Tag Archives: Rothamsted Research

Pick and mix 16 – more links to check out

Wise words from the Oxford University Museum of Natural History

If you live in the UK and like trees in your garden, here are some suggestions of native species to plant – all are good for insects and birds

On managing your urban garden as a productive ecosystem

An excellent resource of historical research done at Rothamsted Research Station – this section all about bees

Still more on bees, this time how bees that are feeling unwell change their diets to fight of infection

More and more species being discovered yet taxonomists are an endangered species themselves; they deserve our respect and more funding

They may be unwanted neighbours but these are beautiful pictures from Gil Wizen

Maria Sibylla Merian, a prodigy from the 17th Century; artist, naturalist and entomologist – remarkable achievements

Many animals, including insects, can count

If you have ever wondered why entomologist kill insects and have 28 minutes to spare listen to this

An irreverent obituary of legendary French chef Paul Bocuse

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Insects in flight – whatever happened to the splatometer?

I have been musing about extinctions and shifting baselines for a while now; BREXIT and an article by Simon Barnes in the Sunday Times magazine (3rd September 2016) finally prompted me to actually put fingers to keyboard.  I fear that BREXIT will result in even more environmental damage than our successive governments have caused already.  They have done a pretty good job of ignoring environmental issues and scientific advice (badgers) even when ‘hindered’ by what they have considered restrictive European legislation and now that we head into BREXIT with a government not renowned for its care for the environment I become increasing fearful for the environment. Remember who it was who restructured English Nature into the now fairly toothless Natural England, because they didn’t like the advice they were being given and whose government was it who, rather than keep beaches up to Blue Flag standard decided to reclassify long-established resort beaches as not officially designated swimming beaches?  And, just to add this list of atrocities against the environment, we now see our precious ‘green belt’ being attacked.

My generation is liable to wax lyrical about the clouds of butterflies that surrounded us as we played very non PC cowboys and Indians outside with our friends in the glorious sunshine.  We can also fondly reminisce about the hordes of moths that used to commit suicide in the lamp fittings or beat fruitlessly against the sitting room windows at night.  The emptying of the lamp bowl was a weekly ceremony in our house.  We also remember, less fondly, having to earn our pocket-money by cleaning our father’s cars, laboriously scraping the smeared bodies of small flies from windscreens, headlamps and radiator grilles on a Saturday morning.  A few years later as students, those of us lucky enough to own a car, remember the hard to wash away red smears left by the eyes of countless Bibionid (St Mark’s) flies, as they crashed into our windscreens.


Typical Bibionid – note the red eyes; designed specially to make a mess on your windscreen 🙂 https://picasaweb.google.com/lh/photo/GBgoGHhRbj-eUUF9SxZ4s9MTjNZETYmyPJy0liipFm0?feat=embedwebsite

Are these memories real or are we looking back at the past through those rose-tinted glasses that only show the sunny days when we lounged on grassy banks listening to In the Summertime and blank out the days we were confined to the sitting room table playing board games?

We have reliable and robust long-term data sets showing the declines of butterflies and moths over the last half-century or so (Thomas, 2005; Fox, 2013) and stories about this worrying trend attract a lot of media attention. On a less scientific note, I certainly do not find myself sweeping up piles of dead moths from around bedside lamps or extricating them from the many spider webs that decorate our house.  Other charismatic groups, such as the dragonflies and damselflies are also in decline (Clausnitzer et al., 2009) as are the ubiquitous, and equally charismatic ground beetles (carabids) (Brooks et al., 2012).  But what about other insects, are they too on the way out?  A remarkable 42-year data set looking at the invertebrates found in cereal fields in southern England (Ewald et al., 2015) found that of the 26 invertebrate taxa studied less than half showed a decrease in abundance; e.g. spiders, Braconid parasitic wasps, carabid beetles, Tachyporus beetles, Enicmus (scavenger beetles), Cryptophagid fungus beetles, leaf mining flies (Agromyzids), Drosophila, Lonchopteridae (pointed wing flies), and surprisingly, or perhaps not, aphids.  The others showed no consistent patterns although bugs, excluding aphids, increased over the study period.  Cereal fields are of course not a natural habitat and are intensely managed, with various pesticides being applied, so are perhaps not likely to be the most biodiverse or representative habitats to be found in the UK.

But what about the car-smearing insects, the flies, aphids and other flying insects?  Have they declined as dramatically?  My first thought was that I certainly don’t ‘collect’ as many insects on my car as I used to, but is there any concrete evidence to support the idea of a decline in their abundance.  After all, there has been a big change in the shape of cars since the 1970s.


Top row – cars from 1970, including the classic Morris 1000 Traveller that my Dad owned and I had to wash on Saturdays.

Bottom row the cars of today, sleek rounded and all looking the same.


Cars were  much more angular then, than they are now, so perhaps the aerodynamics of today’s cars filter the insects away from the windscreen to safety? But how do you test that?  Then I remembered that the RSPB had once run a survey to address this very point.  Sure enough I found it on the internet, the Big Bug Count 2004, organised by the RSPB.  I was very surprised to find that it happened more than a decade ago, I hadn’t thought it was that long ago, but that is what age does to you 🙂


The “Splatometer” as designed by the RSPB

The idea, which was quite cool, was to get standardised counts of insect impacts on car number platesThe results were thought to be very low as the quote below shows, but on what evidence was this based?

“Using a cardboard counting-grid dubbed the “splatometer”, they recorded 324,814 “splats”, an average of only one squashed insect every five miles. In the summers of 30-odd years ago, car bonnets and windscreens would quickly become encrusted with tiny bodies.”  “Many people were astonished by how few insects they splatted,” the survey’s co-ordinator Richard Bashford, said.

Unfortunately despite the wide reporting in the press at the time, the RSPB did not repeat the exercise.  A great shame, as their Big Garden Birdwatch is very successful and gathers useful data.   So what scientific evidence do we have for a decline in these less charismatic insects?  Almost a hundred years ago, Bibionid flies were regarded as a major pest (Morris, 1921) and forty years ago it was possible to catch almost 70 000 adults in a four week period from one field in southern England (Darcy-Burt & Blackshaw, 1987).   Both these observations suggest that in the past Bibionids were very common.  It is still possible to pluck adult Bibionids out of the air (they are very slow, clumsy fliers) in Spring, but if asked I would definitely say that they are not as common as they were when I was a student.  But as Deming once said, “Without data, you’re just another person with an opinion.”  In the UK we are fortunate that a long-term source of insect data exists, courtesy of Rothamsted Research, the longest running agricultural research station in the world.  Data have been collected from a nationwide network of suction and light traps for more than 50 years (Storkey et al., 2016).   Most of the publications arising from the survey have tended to focus on aphids (Bell et al., 2015) and moths (Conrad et al., 2004), although the traps, do of course, catch many other types of insect (Knowler et al., 2016).  Fortuitously, since I was interested in the Bibionids I came across a paper that dealt with them, and other insects likely to make an impact on cars and splatometers (Shortall et al., 2009).  The only downside of their paper was that they only looked at data from four of the Rothamsted Suction Traps, all from the southern part of the UK, which was a little disappointing.


Location and results of the suction traps analysed by Shortall et al. (2009).

Only three of the trap showed downward trends in insect biomass over the 30 years (1973-2002) analysed of which only the Hereford trap showed a significant decline.  So we are really none the wiser; the two studies that focus on a wider range of insect groups (Shortall et al., 2009; Ewald et al., 2015) do not give us a clear indication of insect decline.   On the other hand, both studies are limited in their geographic coverage; we do not know how representative the results are of the whole country.

What a shame the RSPB stopped collecting ‘splatometer’ data, we would now have a half-decent time series on which to back-up or contradict our memories of those buzzing summers of the past.

Post script

After posting this I came across this paper based on Canadian research which shows that many pollinators, possibly billions are killed by vehicles every year.  This reduction in insect numbers and biomass has also been reported in Germany.


Bell, J.R., Alderson, L., Izera, D., Kruger, T., Parker, S., Pickup, J., Shortall, C.R., Taylor, M.S., Verrier, P. & Harrington, R. (2015) Long-term phenological trends, species accumulation rates, aphid traits and climate: five decades of change in migrating aphids.  Journal of Animal Ecology, 84, 21-34.

Brooks, D.R., Bater, J.E., Clark, S.J., Montoth, D.J., Andrews, C., Corbett, S.J., Beaumont, D.A., & Chapman, J.W. (2012) Large carabid beetle declines in a United Kingdom monitoring network increases evidence for a widespread loss of insect biodiversity. Journal of Applied Ecology, 49, 1009-1019.

Clausnitzer, V., Kalkman, V.J., Ram, M., Collen, B., Baillie, J.E.M., Bedjanic, M., Darwall, W.R.T., Dijkstra, K.D.B., Dow, R., Hawking, J., Karube, H., Malikova, E., Paulson, D., Schutte, K., Suhling, F., Villaneuva, R.J., von Ellenrieder, N. & Wilson, K. (2009)  Odonata enter the biodiversity crisis debate: the first global assessment of an insect group.  Biological Conservation, 142, 1864-1869.

Conrad, K.F., Woiwod, I.P., Parsons, M., Fox, R. & Warren, M.S. (2004) Long-term population trends in widespread British moths.  Journal of Insect Conservation, 8, 119-136.

Darcy-Burt, S. & Blackshaw, R.P. (1987) Effects of trap design on catches of grassland Bibionidae (Diptera: Nematocera).  Bulletin of Entomological Research, 77, 309-315.

Ewald, J., Wheatley, C.J., Aebsicher, N.J., Moreby, S.J., Duffield, S.J., Crick, H.Q.P., & Morecroft, M.B. (2015) Influences of extreme weather, climate and pesticide use on invertebrates in cereal fields over 42 years. Global Change Biology, 21, 3931-3950.

Fox, R. (2013) The decline of moths in Great Britain: a review of possible causes. Insect Conservation & Diversity, 6, 5-19.

Knowler, J.T., Flint, P.W.H., & Flint, S. (2016) Trichoptera (Caddisflies) caught by the Rothamsted Light Trap at Rowardennan, Loch Lomondside throughout 2009. The Glasgow Naturalist, 26, 35-42.

Morris, H.M. (1921)  The larval and pupal stages of the Bibionidae.  Bulletin of Entomological Research, 12, 221-232.

Shortall, C.R., Moore, A., Smith, E., Hall, M.J. Woiwod, I.P. & Harrington, R. (2009)  Long-term changes in the abundance of flying insects.  Insect Conservation & Diversity, 2, 251-260.

Storkey, J., MacDonald, A.J., Bell, J.R., Clark, I.M., Gregory, A.S., Hawkins, N. J., Hirsch, P.R., Todman, L.C. & Whitmore, A.P. (2016)  Chapter One – the unique contribution of Rothamsted to ecological research at large temporal scales Advances in Ecological Research, 55, 3-42.

Thomas, J.A. (2005) Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups.  Philosophical Transactions of the Royal Society B, 360, 339-357


Filed under EntoNotes, Uncategorized

The Scent of Fear – the aphid alarm pheromone

We are all familiar with the effects of epinephrine (adrenaline) and norepinephrine (noradrenaline) on us when placed in a position of stress, such as public speaking or even worse danger.  We flush, shake, our heart rate accelerates and many of us we begin to sweat profusely, thus visibly advertising our distress; sometimes embarrassingly so

Sweating nervously


if we have an antiperspirant  fail and happen to be wearing a dark shirt.



Those seeing these symptoms may feel a degree of sympathy for the victim, but do not usually flee the scene, although they may sometimes feel tempted to do so.

The case with aphids is very different.   Aphids, when perceiving a threat to their neighbours by a predator or parasite, flee the scene rapidly, by flight, if winged, on foot if not, or even by leaping from their host-plant to the ground below.  The pea aphid, Acyrthosiphon pisum walks away or drops from their plant (Clegg & Barlow, 1982) as does the rose-grain aphid, Metopolophium dirhodum (Larsen, 1988).  This may seem a risky move since it seems that about 10% of all aphids that fall from their host plant don’t manage to get back (Sunderland et al., 1986), but for a clonal organism the risk is obviously worth it.

So how does this communal fright and flight response come about?  Most aphids have a pair of siphunculi or cornicles at the rear of their abdomen.  These vary in size and shape, in some aphids being long and slender, in others short and stubby and in yet others reduced to a shallow indentation (pore) or in a few species, totally absent.


The role of aphid siphunculi has been debated since the days of Linneaus and Reaumur who considered them to be the source of honeydew (Hottes, 1928).  Hottes himself in a comprehensive review of the various theories put forward for the function of the siphunculi, dismissed the defence theory of Busgen (1891) and plumped for an excretory role, although he did suggest that volatile substances were produced by the siphunculi in addition to the waxy visible drops.    By the middle of the last century it was generally accepted that the siphunculi were involved in defence, but in a purely physical way, in that the waxy exudate was used to deter or disable the attacking predators or parasites (e.g. Dixon, 1958; Edwards, 1966).  At about the same time, the chemical composition of the visible exudate was confirmed as being primarily triglycerides with myristic acid being the major fatty acid present (Strong, 1966).

Oleander Aphid- Milkweed-- Mark Bower - 1 - 1

Aphis nerii siphuncular exudate.  http://springfieldmn.blogspot.co.uk/2014/08/aphid-cornicles.html



Hawthorn-parsley aphid Dysaphis apiifolia producing sipuncular exudates whilst under attack by a parasitic wasp.  Many thanks to Tom Pope for permission to use this clip.

In 1968 an alarm pheromone was identified and isolated from the cotton stainer, Dysdercus intermedius (Calam & Youdeowei, 1968) so it was not surprising that attention should be focused on aphids, many of which show a similar group dispersive behaviour when a predator approaches them.   The aphid alarm pheromone (E)-β-farnesene was, however, not formally identified until  1972 (Bowers et al, 1971), although Maria Dahl had demonstrated the  previous year that a solution made from crushed aphids would cause an alarm response in other aphids of the same and different species (Dahl, 1971). Unsurprisingly, as during the 1970s and 1980s scientists from the USA were notorious for only citing papers written in English, Bowers et al. (1972), failed to cite her in their paper, instead citing two other American authors (Kislow & Edwards, 1972).

This discovery resulted in a flurry of papers from around the world as insect physiologists vied to be the first to isolate alarm pheromone from different aphid species (e.g. Weintjens et al., 1973; Montgomery & Nault, 1977; Wohlers, 1980).  There were also more ecological studies such as that examining the way alarm pheromone in ant-attended aphids enhances the relationship between them and their ant farmers (Nault et al., 1976) thus acting as a synomone (Nordlund & Lewis, 1976).  As time has gone on the interest in aphid alarm pheromone has remained unabated with new twists and surprises being discovered.  For example, as well as stimulating the escape response, the alarm pheromone also stimulates those surviving pea aphids to produce winged offspring thus facilitating future long-distance dispersal away from the predators (Kunert et al., 2005).  Aphid alarm pheromone can also act to help natural enemies find their aphid prey (e.g. Micha & Wyss, 1996), in this case acting as a kairomone.

The use of sex pheromones in integrated pest management is well established (Witzgall et al., 2010) and works very effectively in most cases. More recently, researchers at Rothamsted Research have courted controversy by trialing GM wheat that has been engineered to produce aphid alarm pheromone.  Many entomologists, including me, although finding the concept (Yu et al., 2012) interesting, doubt that it will work in a field situation.  I can certainly see a role for using alarm pheromones as an alternative to conventional chemical control of insect pests and it will be interesting to see if it will prove as effective as using sex pheromones.



Bowers, W. S., Nault, L. R., Webb, R. E. & Dutky, S. R. (1972). Aphid alarm pheromone: isolation, identification, synthesis. Science 177, 1121-1122.

Busgen, M. (1891)  Der Honigtau. Biologische Studien an Pflanzen und Pflanzenläusen.  Jenaische Zeitschrift für Naturwissenschaft, 25, 339-428

Calam, D.H. & Youdeowei, A. (1968) Identification and functions of secretion from the posterior scent gland of fifth instar larva of the bug Dysdercus intermedius. Journal of Insect Physiology, 14, 1147-1158

Clegg, J.M. & Barlow, C.A. (1982) Escape behaviour of the pea aphid, Acyrthosiphon pisum (Harris) in response to alarm pheromone and vibration. Canadian Journal of Zoology, 60, 2245-2252.

Dahl, M. L. (1971). Über einen Schreckstoft bei Aphiden. Deutsche Entomologische Zeitschrift 18, 121-128.

Dixon, A. F. G. (1958). The escape responses shown by certain aphids to the presence of the coccinellid Adalia decempunctata (L.). Transactions of the Royal Entomological Society London,110, 319-334.

Dixon, A. F. G. (1958). The protective function of the siphunculi of the nettle aphid, Microlophium evansi (Theob.). Entomologist’s Monthly Magazine, 94, 8.

Edwards, J.S. (1966) Defence by smear: supercooling in the cornicle wax of aphids.  Nature,  211, 73-74.

Hottes, F. C. (1928). Concerning the structure, function, and origin of the cornicles of the family Aphididae. Proceedings of the Biological Society of Washington 41, 71-84.

Kislow, C.J. & Edwards, L.J. (1972)  Repellent odour in aphids.  Nature, 235, 108-109.

Kunert, G., Otto, S., Rose, U.S.R., Gershenzon, J., & Weisser, W.W. (2005) Alarm pheromone mediates production of winged dispersal morphs in aphids. Ecology Letters, 8, 596-603.

Larsen, K.S. (1988) Responses of different age classes of the rose-grain aphid, Metopolophium dirhodum (Wlk.) to attack by a simulated predator.  Journal of Applied Entomology, 105, 455-459.

Montgomery, M. E. & Nault, L. R. (1977). Comparative response of aphids to the alarm pheromone, (E)-B-farnesene. Entomologia experimentalis et applicata 22, 236-242.

Micha, S.G. & Wyss, U. (1996)  Aphid alarm pheromone (E)-B-farnesene: a host finding kairomone for the aphid primary parasitoid Aphidius uzbekistanicus (Hymenoptera: Aphidinae).  Chemoecology, 7, 132-139

Nault, L. R., Montgomery, M. E. & Bowers, W. S. (1976). Ant-aphid associations: role of aphid alarm pheromone. Science 192, 1349-1351.

Nordlund, D. A. & Lewis, W. J. (1976). Terminology of chemical releasing stimuli in intraspecific and interspecific interactions. Journal of Chemical Ecology 2, 211-220.

Strong, F.E. (1966)  Observations on aphid cornicle secretions.  Annals of the Entomological Society of America, 60, 668-673.

Sunderland, K.D., Fraser, A.M., & Dixon, A.F.G. (1986) Field and laboratory studies on money spiders (Linyphiids) as predators of cereal aphids. Journal of Applied Ecology, 23, 433-447.

Wientjens, W. H., Lakwijk, C. J. M., & Van Der Marel, T. (1973). Alarm pheromone of grain aphids. Experientia, 29, 658-660.

Wohlers, P. (1980). Die fluchtreaktion der erbsenlaus Acyrthosiphon pisum Aausgelöst durch alarmpheromon und zusätzliche reize. Entomologia experimentalis et applicata 27, 156-168.

Witzgall, P., Kirsch, P. & Cork, A. (2010) Sex pheromones and their impact on pest management. Journal of Chemical Ecology, 36, 80-100.

Yu, X.D, Pickett, J., Ma, Y.Z., Bruce, T., Napier, J., Jones, H.D. & Xia, L.Q. (2012)  Metabolic engineering of plant-derived (E)-β-farnesene synthase genes for a novel type of aphid-resistant genetically modified crop plants.  Journal of Integrative Plant Biology, 54, 282-299.


Post Script

A brief guide to mones

An allomone is any chemical substance produced and released by an individual of one species that affects the behaviour of a member of another species to the benefit of the originator but not the receiver e.g. the ability of some plants to release aphid alarm pheromen and thus deter aphids form landing on them.

An apneumone is any substance produced by nonliving material that benefits a recipient species but is detrimental to a different species associated with the non-living material

A kairomone is a semiochemical, emitted by an organism, which mediates interspecific interactions in a way that benefits an individual of another species which receives it, without benefiting the emitter.  For a detailed critique of the term kairomone see Ruther et al. (2002).

A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behaviour of the receiving individual e.g. alarm pheromones, food trail pheromones and sex pheromones.

A synomone is a substance produced by an individual of one species that benefits both the producer and the recipient which is of a different species.  An example is the release of chemical elicitors by plants that attract entomophagous insects when they are attacked by herbivores.



Filed under Aphidology, Aphids