Tag Archives: Panolis flammea

Ideas I had and never followed up

“When I was younger, so much younger than before” I never needed any help to come up with ideas for research topics or papers.   When I was doing my PhD and later as a post-doc, I used to keep a note pad next to my bed so that when I woke up in the middle of night with an idea (which I often did) I could scribble it down and go back to sleep.  (These days sadly, it is my bladder and not ideas that wake me up in the wee small hours 🙂*)

On waking up properly, these ideas, if they still seemed sensible, would  move onto Stage 2, the literature search.  In those days, this was much more difficult than it is now, no Google Scholar or Web of Science then, instead you had to wade though the many hard-copy Abstract series and then get hard copies of the papers of interest.  Once in my hands, either via Inter-library loans or direct from the author, or even photocopied from the journal issue (we did have photocopiers in those days), the papers would be shoved into a handy see-through plastic folder (Stage 3).  Depending on how enthusiastic I was about the idea, I would then either mock-up a paper title page or put the folder in the ‘to deal with later’ pile (Stage 4).   Many of these eventually led on to Stage 5, experiments and published papers.  Others have languished in their folders for twenty or thirty years.

As part of my phased run up to retirement (2021), I have started farming out my long-term publishable (hopefully) data-sets to younger, more statistically astute colleagues and ‘publishing’ less robust, but possibly useful data on my blog site.  I have also, somewhat halfheartedly since the task is monumental, started to go through my old field and lab books that


A monumental collection of data.  The top right picture is my 20-year sycamore data set.  I estimate that there are about 7 million data points in it; of which to date only 1.6 million, give or take a million, are computerised.  I also have a ten-year bird cherry aphid data set from Scotland, waiting to go on the computer, any volunteers?

are not yet computerised.  Whilst doing this I came across some Stage 3 folders, which as you can see from the colour of the paper have languished for some time.


The Forgotten Nine


There were nine forgotten/dismissed proto-papers, the oldest of which, judging by the browning of the paper and my corresponding address, dates from the early 1980s, and is simply titled “What are the costs of reproduction?”.  This appears to have been inspired by a talk given by Graham Bell at a British Ecological Society, Mathematical Ecology Group meeting in 1983.  In case you are wondering, this was one of those meetings supposed to bring theorists and empiricists together.   It didn’t work, neither group felt able to talk to each other 🙂  The idea, inevitably based on aphid data, didn’t bear any fruit, although I do have this graph as a souvenir.  If anyone wants


In those days we used graph paper 🙂

 the data, do let me know.

Slightly later, we find the grandly titled, “Size and phylogeny – factors affecting covariation in the life history traits of aphids”.  This had apparently been worked up from an earlier version of a paper, less grandly, but no less ponderously, titled, “Size and weight: factors affecting the level of reproductive investment in aphids”.  This is based on some basic dissection data from eight aphid species and presents the relationships, or lack of, between adult weight (or surrogate measure), ovariole number, potential fecundity and the number of pigmented embryos.  As far as I can remember these are data that Paul Wellings** and I collected as a follow-up to work we had published from a side project when we were doing our PhDs at the University of East Anglia (Wellings et al., 1980).  The second title was inspired by a paper by Stephen Stearns (Stearns, 1984), who was something of a hero of mine at the time, and was, I guess, an attempt to publish pretty simple data somewhere classier than it deserved 🙂  So this one seems to be a Stage 4, almost Stage 5 idea, and may, if I have time or someone volunteers, actually get published, although I suspect it may only make it to a very minor journal under its original title.

Then we have a real oddity, “Aphids, elephants and oaks: life history strategies re-examined”.  This one as far as I remember, is based on an idea that I had about r- and k-selection being looked at from a human point of view and not the organism’s point of view.  My thesis was that an oak tree was actually r-selected as over its life-time it was more fecund than an aphid 🙂  I suspect this was going to be aimed at the Forum section of Oikos.

The next one, dates from the late-1980s, “Protandry versus protogyny: patterns of occurrence within the Lepidoptera”, and reflects the fact that females of the pine beauty moth, Panolis flammea, on which I was then working, emerge before the males (Leather & Barbour, 1983; Leather, 1984), something not often reported in Lepidoptera.  I wondered what advantage (if any) this gave P. flammea.  I planned this one as a review or forum type paper but never got beyond the title and collecting two references (Robertson, 1987; Zonneveld & Metz, 1991).  I still think this is an interesting idea, but do feel free to have a go yourselves, as again, I suspect that I won’t actually get round to it.

Finishing off my time in Scotland, is a paper simply entitled, “Egg hatch in the bird cherry aphid, Rhopalosiphum padi.” I have ten years of egg hatch data from eight trees waiting to be analysed.  This is almost certainly not worth more than a short note unless I (or a willing volunteer) tie it in with the ten years data on spring and autumn populations on the same trees 🙂 Aphid egg data although not very abundant, is probably not in great demand.  My first published paper (Leather, 1980) was about egg mortality in the bird cherry aphid and 36 years later has only managed to accrue 32 citations, so I guess not an area where one is likely to become famous 🙂

I then have four papers dating from my time as an Associate Member of the NERC Centre for Population Biology at Silwood Park.   The first is titled, “The suitability of British Prunus species as insect host plants” and was definitely inspired by my foray into counting host plant dots as exemplified by the late great Richard Southwood (Leather, 1985, 1986).  I think I was going to look at palatability measures of some sort.

The next is called ‘Realising their full potential: is it important and how many insects achieve it?”  I’m guessing that this was a sort of follow-up to my second most-cited paper ever (Leather, 1988), the story of which you can read here, if at all interested.  Most insects, even those that are pests, die before achieving anywhere near their full reproductive potential, but then so do we humans, and our population continues to grow.  So in answer to the question, I guess not and no it doesn’t matter 🙂

Also linked to insect reproduction is the next paper, which I have followed up with the help of a PhD student, and do hope to submit in the near future, “Queue positions, do they matter”.  As this one may actually see the light of day, I won’t say anything further about it.

And finally, another one about aphid eggs, “Bud burst and egg hatch synchrony in aphids”.  This one was going to be based on my then ten-year sycamore aphid data but is now based on my twenty-year data set and is now in the very capable hands of a PhD student and hopefully will see the light of day next year.

There are also a number of other folders with no titles that are just full of collections of reprints.  I can only guess at what these ideas were so won’t burden you with them.

I mentioned at the beginning of this piece that I don’t wake up in the middle of the night with ideas any more.  As we get older I think there is a tendency to worry that we might run out of ideas, especially when, as we do in the UK, suffer from ludicrously underfunded research councils with very high rejection rates that don’t allow you to resubmit failed grant applications.  It was thus reassuring to see this recent paper that suggests that all is not lost after you hit the grand old age of 30.  That said, I do believe that as you move away from the bench or field, the opportunity to be struck by what you see, does inevitably reduce.  As a PhD student and post-doc you are busy doing whatever it is you do, in my case as an ecological entomologist, counting things, and inevitably you see other things going on within and around your study system, that spark off other ideas.  It was the fear of losing these opportunities as I moved up the academic ladder, which inevitably means, less field and bench time and more time writing grant applications and sitting on committees, that I specifically set aside Monday mornings (very early mornings) to my bird cherry plots and even earlier Thursday mornings to survey my sycamore trees.   Without those sacrosanct mornings I am pretty certain I would have totally lost sight of what is humanly possible to do as a PhD student or post-doc.  This, thankfully for my research group, means that I had, and have, realistic expectations of what their output should be, thus reducing stress levels all round.   As a side benefit I got to go out in the fresh air at least twice a week and do some exercise and at the same time see the wonderful things that were going on around and about my study areas and as a bonus had the chance to get some new ideas.



Leather, S.R. (1984) Factors affecting pupal survival and eclosion in the pine beauty moth, Panolis flammea (D&S). Oecologia, 63, 75-79.

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 host range, plant architecture, age of establishment, taxonomic isolation and species-area relationships. Journal of Animal Ecology, 55, 841-860.

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. & Barbour, D.A. (1983) The effect of temperature on the emergence of pine beauty moth, Panolis flammea Schiff. Zeitschrift fur Angewandte Entomologie, 96, 445-448.

Robertson, H.G. (1987) Oviposition and site selection in Cactoblastis cactorum (Lepidoptera): constraints and compromises. Oecologia, 73, 601-608.

Stearns, S.C. (1984) The effects of size and phylogeny on patterns of covariation inthe life history traits of lizards and snakes. American Naturalist, 123, 56-72.

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.

Zonneveld, C. & Metz, J.A.J. (1991) Models on butterfly protandry – virgin females are at risk to die. Theoretical  Population Biology, 40, 308-321.


*I hasten to add that I do still have new ideas, they just don’t seem to wake me up any more 🙂

**Now Vice-Chancellor of the University of Wollongong



Filed under Science writing, Uncategorized

When frustration becomes serendipitous – My second most cited paper

For most of the 1980s and the early 1990s I worked for the UK Forestry Commission as a research and advisory entomologist. As a civil servant I was subjected to a lot more rules than I am now as a university academic. The most frustrating set of rules in my mind, were those associated with publishing papers. The initial consultation with a statistician before your experiment was planned and any subsequent collaboration with the analysis was very sensible, and I had no problems with that part of the process at all. Our statisticians were very good in that they helped you decide the analysis but expected you to learn GenStat (the Forestry Commission standard statistics programme) and do it yourself unless you were really stuck.

The next bit was the frustrating part. When it came to writing papers you first submitted your paper to your line manager. They then read your paper, very frustrating indeed for me, as my immediate boss considered papers a very low priority and it could be several months before he got around to passing it back with comments and suggestions. Then it was passed to a member of one of the other department such as silviculture, tree breeding or pathology for them to read and make comments. The idea behind this being that it helped make the paper accessible to a wide audience, again a good idea. The problem at this stage was that once again your paper was likely to be a low priority, so yet more delay. Once that was done you then had to submit your paper to the Chief Research Office for him to read and comment on, so once again yet more delay. This meant that quite often it was a year before you actually were able to submit your paper to a journal, which could be deeply frustrating to say the least.


In 1986 a new journal to be published by the British Ecological Society was announced, Functional Ecology. In those days, the dreaded Impact Factors had not yet raised their ugly heads, and one tended to publish in journals relevant to your discipline, or, as in this case, the fancy took you.  I thought it would be cool to publish in the first issue of the first volume of this new journal.  I therefore set to work, with the help of one of our statisticians to produce a paper about life history parameters of the pine beauty moth, from a more ecological point of view and not from the more applied view-point of it as a forest pest (my job remit). I was very proud of the paper and confess to having got somewhat carried away in the discussion, so much so, that it was suggested by all who read it in the very lengthy internal appraisal process, that most of the discussion should be cut as being too far away from the main story. As the process had taken so long already I decided to go with the flow and eventually submitted my paper about a year after first writing it, incidentally giving my statistician a co-authorship. It was accepted and did indeed appear in the first volume of Functional Ecology, albeit the last of the year (Leather & Burnand, 1987)! It has to date (14th October 2015) being cited 53 times, by no means a disgrace, but certainly not my second-most cited paper.

I mentioned earlier that I was really proud of my discussion and I decided that I was going to publish it regardless. I reworked it slightly and submitted it to Oikos as a Forum piece, taking the calculated risk of not submitting it through the official Forestry Commission system. My reasoning was, that a), it was unlikely to be read by anyone in the Forestry Commission, being a very ecological journal, and b), if challenged I would say that it had already been seen by the powers that be, albeit not officially. To my relief it was accepted as is (Leather, 1988) and my immediate boss never mentioned it. To my surprise and delight this is now my second-most cited paper, having so far acquired 207 citations and still picks up a reasonable number of cites every year. I guess that I should actually be grateful to all those internal referees who insisted that I cut my discussion down so drastically.


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. & Burnand, A.C. (1987) Factors affecting life-history parameters of the pine beauty moth, Panolis flammea (D&S): the hidden costs of reproduction. Functional Ecology, 1, 331-338.


Post script

In case you wondered, my most cited paper is an Annual Review paper, written with one of my former PhD students, Caroline Awmack, and now has almost a thousand citations (994 as of today).


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



Filed under Bugbears, Science writing, Uncategorized

Midwinter Madness – The Snow flea

Between 1982 and 1992 I worked as a research and advisory entomologist for the UK Forestry Commission based at their Northern Research Station just outside Edinburgh. For the first five years of my time there I worked almost exclusively on the pine beauty moth, Panolis flammea. The pine beauty moth is

snowflea 1

a native insect that became a pest of a non-native tree, Pinus contorta, then a tree that was widely planted over northern Britain. The majority of planting in Scotland was in the north and this meant that my study sites were in Sutherland and Caithness and Aberdeenshire. My main experimental forest was west of Aberdeen in the Spey valley (very handy for the whisky trail) in the Elchies block of Criagellalchie Forest.

snow flea 2

My experimental forest with nearby distillery marked 😉

In Mid-January 1984, I headed north to do some maintenance on my head capsule collecting funnel traps.

snowflea 3

In those days, snow was a perennial hazard, even in the south of Scotland and as I progressed northwards the drifts at the side of the road became increasingly higher. When I reached the forest gates, it was obvious that I was not going to be able to drive to my site. The sun was shining, the sky was blue and the snow glistened. A perfect day for a walk, albeit one of 10 km. Luckily, the weather had been sunny for the last couple of days so the snow was mostly hard enough to walk on. Only in a few places did I break the surface and find that I was standing on about a metre depth of snow. Two hours later as I was approaching my field site, squinting against the sun bouncing off the white untouched snow, I saw black spots moving on the surface. My immediate thought was that I was suffering the first stages of snow-blindness, but as I got nearer I saw that the black dots were actually insects. At first sight I thought I was hallucinating, was this some strange bizarre form of life perhaps an aphid-fly hybridization experiment gone wrong? On closer examination I realised that I was looking at wingless Mecopterans.

snow flea 4

Male snow flea, Boreus hyemalis http://mecoptera.free.fr/Boreus-hyemalis.html


snow flea 5

Female Boreus hyemalis, note the sting-like ovipositor. http://www.wbrc.org.uk/WORCRECD/32/Bingham–John–Snow_Flea_Boreus_hyemalis.html

Although I was familiar with Scorpion-flies, I had never seen these critters before.

snow flea 6

The aptly named Scorpion fly Panorpa communis : https://commons.wikimedia.org/wiki/File:Scorpion_Fly._Panorpa_communis._Mecoptera_(7837166610).jpg

I collected a few to send off for identification and confirmation and carried on into the depths of the forest to check on my funnels. On returning to civilization a day or so later I sent my specimens off to the Natural History Museum and shortly after was informed that I had they were the snow flea, Boreus hyemalis and that I had extended the recorded range of this particular species, albeit only by a few miles.

snow flea 7

My record – it lasted 10 years as the furthest north before M.S.C. Elliott recorded it in February 1994 in Easter Fearn in the north-west Highlands.

Boreus my record

Distribution of Boreus hyemalis in 1994; my record, then the furthest North.


snow flea 9

Current recorded distribution of Boreus hyemalis – obviously widespread – just lacking people willing to go and look for it in the winter 🙂

So what is a snow flea. It is of course, not a flea, being a Mecopteran or Scorpion fly, albeit non-winged.  In Britain there are three species with wings (in the genus Panorpa), the larvae and adults both being predatory on other insects. The adult snow flea is about 5mm long, and lives among moss on which it feeds as both a larva and adult (Withycombe, 1922, 1926). Interestingly, the BugLife site states that they are predatory in both the larval and adult stage. I am not sure where they got this information as they do not cite a reference and all the published literature I have seen indicates that they are moss feeders (Withycombe, 1922, 1926; Fraser, 1943; Hågvar, 2010). Indeed, Wthycombe (1922) conducted a series of experiments on the larvae and conclusively demonstrated that they were unable to complete their development unless fed on moss, although the adults will apparently also feed on dead insects.

These are true winter-active insects, adults emerging in October and November when they mate and lay their eggs the eggs at the base of moss plants), Polytrichium commune being the preferred host (Fraser, 1943). The eggs start to hatch in November and the larvae forage within the moss clumps, pupating towards the end of the summer, emerging as adults after 6-8 weeks.  The adults, which are wingless, thus come out in the coldest months of the year, usually between October and April.  They are most easily seen when walking or jumping on the snow surface. Considering that the adults are winter-active they have a surprisingly high super-cooling point (-6.5oC) (Sömme & Östbye, 1969), especially when compared with the cereal aphid, Sitobion avenae, which has a super-cooling point of -24oC but rarely survives English winters (Knight & Bale, 1986). The BugLife site wonders “how they (snow fleas) manage to jump up to 5 cm without muscular hind legs” but Burrows (2011) found that their jumping prowess is by virtue of large depressor muscles within the thorax which enables them to jump distances of up to 10 cm with a take-off velocity of 1 m s-1, indicating a force of about 16 times their body weight.  So aptly named in this respect too.

The Snow flea is not found (or at least has not been recorded) in the mild south-west of Britain, seeming to prefer areas with a harsher winter. Climate warming may thus pose a threat for this intriguing and little studied insect. Perhaps it is time for us all to venture out in mid-winter and start scanning the surface of snow drifts in heathland areas for these elusive creatures before it is too late.



Burrows, M (2011) Jumping mechanism and performance of snow fleas (Mecoptera, Boreidae). Journal of Experimental Biology, 214, 2362-2374.

Fraser, F.C. (1943) Ecological and biological notes on Boreus hyemalis (L.) (Mecopt., Boreidae). Journal of the Society for British Entomology, 2, 125-129

Knight, J. D. & Bale, J. S. (1986). Cold hardiness and overwintering of the grain aphid Sitobion avenae. Ecological Entomology 11, 189-197.

Sömme, L. & Östbye, O. (1969) Cold-hardiness in some winter active isnects. Norsk Entomologisk Tidsskrift, 16, 45-48

Withycombe, C. L. (1922). On the life history of Boreus hyemalis L. Transactions of the Entomological Society of London, 1921, 312-318.

Withycombe, C. L. (1926). Additional remarks upon Boreus hyemalis L. Entomologist’s Monthly Magazine, 62, 81-83.


Useful link

For more images and observations see http://www.wbrc.org.uk/WORCRECD/32/Bingham–John–Snow_Flea_Boreus_hyemalis.html



Filed under EntoNotes

Insect egg mimics – plant parts that pretend to be insect eggs

Back in the 1980s I was a forest entomologist working for the UK Forestry Commission at their Northern Research Station based just outside Edinburgh.  I was working on two important pests of Lodgepole pine (Pinus contorta), the pine beauty moth, Panolis flammea and the European pine sawfly, Neodiprion sertifer.  The pine beauty moth lays its eggs in short rows on the upper surface of pine needles in late spring/early summer.

Panolis eggs

Eggs of the pine beauty moth, Panolis flammea  (Image courtesy of Stanislaw Kinelski, Bugwood.org http://www.invasive.org/browse/detail.cfm?imgnum=1258002).

They are pale yellow when first laid and gradually darken as they mature becoming a deep violet colour just before they hatch.  The eggs of Neodiprion sertifer are also laid on the upper part of the pine needles, but are ‘injected’ just under the cuticle of the needle.  After a few days a small necrotic patch develops at the oviposition site.

Neodiprion eggs

Eggs of the European pine sawfly, Neodiprion sertifer (image courtesy of A. Steven Munson, USDA Forest Service, Bugwood.org http://www.forestryimages.org/browse/detail.cfm?imgnum=1470178)

Spring field work for me was several days of rather tedious egg counting and as I scrutinised hundreds of pine needles, I noticed that some of the needles had little flecks or balls of resin on them,

Resin flecks

Resin flecks on bristlecone pine, Pinus arsitata – often confused with scale insect infestations (Photo by Hans G. Oberlack via Wikipedia).

which were, especially on gloomy days in the depths of the forest, quite easy to confuse with pine beauty moth eggs.  Other needles had discoloured areas that looked like pine sawfly eggs or also a bit like pine beauty moth eggs, depending on how they were arranged.

Egg mimics

Possible insect egg mimics on pine needles

Long days working alone in a forest allow one the time to think and it occurred to me one day that if I was being fooled by these ‘pseudo eggs’ then perhaps egg-laying pine beauty moths and pine sawflies might also be getting confused and avoiding laying eggs on these apparently already infested needles.   I wondered if there was any evidence to support my far-fetched hypothesis and to my delight found a paper by (Williams & Gilbert, 1981) that demonstrated quite convincingly that passion-fruit vines, produce structures resembling eggs of Heliconius butterflies and that these deter them from laying eggs on them.

Egg mimics 2

Egg mimics on passion flower leaf – Photo by Lawrence Gilbert http://plantmimicrybz2820.blogspot.co.uk/2015/04/the-passiflora-genus.html

I also found papers that showed that other Lepidoptera (Rothschild & Schoonhoven, 1977; Nomakuchi et al., 2001) and beetles (Mappes & Mäkelä, 1993), are able to discriminate between leaves that already have eggs laid on them and avoid laying more eggs on those leaves, thus reducing larval completion.

Although I never formally checked it, I got the impression that needles bearing ‘egg mimics’ had fewer pine beauty moth eggs or pine sawfly eggs laid on them than those without.  Another question that could be easily looked at is whether pine trees in areas that have had outbreaks have more speckled needles than those in non-outbreak areas.  I always meant to do some formal sampling and a proper experiment to back up my feelings, but never found the time to do it.  I am pretty certain that I am unlikely to get round to doing this in the near future (if ever), but I would like to know if this is indeed another example of  a plant mimicking insect eggs.  I would be very happy indeed if any of you feel like testing my hypothesis and look forward to seeing the results in print.



MacDougal, J.M. (2003)  Passiflora boenderi (Passifloraceae): a new egg mimic passionflower from Costa Rica.  Novon, 13, 454-458

Mappes. J. & Mäkelä, I. (1993)  Egg and larval load assessment and its influence on oviposition behaviour of the leaf beetle Galerucella nymphaeae.  Oecologia, 93, 38-41

Nomakuchi, S., Masumoto, T., Sawada, K., Sunahra, T., Itakura, N. & Suzuki, N. (2001) Possible Age-Dependent Variation in Egg-Loaded Host Selectivity of the Pierid Butterfly, Anthocharis scolymus (Lepidoptera: Pieridae): A Field Observation .  Journal of Insect Behavior, 14, 451-458.

Rothschild, M. & Schoonhoven, L.M. (1977) Assessment of egg load by Pieris brassicae (Lepidoptera: Pieridae). Nature, 266, 352-355.

Williams, K.S. & Gilbert, L.E. (1981) Insects as selective agents on plant vegetative morphology: egg mimicry reduces egg laying by butterflies. Science, 212, 467-469.


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