Woodpigeons calls and displays

Although the woodpigeon is the bird I come across most often and the one I record the most, it is still one of my favourite birds, the species that made me a birdwatcher. I find them truly stunning and very resourceful, feeling at ease in gardens, parks and woodland alike. Despite their reputation as grain guzzlers their diet includes leaves, nuts (acorns and beech mast), buds, flowers and berries, including rowan, ivy, holly, cotoneaster and alder, which they often reach with great acrobatics. Although Woodpigeons have featured often in this blog, here I am compiling and documenting their calls and displays, especially those related to reproductive behaviour. Although in his article on woodpigeon behaviour Cramp says that 'they mark their ownership of a territory primarily by just being there' as indeed 'a Woodpigeon perched in a tree bare of leaves is a large and conspicuous object' Woodpigeons have a simple, but interesting behavioural repertoire.

Advertisement call. A deep, repeated 5 note phrase that serves as a territorial song. This is the well know Woodpigeon song, often heard early in the morning.

Display flight. This is another territorial display most frequent in February and March. The male flies around the territory rising in a broad arc, and then gliding down with wings held stiffly horizontal advertising ownership. They can do the rising and falling a few times before alighting. Although mainly a visual advertising display, as the bird does not call, sometimes the bird will make a clapping noise with its wings at the apex of the arc.

The Bow display. It is a courtship display that territorial males perform when still unpaired and trying to attract a mate and also when the pair bond is becoming cemented. The male approaches the female and with his neck enlarged he calls a three note call as he alternatively stretches up and then lowers its head, displaying the white and iridescent neck markings, and fanning the tail when it's at its highest. Check out this wonderful photo by Richard Hawley showing the display when fanning is at its maximum. Females can display some aggressiveness to the courting male, move away or flee if they are not interested. This display often happens on trees, roofs or aerials, but occasionally on the ground. When on the ground the male will try and get closer and follow the female hopping to bow just in front of her, one of the few occasions that woodpigeons will hop instead of walking.

Allopreening, billing and mating. Feeding the mate, gentle preening and billing precedes mating. I documented this behaviour in a series of photos here.

Slightly opened wings is a threat posture.

Fighting. Woodpigeons have large chest muscles, which power their explosive take off and fast flight. They have small bills and legs, so they also use their wings as offensive weapons in their territorial fights. Males will fly towards intruders landing in their territory and land adopting an aggressive stance, with neck stretched, flattened plumage and semi-open wings, and often this is enough to drive the intruder away. Occasionally, this develops into a fight, which sometimes occur in the depths of a tree. The pigeons will land near each other, alight their bodies and flap trying to hit each other (above).      
                                         
Nest calling. This is a call by the male signalling to his partner a suitable nest site. I've covered this on other posts (here and here) and it is described in detail in the article by Cramp cited below. The call is harsh, and guttural composed of two notes, here is a recording of the call. I want to share here a video I took this morning, in which I got a good angle. You can see pecking or nodding movements to the nest floor in between calls:




More information
Cramp, S. Territorial and other Behaviour of the Woodpigeon. Bird Study 5, 55–66 (1958). here.

Woodpigeon mating

Although the Woodpigeon must be one of the most abundant and familiar British birds, a few days ago I was lucky to watch their courtship and mating at very close quarters, when a pair landed on the garden fence. I should have taken a video, but instead I ended taking a series of photos, which don't cover the full sequence. I saw, but did not photograph, the initial courtship feeding.

1. The male (on the right) had just fed the female, both move the bills, like swallowing. They are right next to each other and excited, note the contracted pupil in the male. The female sits down, crouching, inviting copulation.

2. The male mounts the female, balancing with wings open.

3. The male stabilises on top of the female, which remains motionless, with  wings spread (to make male balancing easier?).

4. Copulation, male flaps for balance, female leans forward lifting tail.

5. The male jumps onto the left of the photo, both partners (which are extremely alike in colour and size) fluff their neck patch and bow their heads in what looks like an aggressive display. These are woodpigeons in all their splendour, what amazingly beautiful birds they are!

6. The female actually turns away from the male, not a brilliant shot but shows how the neck patch feathers are raised. The pupil is very contracted.

7. The male remains in the post-copulation display.

8. Both pair members come together again and preen (I think the female is now on the right, and shows the initiative in caressing the male neck and head area).

9. The caressing carried on for a while.

Stock doves courting

I was lucky this morning to watch a pair of Stock Doves courting from a good vantage point. They usually stick to high branches in the trees in the park, only rarely coming to the ground. Today I heard a male's booming call from the trees opposite the pond, and I climbed to a small hill on the other side. My new camera is heavy and I still need to get used to its weight, so I eventually leaned against a tree trunk to steady myself. The male crouched on a branch bowing parsimoniously at the same time that fanned its tail - very similar to woodpigeon's courtship -, facing the female. Its neck, inflated with air showed while cooing showed its green iridescent neck patch, which looks much larger than when the dove is at rest (bottom photo).

A side view of the male singing

The male looks at the female about to land

She greets her with this lovely display

She eventually lands next to him. This photo is not brilliant, but I like how the neck patch is actually ruffed, something I had not seen before.

More male bowing, fanning tail, showing its iridescent neck patch.

A stock dove at rest, from 27th Feb. Compare the size of the neck path.

Four calling pigeons

I had this little project in my mind for a long time, to get photos of all four local pigeon species as they call. What I had not anticipated was that I would do in in a single day in the space of 15 minutes, all in a very small area. All pigeons are now in full blown song and courtship. When I spotted the Woodpigeon above, he wasn't calling, it appeared to be just enjoying the sun, but as I walked away, I heard it calling and I turned back. The noise of the roadworks didn't stop it calling.

It is unusual to see a Collared Dove calling in the park.

Not much further away, a Stock Dove started calling. This ash is their favourite tree, and most of the time there are two or three there. Stock doves are quite timid, and they call from high vantage points.

To end the series, it wasn't too hard to find the courting Feral Pigeons, there were plenty around. This male and watchful female were on the roof of the cafe at Pearson Park. 

Carrion Crow begging to its partner

A Carrion Crow collected food in its bill and gular pouch on the University playing fields. Sitting in the nest atop a still bare ash, its mate called repeatedly, a begging, hoarse, nasal call, kaaaa! similar to a Rook's. The crow collecting food - most likely the male - looked up to the nest and flew heavily upwards in circles, stopping on a roof on the way. The female called again and flew off the nest to meet him, calling like a fledgling would do, crouching and wing flapping. The begging and feeding carried on out of sight on the roof.
In Carrion Crows, the female incubates and is fed by the male during this time, so it is likely that the nest on the ash tree already has eggs.

male crow collecting food for its mate

The female returns to sit on the nest (maybe already has eggs?)

The female's tail sticks up from the nest.

Swinging goldfinch

A trio of goldfinches came to the feeders today. One of them, while it waited on nearby branch to feed, carried out a curious 'dance', repeatedly swinging its body from one side to the other while it called. I looked into this and found out the dance it's got a name: 'pivoting display'. In a courtship context, the display is carried out by the male and he swivels towards and away the female as he swings. The wings are somewhat lowered - which makes the golden bars more visible - and the tail spread, showing the white patches. Female also pivots, and when both sexes carry out the display it may end in copulation.
 The pivoting display is also used in aggressive contexts, and maybe the goldfinch was becoming impatient waiting by the feeder. I shall keep an eye on my visiting goldfinches and see if I can record this behaviour. I found a little video showing the behaviour in a charm of goldfinches. Check the individual at the bottom of the image from s 36.



And a great video of a pair of captive goldfinches courting.

Mallards grunt-whistling

The drake mallards at the park have now almost completely finished their summer moult and are sporting their dazzling fresh plumage. Mallards start the breeding season quite early. The first stage is pair formation, which starts in September-October. Any unpaired females attract the attention of drakes and a flock of suitors surrounds her, parading. A trademark display of many ducks is the 'grunt-whistle'. In mallards this consists of a sequence of behaviours including a whistle, at the same time that the duck lifts both his head and tail.
 The fact that there are usually many males displaying at once and any of them can carry out the behaviour, means that I've had trouble recording in it a photo previously. By the time I head a whistle, or several, I have missed the behaviour. This week in the park, when I noticed the mallards courting, I decided to focus on a particular male and try and catch him doing it.
 The first step of the grunt-whistle sequence looks like the duck is about to dive. Instead, he arches his head forward forcefully, and his bill tip scoops an arc of water droplets toward the female. I managed to get two shots of this stage. The second shows the droplets quite well.

Then is the whistle, while lifting the tail and head (note the bill is a little open).

After that, the drake turns to the female (top shot), and then swims fast with head forward, almost touching the water.

I will try and get a video of the whole sequence in the next few weeks and upload it here.

More information

Wolfgang M. Schleidt & Erhard Oeser (2011) Konrad Lorenz’s use of cinematic film for studying dabbling duck courtship behaviour and the availability of historic film materials. Wildfowl 61: 45–51. Here.

House Sparrow communal courtship

 House Sparrows are quite vocal birds. In spring, a house sparrow colony is hard to miss, with males advertising their chosen nest sites to potential males by chirping and posturing, but mainly by their peculiar communal courtship. Communal courtships start when a fertile female not guarded by her mate flies by an unmated male, the male then will pursue her while chirping and displaying persistently soliciting copulation. More males usually join in the female chase and courtship in a cacophony of strident chirps. The female is far from passive to the male's attentions, and often lunges and pecks at the males (photo below), which seem quite unfazed. Forced copulations - and also accepted copulations - occur during these communal courtship displays.
 Male sparrows with large black chest badges are more dominant in the sparrow colony. They guard their mates during the fertile period - around the period when egg-laying is taking place - and copulate with her frequently. But males with large, in addition, join communal courtships more often than males with small bibs, so these dominant males may achieve paternity both though their mate's offspring, and through extra-pair copulations.
 Today, in a local farm, I managed to get these shots of communal courtship in House Sparrows. Two or three males were harassing a female, and she managed to peck one of the males (below).

ouch!

More information

Moller, A. (1990). Sexual behavior is related to badge size in the house sparrow Passer domesticus Behavioral Ecology and Sociobiology, 27 (1) DOI: 10.1007/BF00183309

Frisky Pigeons

All of January, Collared Doves have been playing their the three note song in a repetitive, mesmerizing soundtrack. Outside, display flights, in which the doves fly across the street of over roofs with wings held low as they descend, indicate that Collared Doves are starting to gear up for the breeding season.

Woodpigeons too. In fact, they seem to not have stopped breeding. In early November, I watched a fledgling begging for food in the park (above), and yesterday, I saw an immature with no signs of white feathers, feeding on the grass (below). The individual on the top shot appears to be a young pigeon growing its neck and head feathers.

Today, in the lime tree just outside my house a male Woodpigeon nest-called persistently while the female checked a pile of loose sticks on the first fork of the tree.

Even the Stock Doves, a normally quite discreet bird, also showed a lot of exuberant activity in the park. High flights, chases, courting and persistent song. A male tried to approach a female on a brach and she flew off (below).

The - so far - mild winter might be fooling them into an early breeding season.

The fascinating love life of the Dunnock

I watched a Dunnock today, feeding under the garden table, with that characteristic half hopping half walking way Dunnocks have, pecking here and there things too small to be seen at a distance, maybe seeds or small invertebrates. Dunnocks, or Hedge Sparrows (Prunella modularis) are little birds, which live their lives mostly unnoticed amongst the undergrowth and are easily overlooked or taken for House Sparrows. They have a grey chest and head and chestnut backs with dark stripes, a thin beak and orange legs. Both males and females look similar, females just a bit smaller than males. In gardens they often feed on the ground, under bird tables when there is some cover, and they prefer to skulk than to sit out in the open. Only in the spring, where males sing their weak, warbling song from a prominent perch they are somewhat more likely to be noticed (above). Contrasting with their modest attire and retiring habits the Dunnock shows a variable mating system - including a common arrangement of two males and a female, a system called polyandry- , and a courtship behaviour that can only be described as peculiar. I was lucky enough to witness courting Dunnocks a few years back. This is a sketch of what I saw and my description.

Female appears paler, less extensive grey markings than the male. Both individuals are on the ground. The male hops behind the female and she stops a moment with her tail slightly cocked and vibrating her wings, that are dropped.  The male pecks her cloaca repeatedly. Female hops away a little, male follows and the same behaviour starts again. Then the intensity increases, the male pecks her cloaca again and the female stops, then the male jumps on the female and there is a flutter

 What could be the purpose of this bizarre cloaca pecking behaviour? Has it got anything to do with the presence of polyandry? Nick Davies, in a classic paper published almost 30 years ago, provided some answers. He followed a Dunnock population in Cambridge Botanical Gardens. Males outnumbered females due to higher female mortality during winter, and therefore, there was intense male-male competition. He observed several combinations of breeding partners per territory, including male-female pairs, two males and a female (what he called trios), and some more rare cases including two males defending jointly the territories of several females.
Monogamous males guarded the female, following her around closely for a few days before she was due to lay and gaining almost exclusive copulations with her. When there are more than one male in a territory the larger male was dominant to the smaller one and fights were common, with the dominant male trying to chase the subordinate away from the female. Although the dominant male got the best share of copulations, the subordinate also got them as he was usually very persistent and mated with the female unnoticed by the dominant male, or when the male lost track of her - often after a fight. In all events of courtship, be monogamous males or not, the female always exposed her cloaca, often a pumping action was noticed and the male pecked it. After cloaca pecking, the female was seen to eject a droplet of fluid. Davies managed to collect three of these droplets and when he examined them under the microscope he found them to contain bundles of sperm. When he watched the female ejecting the drop of sperm, the male copulated with her immediately after. The purpose of the cloaca pecking behaviour appeared clearer: the male stimulates the female to eject stored sperm from the previous mating, allowing the suitor a shot at paternity. This was confirmed by the fact that the more other male spent near his female, the more a male pecked the female and copulated with her, as the male pursued to increase their chances of paternity. Interestingly, the female played an active role in being part of a trio:  she tries to obtain copulations from the subordinate male, escaping the guarding of the dominant, despite his efforts: obviously ejecting sperm from the previous mating will offer both males a share of paternity. When a female was observed to have mated with two males, the brood raised was fed by both males. In contrast, if the subordinate male failed to mate with the female, he did not contribute subsequently to raise the chicks. As nestlings fed by two males have a better chance of survival, it appears that is in the female interest to mate with both males.

A subsequent study using DNA fingerprinting confirmed what behavioural observations had strongly hinted: monogamous males got 100% paternity, and both dominant and subordinate males fathered chicks (surprisingly more or less equally). The observant reader will notice that the mean paternity of dominant and subordinate do not add to 100% in the polyandry system. This was because an outsider gained access to the female and fathered a chick.

(modified from Table 1 from Burke et al 1989).

Furthermore, this study showed that the chances of males helping the female rear the brood were dependent on them having sired some of the brood. It appears that the male is able to judge if he has had sufficient access to female to gain some paternity and to be worth the effort of helping her rear the chicks.

Although cloaca pecking can be seen as the male bird trying to ensure his paternity, the elaborate courtship of the dunnock also reflects that females are active participants and, that, as they need more than a single male to rear her chicks, this unusual courtship is the way females ensures that both males help her raise her chicks and that she achieves more reproductive success.

A pair of Dunnocks in the garden. What would they be up to?

More information
Davies, N. (1983). Polyandry, cloaca-pecking and sperm competition in dunnocks Nature, 302 (5906), 334-336 DOI: 10.1038/302334a0

Burke, T., Davies, N., Bruford, M., & Hatchwell, B. (1989). Parental care and mating behaviour of polyandrous dunnocks Prunella modularis related to paternity by DNA fingerprinting Nature, 338 (6212), 249-251 DOI: 10.1038/338249a0

Take a bow

There is a lot of Woodpigeon activity these days. I can hear the slow, five-syllable song continuously repeated from the aerials and trees in the street in the morning. The clapping of the territorial display, an exhibition flight, in which the Woodpigeon rises high noisily clapping its wings and then glides down in a circle with wings outstretched and kept horizontal. On a roof yesterday, a male and female were engaged in more active courtship. The pair makes these little jumps as they get closer to each other and they engaged in feeding, usually the male feeding the female.

This couple sat together for a while, but if courtship proceeds the male engages in a 'bowing display, with exaggerated, ritualised slow bowing until its beak touches the ground while lifting its tail and opening and closing it like a fan facing the female. The male calls at the same time. While singing the male ruffles his neck feathers showing off its side white and iridescent patches. If the female is ready she crouches opening its wings and the male quickly mounts her, otherwise, she will move away or try and peck the male or even fight him off with wing strokes.
 The following YouTube video shows a male courting a reluctant female.