Mellow Yellow – Not all aphids live on green leaves

I have written before about aphids and how their quest for the ideal food plant may explain the evolution of host alternation; we find that most aphid species tend to be associated with rapidly growing meristems, or newly flushing leaves (Dixon, 2005). Some aphids are so keen on young plant tissue that they ‘engineer’ youth in their host plants, injecting salivary compounds and forming leaf–rolls, pseudo-galls and galls, all of which act as nutrient sinks and lengthen the time that the modified leaves stay green and nutrient-rich

leaf roll Rhopalosiphum

 Leaf-roll caused by Rhopalosiphum padi on bird cherry, Prunus padus.

Leaf roll Myzus cerasi

Pronounced leaf roll pseudo-gall caused by Myzus cerasi on Prunus avium.

Non host-alternating (autoecious) aphids, such as the sycamore aphid Drepanosiphum platanoidis, the maple aphid, Periphyllus testudinaceus, or the birch aphid, Euceraphis punctipennis, have no such escape route; they are confined to their tree host for the year, albeit, they can, if they ‘wish’, fly to another tree of the same species, but essentially they are held hostage by the their host plant. As the season progresses, leaf nutritional and physical properties change; going from young tender green leaves, with high nitrogen and water contents, to mature, tough leaves, low in nitrogen and water to yellow senescing leaves with again, higher nitrogen levels (Awmack & Leather, 2002) and finally of course, dead brown leaves of no nutritional value.

Seasonal changes

Sycamore and maple aphids, enter a state of suspended animation ‘summer aestivation’ (Essig, 1952; Dixon, 1963), whilst birch and poplar aphids, whose hosts plants often produce new growth during the year, ‘track’ these new leaves (Wratten, 1974; Gould et al., 2007). As far as these aphids are concerned young tissue is their best food source, with senescent tissue being second best and mature leaves being least favoured. During the summer they will, however, take advantage of mature leaves that are prematurely senescing, such as those attacked by leaf diseases such as tar spot. I have often found sycamore aphids feeding and reproducing on these infected leaves whilst those aphids on neighbouring mature leaves remain in aestivation.

Tar spot 2

Effects of tar spot on sycamore leaves

Host-alternating (heteroecious) aphids on the other hand are somewhat different. As their life cycle includes a programmed migration back to their primary tree host in autumn, those autumn morphs (oviparae) are adapted to senescent tissue (Leather & Dixon, 1982, Kundu & Dixon, 1993, 1994). Similarly, the spring morphs (fundatrices and fundatrigeniae) are adapted to young leaves and find it difficult or impossible, to make a living on senescent leaves.
Morphs and host age

There are yet other aphids, such as the green spruce aphid Elatobium abietinum, the pine aphid, Eulachnus agilis and the black pecan aphid, Melanocallis caryaefoliae, that are senescence specialists. In contrast to the flush specialists, these aphids engineer senescence, also using salivary compounds,  and are unable to survive on young foliage (Bliss, 1973; Fisher, 1987; Cottrell et al., 2009).

Elatobium in action

Elatobium abietinum ‘engineering’ senescence on spruce needles and avoiding young flushing tissue.

It is interesting to speculate that perhaps these tree-dwelling non host-alternating aphids are secondarily derived from the autumn part of the life-cycle of host-alternating aphids. After all, if non host-alternating aphids on herbaceous host plants are off-shoots of the summer part of the host-alternating life-cycle why not the other way round. There is just so much more to learn about aphids. Yet another reason why I love aphids so much ;-)

References

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

Bliss, M., Yendol, W.G., & Kearby, W.H. (1973) Probing behaviour of Eulachnus agilis and injury to Scotch pine. Journal of Economic Entomology, 66, 651-655.

Cottrell, T.E., Wood, B.W. & Ni, X. (2009) Chlorotic feeding injury by the Black Pecan Aphid (Hemiptera: Aphididae) to pecan foliage promotes aphid settling and nymphal development. Environmental Entomology, 38, 411-416.

Dixon, A.F.G. (1963) Reproductive activity of the sycamore aphid, Drepanosiphum platanoides (Schr) (Hemiptera, Aphididae). Journal of Animal Ecology, 32, 33-48.

Dixon, A.F.G. (2005) Insect Herbivore-Host Dynamics. Cambridge University Press, Cambridge.

Fisher, M. (1987) The effect of previously infested spruce needles on the growth of the green spruce aphid, Elatobium abietinum. Annals of Applied Biology, 111, 33-41.

Gould, G.G., Jones, C.G., Rifleman, P., Perez, A., & Coelman, J.S. (2007) Variation in Eastern cottonwood (Populus deltoides Bartr.) phloem sap content caused by leaf development may affect feeding site selection behaviour of the aphid, Chaitophorous populicola Thomas (Homoptera: Aphididae). Environmental Entomology, 36, 1212-1225.

Kundu, R. & Dixon, A.F.G. (1993) Do host alternating aphids know which plant they are on? Ecological Entomology, 18, 61-66.

Kundu, R. & Dixon, A.F.G. (1994) Feeding on their primary host by return migrants of the host alternating aphid, Cavariella aegopodii. Ecological Entomology, 19, 83-86.

Leather, S.R. & Dixon, A.F.G. (1981) Growth, survival and reproduction of the bird-cherry aphid, Rhopalosiphum padi, on it’s primary host. Annals of applied Biology, 99, 115-118.

Wratten, S.D. (1974) Aggregation in the birch aphid, Euceraphis punctipennis (Zett.) in relation to food quality. Journal of Animal Ecology, 43, 191-198.

 

Post script

A lot of what I describe comes from a talk I gave in 2009 at a workshop in Oxford on autumn colours (the output of which was Archetti, M., Döring, T.F., Hagen, S.B., Hughes, N.M., Leather, S.R., Lee, D.W., Lev-Yadun, S., Manetas, Y., Ougham, H.J., Schaberg, P.G., & Thomas, H. (2009) Unravelling the evolution of autumn colours: an interdisciplinary approach. Trends in Ecology & Evolution, 24, 166-173. I always meant to write the talk up as an Opinion piece but procrastination set in badly. I was somewhat annoyed with myself when earlier this year this excellent piece by the legendary ecologist and entomologist, Tom White, appeared; I have only myself to blame, six years is a very long bit of procrastination ;-)

White, T.C.R. (2015) Senescence-feeders: a new trophic sub-guild of insect herbivores Journal of Applied Entomology, 139, 11-22.

 

Post post script

This post is dedicated to my eldest son, Sam, who died quietly in his sleep, at a tragically young age, December 23rd 2010.   It would have been his birthday on the 21st May.  Despite being a molecular biologist, (he worked at the Sanger Institute), he was as green as you can get, a great naturalist and conservationist, with an incredibly gentle soul. He strongly believed in conserving the World’s natural resources and amused colleagues by sticking up signs in the toilets at the Sanger, which read “If its yellow let it mellow, if its brown flush it down”.

Sampsa

 

He is sorely missed by us all. He also had more Nature papers than me ;-)

Parkhill, J., Achtman, M., James, K.D. et al., (2000) Complete DNA sequence of a serogroup A strain of Neisseria meningitides. Nature, 404, 502-506

Parkhill, J., Dougan, G. , James, K.D. (2001) Complete genome sequence of a multiple drug resistant Salmonella enterica serovar Typhi CT18. Nature, 413, 848-852.

Parkhill, J., Wren, B.W., Thomson, N.R. et al., (2001) Genome sequence of Yersinia pestis, the causative agent of plague. Nature, 413, 523-527.

Parkhill, J., Sebaihia, M., Preston, A. et al., (2003) Comparative analysis of the genome sequences of Bordetella pertussis,   Bordetella parapertussis and Bordetella bronchiseptica. Nature Genetics, 35, 32-40

Wood, V., Gwilliam, R. Rajandream, M.A. et al., (2002) The genome sequence of Schizosaccharomyces pombe . Nature, 415, 871-880

 

 

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Distant Elephants and Low-Hanging Fruit – confessions of a procrastinator

elephant vsmall

verb; procrastinate – delay or postpone action; put off doing something

I hate writing full-blown grant applications. The thought of sitting down with a blank page in front of me, and justifying why I want to do the research, saying how well it fits the remit of the research council, demonstrating how great I am (track record of applicant), articulating my hypotheses, outlining the work packages, writing an impact statement, getting costings to draw up a budget, justifying the resources requested, and so on and so on, and all with the knowledge that there is an about 1 in 5 chance of getting funded, does not fill me with any great enthusiasm. It also takes up a huge amount of my time and to do it properly you really do need to sit down and exclude all other activities.

I also find it difficult to start writing a paper from scratch, or get to work on a new presentation;  I should, for example, be getting my plenary lecture for ENTO15 ready, but as it only May 11th and I am not speaking until September 2nd I am finding it difficult to get started on that. This is despite the fact that it was only by a superhuman effort before Christmas I got the written version submitted to Ecological Entomology on the required day (now available on Early View if you are interested).  Of course I am steadfastly ignoring the fact that I will be on holiday from July 24th until August 17th and that on my return I will be marking several MSc project reports!

elephant small

I have several delaying (apparently according to Microsoft Word, procrastinatory doesn’t exist) tactics, or low-hanging fruits, that I can pick

Low-hanging fruit

when faced with things I don’t really like doing; I can tidy my desk,

Desk

I can go to check my aphid cultures (a very useful 800 m walk to the glasshouses), I can walk to the Postgraduate Office to check applications (450 m from my office), do a sweep round the PhD offices (also 450 m away from my office) to see how my students are doing, check my email (again and again), accept a request to review a paper, although this then necessitates writing a report ;-) , and as an Editor, I have the very useful excuse that I need to log on to the journal site to check what is happening; the list is long and can certainly fill my day.

elephant medium

In our defence, procrastinators are actually very useful for the academic community; we are the people who agree to review papers, agree to be external examiners, sit on internal and external committees, do a lot of teaching and generally act as good citizens, although perhaps not as altruistically as our colleagues might think, after all we do have an ulterior motive.

Is there a cure?

There are things that we can do; a colleague of mine, does not open her emails on a Monday, I don’t have the will power to do this, I find it difficult enough to ignore my email for two-hour periods, which is my attempt at giving myself a chance to get really into doing some writing. Some people (those who are sociable and find it hard not to go to coffee or chat with colleagues) work from home; again this does not work for me as I then find household chores to distract me. I once went on a time management course in which the use of a daily To Do List was suggested. I dutifully began one of these but found that because of reactive tasks my proactive list just carried on to the next day and never seemed to get completed so that didn’t work for me either, but perhaps I should have said no to the reactive tasks or been less ambitions with the number of things on my To Do List? If anyone has further suggestions please let me know.

So how is that I complete anything substantial? For external tasks this is easy, I promise to deliver, and because I take my solemn promises to other people very seriously indeed, the task gets done, no matter how much I would rather not do it. Unfortunately, promises to myself I find much easier to break, so things that I regard as non-life or career-threatening, do tend to fall by the wayside.

elephant big

 

All in all, I’m very lucky as my procrastinatory (it does exist Microsoft – see here) habits are relatively mild and surmountable, and I have managed to get this far in life without any serious failures.

There are, however, some people who procrastinate because of fear of failure, they set themselves the impossible goal of perfection (unless of course you are a gymnast or diver). This can lead to some very serious failures; for example, students who don’t answer any questions in an exam because they are afraid that their answers will be wrong and hand in a blank answer book, despite having it pointed out to them that if they don’t write anything they will definitely fail. I have also known students who find it almost impossible to hand in written work because they are afraid that it won’t be good enough, despite being told that unless I can see something I can’t help them improve it. To a non-sufferer the solution seems incredibly simple, just get out there and do something, how difficult can it be? It is, however, not simple at all, it can actually be a serious psychological problem that needs very sympathetic and supportive handling, ranging from sharing the task with a friend, colleague or even a suitably sympathetic supervisor, to professional counselling. It is not something to be dismissed lightly as an exasperating foible. Please, please remember this the next time you come across a colleague or student who persistently fails to complete tasks on time. They are not doing it on purpose.  Talk to them about the problem and address the issues sympathetically and with compassion and recommend them to seek psychological help if you feel it is necessary.
Post script

Apropos my latest attempt at writing a major grant proposal, the closing date is 17th May and given that I am writing this on the 11th May and I have, so far only written the opening paragraph of a ten page proposal, I fear that the elephant is about to arrive and trample me ;-)

elephant large

Post post script

It amazes me how many times you hear or see, people using the word prevaricate when they mean procrastinate; they are very different in meaning and not something you would want to own up to in public. Prevaricate means to act or speak in an evasive way (for great examples listen to a politician being interviewed), which to me at least, implies dishonesty.

elephant large

 

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The UK needs more forest health specialists

Last week (April 22nd and 23rd 2015) I had the pleasure of attending the Institute of Chartered Foresters’  National Conference in Cardiff.  The theme of the conference was Tree health, resilience and sustainability.

ICF conference

 The PowerPoint versions of the presentations are available here.

It was very well attended with over 150 delegates and divided into six sessions; Setting the Scene, Overseas Experience, Perspectives on Risk, Searching for Resilience and Sustainability, Practical Responses in the Field and finally Messages for Government and the Profession.  The speakers came from a range of backgrounds; universities, research institutes, the forest industry and others.  Dr John Gibbs, a former colleague of mine from Forest Research opened the formal talks with a masterly review of how forest health problems were tackled in the last century, using Dutch Elm Disease as his focal organism. He was followed by Professor James Brown from the John Innes Institute discussing how lessons from agriculture could be used to develop strategies to combat tree diseases.  Both these speakers pointed out that there was a grave shortage of forest pathologists and entomologists in the UK, particularly in the university sector.   James Brown commented that he had been shocked to discover he had only been able to count seven people in the sector working on tree diseases and added that this did not make them forest pathologists.  We had talks from overseas speakers such as Professor Mike Wingfield from South Africa on global forest health threats, Jim Zwack from the USA speaking on the Emerald Ash Borer as an urban pest problem and Catherine St-Marie highlighting the fact that climate change was aiding and abetting the spread of the Mountain Pine Beetle in Canada.

There was a surprisingly interesting talk on the problems of insuring forests against pests and disease form Phil Cottle of Pardus Underwriting Limited and an enlightening presentation from Professor David Ball from Middlesex University talking about uncertainty and decision-making.  Again both these speakers highlighted the need for further information about pests and diseases.

Day 2 had us searching for resilience and sustainability within the UK forestry sector with a very entertaining talk from Jo O’Hara, Head of Forestry Commission Scotland.  Her talk really drove home to me how much UK forestry has changed over the last 30 years; when I joined the Forestry Commission in 1982 they had only just appointed their first woman District Officer, and now a woman runs FC Scotland – a very welcome sign of change.  Tariq Butt from Swansea University spoke about the use of entomopahogenic fungi as biological control agents in forestry, something increasingly moving higher on the agenda as we face the loss of even more conventional pesticides in the next few years and Martin Ward, the Director-General of EPPO asked us to consider how global plant health arrangements could be improved to protect trees more effectively.  Again the message was that we need more forest health specialist, and not just in the UK.   After the morning coffee break, Joan Webber, the Principal Pathologist for Forest Research UK, spoke about detection and precautionary measures to combat biosecurity threats and yet again highlighted the need for further research and eyes on the ground; in other words more specialist staff are required.  Neil Strong from Network Rail drew our attention to the problems caused by trees to our railway system and then Bill Mason extolled the virtues of increasing species and structural diversity when planting new forests and managing older ones, to improve resilience.

The afternoon session kicked off with Clive Potter from Imperial College talking about understanding what the public’s concerns about tree health are and how certain events can amplify risk perception among the public.  The public outcry about Chalara and Ash Dieback being a particularly good example of the phenomenon.  I followed with a talk about the needs for professional education which gave me the opportunity to point out what subject areas should be covered in an aspiring forester’s education.

Essential skills

I was also able to remind my audience that the number of UK universities providing specific forestry training at undergraduate level had dwindled to less than a handful and that despite offering modules purporting to cover forest health problems, only two employ specialist staff in those areas.  At postgraduate level there is only one course that deals specifically with forest health issues in the UK, the MSc in Conservation & Forest Protection that I run at Harper Adams University.

My take-home messages to a very receptive audience was that students need more emphasis on identification skills and much more practical experience, that current forestry professionals need to keep their eyes open and practice looking for pests and diseases as well as taking any opportunity to refresh their training and that UK universities offering forestry related courses need to employ more forest entomologists and forest pathologists.  Even more importantly, the UK government need to make sure that there are financial incentives to encourage universities to employ more forest entomologists and forest pathologists by increasing targeted research funding in those areas and once increased, maintain those levels of funding.  There also needs to be a clear signposting of career opportunities for the next generation of forest health scientists and if we as a country are serious about safeguarding our native woodlands and forest estate, then more jobs need to be created.

As I have written elsewhere, we cannot afford to sit back and hope that things will get better on their own.  Versions of this slide appeared on the screen several times during the course of the conference.  We are under attack and we need more suitably qualified people to help repel and contain the invaders.

Forest pests

 

Additional reading

Leather, S.R. (2014) Current and future threats to UK forestry. Outlooks on Pest Management, 25, 22-24.

Leather, S.R. (2014) How prepared is the UK to combat future and current threats to forests? Commonwealth Forestry Association Newsletter, 64, 10-11.

 

Post script

I am very grateful indeed to the Institute of Chartered Foresters for giving me the opportunity to speak at the conference and for providing generous hospitality.  It was one of the most engaging and interesting conferences that I have been to for a very long time.  Well done ICF.

 

Post post script

It was also good to see Twitter being used very successfully with the #Treehealth hashtag.  We even had participants from the Canadian Forestry Service!

ICF tweets

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Not all aphids take the same risks

In 1970 an entomologist working on the black bean aphid, Aphis fabae, at Rothamsted Experimental Station (as it then was),  noted that he could categorise the winged individuals as either migrants, flyers or non-flyers; the former flying before they reproduced, the second flying after they reproduced and the final category, never flying (Shaw, 1970).  To describe this phenomenon he used the phrase “migratory urge” a term previously only used in the ornithological literature.

A few years later a group of PhD students in Tony Dixon’s lab at the University of East Anglia started dissecting aphids and counting their ovarioles, finding that unlike most other insects, ovariole number was variable within a species and not related to adult weight (Dixon & Dharma, 1980; Wellings et al., 1980; Leather, 1983).  Generally speaking, in insects, including aphids, the heavier they are, the more fecund they are, although in some instances this is not always true (Leather, 1988).

Ovarioles Fig 1

Figure 1 taken from http://www.aphidsonworldsplants.info/Cloning_Experts_3.htm

Ovarioles Fig 2

Figure 2 What aphid ovarioles really look like Dombrovsky  et al. BMC Research Notes 2009 2:185   doi:10.1186/1756-0500-2-185

What we found then (Wellings et al., 1980), and later (Leather et al., 1988), was that aphids with wings (alatae) even those from the same clone, had much more variability in the number of ovarioles contained within them than those without wings (apterae) (Leather et al., 1988), and that the more ovarioles an aphid contained the more fecund it was, although as mentioned earlier the number of ovarioles appeared to be independent of weight (Leather & Wellings, 1981).

So what does this have to do with migratory urge in Aphis fabae? In the early 1980s Keith Walters was working on migration in cereal aphids (Sitobion avenae and Rhopalosiphum padi) and discovered, that as with Aphis fabae these two species also produced alatae with different flight attributes (Walters & Dixon, 1983).  Building on what we in our group had discovered about ovarioles, Keith was able to show that the degree of migratory urge in aphids was determined by the number of ovarioles they contained. The greater the number of ovarioles the more reluctant they were to take flight (Figure 3ab).

Ovarioles Fig 3a

Figure 3a Relationship between number of ovarioles and time to take-off (minutes) in Sitobion avenae  (Drawn from data in Walters & Dixon, 1983).

Ovarioles Fig 3b

Figure 3b Relationship between number of ovarioles and time to take-off (minutes) in Rhopaloisphum padi  (Drawn from data in Walters & Dixon, 1983).

 He also found that the fewer the number of ovarioles, the steeper the angle of take-off was (Figure 4) i.e. aphids with few ovarioles climbed faster and more steeply and were thus more likely to end  up higher in the air, and thus more likely to travel further than those

Ovarioles Fig 4

Figure 4 Relationship between number of ovarioles and angle of take-off (degrees) in Rhopalosiphum padi (drawn from data in Walters & Dixon, 1983).

taking off at a shallower angle.  He also showed that resistance to starvation was greater in those aphids with fewer ovarioles and that they could also fly for longer periods of time.  Given that alatae of Aphis fabae also have a variable number of ovarioles, 6-12 (Leather et al., 1988), we can see that this fits in very well with Shaw’s classification of migrants, flyers and non-flyers.

This is yet another great example of the flexibility (plasticity) of the aphid clone.  By producing offspring that have different flight capabilities and propensities, the clone is able to hedge its bets in times of adversity; alate aphids in many aphid species are produced in response to crowding and/or poor nutritional quality (Dixon, 1973).  This deterioration in living conditions could be very local i.e. restricted to the plant on which the aphid is feeding or its immediate neighbours, slightly more widespread, i.e. at a field scale or at a much more widespread landscape scale.  Given that long distance aphid migration is very costly (only a tiny proportion survive, Ward et al, 1998) the best option is to spread the risk between the members of your clone.  Those individuals with more ovarioles and greater potential fecundity make the low risk short-distance hops (trivial flights), but take the chance that the next door plant might be just as bad as the one left behind and also within easy reach of natural enemies, but with a higher chance of arriving and reproducing.

Ovarioles Fig 5

A risk taking aphid!

 

At the other end of the scale, those clone members with fewer ovarioles and reduced potential fecundity make the long distance migratory flights, with the risk of not finding a suitable host plant in time, but with the chance that if they do, it will be highly nutritious and natural enemy-free.  A really good example of not putting all your eggs in one basket and yet again a demonstration of what fantastic insects aphids are ;-)

 

References

Dixon, A.F.G. (1973) Biology of Aphids Edward Arnold, London.

Dixon, A.F.G. & Dharma, T.R. (1980) Number of ovarioles and fecundity in the black bean aphid, Aphis fabae. Entomologia Experimentalis et Applicata, 28, 1-14.

Leather, S.R. (1983) Evidence of ovulation after adult moult in the bird cherry-oat aphid, Rhopalosiphum padi. Entomologia experimentalis et applicata, 33, 348-349.

Leather, S. R. (1988). Size, reproductive potential and fecundity in insects: Things aren’t as simple as they seem. Oikos 51: 386-389.

Leather, S.R. & Welllings, P.W. (1981) Ovariole number and fecundity in aphids. Entomologia experimentalis et applicata, 30, 128-133.

Leather, S.R., Wellings, P.W., & Walters, K.F.A. (1988) Variation in ovariole number within the Aphidoidea. Journal of Natural History, 22, 381-393.

Shaw, M.J.P. (1970) Effects of population density on the alienicolae of Aphis fabae Scop.II The effects of crowding on the expression of migratory urge among alatae in the laboratory. Annals of Applied Biology, 65, 197-203.

Walters, K.F.A. & Dixon, A.F.G. (1983) Migratory urge and reproductive investment in aphids: variation within clones. Oecologia, 58, 70-75.

Ward, S.A., Leather, S.R., Pickup, J., & Harrington, R. (1998) Mortality during dispersal and the cost of host-specificity in parasites: how many aphids find hosts? Journal of Animal Ecology, 67, 763-773.

Wellings, P.W., Leather , S.R., & Dixon, A.F.G. (1980) Seasonal variation in reproductive potential: a programmed feature of aphid life cycles. Journal of Animal Ecology, 49, 975-985.

 

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Entomological classics – the pitfall trap

Pitfall arghh I would be amazed if there are any entomologists who have not deployed a pitfall trap or two at some stage in their career. I would also hazard a guess that quite a few non-entomological ecologists have come across the joys of pitfall trap setting and catch sorting as part of their undergraduate training; most field courses seem to include a pitfall trap day, and rightly so.  Pitfall trapping is after all, probably the simplest and most efficient way of collecting data, and not always insects ;-) Pitfall - tapir

Tapir pitfall trap

More seriously though, pitfall traps are a remarkably simple and incredibly versatile way of sampling insects, particularly those that are active on the soil surface (epigeal) e.g carabid beetles. Pitfall forest They can be used in most habitats where you are able to dig into the soil,

Pitfall traps cheap

are very cheap as they can be made from easily obtainable household materials Pitfall traps and can be modified easily depending on your objectives and sampling conditions.  It is very important however, that the lip of the trap is either flush with or below the soil surface.  Not very many beetles or other invertebrates,  are willing to climb up the steep sides  to allow you to capture them. Pitfall - spatial patterns They are also amenable to being deployed in a variety of statistically meaningful ways. (Figure ‘borrowed’ from Woodcock (2005)). Pitfall traps - catch a lot They are of course not perfect.   Some of my students complain that they catch too much!

There has been, and continues to be, much debate about what the catch actually represents.  Are they a measure of activity or of density, i.e. do the trap catches represent the most active and careless beetles, rather than the most abundant?  Southwood (1966) in the first edition of Ecological Methods is fairly dismissive of their use except as a way of studying the activity, seasonal incidence and dispersion of single species and considered them to be of no use whatsoever in comparing communities.  Other authors argue however, that if the trapping is carried out over a long period of time then the data collected can be representative of actual abundance (e.g. Gist & Crossley, 1973; Baars, 1979) and despite Southwood’s comments, they are probably most often used to compare communities (e.g. Rich et al., 2013; Zmihorski et al., 2013;  Wang et al., 2014) For a very thorough account of the use and abuse of pitfall traps see Ben Woodcock’s excellent 2005 article (and I am not just saying that because he is one of my former students). You might expect, given the fact that pitfalls were used by our remote ancestors to trap their vertebrate prey, that entomologists would have adopted this method of trapping very early on, especially given the fact that nature got there first, e.g. as used by larvae of the antlion. Antlion trap

Antlion ‘pitfall traps’.

I was therefore surprised when I started researching this article to find that the earliest reference I could find in the scientific literature was Barber (1931).  I found this very hard to believe so resorted to Twitter.  Richard Jones suggested that a sentence in Pitfall silver sand reference

Notes on Collecting and Preserving Natural History Objects

referring to silver sand pits might be a reference to an early form of pitfall trap.  On further research however, it turned out that sand pits were the results of sand mining operations and were used opportunistically by entomologists.  They worked in a very similar way to Pitfall - St Austell

St Austell Ruddle Moor Sand Pit http://www.cornwall-opc.org/Par_new/a_d/austell_st.php

intercept traps (the subject of a future post).   Interestingly, in some parts of the world, sand pits are now being restored in some places as conservation tools for digger wasp sand bees. Pitfall Bohemia

Sand pit restoration – Bohemia.  http://www.outdoorconservation.eu/project-detail.cfm?projectid=17

  But, I digress.  My next port of call was The Insect Hunter’s Companion (Greene, 1880) which I felt certain would mention pitfall traps.  To my surprise, in the 1880s, entomologists intent on capturing beetles, either pursued them with nets, turned over stones and logs, removed bark from trees, used beating trays or even dug holes in the ground, but never used pitfall traps!  So all very active and energetic methods – no sit and wait in those days ;-) So it seems that Barber’s 1931 description of a pitfall trap does indeed commemorate the first scientific use of a pitfall trap. Barber trap

The Barber trap (Barber, 1931).

Despite their late addition to the entomological armoury and despite the many criticisms levelled at their use, they continue to be perhaps the most widely used method of insect sampling ever; for example if you enter Beetle* AND pitfall* AND trap*  into the Web of Science you will return 1168 hits since 2000, which is more than one a week.  If you further refine your search to exclude beetle but add insect* you can add another 320 hits. If by some chance you have never used a pitfall trap, then I heartily recommend that you set one or two up in a convenient flower bed or even your lawn, and then sit back and wait and see what exciting beasties are roaming your garden.

Post script

Since this post was published I have discovered an earlier reference to the use of pitfall traps (Hertz, 1927).  Many thanks to Jari Niemelä  of Helsinki University for sending me a copy of the reference and many thanks to my eldest daughter for translating the relevant bit, which follows –  “The traps were made of meticulously cleaned tin cans (the rectangle ones used for e.g.  sardines) dug into the ground so deep that the top of the tin was absolutely level with the ground…… it is an ideal way to catch the beetles; with their careless way of running around, they easily fell into the deathtraps, and had no time to use their wings (if they have any)”.  The phrase deathtraps is particularly fine.  The majority of the paper is about the species he caught in different locations and he highlights the fact that he caught seven very rare species using this method.

So this is now the oldest known reference to the use of pitfall traps in the literature, although he does mention that he was using this method to catch beetles in 1914.  But if anyone comes across an earlier reference do let me know.

 

References

Baars, M.A. (1979) Catches in pitfall traps in relation to mean densities of carabid beetles. Oecologia, 41, 25-46.

Barber, H.S. (1931) Traps for cave inhabiting insects.  Journal of the Elisha Mitchell Scientific Society, 46, 259-266.

Gist, C.S. & Crossley, J.D.A. (1973) A method for quantifying pitfall trapsEnvironmental Entomology, 2, 951-952.

Greene, J. (1880) The Insect Hunter’s Companion: Being Instructions for Collecting and Describing Butterflies, Moths, Beetles, Bees, Flies, Etc.  

Hertz, M. (1927) Huomioita petokuoriaisten olinpaikoista.  Luonnon Ystävä, 31, 218-222

Rich, M.C., Gough, L., & Boelman, N.T. (2013) Arctic arthropod assemblages in habitats of differing shrub dominance. Ecography, 36, 994-1003.

Southwood, T.R.E. (1966) Ecological Methods, Chapman & Hall, London.

Wang, X.P., Müller, J., An, L., Ji, L., Liu, Y., Wang, X., & Hao, Z. (2014) Intra-annual variations in abundance and speceis composition of carabid beetles in a temperate forest in Northeast China. Journal of Insect Conservation, 18, 85-98.

Woodcock, B.A. (2005) Pitfall trapping in ecological studies.  Pp 37-57 [In] Insect Sampling in Forest Ecosystems, ed S.R. Leather, Blackwell Publishing, Oxford.

Zmihorski, M., Sienkiewicz, P., & Tryjanowski, P. (2013) Neverending story: a lesson in using sampling efficieny methods with ground beetles. Journal of Insect Conservation, 17, 333-337.

 

Post post script

Pitfall traps are even more versatile than you might think. Mark Telfer has developed a nifty subterranean version http://markgtelfer.co.uk/beetles/techniques-for-studying-beetles/subterranean-pitfall-traps-for-beetles/  and at the opposite end of the spectrum, pitfall traps have also been used in trees to sample spiders (Pinzon & Spence, 2008).

Reference Pinzon, J. & Spence, J. (2008) Performance of two arboreal pitfall trap designs in sampling cursorial spiders from tree trunks.  Journal of Arachnology, 36, 280-286

 

Post post script And for those of you who have had to suffer sitting through the Pokémon movie as I did many years ago, there is also a Pokémon version of the antlion! Pitfall Pokemon

http://bulbapedia.bulbagarden.net/wiki/Trapinch_(Pok%C3%A9mon)

 and don’t forget Winnie the Pooh and his heffalump trap ;-)  Hopefully you will use them more carefully than he did. Pitfall trap - Heffalump

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Celebrating being 60 by walking the Yorkshire Coast – a pictorial record

Those who follow my blog will remember that I turned 60 on March 13th and was pleasantly surprised at lunch time by colleagues, students and friends.

Simon 60 cake

There was however, also another treat in store for me.  My best friend from school (John Pearson) and I used to go walking and camping together in our school and university holidays.

Simon & John T - 21st birthday SRL

Me and John – 21st Birthday Party

 

Family and jobs put an end to this tradition although we and our wives used to (and still do) get together for short walks and visits.  For our 50th birthday however, we diced that it was time to get our walking boots back on in earnest and as good Yorkshiremen we decided that we would walk the Yorkshire Coast (or at least part of it).  In the end we enjoyed a very enjoyable walk from Redcar down to Robin Hood’s Bay, as traditional Yorkshiremen, Redcar, despite boundary changes is still Yorkshire territory as far as we are concerned ;-)

Yorskhire coast 1

 

We stayed in pubs and B&Bs along the way, our camping days being long over.  As a point of honour we walked to the end of every pier (and back) on the way down.

For our 60th birthday, John’s wife Christine paid for three nights in a hotel in Scarborough on the Esplanade for us, The Weston, and we set off to do Robin Hood’s Bay to Bridlington in three days.

Yorkshire coast 2

The weather forecast was truly awful so we were a bit worried, and certainly driving to Scarboroug on Saturday morning, the weather was not promising.  By lunchtime however, the rain had stopped and apart from a short shower we had remarkably good weather for the end of March.  The last two days were, however extremely windy. Each day, we left a car at our finishing point and drove the other car to our start point; that way we were able to stay in comfort in one location and enjoy a well-earned beer at the end of each day and a bottle of very reasonably priced wine at dinner.  Bliss.

 

Day 1 Robin Hood’s Bay to Cloughton

A gentle start as we only had half a day and wanted to break our feet in gently.  Daffodil and Primula garden escapes were very much in evidence on the cliff sides – actually they were present all the way along the coast.

 

Day 1 beer

Staring with a beer – Theakstons Black Bull Bitter

Day 1 Yorkshire best

Yorkshire at its best

Day 1 steep bit

A steep bit (one of many)

Day 1 evening

View from hotel bedroom – evening

Day 2 – Cloughton to Filey

There were some more very steep bits, but despite the gloomy start, it was mainly nice and sunny.  Insects were not much in evidence, but skylarks were very noticeable, and of course there were lots of sea birds.

Day 2 Gloomy morning

First morning – gloomy view from hotel window, but luckily the weather improved as the day progressed.

Day 2 steep bit

There were indeed some steep bits,

Day 2 steep bit Simon

and some very steep bits as well.

Day 2 Slippy bits

Not forgetting the slippy bits!

Day 2 distant objective

Distant objective.

Day 2 dramatic cliffs

Dramatic cliffs.

Day 2 clearing skies

Clearing skies – great views.

Day 3 Filey to Bridlington

An extremely windy day, most of the time we felt like we were walking uphill despite a lot of it being on the flat, on the plus side it was quite sunny.

Day 3 Dawn

Sunrise – a beautiful start to the day

Day 3 - windswept tree

Not just us that was windswept!

Day 3 Bempton

Bempton – lots of Gannets and Guillemots

Day 3 The only puffin

This was the only puffin we saw!

 

Day 3 The sea

The sea, the sea!

Day 3 Footsore

The beach was a bit of a struggle.

Day 2 Flambro

Flamborough Head – we came here for our 3rd form Geography field trip in 1969 – it was joined to the coast then and was a blowhole!

Day 2 Flambro lighthouse

You can’t have a coastal walk without a lighthouse!

Day 4 – Spurn Point

Drove here from Scarborough just to see it as we didn’t feel like slogging along the beach at Bridlington and this is our proposed finishing point for our 65th birthday walk.  We decided that waiting until we were 70 might not be wise ;-)

Spurn Point

Clear(ish) skies but gale force winds.

Spurn Point John

Just like walking in a desert sand storm – we had to turn back after a mile.

Day 3 Spurn Point Brown sea

The sea was brown not blue! Hopefully when we reach this point in 2020 the wind will have abated!

And just to finish a great piece of seaside sculpture.

Freddie Gilroy

Freddie Gilroy on the front at Scarborough.

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Amateur, professional and academic interactions: A day with the British Entomological and Natural History Society

Towards the end of last year I was asked by Claudia Watts, the President of the British Entomological and Natural History Society http://www.benhs.org.uk/site/?q=node/1 if I would like to be nominated to become a member of their Council. I thought about it for a while and then said yes – my resolution to learn to say NO has sadly not been very successful ;-)

There was one snag though, I was not a member of the BENHS; that was, however, easily remedied by completing an application form and being duly approved by the membership secretary and then paying my annual subscription of £19. This process made me think, why wasn’t I already a member of the BENHS? The aims of the Society “are the promotion and advancement of research in entomology with an increasing emphasis now being placed on the conservation of the fauna and flora of the United Kingdom and the protection of wildlife throughout the world”, which is something I have been interested and involved with for almost forty years.

Until I graduated from Leeds the only societies I had belonged to were university ones. On starting my PhD I joined the Association of Applied Biologists, the British Ecological Society, the then Institute of Biology (my father was a founder member) and the Royal Entomological Society. All these had one thing in common, they were, for want of a better description, professional graduate societies, although the Royal Entomological Society did, and does have non-academic and non-professional Members and Fellows. So I guess there was a bit of snobbery involved – the British Entomological and Natural History Society despite its distinguished past, founded 1872, and publishing a quarterly journal and a variety of important identification guides, was in my world view at the time, akin to the Amateur Entomological Society, founded in 1935. A reading of the aims of the AES would however, have told me otherwise, “Our objective is to promote the study of entomology, especially amongst amateurs and the younger generation.”

The objectives of the BENHS, originally the South London Entomological and Natural History Society are remarkably similar to those of the Royal Entomological Society (originally of London), the main differences now, being their financial resources and scale of activities. So having entered an academic career and eventually ending up in one of the UK’s top research intensive universities I had no incentive or inclination to join a society or organisation that did not instantly suggest a professional affiliation, and in the case of the Institute of Biology (now Society of Biology) and the Royal Entomological Society giving me additional letters after my name!

That said, a significant proportion of the UK academic entomological community have published and continue to  publish in the less ‘high impact’ end of the journal spectrum (e.g. Southwood & Johnson, 1957; Dixon, 1958) and indeed me too (Leather & Brotherton, 1987; Leather, 1989). This has been more fully documented elsewhere (Hopkins & Freckleton, 2002). The need for such journals is perhaps not fully appreciated. One of my former students, a Hemipterist of some note, who has published extensively in this journal stratum used to leave these off his job applications until he was asked at a post-doc interview if he had published in these journals as they considered it an important requisite of the job in question as it involved a lot of field work and insect identification. Without journals such as these published here and elsewhere, much of the basic knowledge needed for academic entomologists, ecologists, zoologists etc. to conduct their research would be greatly hampered.

So how does it work in practice? I duly turned up for my first BENHS AGM on Saturday 21st March at the very apposite venue, the Oxford University Museum of Natural History, in time

OUMNH

for the pre-meeting mingle with coffee and biscuits and met some of my former MSc students, now doing PhDs, old friends, current MSc students,  undergraduates, one of whom I had interviewed the day before for a place on the MSc in Entomology that I run, and even a scattering of much younger entomologists.  So a real mixture of amateurs, professionals (those making a living from entomology but not employed by a research institute or university) and current and retired academics.

The programme included a mixture of entomologists, ranging from Matt Shardlow, CEO and founder of Buglife, entomological consultant and saproxylic beetle guru, Keith Alexander, PhD student Tom Wood from Sussex University talking about his pollinator research which has just made the news, author and professional entomologist Richard Jones plugging his latest book and school boy Louis Guillot talking about his research into ant colony development; many of which he stores beneath his bed. We also had an entertaining Presidential Address by Claudia Watts about insects and art through the ages.

Having the AGM in the museum was also of course a very positive plus as in the breaks I was able to have a quick look at some of the exhibits – this one particularly caught my eye, although my camera and photographic skills do not do it justice.

Flea dressed

 

My take home message is that a) I was very foolish not to have got involved with the BENH many years ago and b) given the problems we have in getting the media and general public to understand the importance of entomology and insects to the well-being of the world, that many more of us pre-retirement academic entomologists should be willing to get involved with this and other similar societies.

Insects are important

References

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

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

Leather, S.R. (1989) Phytodecta pallida (L.) (Col.,Chrysomelidae) – a new insect record for bird cherry (Prunus padus). Entomologist’s Monthly Magazine, 125, 17-18.

Leather, S.R. & Brotherton, C.M. (1987) Defensive responses of the pine beauty moth, Panolis flammea (D&S) (Lepidoptera: Noctuidae). Entomologist’s Gazette, 38, 19-24.

Southwood, T.R.E. & Johnson, C.G. (1957) Some records of insect flight activity in May, 1954, with particular reference to the massed flights of Coleoptera and Heteroptera from concealed habitats. Entomologist’s Monthly Magazine, 93, 121–126

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The Seven Ages of an Entomologist – Happy 60th Birthday to Me

Today I turned 60 – an event which has come as a bit of a surprise to me as inside I still feel about 17 ;-) I thought, given the occasion and the fine example set by Jeff Ollerton‘s recent birthday blog post  that it seems a good time to reflect on my career in particular and academic careers in general. Despite there already being at least two other excellent articles about the “Seven Ages”, Jerry Coyne’s, The Seven Ages of the Scientist and Athene Donald’s The Seven Ages of an Academic Scientist, I felt no qualms in adding my own modest contribution to the genre ;-)

Given my own career trajectory it turns out that I need more than seven ages, so as an entomologist I feel justified in adding five larval or nymphal instars to the traditional progression.

 

The Larval Stages

The Infant (first instar)

According to Shakespeare “mewling and puking in the nurse’s arms”, which spending the early part of my childhood in colonial Ghana is actually very apt,

Simon babe in arms

although the photograph below shows a very contented baby indeed.

Simon - baby

I have no entomological memories from this time, although given that then it was normal practice to leave babies outside in their prams, I am sure that I was exposed to the whole range of flying Ghanaian insects. There is some evidence of an early interest in nature and entomology in the picture below where I seem to be investigating a small white butterfly whilst indulging in some early forestry work.

Simon Ghana

My first real biological memory, is however, non-entomological, the blue whale skeleton in the Natural History Museum London in 1958 when my parents were on home leave.

 

The Schoolboy (second instar)

 In 1960 my father was moved to Jamaica to work in the Department of Agriculture as a Plant Pathologist and this is where I started my formal education. Shakespeare describes the schoolboy as “whining schoolboy with his satchel, and shining morning face, creeping like a snail unwillingly to school”.

Simon, Mark & Spences

I certainly had a satchel and it is from this period of my life that I have my first definite entomological memories. We lived in a suburb of Kingston, 32 Gardenia Avenue in Mona Heights. My father kept bees and I spent a lot of time playing with ants, conducting behavioural experiments with crab spiders and having close encounters with wasps and apparently in this picture from 1961, helping with my father’s very luxuriant garden; he grew a great variety of ornamental plants as well as fruit and

Simon 1961      Simon & Pussy cat

 

vegetables, including grapes, bananas, passion fruit, papayas, peanuts and breadfruit as well as coffee and more traditional vegetables. My final school report from my time in Jamaica shows a prescient comment from my biology teacher;

School report

School report bit

 

Secondary school (third instar)

My father’s next posting was to Hong Kong to work for the Ministry of Agriculture; his office was in the New Territories but we lived in Kowloon (Wylie Gardens) where I attended King George V School. Biology was again my favourite subject but apart from cockroaches and ants my entomological experiences were very limited.

Simon - schoolboy           Before braces – 1966

Simon braces

Keeping my mouth shut to hide my orthodontic appliances 1968.

 

Boarding school (fourth instar)

In 1968 my father returned briefly to the UK before his next posting to Fiji and I was sent to a state school, Ripon Grammar School, which had a boarding section. I was to spend five relatively happy years there and despite the competing interests of girls and sports, further developed my interest in invertebrate zoology, due in the main part to my zoology teacher ‘Brian’ Ford. I have many happy memories of pond dipping, searching for Cepea nemoralis and generally fossicking around in hedgerows.

Simon Fiji 1970

When on school holidays in Fiji I found time to investigate the local insect and amphibian fauna; our house seemed to attract toads in huge numbers which my brothers and I used to competitively collect in buckets for later release.

 

Sixth form (final instar)

In my two final years at school sport and girls continued to play a larger part in my life than entomology although I see from the fly-leaf of my books from that time that I owned and had read both volumes of Ralph Bucshbaum’s Life of the Invertebrates and also Darwin’s Origins.

Second fifteen

Ripon Grammar School 2nd XV – I am third from the left on the front row.

 Careers advice when I was at school was not very sophisticated and if you did Biology ‘A’ Level and were a school prefect, it was automatically taken that you were either destined to be a Doctor, a Vet or a Dentist.

School House Prefects1973

I was no different and despite my misgivings, duly applied for and was accepted at Birmingham University to read Medicine. As luck would have it, things did not work out as planned and after a less than happy year at Aston University in Birmingham, in 1974 I left Birmingham and moulted into a proto-entomologist at the University of Leeds.

 

The Undergraduate

The discovery that learning can be fun and that there might actually be a career in doing something that you enjoy.

I did a now extinct degree (although I have plans to exhume it), Agricultural Zoology, essentially a year of vertebrate zoology, with two years of invertebrate zoology, essentially applied entomology, parasitology and nematology. I loved it and thrived on it and grew my hair even longer.

Simon - undergraduate

I decided to become an entomologist in my second year and discovered the wonders of aphids at the same time. It was also round about this time that I decided I was going to become a university academic and started to work a lot harder; the logical end point of someone with a mother who was a secondary school biology teacher and a father who was a research scientist.

 

The Postgraduate

Discovering that being on “the road to find out” (Cat Stevens) is exhilarating

Simon - PhD student

I did my PhD at the University of East Anglia in Norwich – Aspects of the Ecology of the Ecology of the Bird Cherry Aphid, under the supervision of Professor Tony Dixon. A totally fantastic time, despite the ‘second year blues’ which all PhD students seem to go through when they think that they don’t have enough data. I was lucky enough to be in a large research group, at one stage there were thirteen of us in the lab, so there was always plenty of help and advice available. In addition we had the excitement of conferences and the first unsteady steps towards learning to lecture, mainly demonstrating in undergraduate practicals; I spent a lot of time pithing frogs for physiology classes (don’t ask) and also tutoring first year students in mathematics. We also played a lot of squash and enjoyed our social life; for those of you who know Norwich, The Mitre pub on Earlham Road, was our regular haunt.

 

The post-doc

Discovering how to run a research lab

I did two brief post-docs, the first in Finland, under the auspices of the Royal Society and the

Simon Finland 1981

second back at the University of East Anglia funded by the Agriculture and Food Research Council, both working on cereal aphids. At this stage of my career I started to learn how to supervise postgraduate students; the first port of call in a busy lab after the senior PhD student has failed to supply an answer is always the post-doc as the lab head is inevitably very busy. I also got my first real opportunity to lecture undergraduates, which turned out to be a lot harder than I had thought it would be even when talking about my own research.

 

Interlude or host alternation

 The Research Scientist

 Discovering that directed research on its own is not enough

Copy of Simon SSO

In a normal academic career, the next stage after post-doc is an appointment as a University Lecturer. In the early 1980s university lectureships were in short supply and many of us who would normally have gone into an academic career found ourselves either having to go abroad as lecturers at Commonwealth universities (I was offered but turned down a lectureship at Kano University in Nigeria) or joining research institutes. In 1982 I joined the UK Forestry Commission’s Northern Research Station where I spent ten years as a forest entomologist, answering enquiries, conducting directed research and giving the occasional guest lecture. I was however, lucky enough to be able to gain some PhD supervisory experience and after ten years, the last five which were increasingly frustrating, was lucky enough in 1992 to be appointed to a Lectureship at the Silwood Park campus of Imperial College.  In retrospect this was the last time I was able to spend about 90% of my time at the bench and in the field doing ‘hands on’ research, but I have never regretted moving into academia – the opportunity of being able to pass on what you have discovered and hopefully enthuse and motivate a new generation more than makes up for the loss.
Back to the primary host

 

The Lecturer

When I discover that I love teaching

Simon - Lecturer

You may have noticed that I have had a haircut; it was a source of some amusement to me that on joining the university sector I was expected to get my hair cut.

I was appointed as a Lecturer in Pest Management to teach on the world-renowned MSc Entomology course at Silwood Park, and as I was replacing a specific person (Geoff Norton), although not in exactly the same subject area, my ‘grace’ period was shorter that it might have been. Normally at research intensive institutions like Imperial College, new appointments are given two to three years to apply for grants and get their research groups started before being given teaching and departmental jobs. I had a year, but as I discovered that I very much enjoyed teaching (something that many of my colleagues then and later found very strange) I was not dismayed. Unlike some of my colleagues I had read the dictionary definition of the word lecturer: noun. One who delivers lectures, especially professionally.   I have never really understood the mentality of those who aspire to university positions and yet find the idea of having to teach students not only a distraction but in some cases abhorrent and to be avoided at all costs and strive to obtain funding to buy them out of teaching as soon as possible. Some of my senior colleagues at Imperial College (and elsewhere) had and have almost no experience of teaching at all and so have no idea of what is involved in delivering a decent course, a state of affairs that explains some of the very strange decisions that are made at some of the research intensive universities in the UK.   I often felt that they would be much happier in a research institute.

I also discovered that if you take teaching seriously then your ‘bench time’ is much reduced and you begin your career as a research manager, appointing PhD students and post-docs to carry your research ideas forward. I made a decision early on that I would attempt to keep some of my skills extant and set up a long-term field project looking at the insect communities living on sycamores at Silwood Park, especially the aphids. This meant that I had to set a day a week aside to collect data. By doing this it meant that I had a reality check on what was actually possible. I have seen too many colleagues who because of the time they had spent away from the bench or the field, had totally unrealistic expectations of what was actually possible to be achieved by their students and research assistants.

 

The Senior Lecturer

When the Department discovers that I love teaching

In 1996 I was promoted to Senior Lecturer (I think that it is a real shame that some UK universities have decided to adopt North American terminology and introduce the title of Associate Professor, apparently to avoid confusing the rest of the World. At Imperial College promotion to Senior Lecturer was to reward teaching excellence and was usually the kiss of death for any further promotion.

Simon - Lecturera

Senior Lecturer in Applied Ecology

 I was as well as teaching on the MSc Entomology course doing an increasing amount of undergraduate teaching including a final year course in Applied Ecology of which I was very proud, hence the decision to retitle myself. I was also very busy with external activities, being on the Editorial Board of the Bulletin of Entomological Research and just been appointed as Editor-in-Chief of Ecological Entomology, just finished a term on the council of the Royal Entomological Society and been appointed to a slew of Departmental and University committees. My research group was really starting to take off, I was supervising 8 PhD students at the time; given the poor return rate on major grant applications in the UK, I decided early on that going for PhDs was a better use of my limited time and this is a strategy that I have mainly followed to the present day.

Research group

This does not include MSc or BSc students – they would add about 10 to each yearly figure from 1995 onwards

The Reader

 When I discover that it is possible to get even busier

In 2002 I was promoted to Reader one of the definitions of which according to Chambers’s Twentieth Century Dictionary is defined as follows; Old English rǣdere ‘interpreter of dreams, reader’. In the UK university system, it is the rank below full Professor and comes with an endowed title, in my case I chose to become Reader in Applied Ecology to reflect the

Simon - Reader

myriad teaching roles I had accumulated and also to encompass the fact that my research group no longer dealt solely with arthropods, vertebrates had somehow sneaked their way in. Looking at Athene Donald’s list I see that I was pretty much doing a professorial role, serving on external committees, validating degrees for other universities and acting as an external examiner. I was also appointed as Editor-in-Chief of Insect Conservation and Diversity, a new journal for the Royal Entomological Society. My administrative duties had also continued to increase.  It was no wonder that my beard was getting greyer! I was however still preparing my own talks, although I will confess that a lot of my data analysis was being passed on to members of the group, duly acknowledged of course. I am extremely grateful that I have always had a loyal and very supportive research group, without their help life would have been impossible.  My thanks to you all (if any of you are reading this).

 

The Professor

Discovering the joys of being pretty much able to do what I want (with certain restrictions)

It became increasingly obvious that things could not carry on as they were, my teaching and administrative loads were becoming ridiculous; our Director of Teaching calculated that I was actually doing more teaching than anyone else in the Department including the Teaching Fellows. I was seriously considering early retirement although I was reluctant to do this as I was sure that with my retirement the last entomology degree in the UK would quickly disappear. Luckily in 2012 my team and I were miraculously offered the chance to move to a new more supportive location, Harper Adams University in Shropshire.

Simon 2015

So now I have become a Senior Professor, with a new entomology building, with less undergraduate teaching, which I miss, and a role that requires me to sit on more external and internal committees, to meet the great and the good and to make solemn pronouncements.  At the same time however, it does allow me to plough my own furrow and to influence university policy. Most importantly I no longer feel that I am beating my head against a brick wall and that the future of entomology as a degree course in the UK is much safer than it was five years ago.  I think I am at Stage 4 in Jerry Coyne’s list as I now find that I am much more interested in synthesizing and disseminating what I have learnt rather than doing original research – I can feel a book coming on ;-)

My hope is that in five years time when I become a retired Professor and my hair and beard colour are the same, that entomology will be taught at more than one university in the UK and not just at postgraduate level.

A small point of personal satisfaction, is that, despite my elevation, I still do not own a suit ;-)

 

For reference

Jerry A. Coyne’s summary, reproduced from his blog

  1. As student, listens to advisor give talk on student’s own work
  2. As postdoc, gives talks about his/her own work
  3. As professor, gives talks about his/her students’ work
  4. Talks and writes about “the state of the field”
  5. Talks and writes about “the state of the field” eccentrically and incorrectly—always in a self-aggrandizing way.
  6. Gives after-dinner speeches and writes about society and the history of the field
  7. Writes articles about science and religion

 

And the famous original from which the title is borrowed and adapted.

 

Seven Ages Of Man

(from As You Like It by William Shakespeare)

All the world’s a stage,

And all the men and women merely players,

They have their exits and entrances,

And one man in his time plays many parts,

His acts being seven ages. At first the infant,

Mewling and puking in the nurse’s arms.

Then, the whining schoolboy with his satchel

And shining morning face, creeping like snail

Unwillingly to school. And then the lover,

Sighing like furnace, with a woeful ballad

Made to his mistress’ eyebrow. Then a soldier,

Full of strange oaths, and bearded like the pard,

Jealous in honour, sudden, and quick in quarrel,

Seeking the bubble reputation

Even in the cannon’s mouth. And then the justice

In fair round belly, with good capon lin’d,

With eyes severe, and beard of formal cut,

Full of wise saws, and modern instances,

And so he plays his part. The sixth age shifts

Into the lean and slipper’d pantaloon,

With spectacles on nose, and pouch on side,

His youthful hose well sav’d, a world too wide,

For his shrunk shank, and his big manly voice,

Turning again towards childish treble, pipes

And whistles in his sound. Last scene of all,

That ends this strange eventful history,

Is second childishness and mere oblivion,

Sans teeth, sans eyes, sans taste, sans everything.

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The Verrall Supper 2015 – A Photographic Record

Wednesday March 4th 2015 was the occasion of the latest Verrall Supper, an annual event hosted by the Entomological Club, the oldest extant entomological society in the world.  I am, for my sins, the Supper Member, which means that I have to organise the event, for a more detailed description of my role click here.  This year we continued our association with The Rembrandt Hotel as they had done such a good job last year and the year before.  This year almost all the invitations were sent by email and despite the 14% increase in suggested subscription to £48, we had a very good response; as I pointed out to one of the students, show me somewhere else in central London where you can get a three course dinner with coffee and  half a bottle of wine, plus the company of so many entomologists!

In the end we had 181 guests, 55 of whom were female, last year we only had 46 female members so we are definitely moving in the right direction, although I am still keen to get equal numbers.  My impression was that the average age of the membership is definitely decreasing which can only be a good thing.  Enough writing I think, let the photographs speak for themselves.

Jim Hardie & Clive Farrell

Jim Hardie with Clive Farrell of the Entomological Club – once again my thanks to Clive for helping man the Reception Desk

A mixed bag

A mixed bag of entomologists enjoying good conversation whilst waiting for the main course

All ex-students

All ex- or present students of mine, Katy Reed, Lauren Fuller, Jen Banfield-Zanin, Mark Ramsden, Aislinn Pearson

Andrew Salisbury holds forth

Andrew Salisbury, RHS Wisley, holds forth

 

AShleigh & Craig

Ashleigh Whiffin and Craig Perl recreate last year’s photo

Can you find the coleopterist

Can you spot the Coleopterist?

Entomologists with beer

Entomologists with beer

Garth Foster being very definite

Garth Foster making a point

Gemma & James

Gemma Hough and James Hourston

Hagrid

Did you know that Hagrid was an entomologist?  Actually Richard Comont

Helen Roy

Can you spot Helen Roy, the newest member of the Entomological Club?

Jade, Linda & Laurence

Three ex-MSc students – Jade Taylor, Linda Birkin and Laurence Livermore and Hillery Warner.

Mainly current MSc

Mainly current Harper Adams MSc students – Josh Jenkins Shaw, Chris Mackin, Andy Cutts, Aidan Thomas, Richard Prew, Kelleigh Greene, Jordan Ryder (now a PhD student) Dave Stanford-Beale

Marion Gratwick

Marion Gratwick has attended more Verrall Suppers than any other female member.

Mark, Adriana & Jen

Mark Ramsden, Adrian De Palma, Jen Banfield-Zanin, Gemma Hough

More young entomologists

More young entomologists, Ailsa McLean, Paul Manning, Chris Jeffs and James Hourston

Old and young mixing

John Badmin centre stage

Older male entomologists

Some older entomologists

Romantic!

Linda Birkin, Laurence Livermore, Hillery Warner, Aurora Sampson

Some Hymenopterists

Some hymenopterists, including Mark Shaw and Charles Godfray

Some RES worthies

Some Royal Entomological Society worthies including Gordon Port, Jim Hardie and Archie Murchie

Steve Clement & Gill van Emden

On the top table, Stephen Clement who travelled all the way from the USA, speaking to Gill van Emden

Top Table 1

The top table, Gill van Emden, Stephen Clement, Van, Clive Farrell, Chris Lyal, another overseas visitor Junhao Huang, my jacket, Camille Parmesan, Richard Harrington and Tilly Collins out of shot.

Tilly

Has Richard Harrington made Tilly Collins cry?  Are his jokes really that bad?

Sue Hartley, Hugh Loxdale, Peter Leckstein

The Verrall lecturer, Sue Hartley in deep conversation with Hugh Loxdale and Peter Leckstein.  Unfortunately she couldn’t stay for the Supper.

Young female entomologists

Not all entomologists are male and bearded, Ruth Carter, Kirsten Miller, Fran Sconce, Will Nash, Nathan Medd , Hannah Wickenden, Jasper Hubert

Young mixed entomologists

More young entomologists Joe Nunez, Ricahrd Comont, Amo Spooner, Katy Dainton, Molly Carter, Sally-Ann Spence

Older male entomologists

Some of the older entomologists

and finally

Simon stressed

The Supper Organiser looking a little bit stressed!

I look forward to seeing all of you next year on Wednesday 2nd March.

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Ten Papers that Shook My World – Haukioja & Niemelä (1976) – the plant “immune response”

To me this is a landmark paper, both personally and for ecology in general.   I first came across it in the second year of my PhD at the University of East Anglia (1978) and given where it was published, would probably never have seen it if my supervisor, Tony Dixon, hadn’t had a collaborative link with Erkki Haukioja of Turku University (Finland).

That individual plants of the same species are more or less susceptible (constitutive or innate resistance) to pests and diseases has been known for a very long time (e.g. Painter, 1958; Beck, 1965) and has been exploited by plant breeders as part of many pest management programmes.  Despite the stunning footage of the questing bramble in David Attenborough’s classic documentary The Private Life of Plants, plants are often thought of as passive organisms.  The idea that plants might actually respond directly and quickly to insect attack was more in the realms of science fiction than science fact, but this all changed in the 1970s. In 1972 a short paper in Science (Green & Ryan, 1972) suggested that plants might not be as passive as previously thought. Green & Ryan working in the laboratory with the Colorado Potato Beetle, Leptinotarsus decemlineata, showed that when tomato leaves were damaged by beetle feeding the levels of a proteinase inhibitor were raised not just in the wounded leaves but in nearby leaves as well. As proteinase inhibitors were well-known to be part of the plant defence system, they hypothesised that this was a direct response of the plant to repel attack by pests and that it might be a useful tool in developing new pest management approaches. So what does this have to do with two Finnish entomologists?

Erkki Haukioja and his long-term collaborator, Pekka Niemelä were working on an important lepidopteran defoliator of birch, in the far north of Finland, at the Kevo Subarctic Research Station.Kevo

http://www.eu-interact.org/field-sites/finland-4/kevo/

The defoliator that they were working on was the autumnal moth, now Epirrita autumnata, but then Oporinia autumnata.

Epirrita

http://ukmoths.org.uk/show.php?bf=1797

The autumnal moth, as with many tree-feeding Lepidoptera, has a 7-10 year population cycle (Ruohmäki et al., 2000).

Population cycles

Natural enemies are often cited as the causes of these cycles (Turchin et al., 1999) although other factors such as weather (Myers, 1998) or even sunspot activity (Ruohmäki et al., 2000)

Sunspot

have also been suggested. It had also been suggested that the marked population cycles of the larch bud moth, Zeiraphere diniana were caused by changes in the susceptibility of their host trees after defoliation (Benz, 1974). In 1975, Haukioja and his colleague Hakala, attempting to explain the cyclical nature of the E. autumnata population cycles wondered if they were being driven by the insects themselves causing changes in the levels of chemical defence in the trees. To test this Erkki and Pekka did two neat field experiments, remember Green & Ryan’s work was laboratory based and did not test the effects seen on the insects. They first fed Epirrita larvae on foliage from previously defoliated and undefoliated birch trees and found that the pupae that developed from those larvae fed on previously defoliated trees were lighter than those that had fed on previously undefoliated trees (Hauikioja & Niemelä, 1976). At the same time they also did an experiment where they damaged leaves but then rather than feeding the larvae on those leaves, fed them on nearby adjacent undamaged leaves and compared them with larvae feeding on leaves from trees where no damage had occurred. Those larvae feeding on undamaged leaves adjacent to damaged leaves grew significantly more slowly than those feeding on leaves that came from totally undamaged trees (Haukioja & Niemelä, 1977). So pretty convincing evidence that the trees were responding directly to insect damage and altering their chemistry to become more resistant, i.e. an induced defence and not a constitutive one.

Their results had a major impact on the field. The great and the good from around the world found it a fascinating subject area and a plethora of papers investigating the effects of insect feeding on induced defences in birch and willow trees soon followed (e.g. Fowler & Lawton, 1984a; Rhoades, 1985; Hartley & Lawton, 1987) and not forgetting the original researchers (e.g. Haukioja & Hahnimäki, 1984). I, with the aid of colleagues, also added my ‘two pennorth’ (I did say the idea shook my world) by extending the concept to conifers (Leather et al., 1987; Trewhella et al., 1997). The terms rapid induced resistance and delayed induced resistance soon entered the language, the first to describe those changes that occurred within minutes of feeding damage and the second, those that did not take effect until the following year (Haukioja & Hahnmäki, 1984; Ruohmäki et al., 1992) Such was the interest generated by the topic that by 1989 there were enough studies for a major review to be published (Karban & Myers, 1989).

Controversy reared its ugly head early on when Doug Rhoades suggested that not only did plants resist insect attack actively but that they could talk to each other and warn their neighbours that the ‘bad guys’ were in the neighbourhood (Rhoades, 1983, 1985). This sparked a brief but lively debate (e.g. Fowler & Lawton, 1984b, 1985). Ironically it is now taken as axiomatic that plants talk to each other using a range of chemical signals (van Hulten et al., 2006; Heil & Ton, 2008) as well as informing the natural enemies of the pests that a suitable food source is available (e.g. Edwards & Wratten, 1983; Amo et al., 2013; Michereff et al., 2013).

Ton cartoon

A great cartoon from Jurriaan Ton at Sheffield University. https://www.shef.ac.uk/aps/staff-and-students/acadstaff/ton-jurriaan

We now have a greatly increased understanding of the various metabolic pathways that induce these defences against different insect pests (e.g. Smith & Boyko, 2007) and can, by genetically manipulating levels of compounds such as jasmonic and salicyclic acids or even applying them directly to plants affect herbivorous insect communities and their natural enemies thus improving crop protection (e.g. Thaler, 1999; Cao et al., 2014; Mäntyllä, 2014). No wonder this was an idea that shook my world, and yours.

 

Post script

The study of induced plant defences or resistance is now dominated by molecular biologists and current practice is to use the term priming and not induced defence. The increased understanding that this new generation has brought to the field is undeniable but I always feel it is a great shame that they seem to have forgotten those early pioneers in the field.

 

References

Amo, L., Jansen, J.J., Van Dam, N.M., Dicke, M., & Visser, M.E. (2013) Birds exploit herbivore-induced plant volatiles to locate herbivorous prey. Ecology Letters, 16: 1348-1355.

Baldwin, I.T. & Schultz, J.C. (1983) Rapid changes in tree leaf chemistry, induced by damage: evidence for communication between plants. Science, 221, 277-279.

Beck, S.D. (1965) Resistance of plants to insects. Annual Review of Entomology, 10, 207-232.

Benz, G. (1974). Negative Ruckkoppelung durch Raum-und Nahrungskonkurrenz sowie zyklische Veranderung. Zeitschrift für Angewandte Enomologie, 76: 196-228.

Cao, H.H., Wang, S.H., & Liu, T.X. (2014) Jasomante- and salicylate-induced defenses in wheat affect host preference and probing behavior but not performance of the grain aphid, Sitobion avenae. Insect Science, 21, 47-55.

Edwards, P.J. & Wratten, S.D. (1983) Wound induced defences in plants and their consequences for patterns of insect grazing. Oecologia, 59: 88-93.

Fowler, S.V. & Lawton, J.H. (1984a) Foliage preferences of birch herbivores: a field manipulation experiment. Oikos, 42: 239-248.

Fowler, S.V. & Lawton, J.H. (1984b) Trees don’t talk : do they even murmur? Antenna, 8: 69-71.

Fowler, S.V. & Lawton, J.H. (1985) Rapidly induced defences and talking trees: the devils’ advocate position. American Naturalist, 126: 181-195.

Green, T.R. & Ryan, C.A. (1972) Wound induced proteinase inhibitor in plant leaves: a possible defense mechanism against insects. Science: 175: 776-777.

Hartley, S.E. & Lawton, J.H. (1987) Effects of different types of damage on the chemistry of birch foliage and the responses of birch feeding insects. Oecologia, 74: 432-437.

Haukioja, E. & Hakala, T. (1975) Herbivore cycles and periodic outbreaks. Report of the Kevo Subarctic Research Station, 12: 1-9

Haukioja, E. & Hanhimäki, S. (1984) Rapid wound induced resistance in white birch (Betula pubescens) foliage to the geometrid Epirrita autumnata: a comparison of trees and moths within and outside the outbreak range of the moth. Oecologia, 65, 223-228.

Haukioja, E. & Niemelä, P. (1976). Does birch defend itself actively against herbivores? Report of the Kevo Subarctic Research Station 13: 44-47.

Haukioja, E. & Niemelä, P. (1977). Retarded growth of a geometrid larva after mechanical damage to leaves of its host tree. Annales Zoologici Fennici 14: 48-52.

Heil, M. & Ton, J. (2008) Long-distance signalling in plant defence. Trends in Plant Science, 13: 264-272.

Karban, R. & Myers, J.H. (1989) Induced plant responses to herbivory. Annual Review of Ecology & Systematics, 20: 331-348.

Leather, S.R., D., W.A., & Forrest, G.I. (1987) Insect-induced chemical changes in young lodgepole pine (Pinus contorta): the effect of previous defoliation on oviposition, growth and survival of the pine beauty moth, Panolis flammea. Ecological Entomology, 12: 275-281.

Mäntyllä, E., Blande, J.D., & Klemola, T. (2014) Does application of methyl jasmonate to birch mimic herbivory and attract insectivorous birds in nature? Arthropod-Plant Interactions, 8, 143-153.

Michereff, M.F.F., Borges, M., Laumann, R.A., Dinitz, I.R., & Blassioli-Moraes, M.C. (2013) Influence of volatile compounds from herbivore-damaged soybean plants on searching behavior of the egg parasitoid Telonomus podisi. Entomologia experimentalis et applicata, 147: 9-17.

Trewhella, K.E., Leather, S.R., & Day, K.R. (1997) Insect induced resistance in lodgepole pine: effects on two pine feeding insects. Journal of Applied Entomology, 121: 129-136.

Myers, J. H. (1998). Synchrony in outbreaks of forest lepidoptera: a possible example of the Moran effect. Ecology 79: 1111-1117.

Painter, R.H. (1958) Resistance of plants to insects. Annual Review of Entomology, 3: 267-290.

Rhoades, D.F. (1983) Responses of alder and willow to attack by tent caterpillar and webworms: evidence for pheromonal sensitivity of willows. American Chemical Society Symposium Series, 208: 55-68.

Rhoades, D.F. (1985) Offensive-defensive interactions between herbivores and plants: their relevance in herbivore population dynamics and ecological theory. American Naturalist, 125: 205-238.

Ruohomäki, K., Hanhimäki, S., Haukioja, E., Iso-iivari, L., & Neuvonen, S. (1992) Variability in the efficiency of delayed inducible resistanec in mountain birch. Entomologia experimentalis et applicata, 62: 107-116.

Ruohmäki, K., Tanhuanpää, M., Ayres, M.P., Kaitaniemi, P., Tammaru, T. & Haukioja, E. (2000) Causes of cyclicity of Epirrita autumnata (Lepidoptera, Geometridae): grandiose theory and tedious practice. Population Ecology, 42: 211-223

Smith, C.M. & Boyko, E.V. (2007) The molecular basis of plant resistance and defence responses to aphid feeding: current status. Entomologia experimentalis et applicata, 122: 1-16.

Thaler, J. (1999) Induced resistance in agricultural crops: effects of Jasmonic acid on herbivory and yield in tomato plants. Environmental Entomology, 28, 30-37.

Turchin, P., Taylor, A. D. &Reeve, J. D. (1999). Dynamical role of predators in population cycles of a forest insect: an experimental test. Science 285: 1068-1071.

Van Hulten, M., Pelser, M., van Loon, L.C., Pieterse, C.M.J. & Ton, J. (2006) Costs and benefits of priming for defense in Arabidopsis. Proceedings of the National Academy of Sciences USA, 103: 5602-5607.

 

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