramma evanescensJournal of Animal Ecology, 6, 298-317.

Mackauer, M. (1965) Parasitological data as an aid in aphid classification. Canadian Entomologist, 97, 1016-1024.

McNeil, J.N. & Brodeur, J. (1995) Pheromone-mediated mating in the aphid parasitoid, Aphidius nigripesJournal of Chemical Ecology, 21, 959-972.

Monteith, L.G. (1955) Host preferences of Drino bohemica Mesn. (Diptera; Tachnidae) with particular reference to olfactory responses.  Canadian Entomologist, 87, 509-530.

Oliver, T.H., Timms, J.E.L., Taylor, A. & Leather, S.R. (2006) Oviposition responses to patch quality in the larch ladybird Aphidecta obliterata (Coleoptera: Coccinellidae): effects of aphid density, and con- and heterospecific tracks. Bulletin of Entomological Research, 96, 25-34.

Outreman, Y., Le Ralec, A., Plantegenest, M., Chaubet, B, & Pierre, J.S. (2001) Superparasitism limitation in an aphid parasitoid: cornicle secretion avoidance and host discrimination ability. Journal of Insect Physiology, 47, 339-348.

Powell, W. & Zhi-Li, Z. (1983) The reactions of two cereal aphid parasitoids, Aphidius uzbekistanicus and A. ervi to host aphids and their food-plants.  Physiological Entomology, 8, 439-443.

Reuter, O.M. (1913). Lebensgewohnheiten und Instinkte der Insekten (Berlin: Friendlander).

Stary, P. & Rejmanek, M. (1981) Number of parasitoids per host in different systematic groups of aphids: The implications for introduction strategy in biological control (Homoptera: Aphidoidea; Hymenoptera: Aphidiidae). Entomologica Scandinavica, Suppl. 15, 341-351.

Riley, W.A. (1931) Erasmus Darwin and the biologic* control of insects. Science, 73, 475-476.

Sheehan, W. & Shelton, A.M. (1989) The role of experience in plant foraging by the aphid parasitoid Diaeretiella rapae (Hymenoptera: Aphidiidae).  Journal of Insect Behavior, 2, 743-759.

Symondson, W.O.C., Sunderland, K.D., & Greenstone, M.H. (2002) Can generalist predators be effective bicontrol agents? Annual Review of Entomology, 47, 561-594.

Thompson, W.R. (1930) The principles of biological control. Annals of Applied Biology, 17, 306-338.

Thorpe, W.H. & Caudle, H.B. (1938) A study of the olfactory responses of insect parasites to the food plant of their host.  Parasitology, 30, 523-528.

Van Emden, H.F., Spongal, B., Wagner, E., Baker, T., Ganguly, S. & Douloumpaka, S. (1996) Hopkins’ ‘host selection principle’, another nail in its coffin.  Physiological Entomology, 21, 325-328.

Van Lenteren, J.C. (2012) The state of commercial augmentative biological control: plenty of natural enemies, but a frustrating lack of uptake. BioControl, 57, 1-20.

Van Lenteren, J.C. & Woets, J. (1988) Biological and integrated control in greenhouses.  Annual Review of Entomology, 33, 239-269.

Vinson, S.B. (1976) Host selection by insect parasitoids.  Annual Review of Entomology, 21, 109-133.

Wheeler, W.M. (1922). Social life among the insects: Lecture II. Wasps solitary and social. Scientific Monthly, 15, 68-88.

Wheeler, W.M. (1928) Foibles of Insects and Men.  Alfred Knopf, New York

Wickremasinghe, M.G.V. & Van Emden, H.F. (1992) Reactions of adult female parasitoids, particularly Aphidius rhopalosiphi, to volatile chemical cues from the host plants of their aphid prey. Physiological Entomology, 17, 207-304.

*This is how he spelt it; not a mistake on my part J


Filed under Aphidology, Aphids

What use are bedbugs?

As an entomologist, I have, over the years, become used to being asked by non-entomologists “What use are wasps?” My wife and mother-in-law being frequent interrogators. What they actually mean is “What use are Vespids?”, in particular what the Americans call yellow jackets, and their ilk.


Not a bed bug – The European wasp Vespula germanica https://en.wikipedia.org/wiki/Yellow_jacket#/media/File:European_wasp_white_bg.jpg

I now have a well-polished response where I explain that wasps are beneficial insects keeping the caterpillars that eat their garden plants under control and that the occasional hole they make in my interlocutor’s soft fruit is just payment for the job they are doing. I am not saying that this answer always satisfies them, especially if their plums have been devastated, but at least they agree that there is some justification for their existence.

Unfortunately I now have a new question to answer. Last summer (2015) my wife and I took our usual holiday to France. We put our car on the Brittany Ferry, m/V Mont St Michel, and after a drink in the bar retired to our cabin, 9108 in case you want to avoid it, me in Bunk A my wife in Bunk B.


Sailing in comfort?

The next morning my wife was not a happy lady and the question I now have to answer is “What use are bedbugs?”!


The bed bug culprit, Cimex lectularius and victim

I have in the past made a convincing case (well I think so) for the usefulness of mosquitoes compared with pandas, but I suspected that making a case for the usefulness of bed bugs that would satisfy my wife, might be more difficult.

Even the Encyclopaedia of Life has nothing good to say about bed bugs. For evolutionary biologists, ecologists and entomologists, bed bugs are very useful in allowing us to relate horror stories about non-conventional sex. Male bed bugs favour a very robust approach to reproduction, as they indulge in what is somewhat coyly termed ‘traumatic insemination (Reinhardt & Siva-Jothy, 2007). Basically they don’t bother with the female genital opening, they mount the female and search for a pouch (Organ of Ribaga or ectospermalage) on the underside of the female (they have been known to mount other males) which they pierce with their intromittent organ (penis) and into which they release their sperm. The spermatozoa then migrate through the body of the female to the oviduct, taking from 2-10 hours to do so (Cragg, 1920). Apparently males never use the genital tract for insemination (Reinhardt & Siva-Jothy, (2007). This somewhat unconventional approach to mating has, as you might expect, harmful effects on the female, with life spans being reduced by about 30% or even causing death (Morrow & Anrnqvist, 2003), multiple matings can be particularly damaging (Mellanby, 1939).

For their human hosts, bed bugs can have a number of unpleasant effects (Reinhardt & Siva-Jothy, 2007), ranging from psychological distress, allergic reactions as in the case of my wife, secondary infections and economic costs, especially if you are an hotelier. All in all, not a very promising candidate for being useful to us humans 🙂 I was beginning to feel that bed bugs had no redeeming features and that if there was a coordinated campaign calling for the destruction of the entire species, I would be unable to defend them. Then a former student came to their rescue. In desperation, I had emailed Mike Siva-Jothy at Sheffield University who has worked on bed bugs since the late 1990s. He passed my email on to Sophie Evison (a former MSc student of mine) and she came to their rescue as follows

I’ve had a think about this, and I think there are two approaches: 1. dazzle them with the traumatic insemination story or 2. Forensics. I’ve not looked into it, but I’m pretty sure a situation could arise where the contents of a bedbug could drop someone in it, or even perhaps prove an alibi! Do you think your wife would accept that as reasonable?”

I was pretty certain that my wife would not be happy with option 1, but felt that option 2 was definitely worth following up. I very quickly found a paper (Szalanski et al, 2006) in which the possibility of testing the DNA of the blood found in bed bugs was suggested as having a possible forensic application. According to Wikipedia (well if it is good enough for my students) there has already been some success in real life using this very approach. I was now pretty confident that I had found an answer to the question posed by my wife. To be fair, I tested both of them out on her. As predicted, she did not buy the evolutionary biology aspect of the traumatic insemination story as being of any use, but as a great fan of the CSI and NCIS TV shows, she has grudgingly accepted that there is indeed a use for bed bugs! I just hope that all the bed bugs out there appreciate all the effort I have gone to on their behalf 🙂 If anyone else has further suggestions please let me know.


Cragg, F.W. (1920) Further observations on the reproductive system of Cimex, with special reference to the behaviour of the spermatozoa. Indian Journal of Medical Research, 8, 32-79

Mellanby, K. (1939) Fertilization and egg production in the bed-bug Cimex lectularius L. Parasitology, 31, 193-199

Morrow, E.H. & Arnqvist, G. (2003) Costly traumatic insemination and female counter-adaptation in bed bugs. Proceedings of the Royal Society of London Series B, 270, 2377-2381

Reinhardt, K. & Siva-Jothy, M. (2007) Biology of the bed bugs (Cimicidae). Annual Review of Entomology, 52, 351-374

Szalanski, A.L., J.W. Austin, J.A. McKern, C.D. Steelman, D.M. Miller, and R.E. Gold. 2006. Isolation and characterization of human DNA from bed bug, Cimex lectularius L., (Hemiptera: Cimicidae) blood meals. Journal of Agricultural and Urban Entomology, 23, 189-194.




Filed under Bugbears, EntoNotes, Uncategorized

The Curious Case of the Shark-finned Aphid

The large (giant) willow aphid, Tuberlolachnus salignus, is, in my opinion, one of the world’s greatest unsolved mysteries.  This aphid is sometimes regarded as being the largest aphid in the world.  It can reach a length of 5 mm, can weigh up to 13 mg as an adult and the new-born nymphs weigh about 0.25 mg (Hargreaves & Llewellyn, 1978).  You can get an idea of how big it is from the picture below.

willow aphid on finger


This is pretty big for an aphid, although not quite as big as one of my former PhD students (Tilly Collins) liked to pretend!  The picture below used to appear on her website and was the envy of a number of Texan entomologists.  Tuberolachnus salignus, as you might expect, since it feeds through the bark and not on leaves, has rather a long set of stylets, up to  1.8 mm, more than a third of it’s body length (Mittler, 1957).

tilly on aphid

This picture emphasises the first mystery: what is the function of the dorsal tubercle, which so closely resembles a rose thorn, or to me, a shark’s fin.  Nobody knows.  Is it defensive? Unlikely, since T. salignus being a willow feeder is stuffed full of nasty chemicals and very few predators seem to want, or be able to feed on it.  They feed in large aggregations on the stems of their willow tree hosts and can have serious effects on tree growth (Collins et al., 2001).  As the aphids produce a lot of honeydew, they are often ant-attended  (Collins & Leather, 2002) and these also deter potential predators.  In fact the aphid colonies produce so much honeydew in the summer that they attract huge numbers of vespid wasps that are in search of energy-rich sugar sources at that time of year.  These too are likely to make potential predators and parasitoids think twice about approaching the aphids.


Photograph courtesy Dr Tilly Collins

The wasps also cause a problem for researchers and when Tilly was doing her PhD, she used to have to confine her fieldwork to those times of day when the wasps were not around.   In addition, if you crush one of the aphids you will discover that it stains your fingers bright orange and that this stain will last several days if you don’t try too hard to wash it off.  If you get this aphid ‘blood’ on your clothes they will be permanently marked and Tilly used to say that she ought to be paid an extra clothing allowance.

Tuberolachnus salignus, is as far as we can tell, anholocyclic, no males have been recorded and no matter how hard people have tried to induce the formation of males and sexual females, they have not been successful.  This is however, not the second mystery.  The mystery is that every year, in about February, it does a disappearing act and for about four months its whereabouts remain a mystery (Collins et al., 2001).  So we have an aphid that spends a substantial period of the year feeding on willow trees without leaves and then in the spring when most aphids are hatching from their eggs to take advantage of the spring flush, T. salignus disappears!  Does it go underground?  If so, what plant is it feeding on and why leave the willows when their sap is rising and soluble nitrogen is readily available?

So here is a challenge for all entomological detectives out there.  What is the function of the dorsal tubercle and where does T. salignus go for the spring break?

Truly a remarkable aphid and two mysteries that I would dearly love to know the answers to and yet another reason why I love aphids so much.

Collins, C.M. & Leather, S.R. (2002) Ant-mediated dispersal of the black willow aphid Pterocomma salicis L.; does the ant Lasius niger L. judge aphid-host quality. Ecological Entomology, 27, 238-241. http://onlinelibrary.wiley.com/doi/10.1046/j.1365-2311.2002.00390.x/full

Collins, C. M., Rosado, R. G. & Leather, S. R. (2001). The impact of the aphids Tuberloachnus salignus and Pterocomma salicis on willow trees. Annals of Applied Biology 138, 133-140 http://onlinelibrary.wiley.com/doi/10.1111/j.1744-7348.2001.tb00095.x/abstract.

Hargreaves, C. E. M. & Llewellyn, M. (1978). The ecological energetics of the willow aphid, Tuberolachnus salignus:the influence of aphid Journal of Animal Ecology, 47, 605-613. http://www.jstor.org/discover/10.2307/3804?uid=3738032&uid=2&uid=4&sid=21101920521473

Mittler, T. E. (1957). Studies on the feeding and nutrition of Tuberolachnus salignus (Gmehn) (Homoptera, Aphididae). I. The uptake of phloem sap. Journal of  Experimental Biology, 34, 334-341  http://jeb.biologists.org/content/34/3/334.full.pdf

Other resources




Filed under Aphidology, Aphids, EntoNotes, Unsolved mysteries