Wallace, Darwin and the night shift

Last week I had the great honour of attending the Royal Entomological Society’s annual conference, this year hosted by Edge Hill University, to accept the 2017 Alfred Russel Wallace Award. The award is given for the PhD thesis “judged to make the most significant contribution to entomology in the year”, and after several rounds of selection my own thesis, “The role of moths as pollinators, and the effects of environmental change”, was chosen. At the conference, I said that I was particularly proud to receive an award in Wallace’s name, because of his own connections to the subject matter of my thesis – nocturnal pollination by moths. In this blog, I re-tell the story of that connection…

_______

It is 25th January, 1862, and Charles Darwin is writing to his friend and confidante Joseph Dalton Hooker. Earlier that same day, Darwin had taken delivery of a box of specimens of exotic orchids from James Bateman, a Staffordshire-based orchid specialist.1 Among these, Darwin found one in particular (Angraecum sesquipedale, a species from Madagascar) to be remarkable, due to the great length of the nectary, extending 11.5 inches below the flower. “Good Heavens”, wrote Darwin, “what insect can suck it”.

Angraecum_sesquipedale.png
(L-R) a drawing of Angraecum sesquipedale by W.H. Fitch, included in: James Bateman (1876) A second century of orchidaceous plants. Curtis’s Botanical Magazine; two flowers of A. sesquipedale (© Michael Wolf | Wikimedia Commons | CC BY-SA 3.0); a plate commissioned by A.R. Wallace depicting the hypothetical hawkmoth pollinator of A. sesquipedale, drawn by T.W. Wood and included in: Alfred Russel Wallace (1867) Creation by Law. The Quarterly Journal of Science, 477.

Darwin must have acted quickly. In the same letter to Hooker he had described his “infinite satisfaction” at the prospect of correcting the proofs of his next books, Fertilisation of Orchids, “in 2 or 3 weeks”. Yet when Orchids emerged less than four months later on 15th May, Darwin had revised the text, incorporating a discussion of A. sesquipedale and a rough hypothesis as to its insect pollinator. Reasoning that the longest tongues among English insects belonged to hawkmoths (family Sphingidae), he wrote:

in Madagascar there must be moths with probosces [sic] capable of extension to a length of between ten and eleven inches

Darwin’s prediction sparked debate; among those who sprung to his defence was Alfred Russel Wallace. Contrary to popular opinion, correspondence between Darwin and Wallace was friendly and reasonably frequent after they jointly proposed evolution by natural selection (van Wyhe & Rookmaaker 2015). Five years after the publication of Orchids, Wallace’s Creation by Law asserted not only that Darwin’s predicted moth could safely be assumed to exist, but further, that naturalists visiting Madagascar “should search for it with as much confidence as astronomers searched for the planet Neptune”. Wallace picked out one moth in particular to back up this statement – Xanthopan morganii, then known from tropical Africa (as Macrosila morganii), which had an exceptionally long proboscis.

Xanthopan_morgani.png
Xanthopan morgani; a specimen from continental Africa. Malagasy X. m. praedicta have proboscides several inches longer still!

Several decades later, Walter Rothschild (2nd Baron Rothschild) and Karl Jordan described a newly-discovered subspecies of X. morganii, from Madagascar. Remarkably, the subspecies had a tongue that was, by several inches, longer than that of its continental cousins. In honour of the predictions made by first Darwin and later Wallace, the new subspecies was named X. morganii praedicta.

DarwinWallaceRothschild
(L-R) Charles Darwin, Alfred Russel Wallace, and Walter Rothschild

At this point, the trail went cold for the best part of the 20th century. We had the flower, and we had found the moth with a proboscis to match it, but the final proof of their association was lacking because nobody had ever seen the two interact. That state of affairs lasted until the early 1990s, when Professor L.T. Wasserthal (then of the Friedrich-Alexander University Erlangen-Nuremburg) finally photographed, beyond doubt, the pollinator of A. sesquipedale. It was, indeed, X. morgani.

BotanicaActa
Making the front cover, a photograph of Xanthopan morgani praedicta visiting the flower of Angraecum sesquipedale finally resolves the mystery!

As a final chapter, the video below is well worth a watch: both for its remarkable footage, shot in 2004 by Dr Philip DeVries of the University of New Orleans, of this pollination interaction in action, and for the sheer joy and excitement shown by Dr DeVries and his colleagues when their hard work finally paid off!

If you want to learn more about this story, I owe a debt of gratitude to the review of Arditti and colleagues (2012), on which I drew heavily whilst researching this blog.

1: The relationship between Darwin and Bateman must have been complex. Bateman was, at this time, one of Britain’s foremost experts on, and cultivators of, orchids, and it is hard not to assume that he and Darwin corresponded while the latter prepared Fertilisation of Orchids. However, this box of specimens, and a brief subsequent exchange of letters, appears to be the only surviving evidence of direct interaction between the two (Darwin Correspondence Project). Bateman lived at Biddulph Grange in Staffordshire, designing (with his wife Maria) its famous landscape gardens, which are now owned by the National Trust. The gardens featured a Geological Gallery, seen as a direct attempt to refute Darwin’s theory of evolution by natural selection by drawing connections between evidence from the fossil record and the account of creation given in the book of Genesis.

References

Arditti, J., Elliot, J., Kitching, I.J. & Wasserthal, L.T. (2012) ‘Good Heavens what insect can suck it’–Charles Darwin, Angraecum sesquipedale and Xanthopan morganii praedicta. Botanical Journal of the Linnean Society169, 403-432.

Darwin Correspondence Project, “Letter no. 3356,” accessed on 3 April 2018, http://www.darwinproject.ac.uk/DCP-LETT-3356. Also published in The Correspondence of Charles Darwin, vol. 10.

Van Wyhe, J. & Rookmaaker, K. (2015) Alfred Russel Wallace: Letters from the Malay Archipelago. Oxford University Press, Oxford.

Channel-hopping moth plague: our clothes are safe, but mind the cabbages

Callum Macgregor, Newcastle University

The next great plague has descended, and this time it’s moths. Diamondback moths have been crossing the English Channel over the past fortnight in far greater numbers than usual.

Those of you reaching for moth-balls and carpet sprays, stop. There are only two species of British moth (from a cast of thousands) that damage clothes, and the diamondback is not one of them. You, the reader, really don’t need to worry – unless you grow cruciferous vegetables like cabbages or broccoli.

Plutella xylostella – more commonly known as the diamondback – is one of the world’s most widespread and prevalent agricultural pests; so widespread, indeed, that scientists disagree as to where exactly its native range lies.

Their success belongs in part to their preference as caterpillars for eating plants in the Brassicaceae family. This amazing group of crops (notably the species Brassica oleracea) has been selectively bred over hundreds of years to make use of almost every part of the plant, and now contains many popular vegetables such as cabbages, broccoli, Brussels sprouts and turnips, as well as oil-seed rape. The moths lay eggs on brassica leaves; a week later, caterpillars hatch and start eating. An infested plant can quickly be reduced to little more than leaf stems.

Cross-Channel travellers

Despite being one of the smallest moth species to occur in the UK, with a wingspan of under 15mm, diamondbacks are capable of dispersing vast distances, using the wind to make up for their small size. Research has shown that without assistance, diamondbacks are only capable of flying 35 metres – yet when carried along by the wind, they may travel hundreds of miles in a single day.

It is this ability to disperse that leads it to British shores. Every summer, adults cross the Channel and attempt to breed in the UK. A number of butterfly and moth species employ this strategy. In some cases, such as the Painted Lady butterfly, the offspring will return south ahead of the winter. For others, including diamondbacks, the offspring will attempt to remain, but British winters are typically too cold for them to survive.

What’s for dinner? Cabbage, broccoli and Brussels sprouts.
Yuangeng Zhang / shutterstock

For the moths, these fatalities are an unfortunate side effect of a highly dispersive life history; by having offspring constantly travelling beyond the edge of their range and attempting to breed, the moths ensure that no available space is wasted. Should some of the dispersing moths find a new area that is suitable for breeding success, they will benefit enormously from a lack of competition with their peers and, potentially, the absence of specialised predators and parasites. Thus, evolution favours and preserves these intrepid explorers.

Supermoth?

And it is once the moths begin to breed that the problems begin for farmers and gardeners, because these are not your average pest. Diamondback has been referred to in some coverage as a “super-moth”. It’s a title it certainly deserves.

The species has proven itself to be extraordinarily adept at evolving resistance to pesticides. Wild populations have developed significant resistance to almost every significant synthetic insecticide in use, including one of the main classes, pyrethroids.

Perhaps more impressively, diamondback was the first insect recorded to have developed resistance in the field to the bacterial pesticide Bacillus thuringiensis; only one other species is known to have managed this since (the “cabbage looper”, a pest moth named after its favourite food). Interestingly, this has led scientists to target it with the same genetic tools as are employed against the mosquitoes that transmit dengue fever and Zika virus. Field trials are planned in the US for this summer.

Tough times on the veg patch

In the short-term, perhaps. The influx of moths will soon begin to produce larvae, and these may cause substantial damage to crops where they are able to escape death by insecticide. The damage will be most severe to farmers of leaf crops, such as cabbages or kale, as the holes left behind in leaves by caterpillars make the crop unsellable.

Diamondback caterpillars leave cabbage unappetising and unsellable.
Rothamsted Research, CC BY

Looking at the longer term, this year’s invasion will die out over the winter. Next year there will be another, but this is nothing new: these moths come to the UK every year in substantial numbers. I spent the summer of 2014 collecting moths on farms in Oxfordshire for a study on the effects of light pollution on moths. Out of the 203 species of moth I recorded, diamondback was the 26th most common, forming around 1% of my total catch.

Britain’s cabbage-growers have been dealing with diamondback for years and have managed to persist thus far. They may have a hard summer ahead of them but it is not without hope. Although globally diamondbacks have developed resistance to almost all pesticides, most populations are only resistant to a subset of these, and scientists at Rothamsted Research are already studying the current influx to identify their weaknesses. Armed with the results of that research, farmers may be capable of restricting the damage.

The Conversation

Callum Macgregor, PhD Candidate, Newcastle University

This article was originally published on The Conversation. Read the original article.