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

The book!

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

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

Reality strikes!

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

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

I should have known better

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

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

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

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

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

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

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

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

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

References

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

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

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

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

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

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

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

Pick & Mix 65 – beetle based drones, biodiversity, climate change, resilient forests, beavers and rewilding, citizen science and a farewell to Dynamic Ecology blog

A drone with wings inspired by beetle elytra

Interesting take – The case against the concept of biodiversity

Why eye-catching graphics are vital for getting to grips with climate change

Graeme Lyons argues that English names of species should always be capitalised – I agree, do you?

Fantastic project – croplands up close – should be very useful for many fields (pun intended)

How ornithologists figured out how to preserve bird specimens

Is your forest fit for the future? Emily Fensom from the UK Forestry Commission introduces the Climate Matching Tool and suggests how this can be used to build resilient forests

Beavers are back: here’s what this might mean for the UK’s wild spaces

What is and what isn’t citizen science?  I don’t fully agree with some of this but it is an interesting read.

In which the team from Dynamic Ecology announce their semi-retirement.  They will still be keeping us inspired, entertained and stimulated but just not as often. I will miss them and judging by the huge number of comments, I am not the only one. Some fantastic and very well-deserved tributes.

Assignments with benefits – materially rewarding student work

One of the banes (and there have been a few) of my life as an academic, has been the necessity (or is it?) of having to assess student achievement and attainment using exams, tests, essays, quizzes, talks, posters, whatever. I very early on in my academic career decided that students on my undergraduate and postgraduate modules, should not just be tested for memory, knowledge, analysis, evaluation and synthesis but in skills that at Imperial College, London, in the early 1990s were hardly, if ever, mentioned. Yes, you guessed it, posters, talks, debates and group working. These all being things that when I was an undergraduate we received absolutely no training in at all. My final year module in Applied Ecology ran for a solid five weeks in the run-up to the end of the Christmas term. I had a dedicated base room where we could store materials and was not used by anyone else during the module.  In other words, I could timetable independently of the central system – a great boon. As members of the British Ecological Society will know, the week before Christmas heralds the fantastic BES Annual General Meeting, so it was a no-brainer, I would build the coursework part of the assessment for my Applied Ecology module around a mini ‘BES Christmas Meeting’. Why not? It is all part of the scientific process anyway😊

The Mini Conference

I decided that for the actual conference, the students would produce a poster, based on a recently published real paper of their choice, and a talk, which could also be based on the same paper.  Their brief was to adopt the identity of the author and speak and present as that individual.  I also ran a poster making and how to give a talk workshop so that the students knew what was expected from them.  I was, as it turns out, the only module leader in the Department doing this!  I set aside specific slots in the timetable when a dedicated space was available for me and the students to be together for three hours a week where they could work on these and consult me if they came across any problems.

A couple of excerpts from the 1997 Applied Ecology mini-conference – some of you may recognise at least three names that aren’t mine 😊 You may also notice that their interests have changed since then.

The Research Syndicate

The other skill that final year students need to acquire, is how to write scientifically, both in terms of publishing and for their final year projects.

As I have mentioned before, I have huge amounts of unanalysed data which I am always trying to donate to people with more time and modern statistical expertise than me to knock into publication shape, e.g. aphid overwintering, effects of long term herbivory on bird cherry trees, and flowering patterns in sycamore.   For this I provided all the raw data files, the methods and materials so that the students could get a real feel for what it was all about and which data set they would like to analyse.  They had free choice of the data sets and could also choose how much of it to analyse to tell their stories. I then provided five training sessions about scientific paper writing; why we do it, what it is meant to achieve and how papers are structured and read, plus a little rant about the tyranny of impact factors! They were also given five afternoons of access to R training via my PhD students. These were group sessions and I encouraged the students to discuss their approaches between them – after all, as scientists we work in groups.  The output, which was independently done, was a ‘scientific paper’.  The best of which I would help knock into shape for publication to a real journal (see reference list for a few examples).

The Class Debate

Running alongside everything else was the class debate.  You might think that this was a busy module, but the whole point of this module was to get the students to interact – I was, and am not a great believer in the ‘jaw, jaw, jaw’ approach to teaching, especially to final year students.  This exercise involved me stetting a scenario, (my favourite is the one illustrated), and allocating named characters to each student. These ranged from local shop owners, RSPB Wardens, Wildlife Trust employees, Forestry Commission employees, Commercial foresters, Whisky distillery owners, Garden Centre Owners, university academics, ecologists, fishers, local residents (occupations chosen by the student) and, of course, the owner of the peat extraction company and their PR Officer.  I then let the students form alliances and come up with their strategies for the Public Inquiry, for which I allowed a whole day.  Each student produced a short essay as part of their coursework and I allocated extra marks for each contribution made during the debate which I added to their essay marks.

My last ever class debate at Imperial – a sad day, but all good things come to an end.

A few of the characters – anyone recognise the suave city gent in the centre?

The formal part of the course ended with the mini-conference where all the students spoke and some were involved as session chairs. Posters were judged by students and staff as were the talks.  At the end of the two-day conference it was prize giving time.  Top papers, talks, posters and essays were all given copies of books that I thought would be of interest, (Stephen Jay Gould featured a lot), runners up were given book tokens and third placers a bag of chocolate coins (Christmas you know).  Incidentally all winners were given a huge chocolate medal 😊 Plus the students had all their course work marks to take away with them for Christmas. Win-win all round.

Something a little bit different – Cash for assignments – The Antenna article

For our MSc students I thought that it would be nice, that as well as learning a new skill, writing for a general audience, they had the opportunity to be both published and earn a little bit of cash to help pay their way though the course.  Luckily, a solution was close to hand, the Royal Entomological Society, Student Essay Awards.  Even more fortuitously, the submission date for both the coursework and the essay award were very close together.  I provided students with past essay winners and a brief session on the different approaches one could take, from short pieces of fiction, to a ‘news item’, to a factual ‘did you know piece’.  The world was pretty much their oyster. So again, a nice way to make coursework fun.  You will, if you have clicked on the link, seen that we here at Harper Adams University have done quite well over the years.  I also recommend that you spend a few minutes reading some of the winning essays – they are truly wonderful.

Post script

The whole exercise was also very rewarding for me. I got to really know the students and felt that I had done some very useful mentoring. Decades later, I am still in touch with many of the students, including many who are no longer in academia. I was also privileged to win a couple of teaching awards (just certificates, no cash), on the back of this, so definitely worth the effort.

References

Butler, J., Garratt, M.P.D. & Leather, S.R. (2012) Fertilisers and insect herbivores: a meta-analysis. Annals of Applied Biology, 161, 223-233.

Goodwin, C., Keep, B. & Leather, S.R. (2017) Habitat selection and tree species richness of roundabouts: effects on site selection and the prevalence of arboreal caterpillars. Urban Ecosystems, 19, 889-895.

Leahy, M.J.A., Oliver, T.H. & Leather, S.R. (2007) Feeding behaviour of the black pine beetle, Hylastes ater (Coleoptera: Scolytidae). Agricultural and Forest Entomology, 9, 115-124.

Leather, S.R., Ahmed, S.I. & Hogan, L. (1994) Adult feeding preferences of the large pine weevil, Hylobius abietis (Coleoptera: Curculionidae). European Journal of Entomology, 91, 385-389.

Leather, S.R., Small, A.A. & Bogh, S. (1995) Seasonal variation in local abundance of adults of the large pine weevil, Hylobius abietis L. Journal of Applied Entomology, 119, 511-513.

Leather, S.R., Fellowes, M.D.E., Hayman, G.R.F. & Maxen, J.S. (1997) The influence of lodgepole pine (Pinus contorta) provenance on the development and survival of larvae of the pine beauty moth Panolis flammea (Lepidoptera: Noctuidae). Bulletin of Entomological Research, 87, 75-80.

Leather, S.R., Beare, J.A., Cooke, R.C.A. & Fellowes, M.D.E. (1998) Are differences in life history parameters of the pine beauty moth Panolis flammea modified by host plant quality or gender? Entomologia experimentalis et applicata, 87, 237-243.

Scorpion flies – not as scary as they sound or look

I’m very fond of Scorpionflies, in fact, they are almost up there with aphids on my all-time favourite insects list. They, at least to me, are reminiscent of something that one would expect to find in Lewis Carrol’s Alice in Wonderland. They belong to the Order Mecoptera, which until some pesky taxonomists decided that fleas are Mecoptera, was one of the smaller insect Orders with just 600 species, if fleas are indeed Mecoptera then we now have 2600!

Male (left) and female (right) Scorpion flies. Despite the resemblance to the back end of a scorpion it is not a sting, but part of the male genitalia.

Ignoring fleas for the moment, there are nine families of Mecoptera, but only three are common; Scorpion flies (Panorpidae), the Hanging flies (Bittacidae) and Snow fleas (Boreidae) (Byers & Thornhill, 1983). Of these only two occur in the UK, the Scorpion flies and the Snow fleas. Adult Scorpion flies are mostly scavengers, mainly eating dead insects, topping this up with a bit of pollen, nectar and fruit juice and for a special treat, bird faeces. Their larvae live in the soil and mop up whatever dead things they come across.

The Snow fleas feed on moss and are only getting a mention here because they are quite cute and for a few years I held the record for the furthest north record 😊

Cute Snow fleas 😊

The Hanging flies are carnivores capturing live prey as adults and larvae and deserve a special mention as they (and although it shouldn’t, but it appeals to the ten-year old in me, makes me giggle) have a penisfilum.

Male (left) and female of Bittacus planus. Photo provided by Dr. Baozhen Hua. Note the knob in the male!

Anyway, back to the scorpion flies. They are found in temperate regions, worldwide and as of 2018 there were 280 species. The males are highly competitive, as are many of the Mecoptera. Males will fight over their food, which as I mentioned earlier is quite high in dead flies, which they often steal from spider webs.  They have no fear of spiders as they can dissolve the web if they do get caught.

Another cool thing about scorpion flies is that they, like some spiders, use nuptial gifts to increase their chance of mating. They first use a pheromone which as pheromones go, is pretty short range, 10 -15 metres. Once a female has been enticed by the pheromone, the males than flash their wings, which are striped and do a bit of a dance. Depending on species, what happens next could be one of three tactics.

Nuptial gifts and mating of Dicerapanorpa magna Photo provided by Dr. Baozhen Hua.

male gives female food which she eats during copulation either a salivary deposit from enlarged salivary glands or a dead insect, and waits female arrival.  Another tactic is to find a suitable dead insect which he then stands by, waits for a female to arrive, and then copulates with her while she eats it. Some males are less generous and will force themselves on a female without any presents or even pheromones, holding their chosen mate in place with his abdominal clamp (Tong et al., 2018). The size of the gift is related to the duration of copulation and to how long it will be before the female mates with a different male (Byers & Thornhill, 1983); females that were subjected to forced copulation have a very short inhibition time – the more the males invest in their nuptial gifts, the more offspring they sire. Basically, they get what they pay for! The eggs, usually no more than ten per clutch, are laid into damp soil.

When I introduce Scorpionflies to a new audience, I am, as I find frequently with other insects, faced with the usual human exceptionalism question “

“Mecoptera are most often defined by the characters they do not possess” Penny (2016)

They are not pollinators generally regarded as pollinators (thanks Jeff Ollerton for reminding me that some do visit flowers for nectar), but they are not crop pests and nor are they vectors. We don’t eat them and most of them most are not biological control agents. Bittacids are, however, predators. Panorpids are recyclers, they feed on carrion. The Nannochoristids could be seen as s bio-indicators; their larvae need clean water and Boreids could act as climate change ‘canaries’ because of their limited dispersal ability and their need for cold.

Scorpionflies have appeared in video games (Shelomi, 2019) so I guess are helping the economy and keeping people entertained.

In the long distance past (170 MYA), before angiosperms made their appearance and allowed the explosion in insect diversity possible, three groups of scorpionfly, now extinct, fed on the nectar of gymnosperms and in return pollinated them (Ren et al., 2009).

References

Byers, G.W. & Thornhill, R. (1983) Biology of the Mecoptera. Annual Review of Entomology, 28, 203-228.

Palmer, C. (2010) Diversity of feeding strategies in adult Mecoptera. Terrestrial Arthropod Reviews, 3, 111-128.

Ren, D., Labandeira, C.C., et al., (2009) A probable pollination mode before angiosperms: Eurasian long-proboscid scorpionflies. Science, 326, 840-847.

Shelomi, M. (2019). Entomoludology: Arthropods in Video Games. American Entomologist, 65, 97–106

Tong, X., Zhong, W. & Hua, B.Z. (2018) Copulatory mechanism and functional morphology of genitalia and anal horn of the scorpionfly Cerapanorpa dubia (Mecoptera: Panorpidae). Journal of Morphology, 279, 1532-1539.

Pick & Mix 64 – big questions in ecology, entomological colonialism, the return of the Nature Table, ID apps, saving insects, rewilding, making your garden insect friendly and how big?!!

Charley Krebs on what are the big questions in ecology and how we might go about addressing them –  Whither the Big Questions in Ecology?

Colonialism in entomology – a historical problem that still persists today

Totally agree – Nature Tables should make a return to the classroom as in my opinion, should the weekly Nature walk. Even in an urban environment there are plenty of opportunities to introduce children to the natural world.

Why planting tons of trees isn’t enough to solve climate change – Massive projects need much more planning and follow-through to succeed – and other tree protections need to happen too

A warning note that iNaturalist and other ID apps are not perfect – sometimes they can be dangerous, but thankfully, not often

Scary stuff, a long read, but worth it “The older the article, the less likely it is that the links work. If you go back to 1998, 72 percent of the links are dead. Overall, more than half of all articles in The New York Times that contain deep links have at least one rotted link.”

Why I hate pandas and other entomologists hate koalas! Forget charisma, save our insects! Never underestimate the politics swirling around charismatic megafauna

A rewilding experiment set up before the term existed – Monks Wood Wilderness

Life lessons from beekeepers – stop mowing the lawn, don’t pave the driveway and get used to bugs in your salad

A bit of fun – visual comparisons of extinct megafauna and their living relatives – some really startling size differences

Foreign holiday 2021 – over the border to Scotland 😊

Normally at this time of year we would be ending the third week of our summer holiday at our French house in the Languedoc.  Sadly, because of the pandemic and a reluctance on our part to put money into the hands of the PCR testing companies owned by Tory cronies, we are still in the UK.  Fortuitously, as a consequence of the easing of covid restrictions it was decided that the Scottish Forestry Trust August Trustees meeting would be held IRL (which I only discovered recently means In Real Life) and not via Zoom.  My wife, who had not been back to Scotland since we moved to Bracknell 29 years ago, decided that this would be a great opportunity to revisit Edinburgh and our old stamping ground, Peebles and booked us a hotel in Leith (she wanted to see how it had changed).

To give us more flexibility we decided that we would drive and not take the train (next time we will definitely let the train take the strain) and on Monday morning set off in high spirits (and in some pain on my part as I managed to trap a nerve at the weekend) from Shropshire with Mrs Garmin predicting, after insisting that we change country to Scotland, an estimated time of arrival of early afternoon.  Sadly, traffic and weather conspired to defeat her and we were somewhat later than predicted and our plan to spend a few hours in Peebles (where we lived for ten years) was thwarted, although we did manage to get there in time for a pleasant evening with my old colleague Allan Watt and his wife Katy.

Dockside views from Ocean Terminal, including HMS Britannia

View from hotel, en route to Princes Street with bonus butterflies

Iconic Edinburgh views including the reintroduced trams

More iconic Edinburgh including the spectacularly hideous Scott Monument

The Monarch of the Glen (National Gallery of Scotland), The Royal Botanic Gardens taking plant health seriously and the afterlife of a sweet chestnut tree.

National Museum of Scotland, Chambers Street

Scottish Parliament and a stunning flower (artificial display) at the Cold Town House restaurant in the Grassmarket

Peebles, our old house on the High Street, and Tweed Green – it was a very grey day. Little known fact, our house (80 High Street) was previously The Smallest Little Restaurant in the World, (it sat 6) owned by one of the Maxtone-Graham family of Mrs Miniver fame. We got the occasional phone call asking to book a table.

Peebles, Cuddyside walk, the couchée righ (totally unchanged outside and inside despite almost three decades) and Neidpath Castle (yet another place that Mary Queen of Scots visited).

We only had four days in the Borders, heading back to Shropshire on the Friday, via Moffatt (to visit the famous toffee shop) and thanks to an accident on the M6 near Manchester, we had a very convoluted and tedious trip back, via the Mersey Crossing (£2), arriving after a nine-hour journey, by which time I was in absolute agony from my trapped nerve ☹ Definitely the train for us next time!

Reflection – a much needed (and missed) aspect of academic life

I’ve been in Academia a long time, I started my PhD in 1977, and things have changed quite a lot over those forty odd years.  In those heady days of the 1970s and 1980s, an academic taught, did research, wrote papers, reviewed papers wrote grant proposals and even found time to write books. There was also a valuable commodity, time; time to sit back and reflect during work hours. This could involve sitting at your desk with your feet up and your eyes closed or like

Charles Darwin and his ‘thinking path’, go for a walk or a run in the fresh air or just sit under a tree or lie back in the grass and watch the clouds go by as new ideas bubbled up in your mind.

Reflecting on life

When I was in my last year as an undergraduate desperately preparing for my final exams, I would, in between revision bouts, go and sit under a cherry tree outside the Agriculture Building and just let my brain rearrange all the facts that I had accumulated over the past four years of study into some sort of coherent order. It worked and much to my surprise I got a First Class degree. I’m a wee bit older now but it still works.

The important thing when I began my academic career was that there was an opportunity within the working day to gather one’s thoughts and let connections form.  It didn’t necessarily have to be blue sky thinking, just a chance

Blue sky thinking or daydreaming?

to clear the turbid and muddied thoughts and get them into some form of order and allow you to process them into something worthwhile and hopefully clear your mind so that new exciting ideas break through and bubble out into the light.

Struggling to clear those turbid thoughts.

It’s all becoming clearer

In those halcyon days there were, in all the places I worked, well established and popular morning and afternoon coffee/tea breaks.  In fact in Finland where I worked at the Agricultural Research Station just outside Helsinki, we even had breakfast together (early starters those Finns), and in my early days at Silwood Park not only was there morning coffee in the Refectory (Paddy’s to some of us), but in the afternoon we repaired to the Conservatory and Orangery where the redoubtable and doughty Pearl wheeled in her tea trolley and we, depending on the season and weather, either sat inside or reclined outside on the grass chatting and imbibing our drinks of choice. 😊

Back in the 1980s, and this may come as a surprise to the modern academic/researcher, we had typists (I met my wife in the typing pool) to wade through our hand-written drafts and type our papers for us. The along came technology and things began to change and not for the better.  Personal computers started to appear on everyone’s desk, not just in the computer rooms, the tyranny of email replaced the paper mail (finding your post tray full of envelopes was much more satisfying than logging on and finding your email folder telling that you have 120 unread messages) and worst of all, along came electronic ordering and costing. In the old days, if you wanted consumable you asked the Departmental Technician for them, or if not in stock they would order them for you. Similarly, for quotes for equipment for grants etc.  It makes no sense to me that academics should be responsible for ordering stuff themselves (Dreamweb?, Nightmareweb more like). If you don’t use a system daily then every time you do use it, it is a whole new time-consuming learning experience. Likewise, health and safety issues, surely much more efficient and cost-effective use of time to let the H&S Officers access the forms and do the assessments rather than the academic?  I could go on, but I think you know where I am coming from.

All of the above and the huge increase in student numbers (in the UK at any rate) with the concomitant increase in marking and teaching related administration meant yet another erosion of reflective time and as the years progressed a noticeable decline in the number of people finding or making time to venture out of their research silos to have coffee breaks away from their desks and labs.  Many academics now lunch in their offices.  This has meant a reduction in collegiality and the very valuable opportunity to talk and listen to colleagues from other fields.  When, after twenty years, I left Silwood Park, morning coffee in the Refectory had dwindled to just a handful of us entomologists☹ One of the many positive things of moving to Harper Adams was that there was (hopefully when Covid is controlled we can all get back there) a vibrant and very buzzy Staff Common Room which reminded me very much of my early days in academia. Until you experience it at first hand, you don’t realise how important a central, informal gathering place is to working life. A vibrant common meeting place has huge benefits for creative thinking, after all what is the most important part of a conference? Very rarely the talks; the bar, coffee and meal breaks are where it is at and the new ideas and partnerships are forged and the pot of gold at the end of the rainbow found. The fewer the opportunities for relaxed social interactions, the fewer the good ideas we generate – coffee breaks, no matter how chatty, do, contrary to what some senior managers might think, improve productivity.

Without time to reflect one’s productivity goes down, thoughts are mired down in the turbid waters of toil and home and work life become entangled to a greater degree – I think the majority of us do our marking, paper reviewing, thesis reading, paper writing and in extremis, when working to a submission dead-line, our grant writing.  This is not sustainable and certainly not good for our well-being.  Do people still get proper sabbaticals, i.e. ones that the Department funds rather than having to apply to a grant body for one?  In my thirty odd years of university life I never had a sabbatical.

Now that I am Emeritus I am discovering a whole new world of time; time to walk and think, time to sit and think and time to read and write. If it were not for the fact that I am not in my office every day and thus missing out on the coffee culture I could imagine myself on sabbatical. Having the space and the ambiance to think and interact is hugely important. As a concrete example, a couple of years ago I invited two colleagues of mine with whom I am writing a book down to our French house in the Languedoc.  We had a very productive week, every morning working in separate rooms on the book, meeting up for lunch and then spending the afternoon relaxing and thinking.

Being productive away from the office

Since then, back in our respective work worlds, progress on the book has been glacial. Once international travel is back on the cards we plan to repeat the process and hopefully get the book back on course for publication next year.   I have, not quite tongue in cheek, suggested to my Head of Department that he might like to provide the funds to send members of staff down to our French house where I will provide paper and grant writing workshops 😊

Time spent in reflection is not time wasted, like plants, when given space and the right conditions, ideas flourish and bloom.

Pick & Mix 63 – haikus, red leaves, insect opera, insect declines (again), nature friendly cities, dung beetles, bird names and pollinators

Ever wondered why some young leaves are red?  Ray Cannon explains

Winners of the 2021 Hexapod haiku contest – I did enter but……..  ☹

Really interesting article (lots of graphics) about how eating habits have changed in the USA since 1970

Like Opera, interested in nature then you might want to catch this – Locust: The Opera

Providing regular water supplies for humans may be causing insect declines in the tropics

There are over 7,000 English names for birds – here’s what they teach us about our changing relationship with nature

Introducing new dung beetles to Australia: battling the cane toad’s legacy

What would a truly Nature-friendly city look like?

Ecomimicry: the nature-inspired approach to design that could be the antidote to urban ‘blandscapes’

Disentangling the facts from the myths about pollinators – Einstein’s bees, sound bites and vitamins

Citizen Science Rules – The ‘humble’ amateur is the backbone of entomology – laud them, don’t scorn them

I know we deal with invertebrates but the exoskeleton of entomology doesn’t quite hack it 😊 There is a tendency within academia, perhaps not as marked as it was when I entered it, to be somewhat dismissive, even scornful, when it comes to natural historians and amateurs. I once had a bit of an in-print argument with the late Denis Owen about the validity of data collected by ‘amateurs’ (Leather, 1990), which I found a bit surprising considering his wife Jennifer’s deep and life-long involvement with that type of data (Owen, 2010).

I have been a ‘professional’ entomologist for more than forty years and although I may, in the past, when I was imbued with the arrogance of youth, have made remarks about stamp collecting and lack of scientific method, I have always been in awe of the taxonomic expert, whom at a glance can accurately (most of the time) identify an insect to species. Me, I’m pleased when I get the family right.  I remember when I was an undergraduate sitting round a light trap on our field course, being stunned by Judy Honecker (where are you now?) shouting out the names of the moths as they flew into it – all done by the mysterious jizz. She may of course, have been taking advantage of our ignorance, but I don’t think so.

I don’t think, even now, that many academic, research-active, professional entomologists, or ecologists really appreciate the service that the amateur entomologist or in a broader sense, natural historian, provides to the professional community.  I am a long-time member of the British Entomological and Natural History Society (BENHS) which means that a copy of the British Journal of Entomology and Natural History pops through my letter box every three months. Every time I open a copy of this little journal I am humbled by

Latest issue of the British Journal of Entomology and Natural History

the erudition displayed by the contents.  You won’t find professionally crafted accounts of large-scale field studies/experiment or complex laboratory experiments, analysed using the most complicated analysis that packages such as R or SPSS can spit out by any means.  What you will find, and this is perhaps where the slightly scornful attitude to the ‘amateur’ has its roots, are reports of insects found while on holiday in Spain or closer to home* and accounts of the annual exhibition. You will also find note of the first records of species in the UK, new host records, be they plants or other animals, and yes, also reports of experimental work.  The style may not be as ‘scientific’ as that found in mainstream scientific journals, but that does not detract from their value. The thing that really blows my mind about the BJENHS and others of its ilk such as the Entomologist’s Monthly Magazine, (in both of which I have published), are the numbers of new records reported and the identification skills that these demonstrate of the authors. It struck me that whereas those entomologists with similar skills that work within academia, either in museums or universities, are described as systematists (naturalists engaged in classification) or taxonomists (biologists that groups organisms into categories) these expert amateurs didn’t seem to have a single word to describe them. I of course resorted to Twitter to see if anyone out there knew better.

I received a number of responses, the one below summing up the most common answers.

Pretty much the status quo.

One person suggested parataxonomist, which I feel means an entirely different thing, being someone who has been trained to have “expertise is in collecting specimens, mounting them, and performing preliminary sorting of the specimens to morphospecies” (Basset et al., 2000).

What I do think we can all agree on, is that the county recorders, the various group specialists and more general natural historians, are what I would term professional amateurs. They are not paid for what they do, most have jobs outside entomology, or are, in many cases, retired, but they approach their subject in a thoroughly professional way, keeping impeccable records and disseminating their findings through publications, talks at meetings and running (or helping run) training courses.  They do a huge service for entomology, not just by providing data for the ‘professionals’ to mine and analyse, but also by encouraging others to enter the field. The professional and the professional amateur can, sometimes exist in the same person, I was for example, once President of the Amateur Entomologist’s Society and the current President, Erica McAlister (@flygirlNHM, for those of you on Twitter), is also a professional entomologist.

Richard Jones rather neatly summing it all up.

There is now, however, a whole new category of amateur. Citizen science has unleashed the ‘amateur amateur’ into the wild, albeit many stray no further than their gardens. Citizen science, once a bit marginalised, is now pretty much mainstream is fully recognised by the professional ecologists and entomologists as a hugely important contribution to their disciplines Bates et al., 2013; Pernat et al., 2021). The two most publicised UK examples are the Big Butterfly Count (incidentally this year’s launch coinciding with the publication of this post) and the Big Garden Birdwatch, both of which my wife and I do. Despite having my name on ten papers dealing with birds, I count myself when it comes to the latter, as being a true ‘amateur amateur’ 😊

Citizen science projects span disciplines and the globe.  You can take part in a ‘bioblitz’, record the timing of budburst, the number of plants in flower on a particular date, join in the UK ladybird survey, see how many birds you can count over the Christmas period, and many, many others.

I think it is extremely important to take note of the quote below. It explains to some extent, why in the past, and sadly, to a certain extent, why some (not many thankfully) professional scientists still tend to treat citizen science with less respect than it deserves.

 “Traditionally, we think in terms of a data-gathering component of science, a data analysis component of science, and an interpretive “discussion” component of science. As well, the scientific method is typically thought of as being hypothesis driven (with the hypothesis or “question”preceding data collection), and in mainstream science the process of generating and testing hypotheses is the domain of the researchers – those people who are unequivocally “scientists”. In citizen science, the participants are almost exclusively involved in data gathering alone, but most projects include the promise that anyone is welcome to follow through with their own analyses and interpretations. This is the key element that makes citizen science “democratic”, even if a few participants follow up with analyses of their own. “Citizen Scientist” is, therefore, best understood as an honorary title, given to anyone who participates in any level of the scientific enterprise, on a voluntary basis, with the proviso that that most participants are involved only in data collection.” Acorn (2017)

The data that we collect as citizen scientists is not wasted, and compares well with the data collected by the professionals (Pocock et al., 2015; Pernat et al., 2021) and as the picture below illustrates, is of immense value to entomologists and ecologists.

Roger Morris at Dipterist’s Forum June 27^th 2021 ‘When I started it was as rare as rocking horse faeces’ – Roger Morris talking about recording Rhingia rostrata. Roger is highlighting why recording schemes are so useful. Picture from Twitter via @FlygirlNHM)

Just to reiterate the importance of these glorious amateurs.  Just as those insect host records collected and published by the professional amateurs enabled the late Sir Richard Southwood to restart the species-area concept (Southwood, 1961) and give many of us an opportunity to lengthen our publication lists, so today’s army of professional and amateur amateurs are providing data for a new generation of entomologists and ecologists to help understand and explain the changes we are seeing in insect abundance and distribution (e.g. Werenkraut et al., 2020).  It is very important that their contribution is neither overlooked nor unrecognised.  Without them we would be lost.

Did you know that the oldest (unless you know of an older one?) Citizen Science project in the world is the Christmas Bird Count in the USA, which was started in 1900?

References

Acorn, J. (2017). Entomological citizen science in Canada. The Canadian Entomologist, 149, 774-785.

Basset, Y., Novotny, V., Miller, S.E. & Pyle, R. (2000) Quantifying biodiversity: experience with parataxonomists and digital photography in Papua New Guinea and Guyana. BioScience, 50, 899-908.

Bates, A.J., Sadler, J.P., Everett, G., Grundy, D., Lowe, N., Davis, G., Baker, D., Bridge, M., Clifton, J., Freestone, R., Gardner, D., Gibson, C.W.D., Hemming, R., Howarth, S., Orridge, S., Shaw, M., Tams, T., & Young, H. (2013) Assessing the value of the Garden Moth Scheme citizen science dataset: how does light trap type affect catch? Entomologia experimentalis et applicata, 146, 386-397.

Leather, S.R. (1990) The analysis of species-area relationships, with particular reference to macrolepidoptera on Rosaceae: how important is insect data-set quality? The Entomologist, 109, 8-16.

Owen, J. (2010) Wildlife of a Garden; A Thirty-year Study, Royal Horticultural Society, London.

Pernat, N., Kampen, H., Jeschke, J.M. & Werner, D. (2021) Citizen science versus professional data collection: Comparison of approaches to mosquito monitoring in Germany. Journal of Applied Ecology, 58, 214-223.

Pocock, M.J.O., Roy, H.E., Preston, C.D., & Roy, D.B. (2015) The Biological Records Centre: a pioneer of citizen science. Biological Journal of the Linnaean Society, 115, 475-493.

Southwood, T.R.E. (1961) The number of species of insect associated with various trees. Journal of Animal Ecology, 30, 1-8.

Werenkraut, V., Baudino, F., & Roy, H.E. (2020) Citizen science reveals the distribution of the invasive harlequin ladybird (Harmonia axyridis Pallas) in Argentina. Biological Invasions, 22, 2915-2921.

*I remember reading with great amusement a short article in the Entomologist’s Monthly Magazine from the early part of the 20^th Century describing the ‘discovery’ of a species new to Britain that had flown into the bathroom of the author (a retired Lt. Colonel if I recall correctly) while he was having a bath.

Planes, trains and automobiles – insect killers?

I couldn’t not use this – it is (sadly) one of my favourite films 😊

Anyone who has driven (or walked) along a road will have come across roadkill, be it squirrels, pheasants, badgers, deer or even something more exotic, perhaps it us only us entomologists who notice the squashed invertebrates ☹

Dead carabids and mayflies Shay Lane, Staffordshire, 8^th June 2021

But, lets leave the roadkill for a moment, and in the spirit of the title of the film, start in the air. The first thing I discovered when I started to search for the effects of aircraft on insects is the paucity of literature on the subject – it turns out that people are much more interested in stopping disease carrying insects being transported by air or, and coming as a bit of a surprise to me, stopping insects causing plane crashes (House et al., 2020; Grout & Russell, 2021). The aircraft industry is so concerned about the physical dangers posed to ‘planes by insects that NASA actually have a Bug Team dedicated to developing insect proof aircraft.

I am, however, more concerned about how dangerous aircraft are to insects. First, we need to know how many insects are up there and what the probability of them being struck and killed by aircraft is. I’m guessing that bug strike is pretty common, otherwise NASA wouldn’t have a Bug Team. The majority of insects in the air are found at 300-600 m, although this does vary in relation to time of day (Reynolds et al., 2005). Getting a figure for the actual number of insects in the air is as you might expect, actually quite difficult.  The first attempt to trap and collect insects using an aircraft was in 1926 in Louisiana (USA) using a specially designed trap (Glick, 1939).  These do not seem to have been particularly effective as 5 years of trapping, involving 1528 hours of flying, caught just under 30 000 insects (Glick, 1939).  Those of us who have operated pitfall traps for any length of time would consider this a very modest haul 😊

Glick (1939) The aircraft insect trap

That said, the exercise was obviously more hazardous than even collecting insects from roundabouts as this very laconic extract highlights:

 “The skill of the pilots who flew the collecting airplanes is evidenced by the fact that no fatalities occurred.  Only one major accident occurred, when a forced landing resulted in the destruction of the craft and injury to both the pilot (McGinley) and the writer. Such mishaps must be expected in a more or less hazardous undertaking.”

The distribution of catch number was very similar to that reported from the more recent UK study using radar (Reynolds et al., 2005) and is reinforced by this statement from the NASA Bug Team; “The reason we do these tests at low altitudes or do a lot of takeoffs and landings is because bug accumulation occurs at anywhere from the ground to less than 1,000 feet,” said Mia Siochi, a materials researcher at NASA Langley”.

Given the number of flights made globally and the investment being made into protecting aircraft from bug strike, I would assume that the number of insects being killed by aircraft worldwide is probably very high. I am sure that someone with the skill, time and inclination, can probably come up with a fairly realistic figure.  Over to you Dear Readers.

Next up, if we keep to the film title, are trains.  There has been a bit more work looking at the damage that trains do to insects, not a lot, but something is better than nothing.  Work collecting train kill from railway lines showed that snails were particularly vulnerable to being run over, similar to the effects on trail-following ermine moth caterpillars that I observed in Finland in 1981, with Ephemeroptera (Mayflies) in second place (Pop et al., 2020). This, as the authors suggest, was almost certainly due to the time of year and the presence of a lake nearby. Unfortunately no one has done the equivalent of a train splatometer which might be rewarding as these observations from correspondence in British Birds magazine suggest that locomotive engines are causing some mortality to flying insects.  Over to you Bug Life. How about getting the train companies to fit splatometers?

Finally, cars and their effect on insect life. There is anecdotal evidence out there, after all as drivers we have all seen moths in our headlights at night and used our windscreen washers and wipers to try and remove dried on insect corpses and their haemolymph from our front windscreens.

An observation by Ian Bedford

My front bumper – sadly (or perhaps not) much less insect spattered than in the past

Yes, anecdotally we know that insects are being hit by cars (see above) and on my front number plate, a couple of weeks ago (beginning of June) I counted 73 insects, mainly aphids after a 245 km trip. The problem as I see it, is quantifying the numbers killed and calculating the effect that this has on insect abundance. I have mentioned the splatometer in an earlier post which attempts to standardise the number plate counts and I am pleased to see that this has now been revived by Bug Life, and will hopefully carry on for many years. The idea behind this is that over the years we will be able to see if insect numbers as reflected by the change in numbers of splats are increasing, decreasing of remaining the same.  This will not, certainly as described, tell us how many insects are being killed by road using vehicles, although it would be possible if the data were collected over delineated stretches of road (Baxter-Gilbert et al., 2015).  It is not just flying insects that are killed by cars; not all flying insects fly across roads, many seem happy to walk to the other side, reckless as that may seem.

A brave, or possibly fool-hardy carabid beetle crossing the road – Guild Lane, Sutton, Staffordshire, 9^th June 2021.

There have been enough studies done looking at the interactions between roads and insects for a review article to have been published fairly recently, although not all the papers deal directly with mortality effects (Munõz et al., 2015). Many studies have recorded the species affected and the number of dead individuals found but few have attempted to calculate what this means in total. Most studies, as we might expect, have been on large, easily identifiable charismatic species (Munõz et al., 2015) and it from these that we do have some idea of the magnitude of the mayhem caused by road traffic. Some of the figures are incredibly high. A survey of Odonata road kill, albeit near a wetland, of two 500 m stretches of dual carriageway in the Great Lakes region of the USA revealed that at least 88/km/day were being hit and killed by vehicles (Riffell, 1969).  Another study in the USA, this time on Lepidoptera, calculated that about 20 000 000 butterflies (mainly Pieridae) were killed in one week in September (McKenna et al., 2001). The most dramatic figures however, are those from a study in Canada which estimated that 187 billion pollinators (mainly Hymenoptera) are killed over the summer in North America (Baxter-Gilbert et al., 2015).  An unpublished study by Roger Morris (thank you Richard Wilson @ecology_digest for bringing this to my attention) also highlights the dangerous effects of cars on Hymenoptera). Despite the mounting evidence of the harm that road traffic does to insects there is remarkably little information about how this can be reduced, although I did find a paper that noted that if insects are struck by cars driving at speeds of 30-40 km/h they survive the crash whereas speeds greater than this prove fatal (Rao & Girish, 2007).  It might be possible to impose insect safe speed limits along stretches of road that go through sites of special insect interest (perhaps I should try and coin that acronym, SSII, as an additional/alternative term to SSSI (Sites of Special Scientific Interest), but I am not sure how amenable drivers would be to signs telling them to slow down because of insects😊, considering how few drivers slow down in response to the signs warning them about deer and other vertebrate hazards. Another option would be to design road vehicles so that the air flow across them pushes insects away rather than into them; this may already be fortuitously happening as Manu Saunders points in her interesting post about the ‘windscreen anecdote’.  That said, even if cars are more aerodynamic and less likely to splatter insects, the levels of road kill reported in the papers I have cited earlier, still imply that insects are being killed by traffic in huge numbers.

This one didn’t get stuck on a car, but died just the same – A519 outside Forton, Staffordshire, 15^th June 2021

Even if we do accept that deaths down to direct impact with vehicles is lower than in the past, the roads on which we drive our cars are also having a negative effect on insect numbers. Roads, particularly those surfaced with tarmacadam, present an inhospitable surface to some insects which may make them reluctant to fly or walk across. It has been shown that bee and was communities can be different on different sides of a road (Andersson et al., 2017) as the road act as barriers, particularly for smaller species of bees (Fitch & Vaidya, 2021).

Despite the mortality that vehicles impose on insects, roads are not necessarily a totally bad thing for invertebrates; road verges, when sympathetically managed, can provide overwintering sites for a range of arthropod species (Saarinen et al., 2005; Schaffers et al., 2012) and some insect species seem to enjoy feeding on roadside vegetation because of the increased nitrogen content of the plants living alongside traffic (Jones & Leather, 2012).

Overall however, given the very high mortality rates directly associated with cars and other road traffic and the very real indirect effects caused by habitat fragmentation, it would seem that we have much to do to make roads safer for insects and other animals.

References

Andersson, P., Koffman, A., Sjödin, N.E. & Johansson, V. (2017) Roads may act as barriers to flying insects: species composition if bees and wasps differs on two sides of a large highway.  Nature Conservation, 18, 41-59.

Baxter-Gilbert, J.H., Riley, J.L., Neufeld, C.J.H., Litzgus, J.D., & Lesbarreres, D. (2015) Road mortality potentially responsible for billions of pollinating insect deaths annually. Journal of Insect Conservation, 19, 1029-1035.

Fitch, G. & Vaidya, C. (2021) Roads pose a significant barrier to bee movement, mediated by road size, traffic and bee identity. Journal of Applied Ecology, 58,1177–1186.

Glick, P.A. (1939) The Distribution of Insects, Spiders, and Mites in the air.  Technical Bulletin no. 673, USDA. https://naldc.nal.usda.gov/download/CAT86200667/PDF

Grout, A. & Russell, R.C. (2021)H Aircraft disinsection: what is the usefulness as a public health measure? Journal of Travel Medicine, 28, taaa124.

House, A.P.N., Ring, J.G., Hill, M.J. & Shaw, P.P. (2020) Insects and aviation safety: The case of the keyhole wasp Pachodynerus nasidens (Hymenoptera: Vespidae) in Australia. Transportation Research Interdisciplinary Perspectives, 4, 100096.

Jones, E.L. & Leather, S.R. (2012) Invertebrates in urban areas: a review. European Journal of Entomology, 109, 463-478.

McKenna, D.D., McKenna, K., Malcolm, S.B. & Berenbaum, M.R. (2001) Mortality of lepidoptera along roadways in Central Illinois. Journal of the Lepidopterist’s Society, 55, 63-68.

Melis, C., Olsen, C.B., Hyllvang, M., Gobbi, M., Stokke, B.G., & Røskaft, E. (2010) The effect of traffic intensity on ground beetle (Coleoptera: Carabidae) assemblages in central Sweden. Journal of Insect Conservation, 14, 159-168.

Munõz, P.T., Torres, F.P. & Megias, A.G. (2015) Effect of roads on insects: a review. Biodiversity & Conservation, 24, 659-682.

Pop, D.R., Maier, A.R.M., Cadar, A.M., Cicort-Lucaciu, A.S., Ferenți, S. & Cupșa, D. (2020) Slower than the trains! Railway mortality impacts especially snails on a railway in the Apuseni Mountains, Romania. Annales Zoologici Fennici, 57, 225-235.

Rao, R.S.P & Girish, M.K.S. (2007) Road kills: assessing insect casualties using flagship taxon. Current Science, 92, 830-837.

Reynolds, D.R., Chapman, J.W., Edwards, A.S., Smith, A.D., Wood, C. R., Barlow, J. F. and Woiwod, I.P. (2005) Radar studies of the vertical distribution of insects migrating over southern Britain: the influence of temperature inversions on nocturnal layer concentrations. Bulletin of Entomological Research, 95, 259-274.

Riffell, S.K. (1999) Road mortality of dragonflies (Odonata) in a Great Lakes coastal wetland. Great Lakes Entomologist, 32, 63-74.

Saarinen, K., Valtonen, A., Jantunen, J. & Saarnio, J. (2005) Butterflies and diurnal moths along road verges: does road type affect diversity and abundance? Biological Conservation, 123, 403-412.

Schaffers, A.P., Raemakers, I.P., & Sýkora, K.V. (2012) Successful overwintering of arthropods in roadside verges. Journal of Insect Conservation, 16, 511-522.