Hungary and the EIG Annual Research Bursary

Some childhood experiences can be strangely formative – learning to ride a bike, losing a tooth, or perhaps facing the occasionally despicable nature of your fellow man full in the face for the first time. This happened for me on my first trip to Hungary aged 11, when our accommodation was broken into and my Ipod nano, and more importantly our flapjack was stolen. This cosmic episode stuck with me in the ensuing years, but so did the extraordinary wealth of wildlife (at the time, avian was the main interest)  is such seemingly commonplace settings – in gardens, little woods and all the farmland – here was a place that hadn’t been totally put under the plough in a post-war race to the bottom and still retained a large proportion of its seething, heaving mass of ancestral biodiversity – these thoughts lay dormant in my young mind, and as I became more interested in butterflies and moths, germinated into a cold and perhaps unrealistic certainty – I had to return.

And so it was that 8 years later I took up Butterfly Conservation’s European Interest Group Annual Research Bursary (http://www.bc-eig.org.uk/downloads/EIG_Annual_Research_Bursary.pdf) and did just that.  The grant is offered every year and put simply is both extraordinarily generous and extraordinarily helpful – up to £500 to hit the continent and study butterflies – any butterflies, anywhere in Europe, the opportunity of a lifetime. As one of this year’s lucky recipients, I used mine to head to the Ferto-Hansag National Park in the North-West of Hungary for 6 weeks this summer.

The park is around the Hungarian part of the Neusidlersee, the lake that straddles the border with Austria and whose Hungarian part is known as the ‘Ferto-lake’, and is of great importance for wetland birds (think droves of herons, White-Tailed Eagles, waders, terns, Moustached Warblers etc.), also taking in areas of classic Hungarian steppe habitat with species like Saker Falcon and Eastern Imperial Eagle (both satellite-tagged as part of EU LIFE projects and yielding fascinating results), and extensive areas of wet woodland and meadows in the Hansag region. For invertebrates the Ferto-Hansag perhaps lacks the reputation of some other areas of Hungary – the Zemplen Mountains and the Buuk Hills with their beasts from the east, and the chalk of the Balaton uplands with their Lesser Fiery Coppers and Clouded Yellows, but does have internationally important populations of a very special group of butterflies – the Maculineas.

Coming from Somerset, I’m already an acolyte of arion – the Large Blue – the winged victory of British butterfly lovers (returning after a re-introduction project, the most successful of its kind anywhere in the world). Those that know this butterfly well will be familiar with its extraordinarily convoluted life-cycle. Eggs are laid on the flower heads of Wild Thyme (or Marjoram at a couple of sites) and the resultant larvae hatch out and tear through all 4 of their instars in a couple of weeks, dropping off the flower-head in the evening to be adopted by red ants (specifically Myrmica sabuleti) which carry the larva as one of their own into the nest after ‘milking’ its honey gland. Here the larva goes rogue and will attack and eat the ant grubs, puncturing their skin and feeding on their bodily fluids, steadily growing to become a giant, implacable, pale imposter in the nest, before hatching out the following year to dance over the herb-rich slopes on midsummer days.

A Somerset arion

There are several more species of ‘Large Blue’ in Europe, 3,4 or even 5 depending on how you work your taxonomy, Dusky Large Blue (nausithous), Scarce Large Blue (teleius) and Alcon Blue (alcon) are dead certs, while some people produce Mountain Alcon Blue (rebeli) from Alcon and ‘Maculinea ligurica’ from Large Blue. Despite being some of the best studied European butterflies (as a result of their extraordinary life cycles and precipitous declines), a great deal of uncertainty still seems to remain about the classification of Maculineas. Rebeli for example, like alcon, is a cuckoo (feeding on regurgitations from ants rather than larvae, like arion does), but with a different host ant (Myrmica shencki as opposed to ruginodis), and on the whole tends to be slightly darker (something frequently seen in butterflies that fly at higher altitudes, arion does this too, with f. obscura described from the Swiss Alps), but some studies (like Als et al. 2004) suggest very little divergence from alcon. Ligurica too tends to be larger, paler and slower flying than arion, also flying later in the year, but again it’s difficult to be sure how distinct it is from its parent – Berecski et al. (2014) suggest that it’s a species in the making – showing differences in the structure of the wings and the male genitals, but only small ones in the genetics.

Scarce Large Blue

This is all rather complicated, but at least illustrates an important point about maculineas and indeed butterflies in general – like many insects, with their short generation times, they’re able to constantly adapt and evolve to a changing world – to paraphrase Matthew Oates – never underestimate a butterfly, you only have to look at the Quino Checkerspot’s extraordinary re-invention (https://www.theguardian.com/environment/2014/apr/07/endangered-butterfly-species-defies-climate-change-quino-checkerspot) to see the truth of these words. What this means is that the boundaries between forms/species can be even more blurred than they usually are, with fascinating evolutionary experiments happening right before our eyes.

The Ferto-Hansag's Maculineas - alcon, nausithous and teleius.

Anyway, in Hungary, I was helping with mark-release-recapture work on three species: Scarce, Dusky and Alcon (Large and ‘ligurica’ are found in other parts of Hungary too) around the Ferto Hansag from the 10^th July to the 20^th August.  These three species all favour damp grasslands (in contrast to arion’s herb rich slopes), where teleius and nausithous lay on the heads of Great Burnet (Sanguisorba officinalis), using young unopened and old, flowering heads respectively, while alcon scatter their eggs in what seems to be a fairly haphazard way over Marsh Gentians. Why the Maculineas lay in the ways they do seems to still be the subject of on-going debate, alcon appear to fire their eggs over the gentians with all the precision of a small machine gun bound to a couple of jackhammers, often 60 or 70 white pin-pricks on one rather small plant, but Van Dyck et al. (2000) reckoned they were cleverly laying them within the range of host ant (M. ruginodis) nests, gradually shifting to favour ones further away later in the season to reduce competition and cannibalism (the latter is rife amongst young Maculineas). This makes a lot of sense – a young larva only has one shot at adoption and has to be taken in by the correct host, otherwise its chances of survival are pretty low. Still, in order to get to the stage where it dupes innocent ants, a Maculinea larva has to find enough foodplant to get it through the early weeks. This means that the conclusions of Thomas and Elmes (2001) also make a lot of sense, deciding that egg-laying in all the European Maculinea species is governed by host plant phenology – the plants have to be the right age to provide food for the larvae.

Alcon ova scattered with gay abandon over a Marsh Gentian.

Clearly then,  an impasse has been reached, come forth then Patricellia et al. (2015) to change the game completely, deciding that neither conclusion is correct for M. arion, instead proposing an explanation that is in fact far more complex (and interesting!). When under stress, plants will often produce certain VOCs (volatile organic compounds) to try and turn the tables on their stressor. Marjoram and Thyme (the foodplants of arion), for example, produce Thymol and Carvacrol (both isomers of each other), when their roots are damaged by growing ant colonies. These are biocides that will kill the ants, thus saving the plant. Myrmica species (such as our, and arion’s friend, M. sabuleti) are able to upregulate genes to detoxify these compounds, causing the plant to produce them in ever higher concentrations in order to win the battle for survival. Thus, high concentrations of Carvacrol/Thymol around Marjoram/Thyme indicate the presence of Myrmica nests, and as you’ve probably guessed, are highly attractive to female Large Blues, they don’t detect the ants themselves, but a trophic interaction caused by them – intricate, and quite extraordinary!

Scarce and Dusky Large Blues

Back to Hungary then, where the three Maculineas are increasingly threatened by changing farming practices – mowing of meadows tends to happen in July, taking down the gentians and burnet, and with it Maculinea eggs, abandonment of Maculinea country (often already quite dense, tussocky grassland) quickly leads to scrub and no more Maculineas, and introduced Canadian Goldenrod (the cursed Soldago) has swept across Hungary and frequently takes over the damp meadows if not managed correctly. In other words, Maculineas are threatened by just about any change in either direction to their habitat, and their continued survival requires careful management and constant monitoring.

Scarce Large Blues (with a bonus Dusky)!

As it happens, both of these things are happening in the Ferto Hansag, and Maculineas are doing rather well. When I visited, they were having a boom year (there’s some evidence that their abundance is cyclical, following a similar pattern to Holly Blues and Marsh Fritillaries in the UK, though for different reasons) and numbers were very high at all four of the sites we worked at. In fact, we marked record numbers at several of them, with 2500 at one, and my personal tally coming to a respectable 2410 butterflies processed overall. The peak of lazy high summer days for any British butterfly enthusiast is an evening on the chalk, surrounding by little pale flags winking in the grass heads – a communal roost of butterflies is a hard sight to beat, and one indelibly associated with Chalkhill Blues in our butterflying psyche. They’re not infrequent with Common Blues too, and sometimes even Adonis if they’re feeling really enterprising – but Large Blues? No, surely not! To see communal roosts of 20, 30, 40 Maculineas in Hungary was quite mind-blowing, a sight so wrong, but so right, unquestionably ‘how it ought to be’, and a sight that will be one of the abiding memories of the trip.

Marking a Dusky Large Blue

This biblical abundance was no co-incidence, these are some of the strongest Maculinea colonies in Hungary (perhaps Europe as a whole), with much of it due to the hard work of the staff in the Ferto Hansag, constantly seeking to compromise, to balance the needs of the butterflies with the needs of a rapidly intensifying agricultural industry; allowing some areas to be mown for hay with refuges for the butterflies, allowing hunters to clear scrub and burn some areas in return for leaving others. Meeting the needs of these most capricious of butterflies in a fast-changing world is never easy, but it seems it can be done, with just the right amount of sensitivity, patience, and most importantly, hopeless devotion to the cause.

Scarce Large Blue

Andras Ambrus, the park’s resident lepidopterist, one of the most energetic people I’ve ever met (living proof of the adage that butterflies keep you young) with whom I was lucky enough to work, fits that bill perfectly, having carried out the MRR work on some of the sites for over 17 years. While I was struggling with my first words (sadly not ‘butterfly’), he was marking Maculineas, and has continued to do so ever since. This ongoing project provides vital data about population sizes (how are the butterflies responding to management, and wider changes in the countryside and climate?), dispersal (what’s their ability to deal with obstruction and colonise new areas like? It turns out nausithous is much more able to deal with tree-lines than teleius), and phenology (are the Maculineas, like other species, emerging earlier in response to a warming climate?), that in turn feeds into their conservation. Without people like Andras, and their work, young buttefliers like me wouldn’t be able to enjoy such rare butterflies in such extraordinary abundance, we owe them a great deal.

Dusky Large Blue

Not only do I owe a great deal to Andras, but also to EIG – I’ve already said that the Annual Research Bursary is the opportunity of a lifetime, but this cannot be overstated. Being able to spend every day for 6 weeks not only surrounded by Maculineas, but mythic, scarcely-imagined beings like Large Coppers, Lesser Purple Emperors and Common Gliders, seeing Eastern Imperial Eagles passing low over your garden, and Sakers diving on hapless kestrels just down the road – these are memories that will last forever, and are all because someone saw fit to reward my enthusiasm and an A4 page setting out my plans with a whole £500.

If you are a young person with an interest in nature reading this, the message is quite simple – apply!

Maculinea country

References 

Als, T., Vila, R., Kandul, N., Nash, D., Yen. S-H., Hsu, Y-F., Mignault, A., Boomsma, J., Pierce, N. (2004). The evolution of alternative parasitic life histories in large blue butterflies. Nature. 432 (7015), p.386.

Bereczki, J., Tóth, J., Sramkó, G., Varga, Z. (2014). Multilevel studies on the two phenological forms of Large Blue ( Maculinea arion ) (Lepidoptera: Lycaenidae). Journal of Zoological Systematics and Evolutionary Research. 52 (1), pp.32-43.

Hayes, M. (2015). The biology and ecology of the large blue butterfly Phengaris (Maculinea) arion : a review. Journal of Insect Conservation. 19 (6), pp.1037-1051.

Patricelli, D., Barbero, F., Occhipinti, A., Bertea, C., Bonelli, S., Casacci, L., Zebelo, S., Crocoll, C., Gershenzon, J., Maffei, M., Thomas, J., Balletto, E. (2015). Plant defences against ants provide a pathway to social parasitism in butterflies. Proceedings. Biological sciences. 282 (1811).

Thomas J. and Elmes G. (2001). Foodplant niche selection rather than the presence of ant nests explains oviposition patterns in the myrmecophilous butterfly genus Maculinea. Proceedings of the Royal Society B: Biological Sciences. 268 (1466), pp.471-477.

Van Dyck, H., Oostermeijer, J. G., Talloen, W., Feenstra, V., Hidde, A. V. D., Wynhoff, I. (2000). Does the presence of ant nests matter for oviposition to a specialized myrmecophilous Maculinea butterfly? Proceedings of the Royal Society B: Biological Sciences. 267 (1446), pp.861-866.

Larval Doings

After moaning in various forms about the hard time that winter represents for most butterfliers, I’ve now encountered another period of lean pickings, the March – April gap, when the first hibernators appear in early March, injecting the first rays of spring-like hope into the year, before promptly it is promptly removed as the next cold front rolls in from the West. This time of year is arguably more fraught than winter, winter at least brings with it the comforting certainty that there are no butterflies to see, no precious moments of summer to snatch, but in the March – April gap, you can’t help but feel you should be doing more to seek out those first Orange Tips (this is of course foolish, the first Orange Tips always find you), and so it is that a new season of frustration is born.

To combat this, like winter, I have developed a daring strategy to combat this, turning to those shadowy brethren, the immature stages, this year, those orange sports-car butterflies – the Fritillaries.

Somerset is not as blessed with Fritillaries as perhaps it should be, High Brown was last seen at Hurlstone Point in about 2000, victim of extremely challenging terrain for management, and a lack of resources, and Pearl – Bordered Fritillary on Exmoor, up at Mounsey in 1992, whilst Marsh Frit is on the ropes, maintaining a transient presence up on Exmoor, and it would seem, hiding out elsewhere, with the exciting report of a larval web in East Somerset this spring, it remains to be seen if they will re-appear at Ash Priors this spring (fingers and toes firmly crossed).

A 4th instar Heath Fritillary Larva at Haddon Hill - 21/03

Despite this, it’s not all doom and gloom, indeed, one of our trademark species is a Fritillary – the Heath Fritillary. This shouldn’t really be a trademark, a rarity, since it was once a common enough species to earn its own nickname – ‘The Woodman’s Friend’, thanks to its dependence on managed woodland, where it breeds in areas of coppice, typically peaking 2-4 years after coppicing and then tailing off and having to move on to new areas. In coppiced woodland Heath Frits breed on Cow Wheat, laying eggs in small batches (averaging about 40) on and around the foodplant, typically growing in sunnier areas, often over bare ground. The key here – open coppice, sunny spots, reflective bare ground is heat – this butterfly is one of some that like it hot, and can’t survive without it, and so the rapid decline in coppicing sounded the death knell for most of its populations at the turn of the 20^th century, leaving it isolated in Kent (the East Blean Woods complex), and on the Devon-Cornwall Border. This meant then, that its re-discovery on Exmoor on 1984 was one of the great butterflying events of the last century, up there with Fort William’s Chequered Skippers, when strong colonies were found completely defying convention and flying between 200-400m (no other British colonies are above 100m) on moorland edges – they make ‘em hard in Somerset. This was at a time when the loss of the Large Blue had galvanised conservationists, and intensive study by Martin Warren, and some timely management changes ensured its future, though not before a rocky period on Exmoor in the 90s. Now it flies in quite a few areas on Exmoor, and most of its old 70s haunts, as well as several Essex re-introduction sites. Interestingly, its foodplants at the Devon and Cornwall sites (where it flies in old hay meadows and railway cuttings) are Germander Speedwell and Ribwort Plantain – this catholic foodplant choice seems to preclude rarity, but it underlines the key point about Heath Frits – it is the quality of the foodplant, the heat, that is key.

Nowadays a future for this butterfly in our fair isle looks relatively assured (though recent UKBMS results indicate it has declined by 82% over the last decade), with BC’s Two Moors butterfly project just finished, and currently its All the Moor project helping it along on Exmoor (with lots of great management by the National Trust’s Holnicote estate), and I’m lucky enough to have a thriving colony up the road at Haddon Hill, and so it was that I paid them a visit on the 21^st of March. The larvae hatch from their egg batches after a couple of weeks, and feed companionably together in a small web before dispersing in their second instar, and in September, in their third instar, spinning up the edges of dead leaves near the ground and moulting into their fourth instar and going into diapause. They emerge, phoenix-like the following year, basking on dead leaves as their foodplant germinates around them, often in friendly little groups. These larvae were my target on my visit, and after about 20 minutes searching, I struck gold, 2 singletons, and a happy trio all in a small area of bilberry at the bottom of the slope where they fly (about 6 metres from the wood edge, so nice and sunny) where the Cow Wheat was just getting going. They were laid out on dead Birch leaves on the moss through which the plants were growing, on the edge of a trampled Pony track (providing a sheltered little ‘valley’ for them) – sheltered, sunny, like everything with the Heath Frits – it screams heat. These weren’t up to much, and didn’t move in the hour I watched them – living the good life evidently.

4th Instar Heath Frit larvae at Haddon Hill (21/03), singleton in the left circle, and trio in the right
note the warm dead leaves, relatively open bilberry growing through moss (where Cow Wheat is germinating)
and the small path (foreground) creating a sheltered little valley.



A single 4th Instar Heath Fritillary larva on the circled leaf - a warm, open mossy hollow, on 11/04 about 20 larvae were here.

Impressed by these spiny little creatures, I decided to return to see them in their magnificent final form (6^th instar) later on, I made good on this on the 11^th April and was glad I did, for a larval horde awaited, with 36 counted lounging about amongst the Cow Wheat in 45 minutes of not especially intensive searching/photography. When a caterpillar is so common you have to watch your feet, it is a butterfly unashamedly thriving – and it’s good to see. Once again they were all intent on getting warm, basking either on the moss through which the foodplant was germinating, or once more, the dead leaves. I think the key to my lucky strike probably had a lot to do with the area I searched, one of the most open patches of the site, with the largest quantity of these warm moss mats, as opposed to denser areas of bilberry where the cow-wheat is closed out and the ground shaded and things are less suitable for the larvae – heat, heat, heat! I haven’t yet decided what instar these larvae are – they look very similar to the fourth instar larvae of the month before, but with more white spotting (is this just more obvious because of their larger size?) – I’d be interested if someone knows.

A Heath Fritillary larval horde, the two on the right are on a small Cow Wheat seedling (you can see some nibbling to the left of the middle one) - where there is one, there are often more!

Heath Fritillary larva at Haddon Hill 11/04 - not too sure what instar

Heath Fritillary larva basking at Haddon Hill 11/04

This area had about 30 of the larvae I found - note how much more open and mossy it is than the surrounding area, at the bottom of the slope - sheltered and warm, Cow Wheat was germinating in large quantities.



The same area from the other side.

Heath Frits were not the only larval target this spring. More to follow!

Alpine Amble - Part 3

After a reasonable attempt at spring the over the last few days that has seen me clock up all the 5 normal hibernators, the weather has turned and Lepidoptera once more seem a rather distant and frivolous summer thing, and as such, it’s time to burn the swiss midnight oil afresh.

BVW roosting.

The ‘Whites’ is a bit of an amorphous term that I tend to apply fairly freely to anything in the first 80 pages of Tolman & Lewington, hardly a monophyletic group, but convenient. This includes Swallowtails, Apollos, Festoons, Brimstones, Clouded Yellows, and then at some point actual whites. Over the course of the walk, we notched up 17 species in this fairly arbitrary group, with the commonest probably Black-Veined.

Acrobatic copulation from BVWs

Despite their relative abundance, I always find these butterflies strangely thrilling, a little piece of Old England flying by on their papery wings. Black-Veined Whites became extinct in the UK in the 1920s, with the failing of the last Kentish populations, a string of releases has followed since, famously by E. H. Newman in the grounds of Churchill’s Chartwell estate, and most recently, a clandestine introduction to Stockbridge Down resulting in trampling of some of the site’s key habitats by ardent photographers – a cautionary tale about the use of such releases if ever there was one. The causes of extinction are poorly known – Colin Pratt’s ‘Modern review of the Demise of A. crataegi L., the Black-Veined White’ notes a string of wet Septembers tended to precede extinctions on most sites, and the cries of many frustrated breeders of the time who suggested an increase in passerine populations (due to shotgun innovations=fewer raptors=more blue tits=more predation of larval webs and pupae). In essence, it’s far from an open and shut case.

Mountain Green-Veined White at roost.

Following the veined motif, Mountain Green-Veined Whites became one of the commonest butterflies at higher altitudes, and despite being very active in sunny conditions, were relatively easy to find and photograph when at roost on flowerheads. These differ from ‘our’ Green-Veined Whites by having heavier, greyish streaking, and often more of a dark suffusion. Interestingly, they have this in common with Scottish, thomsoni Green-Veined Whites – a nice bit of convergent evolution, adapting to their common struggle against frequent colder spells.

Having looked back through my photos from the trip, it’s clear I didn’t pay perhaps as much attention to the whites as I should have, with only a few photos of Mountain Clouded Yellows and their kin, probably because most of it was lavished on one particular set of broad white wings – Apollos. These are arguably one of Europe’s most impressive butterflies – ardent brits might choose the Purple Emperor for its dazzling iridescence, and aloof, regal nature, yet Apollos are amazing in a different kind of way: ethereal, seemingly detached, gliding past on wings so large and papery it’s possible to locate them as they soar past the back of your head, purely on the basis of the rustling they make as they move, and for such a butterfly, an iconic setting is a must, so snow-capped peaks and sound of music meadows it is – they, like most alpine butterflies, are a sight it’s hard not to be seriously impressed by.