Wing-spreading in Cormorants

The spread-wing posture of cormorants, like the individual above in a local park earlier this month, is one of their most distinctive behaviours. Multiple reasons have been offered to explain this wing-spreading behaviour, is it to dry their plumage? or is it thermoregulation (basking)? Even aiding their balance, signalling successful captures, or help swallowing fish have been put forward as possible explanations for this behaviour. The evidence in the Cormorant overwhelmingly supports the plumage drying hypothesis.
A study by Robin Sellers on wintering cormorants in the river Severn provided strong evidence for wing-drying as the function of this behaviour. Cormorants engage in the open-wing posture almost exclusively after having been diving, when their plumage is wet. After a bout of diving, the cormorant will do some bathing movements in the water, flap its wings and shake its plumage before flying to a perch, where the wing-spreading takes place. Not only the wings, the tail is also spread at the same time. The longer they've been immersed, the longer they will stay open-winged.
Wing-spreading was almost exclusively observed in individuals that had been in water the previous 30 min, and most of the cormorants that had been in the water did engage in wing-spreading shortly afterwards.
Wing-spreading is often followed or preceded by preening the plumage. The duration of the behaviour is also inversely correlated to temperature and wind-speed, supporting the drying hypothesis. Rain tends to inhibit the behaviour.
Wing-spreading in Cormorants is not associated to a successful fishing event, rejecting the 'digestion-aiding' hypothesis.
 Sunny conditions did not trigger wing-spreading, but at low wind speeds cormorants tended to orientate away from the sun.

Super-wettability
So, why don't other diving birds, such as grebes, auks or goosanders engage in this behaviour? The mechanistic, proximate reason is that cormorant feathers are not very water-proof, unlike other diving birds, in fact, they are very become waterlogged after a few minutes in the water. This is not due to a deficiency of their oil glands, but to a different microstructure of their feathers, which is thought to help reduce their buoyancy, and therefore energy requirements, during diving.

Different birds different functions?
Evidence also supports wing drying as the primary reason behind wing-spreading in other studied cormorant species (Shag, Double-crested Cormorant, Galapagos Cormorant, Bank Cormorant and Little Cormorant). However, in the tropical Anhingas, a group related to the cormorant family, wing-spreading takes a much larger proportion of their time than in cormorants, and the behaviour tends to happen for longer in sunny, cool weather, with their back oriented to the sun, and doesn't require wet plumage. Anhingas have a very low metabolic rate, which might favour efficient basking behaviour, and in captive, controlled conditions, oxygen consumption decreased as they engaged in wing-spreading. Therefore, the primary function of the behaviour in anhingas appears to be thermoregulatory ('sunning') rather than drying (although this may also have a secondary role). It is a cautionary tale that apparently identical behaviour in related species can have different functions.

More information
Sellers, R. M. Wing-spreading behaviour of the cormorant Phalacrocorax carbo. Ardea-Wageningen, 83, 27–27 (1995).

Hennemann, W.W. Energetics and Spread-Winged Behavior in Anhingas and Double-Crested Cormorants: The Risks of Generalization. Am. Zool. 28, 845–851 (1988).

Long-tailed tits hanging and feeding

Just a quick field note on foot use. I wasn't aware on Long-tailed tit foot use before, so I was very pleased to watch this behaviour. Sun flower hearts had been placed atop posts on a nature reserve and a flock of Long tailed tits was feeding on them. Individuals will pick a sunflower seed and jump to a branch to hang from it with one foot while holding the seed on the other food to feed on it. Other tits hold seeds between their feet while they feed, perching on a branch. Is it that Long-tailed tits are not able to balance on a branch while bending down on their feet, maybe because of their long tail? I saw the behaviour repeatedly, and I should have taken a video, but I just got this shot from behind that shows the foot holding onto the branch and the other with the seed on it. What call my attention was the fact that they had to jump to a branch to hang from in a premeditated way so that they could feed. The Two in the Bush blog also describes this behaviour, with much better photos. Hanging from one leg poses no issue for these tiny acrobatic birds!

Stonechat pairs in winter territories

Species like the Robin keep territories during the winter, with males and female individuals defending their own separate territories, while in the summer, territories are defended as a pair. There are some exceptions in which pairs defend winter territories, often in different areas that summer ones. One of these species is the Stonechat. In their wintering grounds on the coast and around the Humber estuary, it is frequent to encounter Stonechat pairs in the same area, feeding near each other and following each other, with no aggression (top shot, a pair at Alkborough, Lincolnshire, 11/12/2017). Of course, winter territory defence has nothing to do with reproduction, and often happens in areas that don't even hold breeding pairs in the summer. This unusual behaviour must offer some benefit to persist in the population, maybe allowing individuals to increase their chances of survival to predation or competition and to secure sufficient food resources.
A study of wintering Stonechats in the Negev desert (Israel), shed some light onto the behaviour. A total of 89 individuals were colour banded and followed through two wintering seasons. Their territorial behaviour was noted. The most surprising result of the study is that wintering Stonechat pairs are very fluid. Pairs often form after arrival to winter areas, and individuals rarely stay together all winter, instead pairs change frequently, with an average of just 4 weeks together per pair. In fact, none of the studied pairs stayed together all winter long and individuals arrived in the area and left at different times. This rules out that breeding pairs migrate together or settle on the same winter territory. Up to a third of individual are unpaired at any given time, but all associations are always between a male and a female. There were some aggressive interactions between members of a pair, which were commonly initiated by the male. Both males and females engaged in territory defence, with males more aggressive against intruders than females.

Male Stonechat at Kilnsea, 10/12/18.

Why pair in the winter?
If pairs are not stable, don't migrate together, or don't form in preparation for the winter season, what is the point in defending territories as a pair? Males and female stonechats feed on the same prey, therefore there may be a cost on sharing a territory during the winter, when food resources may be very scarce. A possible benefit outweighing the cost may be joint territory defence: two individuals are better spotting and chasing away conspecific intruders or detecting predators and defending the territory against individuals of other species with similar food resources. In this study, individuals spent a lot of time defending the territory against Mourning Wheatears, but there was no mention of differences when pairs or individual Stonechats were involved.

Another pair at Kilnsea (13/1/2020)

Why male/female pairs?
These results don't clarify why associations are not between two males or two females defending a territory. The authors speculate that as Stonechats are sexually dimorphic, even in the winter, there may be constraints that prevent two males from living peacefully in the same territory, and the increased male aggressiveness may mean two females may be unable to defend a territory against a male/female pair.

More information
Gwinner, E., Rödl, T. and Schwabl, H. Pair Territoriality of Wintering Stonechats: Behaviour, Function and Hormones. Behav. Ecol. Sociobiol. 34, 321–327 (1994).

Loafing gulls

 Gulls spend a good portion of their days doing what is known as 'loafing', in open areas of good visibility: playing fields, beaches, but also on water and on ice in frozen lakes, the same sites are chosen year on year. These loafing areas are different from night roosting sites. At loafing sites gulls appear relaxed: some sleep - head under wing - others sit or stand watching around, others preen. Immatures may play with objects such as sticks or leaves. Loafing areas are common areas, and tend to be quiet, with little or no squabbling or aggressive behaviour, often with several species mixing in the same area. Individuals come and go, commuting to their feeding areas, unless a disturbance provokes a sudden communal flight. On the coast, the number of individuals in a loafing area follows a tidal cycle - as low tides expose food resources the gulls move away the loafing area to feed- and also time of the year. Tidal influence in sleeping is highly prevalent in wading birds, which strongly depend on exposed shores for feeding. A study in Herring Gulls suggest that they have a dual sleeping pattern, with the proportion of sleeping gulls peaking at midnight and at midday. This may be a common pattern with urban gulls or away from the coast, where food is less related to tidal fluctuations.

A mixed flock of Black-headed gulls, Common Gulls and Herring gulls loafing on a rise on a local park.

Loafing gulls allow good opportunities to check individuals for plastic rings, as shown by this preening Common Gull.

A group of Common Gulls loafing on ice on a frozen pond of (16/1/2013)

More information
Shandelle M. Henson, James L. Hayward, Christina M. Burden, Clara J. Logan and Joseph G. Galusha. 2004. Predicting Dynamics of Aggregate Loafing Behavior in Glaucous-Winged Gulls (Larus glaucescens) at a Washington Colony. Auk 121, 380–390.

Cooke, F. and Ross, R. K. 1972. Diurnal and Seasonal Activities of a Post-Breeding Population of Gulls in Southeastern Ontario. Wilson Bull. 84, 164–172.

Galusha, J. G., JR and Amlaner, C. J., JR. 2008. The effects of diurnal and tidal periodicities in the numbers and activities of Herring Gulls, Larus argentatus, in a colony. Ibis 120, 322–328.

Wintering Chiffchaffs

In the last month, I've come across a couple of wintering Chiffchaffs. The last of them this Wednesday on my way to work, the calling bird revealed it's presence atop a maple, busily feeding on the top branches. Although most UK warblers are breeding migrants, four species can be found during winter. Two of these are resident species, the Dartford and Cetti's Warblers. The situation is more complex with the Blackcap, which I've covered before, and the Chiffchaff.

Sewage plants and wetlands
Chiffchaffs are insectivorous year round, so they can't rely on garden feeding like Blackcaps during winter. Overwintering in Chiffchaffs was documented in the early 19th century, but this seem to involve very few birds. In recent decades, however, there seem to have been an increase in numbers, possibly aided by global warming. Ringing and recaptures have allowed estimates of around 100 individuals gathering in suitable sites. Chiffchaff winter survival depends on a steady supply of invertebrates (mainly midges, gnats and aphids) and sheltered sites where to roost. They appear to survive several consecutive nights with frost, so the increased temperatures may have more important effects on insect abundance, not actual survival of the birds. Suitable habitats include sheltered coastal areas, but they also use urban and suburban sites, especially near water, sewage treatment plants (where warmer water means an abundance of insects) woodlands and hedgerows. The Chiffchaff on the top shot, fed low on the marginal vegetation around an urban lake, East Park, Hull on 2/12/2013.

It's complicated...
The Chiffchaff is part of a species complex and its taxonomy has changed quite a lot in recent years. What was considered a single species now includes the Canarian and Iberian Chiffchaff, previously races now elevated to species level. The Chiffchaff proper is now divided into various races: the nominal race, collybita, breeds in the UK, whereas tristis or Siberian Chiffchaff, and Scandinavian Chiffchaff abietinus. The three races are very similar, differing in the tone of the plumage and subtle morphological features, and more notably, song and calls. Given this and the occurrence of intermediate forms between some of the species makes them very difficult to identify in the field.

Residents, migrants and winter visitors
It appears that the bulk of the UK breeding population of Chiffchaffs, belonging to the subspecies collybita, migrate south in the winter. In addition, there is very strong passage through the UK, especially in the autumn, mainly of northern European birds moving to winter quarters in the Iberian Peninsula, and Northern and Western Africa and these include a sprinkling of abietinus and tristis individuals. What about the winter birds? Where do they come from?

Ringing and DNA analysis
Ringing recaptures in the winter don't include locally breeding Chiffchaffs, suggesting that the winter birds are also migrants. Most of them appear to belong to the collybita subspecies, which breeds in western Europe and Southern Scandinavia. Recoveries of birds ringed in winter in Southern England have been made subsequently in N England and Denmank during the breeding season. Recently, DNA analysis has come to the rescue. It is possible to obtain enough material for DNA analysis from tiny feathers, dislodged when ringing birds. Sending these feathers for DNA analysis allows to discriminate between subspecies, as they have distinct DNA. Individuals captured in the south of England include some Siberian chiffchaffs confirmed by DNA. Over 20% of ringed wintering Chiffchaffs in a mixed woodland by a lough in Ireland was confirmed to be tristis and two individuals were confirmed as abietinus. So it appears that wintering Chiffchaffs include individuals from all three European subspecies, that once finding suitable locations during migration they settle for a while to feed. Ringing recaptures have found the same individuals wintering repeated years in the same location. Another intriguing feature of wintering Chiffchaffs is that males, which are larger than females, were the bulk of wintering chiffchaffs in the past, although females are increasingly wintering. There is still so much to learn on Chiffchaff migration, taxonomy and evolution.

More information

Clement, P. 1995. The Chiffchaff. Hamlyn Species Guides. pp126.

Clement, P., Helbig, A. J. and Small, B. 1998. Taxonomy and identification of chiffchaffs in the Western Palearctic. Br. Birds 91, 361–375.

Birds in Cheshire and the Wirral. A breeding and wintering atlas. Wintering Chiffchaffs. 

O’Mahony, B., Farrer, D. and Collinson, M. 2008. Genetic identity of wintering Common Chiffchaffs Phylloscopus collybita trapped in County Kerry in 2015. Ir. Birds 

Ray Meads Ringing Group A review of our wintering chiffchaffs.
Simms, E. 1985. British Warblers. The New Naturalist Series. Collins. 432 pp.

What are goldfinches doing on tree branches?

At this time of year I often see Goldfinches appearing to feed high on tree branches, often sycamores. It is hard to see what exactly they are doing, but definitely there are no seeds involved, it looks like they are pecking the bark or possibly lichens. I checked and they were not holding seeds under their feet (as they often do) or wiping their bills clean. They appear to peck the surface of the branch. I managed to photograph one that appears to lift and hold a bit of bark with lichen on top (top shot).

I am very curious to know what they are doing. During the breeding season Goldfinches use lichen to camouflage their nests, I only see this behaviour during winter. Has anyone seen this? Please comment! Some more photos of the behaviour follow.

3/12/2017.

25/01/2018

25/01/2018. This individual stayed pecking the end of this branch for quite a while.

16/11/2016

UPDATE 11/1/20
I found out a chorusing Goldfinch flock in a local park today. Several of them were feeding on branches in two lime trees. I managed a couple of videos showing how they target areas at the end of broken branches and deftly peel bark and wood, discarding bits. If they are feeding on the bark or wood itself or on some invertebrates living in it is unclear.

Mute Swan acoustic communication

I've said it before, Mute Swans are not really mute, they have a repertoire of calls and sounds. They also use movements of their head and neck to communicate. There is a quick head bowing movement, and a courting movement in which the feathers at the top half of the neck are ruffled, while those at the bottom are not, while the swan bows his head alternatively to left and right; and the aggressive 'busking stance' of the territorial male. In this post I focus on sound communication. I have trawled through Xenocanto, an encyclopaedic website with thousands of bird calls, to compile the following selection of Mute Swan sounds. I was prompted to write the post after listening to an odd distant call, which I didn't recognise to start with, and I was surprised to find out that it was a mute swan's. I heard the call again today, young swans calling to a swan flying overhead. It is not straightforward to distill the 'meaning' of a call, but the context can helps.

1. Snort

A short grunt or snort, often uttered when the swans are relaxed, and appears as a contact call while feeding, between members of a family, but also a greeting call. A lone young swan snorted when I walked by a lake near him.

2. Begging call

A soft repeated whistle typical call of young. Mute Swans don't 'feed' their young by putting food in their bills, but they help the young to obtain their food by paddling with their feet to  disturb the sediments or by pulling underwater plants. Even fully grown immature swans carry on whistling.

  3. Wing-beat Sound

This loud sound produced by the mute swan's wing-beats is really eery, especially when it takes you unawares until you realise that is coming from a swan. It carries very well and appears to replace the trumpeting contact calls of other swans.

4. Hissing

An aggressive sound, usually uttered by the female she has small young and a danger (human, dog) approaches. The bill is open and the swan adopts an upright stance towards the danger source.

5. Contact all

A two-note contact call reminiscent of other swan's flight call. I heard this call when a pair was separated, maybe during landing (one ended up in the river, the other one in a lake). Also when a Mute Swan flew over a family, the juveniles used this distance call.

6. Courtship calls

A range of snorts and grunts during courtship.

 Have I missed any? Let me know in the comments.