Tag Archives: sycamore aphid

Insectageddon, Ecological Armageddon, Global insect Apocalypse – why we need sustained long-term funding

“To him that countryside, largely unspoiled in his early days, was an inexhaustible source of delight and a subject of endless study and mediation…And as the years passed and the countryside faded away under the withering touch of mechanical transport, that knowledge grew more and more precious. Now, the dwindling remnants had to be sought and found with considered judgement and their scanty material eked out with detail from the stores of the remembered past”  R Austin Freeman The Jacob Street Mystery (1942)

The recent release of the IPBES report highlighting the significant global declines in biodiversity has prompted me to revisit the “Insectageddon” debate, some of the ramifications of which I wrote about earlier this year.


Summary from the IPBES report – note that even a well-known group like dragonflies is quite data deficient*.

Insects may be in decline, but papers about their decline have been around for almost twenty years and even more are appearing as we entomologists begin to hope that people may at last be beginning to listen to us.

A selection of some of the many papers that have documented insect declines over the last several years.

Using the now infamous search term “insect decline” in the Google Trends function I was not surprised to see the steep increase since 2016, as 2017 was the year in which the paper reporting  the 75% decline in flying insect biomass appeared (Hallmann et al., 2017), but I was intrigued by what appeared to have been a peak in mentions since 2004.

Google Trends using the phrase insect decline – last data point is 2019 at the time of writing

I wondered what caused the peak in 2004, so using the same key words as Sánchez-Bayo & Wyckhuys (2019), checked Google Scholar and Web of Science to see if I could track down a paper that might have caused a media splash at the time.  I also checked 2003, in case there was a delay in reporting. To my surprise I couldn’t find anything relevant in 2004, but 2003 threw up three papers (Hopkins & Freckleton, 2002; Kotze & O’Hara, 2003; Shreeve & Dennis, 2003).  The first was about the decline of taxonomists, which although a serious problem is unlikely to have generated that much attention, the other two were about long-term declines in Carabid beetles (Kotze & O’Hara, 2003) and the third about the decline of French butterflies (Shreeve & Dennis, 2003) which again, I suspect were probably not high enough profile to generate a big splash.  I was puzzled but then I thought, why not just put it into Google with the date 2004, and sure enough it directed me to a Nature News item with the headline Insect deaths add to extinction fears, which in turn led me to Thomas et al., (2004) which I am pretty certain generated the peak in interest and also highlights the fact that ecologists and entomologists have been worrying about this problem for some time.

Since the appearance of the, now, infamous paper, that sparked the most recent round of Armageddon stories (Sánchez-Bayo & Wyckhuys, 2019), a lot has been, quite justifiably, written about the short-comings of the study both in scientific journals (e.g. Komonen et al., 2019, Simmons et al., 2019; Thomas et al, 2019, Wagner, 2019) and in blog posts, such as this thoughtful piece from Manu Saunders.

What does need to be stressed, is that although these commentators recognise the shortcomings of the paper, none of them, including the most scathing of commentators (Mupepele et al., 2019) dispute the fact, that insects, in general, are in decline. Unfortunately, the climate change deniers and their ilk, have, of course, used the criticisms to try and spread a message of “nothing to fear folks”.

Hopefully a failed attempt at downplaying the insect decline stories, but a great example of how climate change deniers are keen to muddy the waters

For humans with our relatively short lifespans, shifting baselines can be a problem (Leather & Quicke, 2010; Tree, 2018), in that people accept what they have known in their childhoods as the natural state of nature.  It can of course work the other way. I can remember the late great Miriam Rothschild telling me in the early 1990s, how as a “gel” in the 1920s a particular butterfly species that was currently at very low numbers compared with the 1970s which was what I and similar aged colleagues were remarking upon, was 50 years before that, also very low, her message being “populations cycle”.  It is because of this propensity, which is nicely illustrated by some of my 20-year data sets, all from the same 52 trees, that we need access to long-term funding to monitor insect populations.  Chop my data sets into three-year concurrent periods, the time-span of a typical PhD study or research grant, and you end up with some very different pictures of the populations of three common insect species.

The Silwood Park Winter moth, Operophtera brumata – dramatic shifts in population levels

Twenty years of the Sycamore aphid, Drepanosiphum platanoidis, at Silwood Park.  First five years versus last five years – what happened? Does this fit with the recent paper by Stephen Heard and colleagues that species chosen for study because they are common or easy to find, are almost certainly to show declines over the long-term?


The Maple aphid, Periphyllus testudinaceus – twenty-year data run from Silwood Park

Given the above, and the fact that most of the evidence for insect declines is largely based on studies from Europe, the UK heading the list (Wagner, 2019) and on top of that, the evidence from tropical locations is open to different interpretations (e.g.  Willig et al, 2019), there is an urgent need for something to be done.  So, what do we need to do?  I think there are three things that need addressing, sooner, rather than later.


First, we need to build on the work that has been done in Germany (Hallmann et al., 2017) and the UK via the Rothamsted Insect Survey (Bell et al., 2015) and establish active insect monitoring networks using repeatable sampling methods, but on a global scale. New monitoring programs will not help establish past baselines, but they can help us determine trends from this point forward. We can make this truly global by engaging the public through community science. These programs will need to use standardized methods, such as Malaise traps, pitfall traps, light traps, and effort-based counts, with species diversity, abundance and biomass being primary measures. Although biomass is an imperfect estimator because it can be sensitive to changes in abundances of large species, it is still a valuable metric from the ecosystem perspective. Determining biomass trends also does not require fine-scale taxonomic knowledge, which is often lacking in citizen science initiatives. It would, even if it were possible, be incredibly expensive, to try to monitor all insect species from any community with appreciable diversity.  A much better option, and one that will certainly appeal to a wide range of citizen scientists would be to monitor taxa like butterflies, macro-moths, dragonflies, bees, and some beetle groups.  All these can serve as indicator species for other insect groups and, tongue in cheek, many can be observed using binoculars, thus encouraging ornithologists and mammalologists to join in 😊

Innovative use of past data

At national levels, a few long-term monitoring schemes already exist, for example, the UK Environmental Change Network (http://www.ecn.ac.uk/ ) collects biotic and abiotic data, including many insect groups, from 57 different sites across the UK using identical protocols (Rennie, 2016).   Multiple Long-Term Ecological Research projects track different facets of ecosystems in different ways (Magurran et al., 2010). In fact, the LTER network, if expanded to a global scale, could be the natural framework to make a global network proposal feasible, possibly through a targeted step change in funding (Thomas et al., 2019).  This is great for the future, but unfortunately, all the active long-term monitoring schemes are younger than modern agricultural intensification.  A way forward would be to use museum collections and to construct data sets by going through back numbers of those entomological journals that pre-date the 1940s.  There are some long-term historical long-term data that are already accessible, for example the 150 year record pine beauty moth infestations in Germany dating from 1810 (Klimetzek, 1972) and I am sure that others must exist.


Whatever we do, it will need long-term funding. There needs to be a recognition by state research funding agencies that entomological survey and monitoring work, although appearing mundane, should receive a step-change in funding, even if it is at the expense of other taxa  Funding should reflect the diversity and abundance of taxa, not their perceived charisma (Clark & May, 2002; Leather, 2013).  Crowd-funding may draw in some funding, but what is required is stable, substantial and sustained funding that will allow existing and future international collaborations to flourish.  For this to happen and failing sustained state funding, we need to convince philanthropic donors such as the Gates Foundation to turn their attention from insect eradication to insect conservation.

We do, however, need to act quickly, stop talking to just our peers, meet the public, and, if needs be, personally, or via our learned societies, lobby governments; there is no Planet B.



Bell, J.R., Alderson, L., Izera, D., Kruger, T., Parker, S., Pickup, J., Shortal, C.R., Taylor, M.S., Verier, 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.

Cordoso, P. & Leather, S.R. (2019) Predicting a global insect apocalypseInsect Conservation & Diversity, 12, 263-267.

Dennis, R.H.L. & Shreeve, T.G. (2003) Gains and losses of French butterflies: tests of predictions, under-recording and regional extinction from data in a new atlas. Biological Conservation, 110, 131-139.

Hallmann, C.A., Sorg, M., Jongejans, E., Siepel, H., Hoflan, N., Schwan, H., Stenmans, W., Muller, A., Sumser, H., Horren, T., Goulson, D., & De Kroon, H. (2017) More than 75 percent decline over 27 years in total flying insect biomass in protected areas. PLoSONE, 12(10), :e0185809.

Hopkins, G.W. & Freckleton, R.P. (2002) Declines in the numbers of amateur and professional taxonomists: implications for conservation. Animal Conservation, 5, 245-249.

Klimetzek, D. (1972) Die Zeitfolge von Ubervermehrungen nadelfressender kiefernraupen in derPfalz seit 1810 und die Ursachen ihres Ruckanges in neuerer Zeit. Zeitschrift fur Angewandte Entomologie, 71, 414-428.

Kotze, D.J. & O’Hara, R.B. (2003) Species decline – but why?  Explanations of Carabid beetle (Coleoptera, Carabidae) declines in Europe. Oecologia, 135, 138-148.

Leather, S.R. & Quicke, D.J.L. (2010) Do shifting baselines in natural history knowledge threaten the environment?  Environmentalist, 30, 1-2

Magurran, A.E., Baillie, S.R., Buckland, S.T., Dick, J.M., Elston, D.A., Scott, M., Smith, R.I., Somerfiled, P.J. & Watt, A.D. (2010) Long-term datasets in biodiversity research and monitoring: assessing change in ecological communities through time. Trends in Ecology and Evolution, 25, 574-582.

Møller, A.P. (2019) Parallel declines in abundance of insects and insectivorous birds in Denmark over 22 years. Ecology & Evolution, 9, 6581-6587.

Mupepele, A.C., Bruelheide, H., Dauber, J., Krüß, A., Potthast, T., Wägele, W. & Klein, A.M. (2019). Insect decline and its drivers: Unsupported conclusions in a poorly performed meta-analysis on trends—A critique of Sánchez-Bayo and Wyckhuys (2019).  Basic & Applied Ecology, 37, 20-23.

Rennie, S.C. (2016) Providing information on environmental change: Data management, discovery and access in the UK Environmental Change Network data.  Ecological Indicators, 68, 13-20.

Sánchez-Bayo, F. & Wyckhuys, K.A.G. (2019) Worldwide decline of the entomofauna: A review of its drivers. Biological Conservation, 232, 8-27.

Thomas, C.D., Jones, T.H. & Hartley, S.E. (2019) “Insectageddon”: a call for more robust data and rigorous analyses. Global Change Biology, 6, 1891-1892.

Thomas, J.A., Telfer, M.G., Roy, D.B., Preston, C.D., Greenwood, J.J.D., Asher, J., Fox, R., Clarke, R.T. & Lawton, J.H. (2004) Comparative losses of British butterflies, birds, and plants and the global extinction crisis. Science, 303, 1879-1881.

Tree, I. (2018) Wilding, Picador, Pan Macmillan.

Wagner, D.L. (2019) Global insect decline: comments on Sánchez-Bayo and Wyckhuys (2019). Biological Conservation, 233, 332-333.

Willig, M.R., Woolbright, L., Presley, S.J., Schowalter, T.D., Waide, R.B., Heartsill Scalley, T., Zimmerman, J.K.,  González, G. & Lugo, A.E. (2019) Populations are not declining and food webs are not collapsing at the Luquillo Experimental Forest. Proceedings of the National Academy of Sciences, 116, 12143-12144.



Filed under Bugbears, EntoNotes

Not all aphids get eaten – “bottom-up” wins this time

In the lecture that I introduce aphids to our entomology MSc students I show them two quotes that illustrate the prodigious reproductive potential of these fantastic animals.

“In a season the potential descendants of one female aphid contain more substance than 500 million stout men “– Thomas Henry Huxley (1858) and “In a year aphids could form a layer 149 km deep over the surface of the earth.  Thank God for limited resources and natural enemies” – Richard Harrington (1994).

I was a little discomfited whilst researching this article to find that both Huxley and I had been short-changed, although the original quote does hint at the mortality factors that an aphid clone faces during its life.

The original words and the morphed ‘quote’


Both these quotes acknowledge the contribution that both bottom-up and top-down factors have on aphid populations.  For those not familiar with the ecological jargon, ecologists have at times over the last 40 years or so, got quite territorial* about whether herbivorous insect populations are regulated by top-down e.g. predators or bottom-up e.g. host plant quality, factors (e.g. Hunter & Price, 1992).  Who is in charge of an aphid clone’s destiny, natural enemies or the food plant?

Aphids are the favourite food of several insect species; ladybirds (but not all species), lacewing larvae, hoverfly larvae, and also the larvae of some Cecidomyiid flies (Aphidoletes spp.), and Chamaemyiid flies (e.g. Leucopis glyphinivora).  They are also attacked by other Hemipteran species, such as Anthocoris nemorum.   Those insects that make a living almost solely from aphids, are termed aphidophagous and every three years you can, if you feel like it, attend an international conference devoted to the subject 🙂

As well as these specialist predators, aphids are also preyed upon by more generalist predators, such as carabid and staphylinid beetles, harvestmen and spiders. Aphids also provide a nutritious snack for birds and bats.  Faced with all these hungry and voracious predators you might wonder why it is that aphids ever get numerous enough to become pests.  There are two answers, their fantastic reproductive rates and second, aphids, despite appearing soft and squishy, do have anti-predator defence mechanisms.  These range from kicking predators in the face, dropping off the plant, gumming up the jaws of predators by smearing them with wax from their siphunculi, and even jumping out of the way of the predator (Dixon, 1958).  On top of all that,  many are extremely unpalatable and even poisonous.

Some population modelling work from the 1970s explains why aphids can often become pests, as well as introducing us to the concept of population dynamics geography; the endemic and epidemic ridges, and my favourite, the natural enemy ravine (Southwood & Comins, 1976).

The geography of population dynamics from Southwood & Comins (1976)


They suggested that if enough predators are already present in the habitat or arrive shortly after the aphids, then the aphid population either goes extinct or only reaches the “endemic ridge”.  The phenomenal rate at which aphids can reproduce under favourable conditions, usually gets them past the “natural enemy ravine” and up into “epidemic ridge” with only a slight slowdown in population growth.   Evidence for the “natural enemy ravine” is not very convincing and I feel that the suggestion that the dip in population growth at the start of the season is due to intermittent immigration by winged aphids and not the action of polyphagous predators (Carter & Dixon, 1981) is pretty convincing.   That said, later modelling work suggested that the subsequent growth of aphid populations could be slowed down by the action of natural enemies Carter et al., 1982).

Aphids, despite their ability to produce baby aphids extremely quickly, are not equally abundant all year round. Those of us who want to collect aphids know that the best time of year is early in the season, spring and early summer.  This is the time when the plant sap is flowing quickly and is rich in nutrients, especially nitrogen, which aphids need in large quantities.    A characteristic of aphid populations is the way they suddenly disappear during July, a phenomenon known as the “mid-summer or mid-season crash”.  This is not just a phenomenon confined to aphids living on ephemeral herbaceous hosts, it happens to tree-dwelling aphids too e.g. the sycamore aphid, Drepanoisphum platanoidis.  At Silwood Park, where I monitored sycamore aphid populations on fifty-two trees for twenty years**, I saw the same pattern of a rapid build-up followed by an equally rapid collapse every year.  The pattern was the same in both high population and low population years and happened at pretty much the same time every year.  Herbivorous insects are, as you might expect, strongly

High and low population years of sycamore aphid, Drepanosiphum platanoidis at Silwood Park

affected by the quality of their host plant, the availability of nitrogen in the leaves being of most importance (Awmack & Leather, 2002).  Aphids are no exception, and their whole-life cycle is adapted to the ever-changing, but predictable availability of soluble nitrogen and water in their host plants (Dixon, 1977).  Plants become less suitable for aphids as their tissues mature and they lock their nitrogen away in the leaves and other structures, rather than transporting it around in the phloem as they do in spring and autumn (Dixon, 1976).

Aphids respond in two ways to a decline in the nutritional quality of their host plant, they reduce the number of offspring they produce (e.g. Watt, 1979) and those offspring they produce are winged (e.g. Parry, 1977), or if already winged, more likely to take flight and seek new better quality host plants (e.g. Dixon, 1969; Jarosik & Dixon, 1999).  In some aphids there is also an increase in intrinsic mortality (e.g. Kift et al., 1998).

The mid-season crash is not confined to abundant and common aphids, rare aphids show exactly the same changes in their populations, and this is similarly attributed to changes in the nutritional quality of the aphid host plant leading to increased dispersal (e.g. Kean, 2002).

Population crash of the rare aphid Paradoxaphis plagianthi in New Zealand (data from Kean, 2002).

Although some authors, notably Alison Karley and colleagues have suggested that it is the action of natural enemies and not host nutrition that drives the mid-season crash (Karley et al., 2003, 2004), the overwhelming evidence points to the production of winged (alate) morphs and their dispersal, being the major factor in causing the mid-season crash as the graphs below illustrate.

Cereal aphids on wheat showing increased alate production coinciding and subsequent population crash on cereal crops. Data from Wratten, 1975).

Green spruce aphid, Elatobium abietinum on Norway spruce at Silwood Park, showing the population crash and associated increase in the number of winged aphids. Data from Leather & Owuor (1996).

Green spruce aphid in Ireland, population crash associated with marked decline in fecundity and production of winged forms. Data from Day (1984)

Data presented by Way & Banks (1968) might lend some support to the idea that natural enemies cause the mid-season crash.  A close examination of the data however, which might at first glance suggest that keeping natural enemies away, allows aphid populations to prosper, reveals that the process of excluding natural enemies also prevents the dispersal of the winged aphids, which have no choice but to stay on the host plant and reproduce there.

Aphis fabae populations on Spindle bushes from Way & Banks (1968). Top line shows the population kept free of predators until August 2nd, bottom line, exposed to predators.

Moreover, as the authors themselves state “the rise to peak density in each year, coincided with an enormous increase in the proportion of individuals destined to become alatae” (Way & Banks, 1968).   I do not dispute that natural enemies have an effect on aphid populations, but in my opinion, the evidence does not support the hypothesis that they are the driving force behind the mid-season crash.  Rather, the major factor is the reduction in host quality, caused by a decline in the nutritional status of the plant and overcrowding of the aphids, leading to reduced fecundity and an increase in winged dispersers.

I don’t deny that the natural enemies do a very good mopping-up job of those aphids that are left behind, but they are not the force majeure by any stretch of the imagination. Most aphids do not get eaten 🙂



Awmack, C.S. & Leather, S.R. (2002) Host plant quality and fecundity in herbivorous insects. Annual Review of Entomology, 47, 817-844.

Carter, N. & Dixon, A.F.G. (1981) The natural enemy ravine in cereal aphid population dynamics: a consequence of predator activity or aphid biology? Journal of Animal Ecology, 50, 605-611.

Carter, N., Gardner, S.M., Fraser, A.M., & Adams, T.H.L. (1982) The role of natural enemies in cereal aphid population dynamics. Annals of Applied Biology, 101, 190-195.

Day, K.R. (1984) The growth and decline of a population of the spruce aphid Elatobium abietinum during a three  study, and the changing pattern of fecundity, recruitment and alary polymorphism in a Northern Ireland Forest. Oecologia, 64, 118-124.

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. (1969) Population dynamics of the sycamore aphid Drepanosiphum platanoides (Schr) (Hemiptera: Aphididae); migratory and trivial flight activity. Journal of Animal Ecology, 38, 585-606.

Dixon, A.F.G. (1976) Factors determining the distribution of sycamore aphids on sycamore leaves during summer. Ecological Entomology, 1, 275-278.

Dixon, A.F.G. (1977) Aphid Ecology: Life cycles, polymorphism, and population regulation. Annual Review of Ecology & Systematics, 8, 329-353.

Harrington, R. (1994) Aphid layer. Antenna, 18, 50-51.

Hunter, M.D. & Price, P.W. (1992) Playing chutes and ladders – heterogeneity and the relative roles of bottom-up and top-down forces in natural communities. Ecology, 73, 724-732.

Huxley, T.H. (1858) On the agmaic reproduction and morphology of Aphis – Part I. Transactions of the Linnean Society London, 22, 193-219.

Jarosik, V. & Dixon, A.F.G. (1999) Population dynamics of a tree-dwelling aphid: regulation and density-independent processes. Journal of Animal Ecology, 68, 726-732.

Karley, A.J., Parker, W.E., Pitchford, J.W., & Douglas, A.E. (2004) The mid-season crash in aphid populations: why and how does it occur? Ecological Entomology, 29, 383-388.

Karley, A.J., Pitchford, J.W., Douglas, A.E., Parker, W.E., & Howard, J.J. (2003) The causes and processes of the mid-summer population crash of the potato aphids Macrosiphum euphorbiae and Myzus persicae (Hemiptera: Aphididae). Bulletin of Entomological Research, 93, 425-437.

Kean, J.M. (2002) Population patterns of Paradoxaphis plagianthi, a rare New Zealand aphid. New Zealand Journal of Ecology, 26, 171-176.

Kift, N.B., Dewar, A.M. & Dixon, A.F.G. (1998) Onset of a decline in the quality of sugar beet as a host for the aphid Myzus persicaeEntomologia experimentalis et applicata, 88, 155-161.

Leather, S.R. & Owuor, A. (1996) The influence of natural enemies and migration on spring populations of the green spruce aphid, Elatobium abietinum Walker (Hom., Aphididae). Journal of Applied Entomology, 120, 529-536.

Parry, W.H. (1977) The effects of nutrition and density on the production of alate Elatobium abietinum on Sitka spruce. Oecologia, 30, 637-675.

Southwood, T.R.E. & Comins, H.N. (1976) A synoptic population model.  Journal of Animal Ecology, 45, 949-965.

Watt, A.D. (1979) The effect of cereal growth stages on the reproductive activity of Sitobion avenae and Metopolphium dirhodum. Annals of Applied Biology, 91, 147-157.

Way, M.J. & Banks, C.J. (1968) Population studies on the active stages of the black bean aphid, Aphis fabae Scop., on its winter host Euonymus europaeus L. Annals of Applied Biology, 62, 177-197.

Wratten, S.D. (1975) The nature of the effects of the aphids Sitobion avenae and Metopolophium dirhodum on the growth of wheat. Annals of Applied Biology, 79, 27-34.


Post script

For those interested this is how Huxley arrived at his number of potential descendants, and here I quote from his paper,  “In his Lectures, Prof. Owen adopts the calculations taken from Morren (as acknowledged by him) from Tougard that a single impregnated ovum  of Aphis may give rise, without fecundation, to a quintillion of Aphides.” I have not, so far, been able to track down Tougard.

Morren, C.F.A. (1836) sur le Puceron du Pecher, Annales des Sciences Naturelle series 2. vi.

You may not know what a grain is, so to help you visualise it, 7000 grains equals a pound so 2 000 000 grains gives you 286 pounds, or 20 stone or approximately 130 Kg depending on where you come from J


*and generated some magnificent paper titles and quite acrimonious responses J Hassell, M.P., Crawley, M.J., Godfray, H.C.J., & Lawton, J.H. (1998) Top-down versus bottom-up and the Ruritanian bean bug. Proceedings of the National Academy of Sciences USA, 95, 10661-10664.

**A true labour of love as I also counted maple aphids, orange ladybirds, winter moth larvae and any of their predators and parasites that I came across J



Filed under Aphidology, Aphids