Black-rimmed snout hoverfly: the Heineken Fly!

Rhingia campestris Meigen, 1822 showing snout close-up
Rhingia campestris Meigen, 1822 inserting proboscis into Primula flower

The Black-rimmed snout hoverfly, Rhingia campestris Meigen, 1822 (Diptera: Syrphidae) is a common and widespread fly which is often seen visiting flowers or resting on nearby vegetation. The larvae live and develop in cow dung, whilst the adults feed on nectar and pollen. Pollen is required by the females as a protein source for egg development; nectar is needed by both sexes, and as such they are important pollinators of flowers, specialising in species such as bugle (Ajuga reptans), red campion (Silene dioica) and spiked rampion (Phyteuma spicatum). (1, 2).

Rhingia campestris Meigen, 1822 with proboscis inserted into corolla of Primula flower

This distinctive fly has a long, stubby, duck-billed shaped, snout and an extendable proboscis, as long as its body (7–11 mm). It also appears to have been given the name ‘Heineken fly’, after an advert on UK television, because it can supposedly reach the parts of a flower that other hoverflies cannot! I prefer the traditional name: Black-rimmed snout hoverfly. The black rims refer to the black margins of the tergites: the sclerotised plates on the dorsal (upper) sides of each abdominal segment. The similar, conspecific species, R. rostrata, lacks the black margins.

Rhingia campestris cleaning proboscis. Note black-margined tergites on abdomen.

An organ, if that’s what it is, of this complexity and functional importance to the fly must be cleaned and looked after. The proboscis, when not in use, is folded up and stored within the snout, as can be seen in the very high magnification photograph in link number (3). I wonder if anybody has studied the proboscis in detail and worked out exactly how it is extended and folded up? The following photograph is not very good, but is the best one I have of the extended proboscis in side view. The proboscis is clearly a complex structure and one that must have evolved to allow the fly to access deep nectar sources.

Rhingia campestris Meigen, 1822 with proboscis extended
  1. Haslett, J.R. Oecologia (1989) 81: 361. doi:10.1007/BF00377084
  2. Kooi, C. J., Pen, I., Staal, M., Stavenga, D. G., & Elzenga, J. T. M. (2016). Competition for pollinators and intra‐communal spectral dissimilarity of flowers. Plant Biology, 18(1), 56-62.
  3. http://www.microscopy-uk.org.uk/micropolitan/insectportrait/frame5.html

Oak apple galls

Oak apple gall caused by cynipid wasp, Biorhiza pallida

The leaves have only just started to open on this oak tree, a Sessile oak I think, yet it is already covered by many galls. These rounded disfigurations – called Oak apples – are caused by a tiny (5-6 mm) wasp in the family Cynipidae, called Biorhiza pallida. 

Freshly emerging foliage on oak free, 9th April 2017

It is known that the galls are caused by the injection of venom by the wingless, parthenogenetic females, which cause the newly emerged leaves to soften and swell up. These females have emerged from galls growing underground, on the roots, and they have crawled up the tree to start a new generation in the Spring. (1) The eggs hatch and the larvae secrete chemical substances which also cause the tissues to grow and form into a ball; the apple gall.

Oak apple gall caused by the cynipid wasp, Biorhiza pallida

Remarkably, all of the individual wasps developing within a given gall, of which there may be as many as thirty, are of the same sex. (2) Although the gall is made of plant material, because it is induced by the wasp it is said to represent the extended phenotype of gall-wasp genes (Stone and Cook, 1998). (3)

Oak apple galls higher up the tree

The tree was located near the Felmersham Gravel Pits, a Site of Special Scientific Interest between the villages of Felmersham and Sharnbrook, in Bedfordshire.

Oak tree with Oak apple galls, Bedfordshire

The life cycle of these amazing wasps is even more complex than I have outlined here, with individual asexual females able to produce both males and females from unfertilised eggs; alternating sexual and asexual generations and way of life that utilities both the below-ground roots and above-ground shoots of the tree.

  1. https://en.wikipedia.org/wiki/Biorhiza_pallida
  2. http://www.naturespot.org.uk/species/oak-apple-gall-wasp
  3. http://stonegroup.icapb.ed.ac.uk/publications/gallwasps.html

Stabilmenta: spider’s web decorations

Stabilmentum woven by unknown spider, possibly in the genus Cyclosa

Stabilmenta are conspicuous patterns or decorations made by spiders – particularly orb-web spiders – in their webs. Google ‘stabilmenta’ (singular: stabilmentum) and you will see many wonderful examples of these structures, including crosses, spirals, zigzags and so on.

Stabilmentum made by unknown spider species, possibly in the genus Cyclosa

There are a number of different theories as to why spiders make these structures, including: to attract prey; as camouflage; as a moulting platform to stand on; as a way of warming up the web; and as a warning signal for any potential predators which might want to, or just inadvertently, destroy the web (1, 2, 3). It is possible that they have more than one function, although camouflage seems to be the most popular, or agreed upon, theory (2, 4). Nevertheless, some researchers have shown that more flying insects (apart from grasshoppers) are caught, or intercepted, on webs decorated with stabilimenta (5). Which suggest that they might enhance the efficiency of the web; although other researchers came up with a completely different result (see below).

Circular stabilmentum, possible by a Cyclosa species

Spiders in the Araneid spider-genus Argiope often adorn their webs with these structures. I photographed this stabilmentum (below) made by Argiope pulchella in Thailand. The spider positioned itself over the X-shaped stabilmentum, but moved off it to wrap-up any prey caught in the web.

Argiope pulchella on web with stabilmentum

Some experiments have shown that stabilimentum building is a defensive behavior (3), in effect advertising the presence of the spider’s web and preventing birds from flying through the webs. There is no question that they are highly visible and in some situations, actually reduce the number of prey that are caught (3). This ‘cost’ to the spider can presumably be set against the benefit of not having to rebuild the nest every time a bird flies through it by mistake! Unfortunately for the spider making the stabilmentum, other predatory spiders – such as web-invading jumping spiders – have learnt to recognise the patterns and use them to find their prey (6). Perhaps this is why some spider species make silk replicas of themselves! (7). To fool would-be predators! (8)

Argiope pulchella on web with stabilmentum

As many people may have noticed, spiders webs can be highly visible when covered in dew in the morning, or after a rain shower. I photographed this spider’s web in Spain, after a passing shower.

Water droplets on spider’s web after rain

I agree with another blogger (9), that stabilimenta are probably multi-functional structures, and the fact that they are so common in certain species, must mean that they are being selected by evolution. So the overall benefits must out-weigh the costs.

  1. http://www.mol-ecol.uni-halle.de/research/former_topics/stabilimenta/
  2. Cloudsley-Thompson, J. L. (1995). A review of the anti-predator devices of spiders. Bulletin of the british arachnological society, 10(3), 81-96.
  3. Blackledge, T. A., & Wenzel, J. W. (1999). Do stabilimenta in orb webs attract prey or defend spiders?. Behavioral Ecology, 10(4), 372-376.
  4. https://en.wikipedia.org/wiki/Web_decoration
  5. TSO, I. M. (1996). Stabilimentum of the garden spider Argiope trifasciata: a possible prey attractant. Animal Behaviour, 52(1), 183-191.
  6. Seah, W. K., & Li, D. (2001). Stabilimenta attract unwelcome predators to orb–webs. Proceedings of the Royal Society of London B: Biological Sciences, 268(1476), 1553-1558.
  7. https://www.wired.com/2012/12/spider-building-spider/
  8. http://www.popsci.com/article/science/what-i-learned-hunting-amazonian-spiders-weave-fake-spiders
  9. http://www.bugsinthenews.com/stabilimentum_and_some_notions_on%20function.htm

 

Bright iridescent patches are honest signals!

Purple Sapphire (Heliophorus epicles) male showing iridescent blue patches on upper wing surfaces

Males butterflies in the family Lycaenidae, the so-called Blues, typically have brightly coloured, iridescent colours on the upper (dorsal) surfaces of their wings. Vivid blue iridescence such as this on the Purple Sapphire (Heliophorus epicles) shown here, is usually to do with courtship and mate recognition.

The brightly coloured, iridescent males rely on so-called, structural colouration (described below), which is used both in male-to-male interactions (competition), and in attracting females, via flickering or flashing their bright wings. The females are often dark brown and mostly lacking in these bright structural colours. They may – like female Purple Sapphires – have bright pigmentary colours (orange flashes in this case), but these are probably not secondary sexual characters, i.e. used in courtship and mating. I don’t have a picture of the female, but there are many examples on this website (1).

Purple Sapphire (Heliophorus epicles) side view showing brightly coloured, but mostly, non-iridescent under wings

A variety of different types of microscopic ‘nanostructures’ – extremely small regular structures – have been found to generate blue colours in lycaenid butterflies. Many have so-called multilayers – alternating layers of chitin and air – within the individual scales (2, 3).

Purple Sapphire (Heliophorus epicles) male showing iridescent blue patches on upper wing surfaces and antennae

Butterfly wings are covered on both sides by rows of tiny overlapping scales, a bit like very thin, flat roof tiles or shingles. Scales can vary markedly in size and shape across the wing of a butterfly, but depending on the species, there are about 200–600 scales per square millimetre of wing. The scales are very delicate, typically one or two microns (i.e. one thousand times smaller than a millimetre) in thickness, and are denuded by wear and tear as butterflies age.

It has been suggested that the fact that scales detach so easily is an adaptation to allow butterflies (and moths) to escape from spider’s webs. (4). Scales that are attached to the sticky threads of the spider’s web can be sacrificed to allow the butterfly to regain its freedom.  

Each scale consists of two layers held together by a series of tiny pillars. The lower layer of the scale is flat and smooth between 100 to 200 nanometres (one nanometre is a billionth of a metre) in thickness – whilst the upper layer consists of a series of longitudinal ridges or striae – about one or two microns apart – and transverse crossribs which create a three dimensional lattice, or honeycomb structure with windows into the interior of the scale (5). It is the elaborate 3-D nanostructures so-called perforated multilayers – between the lamellae that cause the structural colours and phenomena like iridescence (3).

The reflected iridescence produced by light scattering from the dorsal wing scales of many lycaenids is highly directional, i.e. it is only observable from a narrow angular window. That is why the blue colour is not visible in some photographs (see below), although the scales can also be denuded.

Purple Sapphire (Heliophorus epicles) male showing no iridescent blue patches, probably due to the angle of the wing

The iridescence produced by male wings of butterflies such Heliophorus epicles, and countless other species, appears to be what is called a secondary sexual character. In other words, female butterflies evaluate these colours when choosing which males to mate with. They have also been called ‘colour badges’ and are thought to be honest signals, or reliable information if you will, of the condition of the males (6).  So the theory is that males with a good pedigree (i.e. genes) and a good upbringing (i.e. favourable environmental conditions) will be bright and showy (!), and females will choose them on the basis that they are more likely to be vigorous and fertile.

Presumably because they are ‘costly’ to produce or difficult to generate, and the scales producing the effect are lost, or worn down as the male butterflies age, then structural colours appear to provide a good indication of male quality and vigour in some species. However, even old and worn males – like the individual shown in the following photograph – still have some iridescent scales with which to attract the ladies!

Purple Sapphire (Heliophorus epicles) male showing worn iridescent blue patches

Although there is, as far as I know, no definitive evidence that female butterflies choose between males on the basis of the quality of the intensity, hue or saturation of their reflective colours, the available evidence supports the idea that brilliant male structural colours evolved as a result of sexual selection (7). It seems that sexual selection in butterflies has homed in on the brightness of these structural colours in the same way that it has in terms of the brightness and ornamentation of the peacock’s tail feathers.

I have focused on the blue patches on the upper sides of the males wings in this blog. The bright yellow and red colours on the undersides also clearly have some function, but it is probably not to do with mating (I’m only guessing!) as the males and females look relatively similar on their undersides. Who knows what really goes on in the minds of these butterflies!

Purple Sapphire (Heliophorus epicles) side view showing brightly coloured, but mostly, non-iridescent under wings

All of these photographs were taken in Thailand.

  1. Mazumder, S. 2017. Heliophorus epicles Godart, 1823 – Purple Sapphire. Kunte, K., P. Roy, S. Kalesh and U. Kodandaramaiah (eds.). Butterflies of India, v. 2.24. Indian Foundation for Butterflies.
    http://www.ifoundbutterflies.org/sp/728/Heliophorus-epicles
  2. Vértesy, Z., Bálint, Z., Kertész, K., Vigneron, J. P., Lousse, V., & Biró, L. P. (2006). Wing scale microstructures and nanostructures in butterflies − natural photonic crystals.Journal of microscopy,224(1), 108-110. 
  3. Wilts, B. D., Leertouwer, H. L., & Stavenga, D. G. (2008). Imaging scatterometry and microspectrophotometry of lycaenid butterfly wing scales with perforated multilayers.Journal of The Royal Society Interface, rsif-2008. 
  4. Eisner, T., Alsop, R., & Ettershank, G. (1964). Adhesiveness of spider silk.Science,146(3647), 1058-1061.
  5. Stavenga, D. G. (2014). Thin film and multilayer optics cause structural colors of many insects and birds.Materials Today: Proceedings,1, 109-121.
  6. Kemp, D. J. (2006). Heightened phenotypic variation and age-based fading of ultraviolet butterfly wing coloration. Evolutionary Ecology Research, 8(3), 515-527.
  7. Kemp, D. J., Vukusic, P., & Rutowski, R. L. (2006). Stress‐mediated covariance between nano‐structural architecture and ultraviolet butterfly coloration. Functional Ecology, 20(2), 282-289.

 

 

 

 

Red Admirals – European migrants

Red Admiral (Vanessa atlanata) dorsal side. Galicia, Spain. 11 June.

Migrant Red Admirals Vanessa atalanta (L.), usually arrive in the UK during May and June each year. Like the closely related butterfly, The Painted Lady, Vanessa cardui (L.), these migrations of Red admirals originate from countries around the Mediterranean – possibly as far south as the North African coast. (2) The butterflies fly north on southerly winds to feed on new growth as it becomes available in the Spring (1).

The Painted Lady, Vanessa cardui (L.) on Aesculus californica. Barcelona, Spain. 6th June.

Most European Red Admirals  migrate north in the Spring and – after producing a new generation – migrate south again in the Autumn. (3)  This seasonal movement appears to occur right across Europe and western Asia, although this still needs confirmation from many regions, with waves of migrants moving north, for example up into Finland, northern Norway and northern Russia. (4, 5, 9, 10).

Red Admiral (Vanessa atlanata) feeding on thistle. Galicia, Spain, 11 June.

Red Admirals arriving in the UK, mate and lay their eggs mainly on stinging nettles (Urtica diocia); a new generation emerges sometime over the period, August to October.  A small number of Red Admirals remain to overwinter in the British Isles (mainly in southern England) – although numbers appear to be increasing with climate change – whilst the majority elect to migrate. (3) How does this choice to migrate or not work in practice? “Should I stay or should I go now”?! (6). Perhaps a small proportion of the population are genetically programmed not to migrate?

The Red Admiral (Vanessa atalanta) on bell heather. Galicia, Spain, 28th August.

Of those individuals that remain in the UK, it is not thought that they hibernate in a physiological sense, although many sites state that they do hibernate, I think it is true to say that they merely remain dormant, since they can become active on sunny days throughout the winter. (5)  Some of these remaining butterflies must mate in the autumn, as there are records of V. atalanta larvae developing slowly over winter. In other words, a second generation gradually develops over the period from autumn until the following spring. This is exactly what happens when the migrants arrive back in Spain in October and early November as well; ‘larval development occurs throughout the winter until a first annual generation of adults appears in early spring’ (Stefanescu, 2001). (3)

The Red Admiral (Vanessa atalanta) on bell heather. Galicia, Spain, 28th August.

The small proportion of the UK population which do not migrate south are in effect opportunists, which presumably do well in mild winters but suffer heavy mortality in cold ones. The home-grown adults appear in early spring in the UK, well before the next wave of migrants arrive from southern climes, but the overall contribution of these overwintering individuals is thought to be minimal; populations in northern Europe were considered to be entirely dependent on immigration which determines abundance (8). This situation may however, be changing as the climate warms.

The Red Admiral (Vanessa atalanta) on bell heather. Galicia, Spain, 28th August. Wings held in typical 3/4’s open position (See Link 11).

Red Admirals flying southwards in September, in Finland, were found to migrate on sunny days when cool northern winds were blowing (13). Red Admirals take about 5 weeks to fly the 3,000 km from Northern Europe down to the countries surrounding the Mediterranean (1). Circumstantial evidence from meteorological radar observations suggests that they migrate at high altitudes (up to 2,000m or more), where temperatures may be as low as 2-3 deg C! Once they arrive in the south again, in places such as the Catalonia lowlands in north-east Spain – in October and early November, they start breeding a new generation. (2)

The Red Admiral (Vanessa atalanta) on bell heather. Galicia, Spain, 28 August.

Not all Red Admirals migrate over long distances. Studies in Spain by Stefanescu (2001) have shown that some individuals fly much shorter distances towards nearby locations of a high altitude. The butterflies shown here (e.g. above and below) feeding on bell heather were photographed in late August at one such location, near the peak of a hill in Galicia, Spain.

The Red Admiral (Vanessa atalanta) on bell heather. Galicia, Spain, 28 August.

Citizen science projects, such as the one on Red Admiral migration run by the Insect Migration & Ecology Research Group, at the University of Bern, Switzerland (13), offer enormous potential for gathering information on insect migration. People all over Europe can record sightings on a plethora of citizen science portals – some of which are configured as easy to use Apps – allowing researchers to build up unprecedented data bases of records in time and space. It will be fascinating to see what they can come up with in terms of new findings.

  1. Stefanescu, C., Alarcón, M., & Àvila, A. (2007). Migration of the painted lady butterfly, Vanessa cardui, to north‐eastern Spain is aided by African wind currents. Journal of Animal Ecology, 76(5), 888-898.
  2. Brattström, O., Bensch, S., Wassenaar, L. I., Hobson, K. A., & Åkesson, S. (2010). Understanding the migration ecology of European red admirals Vanessa atalanta using stable hydrogen isotopes. Ecography, 33(4), 720-729.
  3. Stefanescu, C. (2001). The nature of migration in the red admiral butterfly Vanessa atalanta: evidence from the population ecology in its southern range. Ecological Entomology, 26(5), 525-536.
  4. Fox, R. & Dennis, R. L. (2010). Winter survival of Vanessa atalanta (Linnaeus, 1758)(Lepidoptera: Nymphalidae): a new resident butterfly for Britain and Ireland?. Entomologist”s Gazette, 61(2), 94.
  5. Bolotov, I. N., Bochneva, I. A., Podbolotskaya, M. V., Gofarov, M. Y., & Spitsyn, V. M. (2015). Butterflies (Lepidoptera: Papilionoidea and Hesperioidea) from meadows of Vinogradovsky District, Arkhangelsk Region, northern European Russia, with notes on recent intense expansion of the southern species to the north. Check List, 11(5), 1727.
  6. https://www.youtube.com/watch?v=BN1WwnEDWAM (The Clash video).
  7. http://nurturing-nature.co.uk/wildlife-garden-videos/red-admiral-butterflies-do-they-hibernate-in-britain/
  8. Pollard, E., & Greatorex-Davies, J. N. (1998). Increased abundance of the red admiral butterfly Vanessa atalanta in Britain: the roles of immigration, overwintering and breeding within the country. Ecology Letters, 1(2), 77-81.
  9. Brattström, O. (2007). Ecology of red admiral migration. Department of Animal Ecology, Lund University.
  10. Brattström, O., Åkesson, S., & Bensch, S. (2010). AFLP reveals cryptic population structure in migratory European red admirals (Vanessa atalanta). Ecological Entomology, 35(2), 248-252.
  11. Peter B. Hardy. The Butterflies of Greater Manchester. http://www.oocities.org/pgll@btopenworld.com/bgm/bgm.htm
  12. Mikkola, K. (2003). Red admirals Vanessa atalanta (Lepidoptera: Nymphalidae) select northern winds on southward migration. Entomol. Fenn., 14(1), 15-24.
  13. https://insectmigration.wordpress.com/red-admiral-migration/