Meadow brown (Maniola jurtina) butterflies, must, I think, be one of the most common and widespread butterflies in Europe. I am surprised therefore, not to be able to find any photographs of courtship behaviour in this species. The excellent UK Butterflies website states that: “Courtship between male and female is brief. The male showers the female with scent scales from his sex brands which act as an aphrodisiac that seduces the female and mating quickly follows.” Adrian Hoskins (Learn about Butterflies), in his entry for the meadow brown, also describes the courtship ritual as being very brief, “in which the female adopts a fluttering and gliding flight just above the grasses, while the male flutters very closely behind her, almost making physical contact during flight.” This is, I think, the initial phase of courtship. The final phase, which may or may not lead on to mating – it usually does not, for reason described below – is the one in which the male brushes the female’s antennae with his androconia (i.e. sex brands). The long, tufted, androconial scales can be seen on the forewings of newly emerged, male meadow browns, such as the one shown below.
Males are generally more active than females and according to Brakefield (1982), search for mates by making “slow exploratory flights close to the vegetation” with frequent changes in direction. Professor Paul Brakefield FRS – who studied this species for his PhD – mentions that “courtship is short with few preliminaries” and “is similar to that of Hipparchia semele“, the common grayling, which was studied by Niko Tinbergen in the 1940’s, as described by my blog on the subject.
The females often rest in the grass (see above). In very hot countries, e.g. in Southern Europe, the females enter a diapause during the hottest days of summer (often lasting until September) and delay the maturation of their eggs. Most female meadow browns only mate once, although a small proportion (7%) were found to have mated twice, in a study carried out in Spain; determined by counting the number of spermatophores (García-Barros, 1987). It is quite possible therefore, that many of the females sitting in the grass and being approached by courting males, will already have mated, and will not want to do so again; either because they are still laying or eggs, or because the majority of females only mate once.
Since I have got my eye in – meaning, becoming more experienced, for non-English readers! – in terms of observing this behaviour, I have noticed it a number of times when watching meadow browns, both in the UK and whilst on holiday in Spain. As mentioned above, the final courtship encounter is very brief, and is over in a few seconds. So, I was very pleased, and a bit surprised to have captured it in a series of photographs (below). It happened whilst I was walking along a remote pathway through mixed woodland in Galicia, Spain, last month (June 2019). I suddenly noticed a pair of butterflies chasing each other (well, the male was chasing the female) and I spotted the characteristic bowing movements made by the male. At first, I thought I was looking at a pair of grayling butterflies, as both graylings (Hipparchia semele) and rock graylings (Hipparchia hermione) engage in courtship bowing, as described in another blog of mine. The bowing behaviour is so distinctive (see below), that once you have seen it, you never forget it. It is almost as though the male is headbutting the female! In fact what he is doing, is brushing his androconial, scent scales against her antennae: thus, transferring a pheromone signal. This what it looks like in rock graylings (below).
The two butterflies, which as it turned out were meadow browns, flew off the track into dense foliage (much of which was spiky gorse!). I was determined to try and capture this behaviour, so I pushed my way into the gorse and found the female sitting on a small, horizontal twig, not far off the ground (below). The male was also sitting still close by, but slightly out of sight.
I thought to myself, right, I am going to sit here – with my eye pressed up against the camera viewfinder – in an uncomfortable position, for as long as it takes! Fortunately, I did not have to wait too long, and because I had my finger on the button, I managed to take three quick photos before the action was over! In retrospect, I wished I had used a faster shutter speed than the ‘one thousands of a second’ I took the shots with, but sometimes you just have to be happy with what you got! The slight blurring conveys a sense of motion, hopefully.
The images suggest that the male rotates his body forward on the sagittal axis, a bowing movement, a number of times. After the bowing was over, the male fluttered around the female briefly (below), before she took off.
It is possible that these were Maniola jurtina subspecies hispulla, as they had a nice orange colouration?
The female eventually flew off and came to rest on a gorse stem (below).
The courtship bowing shots (above) are rather blurred and I am sure other photographers can greatly improve on them, if they can get themselves into the right position to record this behaviour. I tried a couple of times back in England this year, and failed to capture it. So, I guess I was just lucky, but I intend to try again. N.B. this courtship bowing occurs in a number of other species, including many different graylings and other satyrids. The fine details of the behaviour will, almost certainly, differ between different species, as demonstrated by Manuela Pinzari (see Pinzari, 2009 and Pinzari & Sbordoni, 2013).
Brakefield, P. M. (1979). An experimental study of the maintenance of variation in spot pattern in Maniola jurtina. Unpublished Ph. D. Thesis, University of Liverpool.
Brakefield, P. M. (1982). Ecological studies on the butterfly Maniola jurtina in Britain. I. Adult behaviour, microdistribution and dispersal. The Journal of Animal Ecology, 713-726.
García-Barros, E. (1987). Observations on the biology of Maniola jurtina (L., 1758) in the center of the Iberian Peninsula: general phenology of the biological cycle, duration of the proposed period and potential fecundity of the females (Lep. Nymphalidae). Bol. Asoc. Rsp. Entomol , 11 , pp.235-247.
Pinzari, M. (2009). A comparative analysis of mating recognition signals in graylings: Hipparchia statilinus vs. H. semele (Lepidoptera: Nymphalidae, Satyrinae). Journal of insect behavior, 22(3), 227-244.
Pinzari, M., & Sbordoni, V. (2013). Species and mate recognition in two sympatric Grayling butterflies: Hipparchia fagi and H. hermione genava(Lepidoptera). Ethology Ecology & Evolution, 25(1), 28-51.
great stuff as ever Ray and so closely, minutely observed. Thank you
[…] the butterflies – Meadow Browns as it turned out – into a gorse bush! As I explained in my blog about courtship bowing in Meadow browns, I just waited patiently, until they repeated the […]
[…] but the differences are most apparent on the uppersides (as shown below). I once came across a courting couple of Meadow browns, and documented their courtship […]