Tag Archives: dispersal

The most difficult thing I have ever had to write – Insects – A Very Short Introduction

The book!

I have written a lot of papers (more than 220 according to Web of Science) and quite a few books, two real ones (Leather et al., 1993; Leather & Bland, 1996) and eight edited volumes, over the last forty odd years. Up until now I thought the most exacting piece of writing I had ever done was my entry for the Biological Flora (Leather, 1996).  I mention this because it has a very similar feel to my most recent, most difficult piece of writing, Insects, A Very Short Introduction.

I did my PhD on the bird cherry aphid, Rhopalosiphum padi (Leather, 1980) during which I developed a real love for the bird cherry tree, Prunus padus. Just to illustrate this, my second son’s middle name is Tuomi – Finnish for bird cherry. Over the next decade or so I expanded my studies on to the different insects associated with it and also became quite adept at striking scions, grafting it and manipulating its phenology. I could (and still can), thanks to sampling bird cherry trees in Finland in the depths of winter, identify it in the dark by the smell of the bark 😊 In the mid-1980s, I jumped on to the species-area relationship bandwagon (Leather, 1985, 1986) and discovered the wonderful Biological Flora of the British Isles, hosted by the British Ecological Society. Armed with the arrogance of youth, and obviously at the time, not suffering from ‘imposter syndrome’, I contacted Arthur Willis (the then Editor of the series) and volunteered to write the entry for bird cherry. Pretty cheeky for an entomologist, but hey, both my parents were botanists and me and bird cherry were old pals! Arthur said yes and sent me the instructions for contributors which included all the headings and sub-sections required. It looked pretty straightforward to me; go down the headings, insert the information and write the narrative. Easy peasy lemon squeezy.  Or so I very naively thought ☹ I whipped through the headings, filled in the data that I had, wrote the accompanying narrative and posted it off to Arthur.  Job done! A few weeks later he returned my manuscript telling me that I was also expected to fill in the missing data gaps with data collected by me, not just leave them blank! 

Reality strikes!

So, over the next two years that is exactly what I did.  I checked out which mycorrhizae were associated with bird cherry, collected seed and calculated germination rates, sketched the different seedling stages, did NVC surveys at six different sites, characterised the growth structure of the tree and shrub forms and looked at responses to defoliation. I learnt a lot of botany! Arthur was an incredibly helpful editor and without his encouragement I would never have completed the entry.  I did, however, turn down Arthur’s suggestion that I do the entry for Prunus avium 😊

In 2018 I was contacted by Latha Menon, a Senior Commissioning Editor for OUP, who wanted to know if I would be interested in writing a book for their Very Short Introduction series, in this case, insects.  Having since 1990 grown older and picked up imposter syndrome on the way, I was initially a bit hesitant and asked if it would be possible to have a co-author.  Latha was somewhat lukewarm about this, saying that what the series was about was producing an extended essay (35 000 words) for a general audience reflecting the expertise and enthusiasm of the author.  I pondered about it and thought, well I’ve been a professional entomologist for more than forty years, and an amateur for my whole life, and taught entomology for more than thirty years, so how hard could it be to write 35 000 words about insects? I have also been blogging for what I have envisaged as a mixed audience since 2012, so surely within my capabilities?

I should have known better

It turns out that you can know too much about insects at the same time as you don’t know enough about them! First, I had to decide on a structure for the essay and also what particular aspects of entomology would leap off the page and grab my readers.  Much as I love aphids, other insects would have to feature.  The obvious place to start, I thought, would be to look at how our entomology course introduced the subject to new students, so Chapter 1, In the beginning was born. This was actually not that simple as I had to deal with the evolution of insects, their classification, their anatomy, a lot of their physiology, and also why they were and are so successful. Trying to make that not like a series of Wikipedia sections was not easy.

Now to me, one of the things that make insects so spectacular and evinces amazement in non-entomologists, is how good they are at reproducing themselves, hey presto, Chapter 2, Prolific procreators and the need to discuss sexual selection, mating behaviour, courtship, lekking and much more. I had originally planned to cover host selection and life history strategies in this chapter, but found that it didn’t sit very well in that context.

Instead, I moved on to Chapter 3, On the move, which began with the evolution of flight, much, of which, despite my decades of experience, was quite a revelation to me. From flight I moved onto host selection, and the physiology and ecology of specialist and generalist feeding. These first three chapters were the most difficult to write, bearing in mind that my readers need to understand the basics of how insects work to fully grasp the wonder of what insects are actually achieving. I had to rewrite these first chapters three times before my Commissioning Editor, my non-entomologist lay-reader (my wife, a humanities graduate), a botanist colleague and an entomological colleague, were happy with them.

Those chapters behind me, I now felt it was time to move on to those aspects of entomology that had super wow factor, those facts that would make my readers exclaim things like “I didn’t know that” and encourage them to pass them on to their friends and relatives. Heavily influenced by my childhood love of social insects, Chapter 4, Living together, appeared on the scene. As well as talking about the well-known bees, ants and termites, it also gave me the chance to discuss lesser known examples such as dung beetles, social bugs, insect symbionts, insect-plant interactions, and yes, of course, aphids get more than a passing mention😊

The next three chapters are a bit niche, but gave me the chance to launch Chapter 5, Aquatic Insects, into the mix and talk about the importance of freshwater insects and even more excitingly, wax lyrical about the little known truly marine Ocean Skaters, which I first came across on a work trip to Mauritius in the mid-1990s. Judging by what I see on social media platforms, non-entomologists are, in the English vernacular, totally gob-smacked by how insects can pretend to be something else, so this made Chapter 6, Crypsis, mimicry and blatant advertising, an obvious heading.  I had a lot of fun with this, but was slightly hampered by the restriction placed by OUP on how many illustrations I was allowed to include, and also sadly not being allowed colour.

To me, the ability of insects to live in very inhospitable conditions, from deserts, to ice caves, to mountain tops and to survive arctic winters and other extreme weather conditions without the benefit of fur coats and cold baths, made Chapter 7, Against the odds, a natural.

I have, over the years, been amazed by how limited many people’s appreciation of the positive economic, social, artistic and health benefits of insects to the human world are (Leather, 2015), hence Chapter 8, The good, the bad and the ugly.  Here I dealt with pests, disease vectors, biological control, maggot therapy, pollinators, the role of insects as ecosystem engineers, as waste-disposal specialists and as recyclers.

I began my book by describing the diversity and ubiquity of insects. I discussed their origins and marveled at the ways in which insects have adapted to a wide range of environments and the roles that they play in maintaining ecosystem health. Given that for at least the last twenty years or so, entomologists and ecologists have been warning about the dangers of losing species I felt it fitting to end it by discussing the harm that we are doing to insects and the planet that we, and they, inhabit, and ways in which we might act to halt or reverse our course, if it is not already too late.  I finished with a warning, and the hope that we are not too late to stem disaster, Chapter 9, Ecological Armageddon—insects in decline?

It was, primarily due to the fact that at times, for every word I took out, I seemed to add another ten, both one of the most frustrating but also satisfying pieces of writing that I have ever done. I just hope that when it hits the shops, my readers will find as much to enjoy as I have since I first came across insects just over sixty years ago.

References

Leather, S.R. (1980) Aspects of the Ecology of the Bird Cherry-Oat Aphid, Rhopalosiphum padi L., PhD Thesis University of East Anglia, Norwich.

Leather, S.R. (1985) Does the bird cherry have its ‘fair share’ of insect pests? An appraisal of the species-area relationships of the phytophagous insects associated with British Prunus species. Ecological Entomology, 10, 43-56.

Leather, S.R. (1986) Insect species richness of the British Rosaceae: the importance of hostrange, plant architecture, age of establishment, taxonomic isolation and species-area relationships. Journal of Animal Ecology, 55, 841-860.

Leather, S.R. (1996) Biological flora of the British Isles Prunus padus L. Journal of Ecology, 84, 125-132.

Leather, S.R. (2015) Influential entomology: a short review of the scientific, societal, economic and educational services provided by entomology. Ecological Entomology, 40 (Suppl. 1), 36-44.

Leather, S.R. & Bland, K.P. (1999) Insects on Cherry Trees, Richmond Publishing Co, Ltd, Slough.

Leather, S.R., Walters, K.F.A. & Bale, J.S. (1993) The Ecology of Insect Overwintering, Cambridge University Press, Cambridge.

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Not all moths have wings

Insects really took off when they developed flight (Alexander, 2015), so it is perhaps surprising that so many have lost the ability subsequently.  Nearly all the winged Orders have developed flightless members, with beetles of course, topping the list (Wagner & Liebherr, 1992).  A number of reasons for why flightlessness made a reappearance have been put forward. The eminent coloepterist, Thomas Vernon Wollaston, noted that the island of Madeira had an unusually high number of wingless (apterous) beetles. His friend, Charles Darwin, suggested that for island dwelling animals, it was a disadvantage to be winged especially if you were small or subjected to high winds (Darwin, 1859). Many years later, Derek Roff reviewed the literature, and found that there was no difference in the proportion of non-winged insects on islands compared with those on continental areas (Roff, 1990).  Winglessness is also common in insects living at high altitudes, in cold climates or in those that are autumn or winter active (Hackman, 1966).  It might be that wings are energetically costly in those environments (Mani, 1962), but why then is it that in many cases, it is only the females that are wingless?  To explain this we can hypothesise that eggs are energetically more expensive than sperm (Hayward & Gilooly, 2011), so that males can ‘afford’ to be winged and travel to find a mate. For this to work, the females need to be able to attract males from a distance, something moths are renowned for (Greenfield, 1981).

Male Thyridopteryx ephemeraeformis – note the well-developed antenna – ideal for picking up female sex pheromones. https://upload.wikimedia.org/wikipedia/commons/4/4d/-_0457_%E2%80%93_Thyridopteryx_ephemeraeformis_%E2%80%93_Evergreen_Bagworm_Moth_%2814869905567%29.jpg By Andy Reago & Chrissy McClarren [CC BY 2.0 (http://creativecommons.org/licenses/by/2.0)%5D, via Wikimedia Commons

We also know that in those insects with wing dimorphism, the apterous forms are more fecund compared with those with wings (Dixon, 1972; Mackay & Wellington, 1975). In those insects that retain their wings, many resorb their wing muscles once they have found suitable egg laying sites (Stjernholm et al., 2005; Tan et al., 2010), further proof that wings are costly. Winglessness is also common in those insects that are parasitic on vertebrates, bedbugs, fleas and lice for example.  Those that do start with wings, such as the Hippoboscid flies, lose their wings once they have found a suitable host. Finally, winglessness is often associated with stable and extensive habitats, such as forests, or surprisingly to me at any rate, mountains, where dispersal is not a high priority (Roff, 1990).

Thyridopteryx ephemaeraeformis https://ideastations.org/sites/default/files/storage/secondary-images/bagworm-case.jpg  

I first saw bagworms as a child in Jamaica but of course at the time had no idea what species they were. I was however, fascinated by the sight of the cunningly constructed cases in which the larvae lived and eventually pupated within.  To me, case bearer moths and caddisflies were the insect equivalent of hermit crabs, which were and are one of my favourite non-insect animals*.  Little did I know that one day I would write about these very same bagworms (Rhainds et al., 2008). Bagworms, of which just over half have wingless females, immediately contradict the cold climate hypothesis of winglessness, as many of them are tropical and there are just as many wingless species in the tropics as there are elsewhere (Rhainds et al., 2009). The bagworms belong to the family Pyschidae, which contain a 1000 species or so.  Not only do over half of these have wingless females, some also have females which are legless and never leave their pupal case, even mating in it.

Male Thyridopteryx ephemeraeformis bagworm mating with bag-enclosed female (Jones, 1927)

Even though the more primitive (less derived) members of the Psychidae have wings, the winged females are less active than the males (Rhainds et al. 2009).  As you might expect, host plant selection is by the larval stage, which on hatching, throw out a silk thread and float off with great expectations (Moore & Hanks, 2004).  Once they find a suitable host plant, which is not as difficult as you might expect, as they extremely polygamous, they begin to feed and construct their cases.  Some of the larval cases that Psychids construct are truly magnificent.  A great example is Eumeta crameri, the large faggot worm, so called because it looks like it is carrying a pile of firewood on its back 🙂

Eumeta crameri, the large faggot worm, so called because of the twigs its carries around on its back Melvyn Yeo

In case you are wondering about the ornate cases, they are not decorative, but more likely to be anti-predator devices (Khan, 2020).

Although the Psychids have the largest number of species with wingless females, there are 18 other moth families with species with wingless females.  Species that are found at high altitudes and northern latitudes have the most flightless species (Hackmann, 1966) or, like the Psychids, inhabit stable forest and woodland habitats (Barbosa et al., 1989).  Another characteristic of wingless moth species is that they overwinter as eggs or first instar larvae (Barbosa et al., 1989, although there are of course, many moths that have similar habits and are not wingless, such as the small ermine moths (Leather, 1986a).

After the Psychids, the families with the greatest number of species with wingless females are the Geometridae (loopers) and the Lymantridae (tussock moths). In the Lymantridae some are wingless and many have non-functional wings (Hackman, 1966).  The Arctidae and the Lasiocampidae also have some flightless species, the genus Chondrostega, endemic to the Iberian Peninsula having some notable examples, (Hackman, 1966). An oddity, as they are not strictly flightless, are females of the tortricid Choristoneura fumiferana, which have functional wings, but are behaviourally flightless, only taking flight under particular environmental conditions (Barbosa et al., 1989).

Moth species that have flightless females all have one thing in common, they aren’t picky about their diet, they are polyphagous and live in forests and woodlands. They also tend to have larvae that can disperse by ballooning, although not all moths with ballooning larvae have flightless females.  First instar larvae of the pine beauty moth, Panolis flammea, which readily balloon in outbreak situations, and usefully, can survive several days without food (Leather, 1986b).

In the UK there are two very common moths with wingless females, the winter moth, Operphthera brumata and the Vapourer moth, Orgyia antiqua, the former a Geoemtrid, the latter, a Lymantrid. Both are extremely polyphagous, usually feeding on broadleaf trees and shrubs, but both have recently added conifer species to their diets.  The Vapourer moth ‘decided’ that the introduced lodgepole pine, Pinus contorta growing in Sutherland and Caithness, would make a suitable alternative food plant (Leather, 1986) and the winter moth opted for another introduced conifer, Sitka spruce, Picea sitchensis, in the Scottish Borders (Hunter et al., 1991). Why both these host shifts happened in the early 1980s and in Scotland, remains a mystery, although it is possible that they moved onto conifers via heather (Hewson & Mardon, 1970; Kerslake et al., 1996).

They do, however, have some striking differences in their approach to life. Larvae of the Winter moth are spring flush feeders, and very dependent on egg hatch coinciding with bud burst (Wint, 1983), Vapourers are summer foliage feeders so are adapted to feeding on mature leaves. The adults of the Winter moth, as its name suggests are active in the winter months, laying their eggs on the bark or in crevices of their host trees in November and December and even January. Vapourer adults on the other hand are summer active, the eggs being laid on their pupal cases on the leaves of their host trees from July to September.

Female Vapourer moth and her egg mass – note the short legs and much reduced wings

 

Long legged female winter moth Operphtera brumata https://butterfly-conservation.org/sites/default/files/styles/large/public/2019-01/8179909985_76865dd047_o.jpg

Hackman (1966) distinguishes two types of wingless females, those with reduced locomotion, very heavy, filled with eggs and what I describe in class as splurgers, i.e. all their eggs laid in one go.  The female Vapourer with short legs and much-reduced wings is an ideal example.  The female winter moth is a good example of the second type, those possessing good strong legs which after copulation seek out suitable egg-laying sites.  Despite the difference in oviposition tactics, the first instar larvae of both species are adept ballooners, and it is they who ‘decide’ whether to stay or go (Tikkanen et al., 1999).

First instar Vapourer moth larvae in the process of dispersing.

Understandably, they have very little control of where they land, although presumably, they can reject the plant they land on and launch themselves into space again. How many times they can do this and how long they can live for without feeding, is something that needs research, but given that the first instar larvae of the pine specialist P. flammea can live several days without feeding, I would expect that the Winter moth and Vapourer moth larvae are equally capable of resisting starvation.

Moths without wings, but highly successful and many are pests, so not such a dumb approach to life after all?

And while we’re at it, here is the lymantriid Teia anartoides. With hamsterlike apterous females! AinsleyS @americanbeetles

References

Alexander, D.E. (2015) On the Wing, Oxford University Press. (This is an excellent book).

Barbosa, P., Krischik, V. & Lance, D. (1989) Life history traits of forest-inhabiting flightless lepidoptera. American Midland Naturalist, 122, 262-274.

Darwin, C. (1859) 0n the Origin of Species, JohnMurray, \lodnon.

Dixon, A.F.G. (1972) Fecundity of brachypterus and macropterous alatae in Drepanosiphum dixoni (Callaphididae, Aphididae). Entomologia experimentalis et applicata, 15, 335-340.

Greenfield, M.D. (1981) Moth sex pheromones: an evolutionary perspective. Florida Entomologist, 64, 4-17.

Hackman, W. (1966) On wing reduction and loss of wings in Lepidoptera. Notulae Entomologicae, 46, 1-16.

Hayward, A. & Gillooly, J.F. (2011) The cost of sex: quantifying energetic investment in gamete production by males and females. PLoS One, 6, e16557

Hewson, R. & Mardon, D.K. (1970) Damage to heather moorland by caterpillars of the vapourer moth Orgyia antiqua L. (Lep., Lymantridae). Entomologist’s Monthly Magazine, 106, 82-84.

Hunter, M.D., Watt, A.D. & Docherty, M. (1991) Outbreaks of the winter moth on Sitka spruce are not influenced by nutrient deficiencies of trees. Oecologia, 86, 62-69.

Jones, F.M. (1927) Mating of the Psychidae (Lepidoptera). Transactions of the Entomological Society of America, 53, 293-312.

Kerslake, J.E., Kruuk, L.E.B., Hartley, S.E. & Woodin, S.J. (1996) Winter moth (Operophtera brumata (Lepidoptera: Geometridae)) outbreaks on Scottish  moorlands; effects of host plant and parasitoids on larval survival and development. Bulletin of Entomological Research, 86, 155-164.

Khan, M.K. (2020) Bagworm decorations are an anti-predator structure.  Ecological Entomology https://onlinelibrary.wiley.com/doi/epdf/10.1111/een.12876

Leather, S.R. (1986a) Insects on bird cherry I The bird cherry ermine moth, Yponomeuta evonymellus (L.). Entomologist’s Gazette, 37, 209-213.

Leather, S.R. (1986b) The effect of neonatal starvation on the growth, development and survival of larvae of the pine beauty moth Panolis flammea. Oecologia, 71, 90-93.

Leather, S.R. (1986c) Keep an eye out for the vapourer moth. Forestry & British Timber, 15, 13.

Mackay, P.A. & Wellington, W.G. (1975) A comparison of the reproductive patterns of apterous and alate virginoparous Acyrthosiphon pisum (Homoptera: Aphididae). Canadian Entomologist, 107, 1161-166.

Mani, M.S. (1962) Introduction to High Altitude Entomology: Insect Life above the Timber-line in the Northwest Himalaya. Methuen, London.

Moore, R.G. & Hanks, L.M. (2004) Aerial dispersal and host plant selection by neonate Thyridopteryx ephemeraeformis (Lepidoptera: Psychidae). Ecological Entomology, 29, 327-335.

Rhainds, M., Leather, S.R. & Sadoff, C. (2008) Polyphagy, flightlessness and reproductive output of females: a case study with bagworms (Lepidoptera: Psychidae). Ecological Entomology, 33, 663-672.

Rhains, M., Davis, D.R. & Price, P.W.(2009) Bionomics of Bagworms (Lepidoptera: Psychidae). Annual Review of Entomology, 54, 209-226.

Roff, D.A. (1990) The evolution of flightlessness in insects. Ecological Monographs, 60, 389-422.

Stjernholm, F., Karlsson, B. & Boggs, C.A. (2005) Age-related changes in thoracic mass: possible reallocation of resources to reproduction in butterflies. Biological Journal of the Linnean Society, 86, 363-380.

Tan, J.Y., Wainhouse, D.W., Day, K.R. & Morgan, G. (2010) Flight ability and reproductive development in newly-emerged pine weevil Hylobius abietis and the potential effects of climate change. Agricultural and Forest Entomology, 12, 427-434.

Tikkanen, O.P., Carr, T.G. & Roininen, H. (1999) Factors influencing the distribution of a generalist spring-feeding moth, Operophtera brumata (Lepidoptera: Geometridae), on host plants. Environmental Entomology, 28, 461-469.

Wagner, D.L. & Liebherr, J.K. (1992) Flightlessness in insects. Trends in Evolution & Ecology, 7, 216-219.

Watt, A.D., Evans, R. & Varley, T. (1992) The egg-laying behaviour of a native insect, the winter moth Operophtera brumata (L.) (Lep., Geometridae), on an introduced tree species, Sitka spruce, Picea sitchensis. Journal of Applied Entomology, 114, 1-4.

Wint, W. (1983) The role of alternative host-plant species in the life of a polyphagous moth, Operophtera brumata (Lepidoptera, Geomtridae). Journal of Animal Ecology, 52, 439-450.

Wollaston, T.V. (1854) Insecta Maderensia, John van Voorst, London.

 

 

 

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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 🙂

 

References

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

 

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Aphids Aweigh – there are sailor aphids too!

In an earlier post I described the fascinating world of soldier aphids.  There are also some aphids that might, with a little bit of imagination, be called sailor aphids.  I first came across the phenomenon of aquatic aphids when I was doing my PhD at the University of East Anglia in the late 1970s.  At the time I was working on a beautiful little aphid, the bird cherry-oat aphid, Rhopalosiphum padi , which characteristically has a rusty-brown bum.

padi

At the time, the foyer of the Biological Sciences building was graced by the presence of a large aquarium. One day whilst waiting for my lab mates to join for the long trek to coffee, I was amusing myself by teasing the fish and noticed a number of aphids feeding on the water-lilies.  Two things piqued my curiosity, one they vaguely looked like my beloved study aphid and secondly they were feeding on the submerged stems, rather than on the leaves.  It turned out that they were indeed related to my aphid, I identified them as Rhopalosiphum nymphaeae the water-lily aphid, or more prosaically, the reddish-brown plum aphid! They are known to be able to feed underwater for some time as they are able to trap air on their bodies using specialised hairs and when large numbers of  aggregate on a submerged stem, the trapped air bubble sometimes covers the entire colony.  They can also apparently walk on water.  Unfortunately I did not have a camera with me and have so far not been able to find a picture demonstrating this phenomenon or their ability to remain submerged whilst feeding.  They are regarded as a bit of pest by some gardeners although they can also be used as a biological control agent.

Another aphid which might more appropriately be called a sailor aphid, is Pemphigus trehernei, which feeds on sea asters in the salt marshes of Western Europe, including those on the Norfolk coast in East Anglia (Foster & Treherne, 1975).  As you might expect given its habitat, it is able to survive being submerged during high tides, but what I really find amazing is that the first instar nymphs disperse by floating away on the sea, hoping to bump into another suitable sea aster on which to establish a new colony (Foster & Treherne, 1978).  Optimistic indeed, although I guess as they are still with us it is effective enough.  There is yet another intertidal aphid, Staticobium limonii, which is also able to withstand being submerged by high tides, but does not have the same ability as P. trehernei, 50%  of them being dead after 20 hours and all being dead after a mere 48 hours under water, compared with the 400 hours plus that some 10% of P. trehernei can manage (Foster & Treherne, 1976).  A pretty good attempt at holding their breath I reckon and a good reason to describe them as sailor aphids, although submariners might be a better description. And of course, yet another reason why I love aphids so much.

Centre, T.D., Dray, F.A., Jubinsky, G.P. & Grodowitz, M.J. (1999).  Insects and Other Arthropods That Feed on Aquatic and Wetland Plants.  United States, Department of Agriculture, Agricultural Research Service Technical Bulletin 1870

Foster, W.A. & Treherne, J.E. (1975).  The distribution of an intertidal aphid, Pemphigus trehernei Foster, on marine saltmarshes.  Oecologia, 21, 141-155.

Foster, W.A. & Treherne, J.E. (1976).  The effects of tidal submergence on an intertidal aphid, Pemphigus trehernei Foster.  Journal of Animal Ecology, 45, 291-301.

Foster, W.A. & Treherne, J.E. (1978) Dispersal mechanism in an intertidal aphid.  Journal of Animal Ecology, 47, 205-217.

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Filed under Aphidology, Aphids, EntoNotes, Uncategorized