Large, complex, beautiful and multi-chambered – Robin’s pincushion, rose bedeguar gall, mossy rose gall

As I wrote a little while ago, thanks to the Covid-19 lockdown I have been roaming the countryside around my rural retreat a lot more than I normally do. One of the things that I have noticed is that there seem to be a lot more of the spectacular galls caused by the gall wasp, Diplolepis rosae, than I can remember seeing in previous years. I have absolutely no empirical evidence for this observation, so it could all be down to shifting baselines (Jones et al., 2020). Nevertheless, to me they have been much more noticeable this year, perhaps confounding the insect apocalypse narrative (Leather, 2018), but then again, perhaps not.

I am, when it comes to galls, more of an expert on those caused by aphids, than on those induced by Hymenoptera, although, as it happens I am a co-author on an oak gall wasp paper (Walker et al., 2002).  Of course, that does not make me an expert. One the rose bushes in our front garden always has at least one of Robin’s spectacular pincushions clamouring for attention.  The others rarely have one and I have idly wondered about the host preferences of the wasp and the suitability of different roses for the development of the larvae.

Our front garden Diplolepis rosae gall

Having noticed that despite the relative abundance of the galls in the hedgerow roses, some bushes were totally gall-free while others supported several, sometimes in close proximity to each other, my thoughts immediately turned to possible student project.

Multiple infestations of Diplolepis rosae on hedgerow roses in the environs of Sutton, Staffordshire

A preliminary bit of research with the aid of Google Scholar quickly disabused me of that idea, but, as is the way with the internet, soon had me delving deep into the past in search of the history of this fascinating manifestation of insect activity.

My first discovery was that is also known as the rose bedeguar gall, something that had, until now, totally passed me by. According to my favourite dictionary (Gordh & Headrich, 2001),

“Bedeguar, Bedegar or Bedequar’ comes from a French word, bédégar, and is ultimately from the Persian, bād-āwar, meaning ‘wind-brought’”.

I’m guessing this meant that the first people to see them had no idea what caused them.

Not just complex galls

My next discovery was that D. rosae and the parasitoids that share its gall is a fairly well studied system (Randolph, 2005; Urban, 2018), although I am sure that it would still be possible to come up with some sort of project.  Diploelpis rosae is a member of the order Hymenoptera, so related to the larger and much more obvious bees, wasps and ants. It belongs to a family of wasps, the Cynipidae, commonly known as gall wasps.  Considering how small they are, 3.8 mm (Urban, 2018), the galls they make are spectacularly huge as are those that their relatives on oak form. Interestingly* more than 80% of gall wasps are associated with oaks, with most of the rest forming galls on members of the rose family (Shorthouse, 1973).  That in itself is, at least to me, an interesting fact; why such a restricted host range?  It is univoltine (one generation a year), overwinters as mature larva or pre-pupa in the galls, and emerges as adults in the spring when it seeks out suitable egg-laying sites. It is mainly parthenogenetic, although males are occasionally found (Callan, 1940; Stille, 1984). Despite being tiny, each wasp has the potential to lay about 500 eggs (Stille & Dävring, 1980). The adults are not very adventurous, usually laying eggs in the developing buds or flowers of the bush they emerged on, or on another close by (Bronner, 1985: Urban, 2018). That said, there must be some sort of host preference and selection going on, as in Sweden and the Czech Republic most galls are found on Rosa canina (Stille, 1984; Urban 2018). They also seem to favour younger bushes, or those that produce long vigorous stems (Stille, 1984).  The potentially high fecundity is presumably an adaptation to the high rates of parasitism that the larvae of D. rosae can experience, up to 70% in some cases (Stille, 1984).  In fact, so varied and numerous are the parasites, that many of the early papers about D. rosae pay more attention to the other inhabitants of the galls than they do the architects (Osten Sacken, 1870; Blair, 1945; Bugbee, 1951).  Females that lay a lot of eggs in the same developing bud produce bigger galls and a greater proportion of the larvae survive (Stille, 1984).

Survival of D. rosae in relation to gall weight (after Stille, 1984)

 It also appears that the closer to the ground the galls are, the lower the parasitism rate (Laszló, 2001).

The ideal strategy would then be for female D. rosae to lay big galls as low down on the plants as possible, but from personal observation this is not always the case , so as is often the case with the “Mother knows best” hypothesis there is something we humans are missing that the insects aren’t (Awmack & Leather, 2002).

Those darned taxonomists!

So, as so much is already known about this tiny wasp and its spectacular gall, I thought I would do a little bit of entomological archaeology and trace the entomological history of this little insect. Now I think taxonomists are wonderful people and have a huge respect for the very often unacknowledged work that they do, and am a great supporter of the campaign to make sure that people cite them in their papers (Packer et al., 2018), but they are a contentious bunch :-).  I mention this because inputting Diplolepis rosae into Google Scholar didn’t get me very far back in time, 1951 to be precise (Bugbee, 1951).  This paper justifies my good-natured jibe at the argumentative nature of taxonomists as he explains his renaming of what was then Rhodites rosae, by citing a 1917 paper (Rohwer & Fagan, 1917) who argued that the French entomologist Étienne Geoffroy (1725-1810) who raised the Genus Diplolepis in 1762, should take precedence over Theodor Hartig’s (1805-1880) 1840 Genus Rhodites. This pointed me back to the 1940s and the discovery that the mossy rose gall was then known as Rhodites rosae and I quote “I have recently published an epitome of  my own experience e in rearing from galls of R. rosae” (Blair, 1945). A bit more delving and I found that in France in the 1930s and in the USA in the 1920s, it was still known as R. rosae (Weld, 1926; da Silva  Tavares, 1930). Wending my back via citations I arrived in 1903 to find it listed as Cynips rosae (Ashmead, 1903).

Historical insights – Monsieur Wirey was ahead of his time

Armed with this knowledge, my journey back into the history of D. roase was much simplified and introduced me to a gem of a book, Insect Architecture by James Rennie  (1787-1867) (Rennie, 1851). Here I found an interesting account of galls in general but a detailed exposition of the Bedeguar gall of rose as he described it, including this rather nice drawing.

Professor Rennie presents some hypotheses on the formation of insect galls in general;

“Many of the processes which we have detailed bear some resemblance to our own operations of building with materials cemented together; but we shall now turn our attention to a class of insect-architects, and who cannot, so far as we know, be matched in prospective skill by any of the higher orders of animals. We refer to the numerous family which have received the name of gall-flies,

1. Wirey says, the gall tubercle is produced by irritation, in the same way as an inflamed tumor in an animal body, by the swelling of the cellular tissue and the flow of liquid matter, which changes the organization, and alters the natural external form. This seems to be the received doctrine at present in France. “

As you can see from the above he has little time for the French explanation (typical English exceptionalism) and puts forward his own idea that the galls are formed because the egg laying process blocks the vessels of the plant and the fluid that would normally flow unimpeded blows up the tissue surrounding the egg like a balloon.  Of course he was wrong and M. Wirey was correct :-).  Considering that he had no access to the sophisticated techniques we have he pretty much hit the nail on the head.

That aside, his book introduced me to an entomologist I had never heard of, Priscilla Wakefield (1751-1832), yet another overlooked and forgotten female scientist.

Possible projects?

Although my plans for lots of great MSc projects were reduced somewhat I have had a lot of educational fun and I think that there are still some things that could usefully be looked at, long term recording across multiple sites which I hope the British Plant Gall Society is doing would be interesting.  On my walks I noticed a lot of variability in size and phenology of gall formation.  At the end of August I was coming across small very fresh looking galls at the same time as I was seeing larger more advanced galls.

A very fresh looking Bedeguar gall, August 26^th 2020, Sutton

As far as I can tell, the timing of gall formation and its effect on final size of the galls has not been looked at in detail; do early galls enter winter larger than later formed galls, or is it entirely due to the number of eggs laid?  Given the huge number of other inhabitants of the galls, at least fourteen different species (Laszló, 2001, there is probably a viable project in looking at the timing of invasion by the different gall parasites and the outcome this may or may not have on the final composition of the gall fauna.

Feel free to suggest additional projects in the comments.

References

Ashmead, W.H. (1903) Classification of the gall-wasps and the parasitic cynipoids, or the superfamily Cynipoidea. IV, Psyche, 10, 210-216.

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

Blair, K.G. (1945) Notes on the economy of the rose‐galls formed by Rhodites (Hymenoptera, Cynipidae). Proceedings of the Royal Entomological Society, Series A, 20, 26-31.

Bronner, R. (1985) Anatomy of the ovipositor and oviposition behavior of the gall wasp Diplolepis rosae (Hymenoptera: Cynipidae). Canadian Entomologist, 117, 849-858.

Bugbee, R. E. (1951). New and described parasites of the genus Eurytoma illiger from rose galls caused by species of the cynipid genus Diplolepis Geoffrey Hymenoptera: Eurytomidae). Annals of the Entomological Society of America, 44, 213–261

Callan, E.Mc. (1940) On the occurrence of males of Rhodites rosae (l.) (Hymenoptera, Cynipidae). Proceedings of the Royal Entomological Society of London, Series A., 15, 21-26.

Da Silva Tavares, J. (1930) Quelques Cécidies du Centre de la France, Publications de la Société Linnéenne de Lyon, 75, 145-167.

Gordh, G & Headrick, D.H. (2001) A Dictionary of Entomology, CABI, Wallingford

Jones, L.P., Turvey, S.T.,  Massimino, D. & Papworth, S. K.(2020) Investigating the implications of shifting baseline syndrome on conservation. People & Nature,

Laszló, Z. (2001) The parasitic complex of Diplolepis rosae (LinnaeuS, 1758) (Hymenoptera, Cynipidae): influencing factors and interspecific relationships. Entomologica Romanica, 6, 133–140.

Leather, S.R. (2018) “Ecological Armageddon” – more evidence for the drastic decline in insect numbers. Annals of Applied Biology, 172, 1-3.

Osten Sacken, R. (1870) Contributions to the natural hstory of the Cynipidæ of the United State and their galls.  Transactions of the American Entomological Society, 3, 54-64.

Packer, L., Monckton, S.K., Onuferko, T.M. & Ferrari, R.R. (2018) Validating taxonomic identifications in entomological research. Insect Conservation & Diversity, 11, 1-12.

Randolph, S. (2005) The Natural History of the Rose Bedeguar Gall and its Insect Community, The British Plant Gall Society.

Rennie, J. (1857) Insect Architecture. John Murray, London.

Rohwer, S. A. & Fagan, M. M. (1917) The type-species of the genera of the Cynipidea, or the gall wasps and parasitic cynipoids. Proceedings of the U.S. National Museum, 53, 357-380.

Shorthouse, J.D. (1973) The insect community associated with rose galls of Diplolepis polita (Cynipidae, Hymenoptera). Quaestiones Entomologicae, 9, 55-98.

Stille, B. (1984) The effect of hosptlant and parasitoids on the reproductive success of the parthenogenetic gall wasp Diplolepis rosae (Hymenoptera, Cynipidae). Oecologia, 63, 364-369.

Stille, B. & Dävring, L. (1980) Meiosis and reproductive strategy in the parthenogenetic gall wasp Diplolepis rosae (L.) (Hymenoptera, Cynipidae), Heriditas, 92, 353-362.

Urban, J. (2018). Diplolepis rosae (L.) Hymenoptera: Cynipidae): development, ecology and galls in the Brno region. Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis, 66, 905-925.

Wakefield, P. (1816) An Introduction to the Natural History and Classification of Insects in a series of Familiar Letters with Illustrative Engravings. Darton, Harvey & Darton, London.

Walker, P., Leather, S.R. & Crawley, M.J. (2002) Differential rates of invasion in three related alien oak gall wasps (Cynipidae: Hymenoptera). Diversity & Distributions, 8, 335-349.

Weld, L. H. (1926) Field notes on gall-inhabiting cynipid wasps with descriptions of new species. Proceedings of the United States National Museum, 68, 1-131.

https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1365-3032.1940.tb00573.x

https://www.britishplantgallsociety.org/bedeguar-keys.pdf

*

my wife tells me that I am always starting sentences like this and more often than not, it is not interesting at all 🙂

 

Buzzing with invention and intrigue – The Bees by Laline Paull

Laline Paulll, The Bees, Harper Collins (2014) ISBN 978-0-00-755774-5

Having suffered the trauma of watching Antz and the Bee Movie, I’m always a tad reluctant to embark on books that feature insects as their main protagonists. Maya Leonard’s Beetle Boy trilogy, which I thoroughly enjoyed, is different, as the insects play a supporting role.  It is probably this prejudice that has allowed this wonderful book to have been unread by me for six long years 🙂

I can’t remember who recommended this book, but I’m glad I took them up on it.  Despite the glowing recommendation and the numerous blurbs inside and out (after all Stephen Heard has recently revealed the truth about book blurbs) I began reading Laline Paull’s debut novel with some trepidation. I was pleasantly surprised, despite the inevitable anthropomorphisation of the heroine*, (I don’t think the novel would have worked without it), I engaged wholeheartedly with the story.

I was a bit dubious at first about the kin group theme, the heroine is a Flora (717 to be exact), and there are Teasels, Clovers and the evil Sages, as I, erroneously as it turned out, had this idea that all the members of a hive were full sisters.  I had, however, misremembered, honey bee queens, unlike many other Hymenoptera, are no strangers to multiple mating**(polyandry), having, in fact, the highest levels of this trait of all the social insects (Strassman, 2001). Biologically, the Queen having access to multiple sperm-donors is highly advantageous, as when disease strikes, as it does in the novel, not all the inhabitants of the colony are equally vulnerable (Tarpy, 2003). British elms would not have been all but exterminated by Dutch Elm Disease, if they had not all been members of a single clone.

The other characteristic of bees that some might feel that Laline Paull plays a little fast and loose with is temporal polyethism (age based division of labour). I had slight misgivings about the rigidity of the division of labour within the hive. It has long been known that honeybee workers exhibit temporal polyethism (age-based division of labour) (Pérez, 1889). Young workers perform brood-nest associated tasks such as brood-cell cleaning and larval feeding, graduating on to food processing, nest construction, and guarding and finally as they enter old age, become foragers (Seeley and Kolmes 1991). Flora 717 does indeed go through these phases, but the rest of her kin group seem to be sanitation workers throughout their lives and the scheming Sage priestesses seem to have no other function than to spread their mantra of “Accept, Obey, Serve” and to direct the action of the sinister police bees. In case you think that police bees are a bit too detached from reality, worker bees do ‘police’ other workers when it comes to ‘unauthorised’ egg laying (Ratnieks & Visscher, 1989). Although it has been shown that different genotypes of bees within a hive do show some variation in the timing of their move from one task to another (Siegel et al., 2013), there is, as far, as I can find, no evidence of genotypes that remain fixed in one job their whole lives.

I guess the biggest issue, without giving the climax of the story away, is the production of a Queen from an egg laid by a worker bee. Worker bees can, and do lay unfertilised eggs, but, with one exception, they are invariably males.  Workers of the Cape honeybees (Apis mellifera capensis), however, can produce female eggs parthenogenetically (Hepburn, 1994), a phenomenon known as thelytoky. If fed the right food during the first 72 hours of their larval life, these eggs, could in theory, develop into Queens, (Pérez, 1889). Although the story is not set in South Africa, I am willing to give this a pass and assume that one of the drones that impregnated the Queen of Flora’s hive was a Cape honeybee.

The many issues facing honeybees are brought to life in this dramatic and believable story.  Experience the effects of pesticides, pollution and ‘phone masts on our heroine and her hive mates at first hand.  Cower as the wasps attack, and when a starving mouse gains entry to the hive in mid-winter, wince as the surplus drones are disposed of by the workers and cheer as our heroine saves the day.  This is a gripping story, and despite my reservations about the ‘hive mind’ Laline has taken the advice of her entomological advisors to heart and made a hugely successful foray into depicting the life style and ecology of the honeybee.

Definitely worth reading, a tour de force.

 

References

Hepburn, H.R. (1994) Reproductive cycling and hierarchical competition in Cape honeybees, Apis mellifera capensis Esch. Apidologie, 25, 38-48.

Pérez J. (1889) Les Abeilles. Paris, France: Hachette et Cie.

Ratnieks, F.L.W. & Visscher, P.K. (1989) Worker policing in the honeybee. Nature, 342, 796-797.

Seeley, T.D. & Kolmes, S.A. (1991) Age polyethism for hive duties in honey bees — illusion or reality? Ethology, 87, 284-297.

Siegel, A. J., Fondrk, M. K., Amdam, G. V., & Page, R. E., Jr (2013). In-hive patterns of temporal polyethism in strains of honey bees (Apis mellifera) with distinct genetic backgrounds. Behavioral Ecology and Sociobiology, 67, 1623–1632.

Strassmann, J. E. (2001) The rarity of multiple mating by females in the social Hymenoptera. Insectes Sociaux, 48, 1–13.

Tarpy, D.R. (2003) Genetic diversity within honeybee colonies prevents severe infections and promotes colony growth. Proceedings of the Royal Society B, 270, 99-103.

*Unlike makers of The Bee Movie, Laline knows what sex worker bees are 🙂

** Note that I did not use the word promiscuous; promiscuity is a human trait, not an insect one.

Shocking News – the truth about electroperception – insects can ‘feel’ electric fields

Static electric fields are common throughout the environment and this has been known for some time (e.g Lund (1929) and back in 1918, the great Jean-Henri Fabre, writing about the dung beetle, Geotrupes stated “They seem to be influenced above all by the electric tension of the atmosphere. On hot and sultry evenings, when a storm is brewing, I see them moving about even more than usual. The morrow is always marked by violent claps of thunder”

Given this, it is surprising that it was not until the 1960s that entomologists started to take a real interest in electroperception, when a Canadian entomologist decided to investigate the phenomenon further, but using flies (Edwards, 1960).  He found that if Drosophila melanogaster and Calliphora vicina exposed to, but not in contact with, an electrical field, they stopped moving. Calliphora vicina needed a stronger voltage to elicit a response than D. melanogaster, which perhaps could be related to their relative sizes. It seemed that their movement was reduced when electrical charge applied and changed, but not if the field was constant.

Responses of two fly species to electrical fields (From Edwards, 1960)

In a follow up experiment with the the Geometrid moth Nepytia phantasmaria he showed that females were less likely to lay eggs when exposed to electrical fields (Edwards, 1961), but the replication was very low and the conditions under which the experiment was run were not very realistic.

In the same year, Maw (1961) working on the Ichneumonid wasp, Itoplectis conquisitor, which is attracted to light, put ten females into a chamber with a light at one end but with parts of the floor charged at different levels.  The poor wasps were strongly attracted to the light but the electrical ‘barrier’ slowed them down; the stronger the charge, the greater the reluctance to enter the field.

On the other hand, some years later, working with the housefly, Musca domestica and the cabbage looper, Trichoplusia ni, across a range of different strength electrical fields, Perumpral et al., (1978)   found no consistent avoidance patterns in where the houseflies preferred to settle, but did find that wing beat frequency of male looper moths was significantly affected, although inconsistently.  Female moths on the other hand were not significantly affected.  This put paid to their intention to develop a non-chemical control method for these two pests.

A more promising results was obtained using the cockroach Periplaneta americana.  Christopher Jackson and colleagues at Southampton University showed that the cockroaches turned away, or were repulsed, when they encountered an electric field and if continuously exposed to one, walked more slowly, turned more often and covered less distance (Jackson et al., 2011).  As an aside, this is similar to the effects one of my PhD students found when she exposed carabid beetles exposed to sub-lethal applications of the insecticide dimethoate*.

Periplaneta americana definitely showing a reluctance to cross an electrical field (Jackson et al., 2011).

Other insect orders have also been shown to respond to electric fields.  Ants, in particular the fire ant, Solenopsis invicta, are apparently a well-known hazard to electrical fittings (MacKay et al., 1992), and a number of species have been found in telephone receivers (Eagleson, 1940), light fittings and switches (Little, 1984), and even televisions (Jolivet, 1986), causing short circuits and presumably, coming to untimely ends 🙂

Rosanna Wijenberg and colleagues at Simon Fraser University in Canada, really went to town and tested the responses of a variety of different insect pests to electric fields. They found that the common earwig, Forficula auricularia, two cockroaches, Blatta germanica, Supella longipalpa, two Thysanurans, the silverfish, Lepisma saccharina and the firebrat Thermobia domestica were attracted to, or at least arrested by electrified coils.  Periplaneta americana, on the other hand, was repulsed (Wijenberg et al., 2013).  They suggested that using electrified coils as non-toxic baits might be an environmentally friendly method of domestic pest control.  I have, however, not been able to find any commercial applications of this idea although perhaps you know better?

Although a number of marine vertebrates generate electricity and electric fields as well as perceiving and communicate using them, there was, until fairly recently, no evidence of electrocommunication within the insect world (Bullock, 1999); after all, they have pheromones 😊

When we look at the interaction between insects and electromagnetic fields there is growing evidence that bees, or at least honey bees, like some birds (Mouritsen et al., 2016) have the wherewithal and ability to navigate using magnetic fields (Lambinet et al., 2017ab).  Interestingly**, honeybees, Apis mellifera have been shown to generate their own electrical fields during their waggle dances which their conspecifics are able to detect (Greggers et al., 2013).  Bumble bees (Bombus terrestris), have also been shown to be able to detect electrical fields.  In this case, those surrounding individual plants.  The bees use the presence or absence of an electrical charge to ‘decide’ whether to visit flowers or not. If charged they are worth visiting, the charge being built up by visitation rates of other pollinating insects  (Clarke et al., 2013)

Since I’m on bees, I can’t leave this topic without mentioning mobile phones and electromagnetic radiation, although it really deserves an article of its own.  The almost ubiquitous presence of mobile phones has for a long time raised concern about the effect that their prolonged use and consequent exposure of their users to electromagnetic radiation in terms of cancer and other health issues (Simkó & Mattson, 2019). Although there is growing evidence that some forms of human cancer can be linked to their use (e.g. Mialon & Nesson, 2020), the overall picture is far from clear (Kim et al., 2016). Given the ways in which bees navigate and the concerns about honeybee populations it is not surprising that some people suggested that electromagnetic radiation as well as neonicitinoids might be responsible for the various ills affecting commercial bee hives (Sharma & Kumar, 2010, Favre, 2011). The evidence is far from convincing (Carreck, 2014) although a study from Greece looking at the intensity of electromagnetic radiation from mobile phone base stations on the abundance of pollinators found that the abundance of beetles, wasps and most hoverflies decreased with proximity to the base stations, but conversely, the abundance of bee-flies and underground nesting wild bees increased, while butterflies were unaffected (Lázaro et al., 2016). A more recent study has shown that exposure to mobile phones resulted in increased pupal mortality in honeybee queens but did not affect their mating success (Odemer & Odemer, 2019).  All in all, the general consensus is that although laboratory studies show that electromagnetic radiation can affect insect behaviour and reproduction the picture remains unclear and that there are few, if any field-based studies that provide reliable evidence one way or the other (Vanbergen et al., 2019).   Much more research is needed before we can truly quantify the likely impacts of electromagnetic radiation on pollinators and insects in general.

 

Acknowledgements

I must confess that I had never really thought about insect electroperception until I was at a conference and came across a poster on the subject by Matthew Wheelwright, then an MRes student at the University of Bristol, so it is only fair to dedicate this to him.

 

References

 

Bullock, T.H. (1999) The future of research on elctroreception and eclectrocommunication.  Journal of Experimental Biology, 10, 1455-1458.

Carreck, N. (2014) Electromagnetic radiation and bees, again…, Bee World, 91, 101-102.

Clarke, D., Whitney, H., Sutton, G. & Robert, D. (2013) Detection and learning of floral electric fields by bumblebees. Science, 340, 66-69.

Eagleson, C. (1940) Fire ants causing damage to telephone equipment.  Journal of Economic  Entomology, 33, 700.

Edwards, D.K. (1960) Effects of artificially produced atmospheric electrical fields upon the activity of some adult Diptera.  Canadian Journal of Zoology, 38, 899-912.

Edwards, D.K. (1961) Influence of electrical field on pupation and oviposition in Nepytia phantasmaria Stykr. (Lepidoptera: Geometridae). Nature, 191, 976.

Fabre, J.H. (1918) The Sacred Beetle and Others. Dodd Mead & Co., New York.

Favre, D. (2011) Mobile phone induced honeybee worker piping. Apidologie, 42, 270-279.

Greggers, U., Koch, G., Schmidt, V., Durr, A., Floriou-Servou, A., Piepenbrock, D., Gopfert, M.C. & Menzel, R. (2013) Reception and learning of electric fields in bees. Proceedings of the Royal Society B, 280, 20130528.

Jackson, C.W., Hunt, E., Sjarkh, S. & Newland, P.L. (20111) Static electric fields modify the locomotory behaviour of cockroaches. Journal of Experimental Biology, 214, 2020-2026.

Jolivet, P. (1986) Les fourmis et la Television. L’Entomologiste, 42,321-323.

Kim, K.H., Kabir, E. & Jahan, S.A. (2016) The use of cell phone and insight into its potential human health impacts. Environmental Monitoring & Assessment, 188, 221.

Lambinet, V., Hayden, M.E., Reigel, C. & Gries, G. (2017a) Honeybees possess a polarity-sensitive magnetoreceptor. Journal of Comparative Physiology A, 203, 1029-1036.

Lambinet V, Hayden ME, Reigl K, Gomis S, Gries G. (2017b) Linking magnetite in the abdomen of honey bees to a magnetoreceptive function. Proceedings of the Royal Society, B., 284, 20162873.

Lazáro, A., Chroni, A., Tscheulin, T., Devalez, J., Matsoukas, C. & Petanidou, T. (2016) Electromagnetic radiation of mobile telecommunication antennas affects the abundance and composition of wild pollinators.  Journal of Insect Conservation, 20, 315-324.

Little, E.C. (1984) Ants in electric switches. New Zealand Entomologist, 8, 47.

Lund, E.J. (1929) Electrical polarity in the Douglas Fir. Publication of the Puget Sound Biological Station University of Washington, 7, 1-28.

MacKay, W.P., Majdi, S., Irving, J., Vinson, S.B. & Messer, C. (1992) Attraction of ants (Hymenoptera: Formicidae) to electric fields. Journal of the Kansas Entomological Society, 65, 39-43.

Maw, M.G. (1961) Behaviour of an insect on an electrically charged surface. Canadian Entomologist, 93, 391-393.

Mialon, H.M. & Nesson, E.T. (2020) The association between mobile phones and the risk of brain cancer mortality: a 25‐year cross‐country analysis. Contemporary Economic Policy, 38, 258-269.

Mouritsen, H., Heyers, D. & Güntürkün, O. (2016) The neural basis of long-distance navigation in birds. Annual Review of Physiology, 78, 33-154.

Odemer, R., & Odemer, F. (2019). Effects of radiofrequency electromagnetic radiation (RF-EMF) on honey bee queen development and mating success. Science of The Total Environment, 661, 553–562.

Perumpral, J.V., Earp, U.F. & Stanley, J.M. (1978) Effects of electrostatic field on locational preference of house flies and flight activities of cabbage loopers. Environmental Entomology, 7, 482-486.

Sharma, V.P. & Kumar, N.R. (2010) Changes in honeybee behaviour and biology under the influence of cellphone radiation. Current Science, 98, 1376-1378.

Simkó, M. & Mattson, M.O. (2019) 5G wireless communication and health effects—A pragmatic review based on available studies regarding 6 to 100 GHz. International Journal of Environmental Research & Public Health, 16, 3406.

Vanbergen, A.J., Potts, S.G., Vian, A., Malkemper, E.P., Young, J. & Tscheulin, T. (2019) Risk to pollinators from anthropogenic electro-magnetic radiation (EMR): Evidence and knowledge gaps. Science of the Total Environment, 695, 133833.

Wijenberg, R., Hayden, M.E., Takáca, S. & Gries, G. (2013) Behavioural responses of diverse insect groups to electric stimuli. Entomoloogia experimentalis et applicata, 147, 132-140.

 

*

yet another entry for my data I am never going to publish series 😊

 

**

My wife really hates it when I start a sentence like this, as she says “You’re always starting sentences like that and it is rarely interesting”

Twisted, hairy, scaly, gnawed and pure – side-tracked by Orders

I’m supposed to be writing a book, well actually two, but you have to be in the right mood to make real progress. Right now, I’m avoiding working on one of the three chapters that I haven’t even started yet* and I really should be on top of them by now as I have already spent the advance, and have less than a year to go to deliver the manuscript 😦 Instead of starting a new chapter I’m tweaking Chapter 1, which includes an overview of Insect Orders.  While doing that I was side tracked by etymology. After all, the word is quite similar to my favourite subject and a lot of people confuse the two. Anyway, after some fun time with my Dictionary of Entomology, (which is much more of an encyclopaedia than a dictionary), and of course Google, I have great pleasure in presenting my one stop shop for those of you who wonder how insect orders got their names.  Here they are, all in one easy to access place with a few fun-filled facts to leaven the mixture.

Wings, beautiful wings (very much not to scale)

First, a little bit of entomological jargon for those not totally au fait with it.  Broadly speaking we are talking bastardised Greek and Latin. I hated Latin at school but once I really got into entomology I realised just how useful it is.  I didn’t do Greek though 😊, which is a shame as Pteron is Greek for wing and this is the root of the Latin ptera, which features all over the place in entomology.

Since I am really only talking about insects and wings, I won’t mention things like the Diplura, Thysanura and other Apterygota.  They don’t have wings, the clue being in the name, which is derived from Greek; A = not, pterygota, derived from the Greek ptérugos = winged, which put together gives us unwinged or wingless. In Entojargon, when we talk about wingless insects we use the term apterous, or if working with aphids, aptera (singular) or apterae (plural).   I’m going to deal with winged insects, the Exopterygota and the Endopterygota. The Exopterygota are insects whose wings develop outside the body and there is a gradual change from immature to adult.  Think of an aphid for example (and why not?); when the nymph (more Entojargon for immature hemimetabolus insects) reaches the third of fourth instar (Entojargon for different moulted stages), they look like they have shoulder pads; these are the wing buds, and the process of going from egg to adult in this way is called incomplete metamorphosis.

Fourth instar alatiform nymph of the Delphiniobium junackianum the Monkshood aphid.  Picture from the fantastic Influential Points site https://influentialpoints.com/Images/Delphiniobium_junackianum_fourth_instar_alate_img_6833ew.jpg (Any excuse for an aphid pciture)

In the Endopterygota, those insects where the wings develop inside the body, e.g butterflies and moths, the adult bears no resemblance to the larva and the process is described as complete metamorphosis and the life cycle type as holometabolous. It is also important to note that the p in A-, Ecto- and Endopterygota is silent.

Now on to the Orders and their names.  A handy tip is to remember is that aptera means no wings and ptera means with wings.  This can be a bit confusing as most of the Orders all look and sound as if they have wings.  This is in part, due to our appalling pronunciation of words; we tend to make the syllables fit our normal speech patterns which doesn’t necessarily mean breaking the words up in their correct component parts. Diptera and Coleoptera are two good examples – we pronounce the former as Dip-tera and informally as Dips.  From a purist’s point of view, we should be pronouncing the word Di-tera – two wings, and similarly, Coleoptera as Coleo-tera, without the p 🙂 Anyway, enough of the grammar lessons and on with the insects.

Exopterygota

Ephemeroptera The Mayflies, lasting a day or winged for a day J The oldest extant group with wings. They are also a bit weird, as unlike other Exopterygota they have a winged sub-adult stage

Odonata              Dragonflies and Damselflies – think dentists, toothed, derived from the Greek for tooth, odoús. Despite their amazing flight capability, the name refers to their toothed mandibles.  The wings do get a mention when we get down to infraorders, the dragonflies, Anisoptera meaning uneven in that the fore and hind wings are a different shape and the damselflies, Zygoptera  meaning even or yoke, both sets of wings being pretty much identical.

Dermaptera       Earwigs, leathery/skin/hide, referring to the fore-wings which as well as being leathery are reduced in size.  Despite this, the much larger membranous hind wings are safely folded away underneath them.

A not very well drawn (by me) earwig wing 😊

Plecoptera          Stoneflies, wickerwork wings – can you see them in the main image?

Orthoptera         Grasshoppers and crickets, straight wings, referring to the sclerotised forewings that cover the membranous, sometimes brightly coloured hind wings.  Many people are surprised the first time they see a grasshopper flying as they have been taken in by the hopper part of the name and the common portrayal of grasshoppers in cartoons and children’s literature; or perhaps not read their bible “And the locusts went up over all the land of Egypt, and rested in all the coasts of Egypt”. I think also that many people don’t realise that locusts are grasshoppers per se.

Grasshopper wings

Dictyoptera        Cockroaches, termites and allies, net wings

Notoptera           The order to which the wingless Ice crawlers (Grylloblattodea) and Gladiators Mantophasmatodea) belong. Despite being wingless, Notoptera translates as back wings. It makes more sense when you realise that the name was coined when only extinct members of this order were known and they were winged.

Mantodea           Mantids, the praying mantis being the one we are all familiar with, hence the name which can be translated as prophet or soothsayer

Phasmotodea    Phasmids, the stick insects and leaf insects – phantom, presumably referring to their ability to blend into the background.

Psocoptera         Bark lice and book lice, gnawed or biting with wings. In this case the adjective is not in reference to the appearance of the wings, but that they are winged insects that can bite and that includes humans, although in my experience, not very painful, just a little itchy. They are also able to take up water directly from the atmosphere which means that they can exploit extremely dry environments.

Embioptera        Web spinners, lively wings. Did you know that Janice Edgerly-Rooks at Santa Clara University has collaborated with musicians to produce a music video of Embiopteran silk spinning? https://www.youtube.com/watch?v=veehbMKjMgw

Zoraptera            Now this is the opposite of the Notoptera, the Angel insects, Zora meaning pure in the sense of not having any wings.  Unfortunately for the taxonomists who named this order, winged forms have now been found 🙂

Thysanoptera    Thrips and yes that is both the plural and singular, thysan meaning tassel wings, although I always think that feather would be a much more appropriate description.

Feathery thrips wing – Photo courtesy of Tom Pope @Ipm_Tom

Hemiptera          True bugs – half wings.  The two former official suborders were very useful descriptions, Homoptera, e.g. aphids, the same. Heteroptera such as Lygaeids, e.g. Chinch bugs, which are often misidentified by non-entomologists as beetles where the prefix Hetero means different, referring to the fact that the fore wings are hardened and often brightly coloured in comparison with the membranous hind wings.

Coreid bug – Gonecerus acuteangulatus – Photo Tristan Banstock https://www.britishbugs.org.uk/heteroptera/Coreidae/gonocerus_acuteangulatus.html

Phthiraptera      The lice, the name translates as wingless louse. I guess as one of the common names for aphids is plant lice they felt the need to make the distinction in the name.

Siphonaptera     Fleas – tube without wings, referring to their mouthparts

 

Endopterygota

Rhapidioptera   Snakeflies – needle with wings, in this case referring to the ovipositor, not to the wings, which are similar to those of dragonflies.

The pointy end of a female snakefly

Megaloptera      Alderflies, Dobsonflies – large wings

Neuroptera        Lacewings – veined wings

Coleoptera         Beetles – sheathed wings, referring to the hardened forewings, elytra, that cover the membranous hind wings. The complex process of unfolding and refolding their hind wings means that many beetles are ‘reluctant’ to fly unless they really need to.

Strepsiptera       These are sometimes referred to as Stylops.  They are endoparasites of other insects. The name translates as twisted wings. Like flies, they have only two pairs of functional wings the other pair being modified into halteres.  Unlike flies, their halteres are modified fore wings.  Their other claim to fame is that they feature on the logo of the Royal Entomological Society.

The Royal Entomological Society Strepsipteran

Mecoptera         Scorpionflies, hanging flies – long wings.  Again, not all Mecoptera are winged, but those that are, do indeed have long wings in relation to their body size.

Male Scorpionfly, Panorpa communis.  Photo David Nicholls https://www.naturespot.org.uk/species/scorpion-fly

Siphonaptera     Fleas – tube no wings. The tube part of the name refers to their mouthparts.

Diptera                 Flies, two wings, the hind pair are reduced to form the halteres, which are a highly complex orientation and balancing device.

Trichoptera         Caddisflies, which are, evolutionarily speaking, very closely related to the Lepidoptera.  Instead of scales, however, their wings are densely cover with small hairs, hence the name hairy wings.  Some species can, at first glance, be mistaken for small moths. If you want to know more about caddisflies I have written about them here.

Lepidoptera       Moths and butterflies, scaly wings; you all know what happens if you pick a moth or butterfly up by its wings.

Moth wing with displaced scales

 

Hymenoptera    Wasps, bees, ants – membrane wings

Wing of a wood wasp, Sirex noctilio

 

And there you have it, all 30 extant insect orders in one easy location.

 

*

Just to remind you that I am a champion procrastinator 🙂

 

Pick and mix 21 – a cornucopia of links

There may actually be more Hymenoptera than there are Coleoptera!

Some book aren’t just for reading – wonderful hidden art

Fighting bats with long tails – moth evolution

Are you working on the right problem?

Bang, crackle, flash – Interesting paper about insect and arthropod names for fireworks

Inspired by the recent World Cup the John Innes Centre held their own version to champion discoveries they have made over the last 70 years 🙂

Insects through the Looking Glass – using Lewis Carroll to foster a love of insects

Victorian entomologists had a lot of fun – great post from Manu Saunders

A great post about science communication via Twitter by Stephen Heard

Spots on butterfly wings – what are they for?  Ray Cannon has some thoughts

My group is bigger, better and more beautiful than yours – The annual MSc Entomology trip to the Natural History Museum, London, 2018

This week we went on one of my favourite trips with the MSc Entomology students.  We visited the Natural History Museum in London.  We got off to fantastic start – all the students, and staff, arrived at the arranged time of 0645, something that had never happened before :-). The weather was fine, although at that time in the morning it was too dark to really appreciate it, and off we set.  I should have known that something would go wrong and sure enough the traffic was awful, and we had to make an unscheduled stop at a motorway service station to make sure our driver didn’t exceed his quota of working hours.

The now much delayed coach basking in the sunshine at a motorway service station.

Some of the MSc students; remaining cheerful despite the delay.

Forty-five minutes later we set off again and despite encountering a few further delays arrived safely, albeit almost an hour and a half late.  Luckily our host for the day Erica McAlister (@flygirlNHM) was ready and waiting and very efficiently got our visit back on track.  This year we were shown Colossal Coleoptera by Michael Geiser, Huge Hymenoptera by Nathalie Dale-Skey, Lustrous Lepidoptera by Alessandro Giusi and Deadly Diptera by Erica McAlister.   All our specialist hosts were, as you would expect, very keen to extol the virtues of their groups, and who can blame them.  I do the same with Awesome aphids 🙂 We are always very appreciative of the time and care that the NHM entomologists give us, especially as they have, sadly, recently had their numbers reduced.  Hopefully, as the realities of the problems associated with insect conservation and identification become even more apparent than they already are, we will see the appointment of more entomologists to this very much-needed global resource.  Here are some pictures to give you a flavour of the day.

Mouse mat for forensic entomologists 🙂

Alessandro Giusti waxing lyrical about the biggest, the smallest and the most beautiful Lepidoptera (moths as far as he is concerned).

 

The large and the small (a really bad photo by yours truly, I am still getting to grips with my new camera)

Natalie Dale-Skey extolling the virtues of Hymenoptera

They don’t have to be big and tropical to be beautiful – these are tiny but gorgeous

I do like a good wasp nest 🙂

Erica McAlister on the sex life of flies

The biggest flies in the world pretending to be wasps

A selection of flies

I was very impressed that the Crane fly still has all its legs attached.  I collected Crane flies for my undergraduate collection and had to resort to sticking their legs on to a piece of card.

Not quite the rarest fly in the World but as its larvae live inside rhinoceroses it could be in trouble 😦

Big beautiful beetles

Cockchafers aren’t really this big, but wouldn’t it be awesome if they were?

MSc Entomology (@Entomasters) at the end of the visit.  Photo courtesy of Heather Campbell (@ScienceHeather), our newest member of staff

Once again, a huge vote of thanks to Erica and colleagues for making this a memorable visit.  We had a fantastic day.

An inordinate fondness for biodiversity – a visit behind the scenes at the Natural History Museum

Last week  (13th February) I traveled with the MSc Entomology students to the Natural History Museum, London.  As part of their course they are taken behind the scenes and meet some of the curators and their favourite beasts.  This one of my favourite course trips and although I have made the pilgrimage for many years I always find something new to marvel at as well as reacquainting myself with some of my old favourites.  After an early start (0645) we arrived exactly on time (for a change), 10.30, at the Museum site in South Kensington.  I always have mixed feelings about South Kensington, having spent twenty years of my life commuting to Imperial College, just up the road from the museum.  I loved teaching on the Applied Ecology course I ran, but over the years the working atmosphere in the Department became really toxic* and I was extremely glad to move to my present location, Harper Adams University.  After signing in, which with twenty students took some time, Erica McAlister (@flygirl) led us through the thronged galleries (it was half term) to the staff

Nostalgia time, my first biological memory, aged 3.

areas, where the research, identification and curating takes place.  Our first port of call was the Diptera where Erica regaled us with lurid tales of flies, big and small, beneficial and pestiferous.

Erica McAlister extolling the virtues of bot flies

Any one fancy cake for tea?  Kungu cake, made from African gnats

Early advisory poster

As we left to move on to the Hymenopteran, hosted by David Notton, I noticed this classic poster warning against mosquitoes.  David chose bees as the main focus of his part of the tour, which as four of the students will be doing bee-based research projects was very apt.

Admiring the bees

Whilst the students were engrossed with the bees I did a bit of fossicking and was amused to find that tobacco boxes were obviously a preferred choice by Scandinavian Hymenopterists in which to send their specimens to the museum.

Finnish and Swedish tobacco boxes being put to good use

Next was that most eminent of Coleopterists, Max @Coleopterist Barclay who as usual enthralled the students and me, with stories of

Max Barclay demonstrating a Lindgren funnel and talking about ‘fossilised’ dung balls

beetles large and small, anecdotes of Darwin and Wallace and the amusing story of how ancient clay-encased dung balls were for many years thought by anthropologists and archaeologists to be remnants of early humankind’s bolas hunting equipment.  It was only when someone accidentally broke one and found a long-dead dung beetle inside that the truth was revealed 🙂

Often overlooked, the Natural History Museum is an exhibit in itself

 As we were leaving to move on to the Lepidoptera section, I felt obliged to point out to the students that not only is the outside of the museum stunningly beautiful but that the interior is also a work of art in itself, something that a lot of visitors tend to overlook. Once in the Lepidoptera section  Geoff Martin proudly displayed his magnificent collection of Lepidoptera, gaudy and otherwise, including the type specimen of the Queen Alexandra’s Birdwing which was captured with the aid of a shotgun!

Lepidopterist, Geoff Martin, vying with his subjects in colourful appearance 🙂

Lunch and a chance to enjoy the galleries was next on the agenda.  Unfortunately, as it was half term this was easier said than done, although I did find a sunny spot to eat my packed lunch, as a Yorkshireman I always find the prices charged for refreshments by museums somewhat a painful.  In an almost deserted gallery I came across this rather nice picture.

A lovely piece of historical entomological art.

Then it was on to the Spirit Collection.  Erica had laid on a special treat, Oliver Crimmen, fish man extraordinaire.  I may be an entomologist but I can sympathise with this branch of vertebrate zoology.  Fish, like insects are undeservedly ranked below the furries, despite being the most speciose vertebrate group.  I have been in the Spirit Room many times but have never seen inside the giant metal tanks.  Some of these, as Ollie demonstrated with a refreshing disregard for health and safety, are filled with giant fish floating in 70% alcohol.

Fish man, Oliver Crimmen, literally getting to grips with his subjects.

A fantastic end to the day culminated with a group photo with a spectacular set of choppers 🙂

Many thanks to Erica McAlister for hosting and organising our visit and to the NHM staff who passionately attempted to convert the students to their respective ‘pets’.

*one day I will write about it.

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.

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.

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 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.

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 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.

 

References

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.

 

Entomological classics – The Malaise Trap

More years ago than I care to remember, my friends and I were playing the now, very non-PC game of Cowboys and Indians, when we saw through the trees, what we thought was a tent. On sneaking up to it we found that, if it was a tent, it wasn’t very watertight!  There were no sides, instead there was a central panel and the whole thing was made of netting.  What we had actually found, was of course a Malaise trap, although of course we did not know this at the time.  It was only later as an undergraduate that I realised what we had found all those years before.

So exactly what is a Malaise trap and how did it come into being? The Malaise Trap is a relatively new invention.  It was invented by the Swedish entomologist, Dr René Malaise in the 1930s (hence the name) and revealed to a more general entomological audience in 1937 (Malaise, 1937).  It was actually designed as a replacement for the traditional hand-held collecting net, which as Malaise states in the introduction to his paper ‘”Since the time of Linneaus, the technique of catching insects has not improved very much, and we are to-day using the same kind of net as then for our main instrument”.

I was amused, when reading on further, to find that my childhood gaffe of confusing a Malaise Trap with a net was fully justified. Malaise, later in the same paper writes, ”During my extensive travels I have repeatedly found that insects happened to enter my tent, and that they always accumulated at the ceiling-corners in vain efforts to escape at that place without paying any attention to the open tent door”. He then goes on to describe how he conjectured that “a trap made as invisible as possible and put up at a place where insect are wont to patrol back and forth, might catch them much better than any tent, and perhaps better than a man with a net, as a trap could catch them all the time, by night as by day, and never be forced to quit catching when it was best because dinner-time was at hand”.

He thus set about constructing a trap based on the idea of an open tent with a collecting device attached to the central end pole to take advantage of the fact that most insects when encountering an obstacle tend to fly upwards. On reaching the apex of the tent, the only way out is into the collecting device which is filled with a killing agent.  It is in effect, a flight intercept trap, but unlike window traps (subject of a later post), the insects instead of falling into a collecting device, head upwards and collect themselves. Malaise tested his first version of the trap on an expedition to Burma and found them to be a great success “every day’s catch from the traps grew larger and larger, and sorting it required more and more time”. He found the traps particularly good for Diptera and Hymenoptera but also very good for Coleoptera and Noctuid and Sphingid moths.  He also mentions catching Hemiptera.

In outward form, the Malaise Trap has remained fairly unchanged since its invention. The first versions were apparently fairly heavy, having a brass insect collecting cylinder and also only had one opening.  Malaise recognised the disadvantages of the single entrance version and suggested in the 1937 paper that a bilateral model would be more effective.  These followed in due course. Modified versions using plastic cylinders and different netting material were  invented in the 1960s (Gressit & Gressit, 1962; Townes, 1962; Butler, 1965).  Townes’s paper gives a very detailed description of the construction and use of modified Malaise traps (90 pages) in contrast to Butler’s three page description of a cheap and cheerful version made from a modified bed-net.

Nowadays, entomologists world-wide, particularly Dipterists and Hymenopterists, use Malaise traps of various designs and colours, and cost.  In the UK they are available from commercial outlets at prices ranging from £60 to £165. They are extremely effective and we use them to collect insects for our practical classes in the Entomology MSc based at Harper Adams University.

    

Malaise trap in operation, Harper Adams University, Shropshire, UK.

 

References

Butler, G.D. 91965) A modified Malaise insect trap. The Pan-Pacific Entomologist, 41, 51-53

Gressitt, J.L. & Gressitt, M.K. (1962) An improved Malaise Trap. Pacific Insects, 4, 87-90

Malaise, R. (1937) A new insect-trap.  Entomologisk Tidskrift, Stockholm, 58, 148-160

Townes, H. (1962) Design for a Malaise trap. Proceedings of the Entomological Society of  Washington, 64, 162-253

 

Post script

Malaise was not just an entomologist; he was an explorer and a passionate believer in the existence of Atlantis. A detailed biography of this extraordinary character can be found here, including a photograph of the original Malaise trap.

 

Post post script

I was amused to find in the 1949 edition of Instructions for Collectors No. 4a, Insects (Smart, 1949), this somewhat dismissive comment about the Malaise Trap “It is a very novel idea and captures large numbers of insects, but as at present designed is rather cumbersome, and since its design will probably be modified with experience it is not described here“

Entomological classics – The insect olfactometer

In 1924, Norman McIndoo (1881-1956) an entomologist based at the Fruit Insect Investigation Department in the USDA Bureau of Entomology in Washington DC was instructed by his boss Dr. A.L. Quaintance, to make a study of insect repellents and attractants.   After two years of frustrated experimentation McIndoo invented a piece of apparatus that would revolutionise the study of insect behaviour, the Y-tube olfactometer (McIndoo, 1926) . He freely admitted in his paper that he had borrowed the name from the Zwaademaker olfactometer (Zwaademaker, 1889) a device used to test the sense of smell in humans.  As you can see however, his apparatus bore no resemblance to that of Zwaademaker.

   

McIndoo ‘s apparatus was first used to find out whether Colorado potato beetles (Leptinotarsa decemlineata) responded to the odour of the potato plants. The beetles were placed in a dark bottle in a light-tight box, the bottle being attached to the stem of the Y-tube by a tube through which the beetles were able to move, at first being attracted to the light. Once they reached the junction of the Y they then had to make a choice between the two forks this time using their sense of smell. A pump was used to draw air from the two forks, one of which was connected to a jar containing a potato plant, the other which held the control substance. In theory, once at the fork the beetles were confronted with two streams of air, one smelling of potato, the other being odourless. McIndoo was indeed able to show that about 70% of the beetles responded positively to the odour produced by the potatoes. He also showed that the beetles responded to extracts made from the foliage of a number of different host plants.  He briefly mentions in the paper that the beetles were able to tell the opposite sex by smell and that the males would follow sexually mature females. He had accidentally discovered insect sex pheromones but did not realise it at the time. In the last part of his paper he provides data showing that other insect species, including Lepidoptera, were also able to respond to host plant odours.  The Y-tube olfactometer and the closely related T-tube olfactometers soon became the accepted way to test insect response to odours and are widely used in laboratories around the world to this day, for example http://weslaco.tamu.edu/research-programs/entomology/subtropical/behavior/ and http://sciencebykathy.wordpress.com/

http://openi.nlm.nih.gov/detailedresult.php?img=3422343_pone.0043607.g005&req=4

They do however have some limitations; there is a tendency for turbulence to occur at the junction of the Y- and T-tubes which means that there is some mixing of the test odours and this means that there is not a clearly delineated odour field into which the insects can enter, leave and re-enter if they so wish. In 1970, Jan Pettersson from the Swedish University of Agricultural Sciences at Uppsala, invented the four-way olfactometer with which to test the existence of a sex pheromone in the aphid Schizaphis borealis (Pettersson, 1970).

  

The four-way olfactometer provides a neutral central zone which is surrounded by four very distinct odour boundaries which the test insects can enter, sample the odour and then either stay or leave and move into another area of the apparatus. Louise Vet and colleagues (Vet et al., 1983) from the University of Leiden added some modifications to the original Pettersson version, with which to study the behaviour of aphids and their parasitoids.

 The four-way olfactometer, whether a Pettersson or Vet version, or a modification of the two, is now regarded as the ‘gold’ standard and is used very widely around the world.

http://www.nrcb.res.in/gallery8.html

It is certainly our research group’s favoured version and we use it for testing the responses of aphids, hymenopteran parasitoids, lepidoptera and beetles to a range of odours (Trewhella et al., 1997; Leahy et al., 2007; Pope et al., 2012). We are currently using mini-versions to test the olfactory responses of predatory mites. Watch this space.

References

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. http://onlinelibrary.wiley.com/doi/10.1111/j.1461-9563.2007.00328.x/full

McIndoo, N.E. (1926) An insect olfactometer. Journal of Economic Entomology, 19, 545-571

Pope, T.W., Girling, R.D., Staley, J.T., Trigodet, B., Wright, D.J., Leather, S.R., Van Emden, H.F., & Poppy, G.M. (2012) Effects of organic and conventional fertilizer treatments on host selection by the aphid parasitoid Diaeretiella rapae. Journal of Applied Entomology, 136, 445-455. http://onlinelibrary.wiley.com/doi/10.1111/j.1439-0418.2011.01667.x/full

Pettersson, J. (1970). An aphid sex attractant I Biological studies. Entomologia Scandinavica 1: 63-73.

Sanford, E.C. (1891) Laboratory course in physiological psychology. American Journal of Psychology, 4, 141-155, http://psychclassics.yorku.ca/Sanford/course2.htm

Trewhella, K.E., Leather, S.R., & Day, K.R. (1997) The effect of constitutive resistance in lodgepole pine (Pinus contorta) and Scots pine (P. sylvestris) on oviposition by three pine feeding herbivores. Bulletin of Entomological Research, 87, 81-88. http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=2497592

Vet, L.E.M., Van Lenteren, J.C., Heymans, M., & Meelis, E. (1983) An airflow olfactometer for measuring olfactory responses of hymenopterous parasitoids and other small insects. Physiological Entomology, 8, 97-106. http://onlinelibrary.wiley.com/doi/10.1111/j.1365-3032.1983.tb00338.x/abstract

Zwaademaker, H. (1889) On measurement of the sense of smell in clinical examination. The Lancet, 133, 1300-1302