www.ukbutterflies.co.uk

Hi Goldie,

thank you for you kind comments, again. Appreciated. Make sure you do take me up on my offer. Might even see you there.

All the best, Nick.

An insight into the life of Chiddingfold Wood's Wood White.

Part four...Courtship of the Wood White

I wrote the following text [changed some wording] in my Personal Diary, [on this forum] nearly three years ago on Wednesday, March 7th 2012 about the courtship of the Wood White. [p]

But are they actually courting? I say this because i have seen this behaviour on many occasions with 1st and 2nd brood Wood Whites but, it has never ended with the pair of Wood Whites mating. Whenever i have witnessed this ritual, with the male whipping the female with his proboscis the male has always flown off without even a sniff, let alone managing to woo the female into mating.

When i wrote the above text i did not believe that the mentioned courtship of the Wood White ever resulted in the male and female joining in the act of copulation. And i have not seen any evidence since in my observations of the Wood White over the last couple of years to change my mind but, only to strengthen my beliefs that the ritual never finishes with the act of mating of this species.

I have witnessed this ritual [courtship] over a hundred times now and, you would think on the law of averages i would have been a witness to this behaviour at least once. In reality with the amount of times i have observed this ritual, i believe i should of had really seen this ritual end with the act of copulation not once, but several times and, have i? No, not once. And why not, is it that this courtship is not a courtship and never ends with the male and female joining in the act of copulation?

On one such occasion whilst observing this behaviour, the male whipping the female on the side of her wings, [i believe this hitting of the side of the wings with his tongue is deliberate and, is actually what the males are trying to achieve, but a lot of the time the males are totally inaccurate with their proboscis and miss, but more of that later] eventually the male flew off after a few minutes of performing this ritual. The female which was sitting on Bitter Vetch at the time the male was performing this ritual, bent her abdomen to one side, only for the female to lay an egg. But it brought a smile to my face. In my observations i have noticed that a very high proportion of females had been mated prior to this ritual.The female in the photo above has clearly been mated.As this female has as well, clearly heavily laden with ovum.

Anyone who has studied the Wood White knows the rarity of seeing the joining of a male and female of this species, and i have been privileged to have witnessed this act on a few occasions, the last time i observed this phenomenon was on the afternoon of 27.7.2014 [the afternoon is the only time i have seen Wood Whites mating] when i was watching a lone female sitting on some vegetation late afternoon. I had been sitting underneath an Oak Tree when after about ten minutes a male happened along, [he probably smelt her first] then he flew over to her and they joined. No courtship of any description, none, nothing at all, the male just flew in and they joined, simple as that. And this has been the case on every occasion i have witnessed the joining of a male and female Wood White in the act of copulation, no courtship. The only difference between them all was, the length of time the females sat waiting, [probably intentionally waiting for a male to mate with] any where from a couple of minutes, to 45 minutes in full sunshine as in this particular case. I remember the day well. The wood was popular with walkers that day, and a lot of them stopped and asked what i was doing. So explained that i was watching a female Wood White butterfly, and i pointed her out to one and all, and that i was hoping a male would come by, seek the female out and mate, [i,m sure some of them thought i must have a screw loose] but all the same, they all wished me luck. And after a 45mins wait, a male happened along, [ again probably smelling her first, before seeing her] and bingo he was in there in a shot and, they joined. No courtship, not a sniff of one, nothing, just straight in there and joined. That was my first experience of seeing a Male and female join in the act of mating, i was convinced that it would happen, all i had to do was wait, so i did.

Copulation normally lasts anywhere between 4o min. and 100 min. depending on the time of day they mate, to how long copulation lasts. I have left mating pairs of Wood Whites as late as 7 o'clock in the evening, [and like a pair of Orange-tips i found on 29th April 2014, which had stayed joined all night] i believe these late pairings stay joined all night as well [this is something i will have to investigate further]. The female Wood White always takes total control when joined in the act of copulation, when the female wants to move about the plant they are sitting on, or to fly to another location if disturbed, then he has no option but to follow, as the female will drag the male behind her even in flight. When the female decides to land again after being disturbed it is quite common to find the male hanging in mid-air. Sometimes when a copulating pair of Wood Whites are disturbed, the female will fly off still joined to the male and, without warning will stop flying and drop like a stone into the vegetation below.



To be continued...

Great observations, Nick, which substantiate statements made by some other authors; this doesn't seem to be a "courtship ritual" - the term typically used to describe this process. It really does make you wonder what the point is! I'm not sure if this is a "bonding ritual", if no bonding actually occurs and the butterflies go their separate ways.

By the way, I don't believe you can copyright text in the way you've outlined; I'm pretty sure it's not breaking any enforceable copyright (no matter what you state) if someone quotes an excerpt with full and clear credit given to you, with a reference to the original content.

Cheers,

- Pete

HI Nick,
Excellent observations of this rare little Butterfly .We observed some of the behaviours you describe with the
second brood last year. As a lot of this action takes place around ' Tea Time ' most visitors have gone home.
I am glad we went to see them in the latter part of the day.
This year it would be good if Wood White and Purple Emperor coincide for a few days.
Hope to see you at the Triangle this season.

TREVOR.

Hi Nick. From your observations, can you confirm that this interaction always takes place between a male and female and is always asymmetrical, the male whipping the female?

If it never leads to mating one could speculate that it only happens after the female has mated. Who is benefiting and what is the benefit? Do males seek out females carrying their own sperm (I don't expect you to be able to answer that!)? Since reading your post, theories have been flying through my mind, but sadly I have to go and teach maths now ... Fascinating stuff.

Guy

Hi! Nick, I have a theory but of course that's all it is,
Last year I photographed the male Speckled Wood who wanted to mate with this Female SW, she of course had just mated and refused him, he was very persistent so she played dead , I was really surprised I thought at first there was some thing wrong with her but as soon has the male flew off she went her own way .
I wonder if the Male Wood White in touching the Female in the way you've described can tell by her stance or the position she's in that she wants to mate, you said in one case the Male flew off, the Female shortly after wards laid her eggs.
You also described a female that wasn't ready to mate so the male flew off then she changed her mind, too late the male had gone!
In your description of a pair actually mating and the Male just flying straight in, it's surely the way the Female was waiting for a male and the actual male knew this by the way she'd positioned herself, just a thought. you know the saying "Position is every thing" it could be for Butterflies Goldie

Thank you all for you comments. Appreciated as always.

As for the copyrights on my text, i know there are no rights to the text, but i thought i would still go ahead and write it. I'm not the only person to do this. You see most people can be trusted to do the right thing but, not everyone is honest as i found out a few weeks ago on Dispar. I discovered something about butterflies in my first year of studying these insects. Three years later i see someone on this forum [an old friend, female, blond bleached hair, Pauline Richards] has taken the credit for something i had told/shown her in the field. How to identify a male from a female Large Skipper/ Small Skipper by the antenna. Pauline told me she would check her photos when she got home. And she did, and found i was right. But Pauline decided to take the applause for my discovery on this forum knowing i didn't follow this web site at the time. People on this forum applauded her for her discovery, but it was my discovery. Will i ever get an apology from Pauline. There is not a thing i can do about it now. Thats life. But its dishonest and dam right low. I have informed Pete Eeles of this.

Trevor you are quite right the afternoon is the best time to study/see and photograph these brilliant little insects. I'm sure we will bump into each other at some point at Chiddingfold Wood later in the year.


Guy. This ritual takes place between male and female, MALE on MALE, so not asymmetrical, but not female on female or female on male. You wrote, do males seek out females that carry their sperm. Thats one of my theories from about 3 years ago, that has gone out the window. I still have more to write on this behaviour, with photos, depicting this act and what i believe they are trying to achieve with this display, but whats the importance of this ritual, i can only surmise .


Goldie. as you quite rightly say, the position of the female Wood White is probably important to this issue of copulation, and if you add to the equation, the female letting off an odour telling the male that she is ready to mate, then you could very well be right.

All the best, Nick.

Nick Broomer wrote:Pete. I call the tongue lashing between two Wood Whites a bonding ritual, not just a bonding between to individuals [male and female] but covering all individuals within this species like a friendship. Humans bond all the time, whether its a gang of mates [all boys] down the pub having a good time or a group of girls and boys having a good time at a fun fair etc. but bonding all the same. Its all i could come up with, when all my previous theories went flying out the window.

I had a similar thought, but couldn't justify it. And that is - the ritual confirms that the species flying together are, indeed, compatible (not at the level of individual), which must have been a real need when different Wood White species flew together and you wanted to ensure that a potential mate is likely to produce fertile offspring. As is believed, the mainland was swamped with Cryptic Wood White before they all disappeared, with the remnant population being confined to Ireland.

Cheers,

- Pete

Pete Eeles wrote:

Nick Broomer wrote:Pete. I call the tongue lashing between two Wood Whites a bonding ritual, not just a bonding between to individuals [male and female] but covering all individuals within this species like a friendship. Humans bond all the time, whether its a gang of mates [all boys] down the pub having a good time or a group of girls and boys having a good time at a fun fair etc. but bonding all the same. Its all i could come up with, when all my previous theories went flying out the window.

I had a similar thought, but couldn't justify it. And that is - the ritual confirms that the species flying together are, indeed, compatible (not at the level of individual), which must have been a real need when different Wood White species flew together and you wanted to ensure that a potential mate is likely to produce fertile offspring. As is believed, the mainland was swamped with Cryptic Wood White before they all disappeared, with the remnant population being confined to Ireland.

Cheers,

- Pete

Thats an interesting theory Pete. I wonder how, if any, this behaviour would differ if it was between a sinapis and a juvernica for example and if it could be tested somehow

I'm really finding these observations totally engrossing Nick. I think I'm going to have to add seeing my first Wood White to my 2015 bucket list now!

Fascinating and detailed information Nick - good luck with the continued observations - I hope they yield an answer

Have a goodun

Wurzel

Hi Buggy and Wurzel,

Thank you for your comments, as always appreciated.

Buggy you should try and visit Chiddiingfold Wood if you can. I see you visit Boxhilll, another 30 min. drive will see you at this wood. If you do make the trip and need information on where to find the W.W., just ask.

Wurzel, i have all the answers i need about the W.W. and it's life, except the real reason behind their ritual, one can only surmise the true reason behind this ritual, and for now i'm sticking with that it is a form of bonding between the species as a whole.

All the best, Nick.

Hi! Nick, I think you could be right about the bonding, may be the Wood White's are just different in their approach to it, my Speckled Wood for instance( in my latest diary) pushed and prodded the Female and she didn't want to bond with him and played dead. It could be the way Wood White's by touching the Female find out what mood she's in and if she's willing to bond. If she's not they fly off, like yours did, where as the Speckled Wood persist's in trying to bond.
It could be this approach in the Wood White why their not as successful and why they've declined so much compared to other species. Just a thought! The other species could be more aggressive in their behaviour compared to the Wood White Goldie

Fascinating and thought provoking stuff Nick, with impressive photos

Looking at Adrian Hoskins' ever-helpful website, he has some interesting thoughts on this ritual http://www.learnaboutbutterflies.com/Br ... inapis.htm
This is based on the observation that, unlike other Whites, mated Wood White females have no rejection signal, so new males continue unsuccessful courtship for lengthy periods before giving up.

Vince

I must admit, I'm confused at the discrepancy between field reports (such as Nick's) from individuals who have seen pairings occur, with no courtship, and what's written in books and on websites (and I can't help thinking that these regurgitate each other's claims and assume that the actions really are a courtship!). Has anyone reading this EVER seen a pairing follow the "courtship" display?

Cheers,

- Pete

Pete Eeles wrote:Has anyone reading this EVER seen a pairing follow the "courtship" display?

To me that's a crucial question. Natural selection does not look favourably on energy expended on doomed missions! Also, if male-on male is not merely anomalous, it would seem information is being communicated here that is of general benefit to a colony (as ants and bees communicate information) rather than specifically reproductive information. Confirming the presence of the appropriate species is possible, but to me unlikely, as more than one of these Leptidea species often share the same habitat - so if it takes all this rigmarole to confirm ID, how come they can zoom in and mate with the right species in seconds?

Guy

Given that two near identical species of Wood Whites occur in certain places, could it be that some form of additional recognition steps are necessary?

Sorry to clog your diary up, Nick - let me know if you want this discussion moved! In the meantime, the following paper demonstrates that this "courtship" does lead to pairing, at least in captivity:

http://www.zoologi.su.se/research/fribe ... al2008.pdf

EDIT: And this related article:
EDIT: Link amended to work!

http://www.zoologi.su.se/research/leima ... al2013.pdf

Cheers,

- Pete

Hi all,

Pete wrote. Has anyone reading this ever seen a pairing following the "courtship" display? I asked the same question three years ago on this forum. I got two replies, one from Dave Millerd and one from Susie, who both stated that they had not seen this courtship end in mating.

Vince wrote. This is based on the observation that, unlike other Whites, mated W.W. females have no rejection signal. So new males continue unsuccessful courtships for lengthy periods, before giving up.

My observations.

In the wild this courtship never seems to last more than a few minutes. Also it seems that a male cannot distinguish between a mated and unmated female. a Why does a male try courting a female thats been mated but, does not try courting a female that has not been mated? When i have witnessed the joining of a male and female, the female just sits there waiting to mate, having already released a body scent that tells any passing male that she is receptive to mating. The male seems to know this instinctively when a female is ready for copulation, [which i believe the male can smell the female from at least a couple of metres] and will fly in like a bullet and join, no courtship. I have never witnessed the courtship ending in the act of mating in at least five years. After a female has mated i believe that there is a possibility that she loses the ability to continue to produce the body scent, as it is no longer needed. On one particular occasion i was watching a sitting female, a male headed towards the female, got within 30cm of the female and carried on flying past her. Then another male came along, saw her, flew down and around her and then flew away. Then third male happened along, he got closer and closer and then flew straight over her head and away. Then i realised, she has probably lost her body scent, as she no longer requires it as she has probably already been mated, . A male butterfly is programmed to mate, thats why it exists and, for no other purpose, so for three males to ignore a female in about 15 min. surely must indicate the female had been mated or, that she had not turned her attracter scent on, which i find to be inconceivable that a female can just turn her body scent on whenever she wants.

And then there is this, male on male. So is this still a courtship?

Nick Broomer wrote:Pete wrote. Has anyone reading this ever seen a pairing following the "courtship" display? I asked the same question three years ago on this forum. I got two replies, one from Dave Millerd and one from Susie, who both stated that they had not seen this courtship end in mating.

If you read the papers I've provided links to, you'll see that courtship did precede mating and that this was achieved in both captivity and the wild (with released females). In addition, "many females ... accepted mating after only a few seconds of courtship".

Cheers,

- Pete

Yes, this complex behaviour is, without doubt, courtship. My own observations of Wood White behaviour, courtship and mating are entirely consistent with the huge body of research performed on this species, by some of our most respected lepidopterists, and particularly Professor Christer Wiklund. The summary provided by Adrian Hoskins is spot-on.

I have personally watched Wood White interactions result in copulation on four occasions since 2007, all at Chiddingfold. In two of those, the courtship display lasted approximately one minute (+/- 20 seconds), and included wing-clapping, in addition to proboscis and antennal waving. On the other two occasions courtship lasted approximately 10 seconds, without the wing-clapping.

Wiklund himself has witnessed almost instantaneous pairing, and short duration courtship has been well documented (Friberg et al., 2008). These short duration courtships still give the females sufficient time to confirm potential mates as belonging to the same species (of Wood White) as they. It would seem that (almost) instantaneous pairings do occur, but quite rarely. There are exceptions to almost everything in nature. Meadow Browns and Small Tortoiseshells should not be having sex with each other!

Males cannot differentiate between females belonging to different species of Wood White, so the decision-making is entirely the prerogative of the female. Note that more than one species of Wood White occur in some regions today (ex UK), and historically would have done so in others. This probably explains the complexity of the courtship, as the various species are so similar (even to Wood Whites!) that a complex combination of signals is required.

As Adrian points out, the mated females find it very difficult to communicate 'rejection' to the males, and hence the courtship ritual extends far beyond the timeframe necessary for a receptive (virgin) female to respond to a male's advances. It is unsurprising, therefore, that the much lengthier, unsuccessful courtship rituals are far more commonly observed.

It is probably the complexity and duration of these displays which lead some to doubt that this is courtship. However, in my opinion, the ratio of observed 'failures' on the part of males, is no different to that seen in many other species. Given sufficiently good weather on emergence, female butterflies remain virgin for a very short period of time. They then spend the rest of their lives spurning the advances of amorous males. In other species, even persistent individuals will 'get the message' quicker than a male Wood White.

If we consider some hypothetical figures, we see that the rarity of an observed, successful conclusion in Wood White is far from unusual. If we consider that a female butterfly of species X lives for 20 days, she will perhaps be active for 50% of that period for 5 hours per day, once averaged out. Assuming she was mated very quickly (often before her wings are fully hardened), she will then be accosted on numerous occasions, every hour, by amorous males. Even at the very conservative figure of two interactions with males per hour, we arrive at a ratio of 100:1 unsuccessful:successful encounters. The asymmetry of outcomes is probably far greater than this.

We see this all the time in common species such as the Brimstone and Orange Tip, where the courtship behaviour is, by comparison, very basic. How often do we see a successful outcome? But we don’t doubt that it is courtship.

It is also worth pointing out that the ratio of male:female Wood Whites is approximately 50:50. However, the females behave differently and are seen less often. This is quite normal, although there are well documented cases where the sex ratio is skewed in some species (in favour of females) by sex-specific parasites.

Also, ‘bonding’ does not occur in Lepidoptera. Bonding, either social bonding or pair bonding, is a behaviour which has developed in animals because it serves a particular purpose, and confers an evolutionary advantage. There can be no such advantage to butterflies in pursuing this type of behaviour. It doesn’t happen.

BWs, Neil