Veli-Matti Pakanen, Kari Koivula and colleagues have been studying Finnish Dunlin for nearly twenty years. These are schinzii birds that breed in coastal grazing meadows around the Baltic Sea. Several papers have been published, as you will see below, some of which are based on information collected using geolocators attached to leg-flags. Dunlin numbers in coastal Finland are dropping quickly, so this research is important to the conservation of the species. In a 2020 paper, Veli-Matti and colleagues ask whether intensive geolocator-based studies are having a negative effect on individual birds in this already-declining study population.
Up to ten races of Dunlin have been identified, which together encircle the globe. Most Dunlin spend the non-breeding season in the northern hemisphere and all migrate north in spring. The breeding and wintering ranges of the various races are summarised at the end of this blog. Here, the focus is on Baltic schinzii Dunlin, a small part of a race that largely winters in coastal North Africa and breeds in southern Scandinavia, Northern Europe, the British Isles, Iceland and southern Greenland.
According to Wetlands International, there were between 4.3 and 6.8 million Dunlin worldwide in 2015, with about one million breeding birds within Europe (BirdLife International). Whilst it is acknowledged that numbers are declining, the large range of the species means that it is still considered to be of ‘least concern’.
Different populations are faring differently. In his description of the changing Dunlin distribution in European Breeding Bird Atlas 2, John Calladine points out that there have been major losses across Europe, including Britain & Ireland, and that Baltic populations ‘are now considered threatened’. The Baltic schinzii population was most recently estimated as between 500 and 640 pairs – less than a fifth of the estimate in the 1980s. John highlights gaps in knowledge that research by the team in Finland and other groups elsewhere are helping to fill.
Migration and survival of schinzii
In pre-geolocator days, Ole Thorup and colleagues analysed recovery information available for Dunlin breeding around the Baltic, using 40 years of ringing data from Finland, Sweden, Denmark and Germany. At that stage there were only six mid-winter recoveries in N and NW Africa. The analysis emphasised the importance of wintering and stop-over sites in the Baltic, the Wadden Sea, SE and S England, the Atlantic seaboard of France, and the Iberian Peninsula. Paper in Ardea (2009).
Pakanen et al investigated changes in the survival rates of schinzii Dunlin nesting in Denmark, based on ringing and recaptures of a marked population. They found that annual apparent survival rates dropped from 0.817 to 0.650 between 1990 and 2006, equivalent to a doubling of the chance of dying in any given year. Paper in Bird Study (2016). The importance of monitoring survival rates is discussed in the WaderTales blog: Measuring shorebird survival.
The use of geolocators enabled Veli-Matti Pakanen to add more detail to the migration story in the 2018 paper, Migration strategies of the Baltic dunlin: rapid jump migration in the autumn but slower skipping type spring migration. He and his colleagues showed that autumn migration is faster than spring migration, characterised by fewer stationary periods, shorter total stopping time and faster flight. The Wadden Sea was found to be an important autumn staging area for all of the tagged birds. Some birds stopped once more before reaching Mauritania. On spring migration, more sites were visited on the way north. The important conservation message from the paper is that Baltic Dunlin may be especially vulnerable to rapid environmental changes at their staging and wintering areas. (In Travel advice for Sanderling there is a suggestion that annual survival is relatively low for birds that winter in Mauritania).
In a 2015 paper in Ornis Fennica, Pakanen et al reported on the results of a single-year analysis of survival rates, concluding that there were no strong effects of leg-flagged geolocators on return rates or reproduction in schinzii Dunlin. However, they did suggest that “long term evaluations that capture the full suite of environmental conditions and assess impact on brood care are needed”. This is a question which leads us neatly on to the 2020 paper: Survival probability in a small shorebird decreases with the time an individual carries a tracking device.
Long-term tracking of individuals
To understand the range of pressures that migratory birds face, one needs to know where individuals spend their time, as was demonstrated in Spoon-billed Sandpiper: Track & Trace and Teenage Waders. Waders of a range of species have been wearing geolocators for ten or more years now, either the same devices on birds which have evaded capture, or a series of tags, when data have been downloaded and replacement tags fitted. These long-term surveys are developing our understanding of the repeatability of migratory behaviour and how birds deal with variable weather patterns, but is there a long-term cost to the individual birds that are tasked with finding out this important information?
Effects of tracking devices on survival are generally considered to be small. However, most studies to date have focused on birds that were caught in one breeding season and recaptured in the following season, to retrieve the geolocator and download the data. In their 2020 paper, Veli-Matti Pakanen and colleagues were able to investigate the possible accumulation of negative effects when individuals have carried the tracking devices for longer periods. Survival rates for tagged birds were compared with 338 colour-ringed birds that were followed for all or part of the period 2002 to 2018.
In the summers of 2013 & 2014, fifty-three adult schinzii Dunlin were fitted with leg-flag mounted geolocators, with a mass equivalent to 1.5 – 2.0% of body-mass. Of these birds, 17 tags were retrieved after one year and 9 after two years. Other marked birds could not be caught and carried their tags for longer periods (3 or 4 years). The research team found that Dunlin carrying a geolocator had reduced chances of survival. Their models suggest that annual survival of colour-ringed males was 0.813. For a bird that carried a geolocator for a year, annual survival probability declined to 0.748 and to 0.581 for birds that carried the geolocator for at least 2 years. Their data suggest that the reduction in survival rates was greater for females than males, even though females are larger than males.
In a thorough Discussion, the authors consider reasons why tags on small waders may be reducing survival, either through ongoing stresses, impacting on things such as feeding efficiency and the energy needed during migration, or because the extra burden means that tagged birds find it harder to cope with occasional periods of tough environmental conditions. They comment on the condition of the skin under removed geolocators – something that other researchers night want to look out for.
As anyone studying breeding waders will know, nest-trapping to retrieve tags is not easy, especially if adults lose their clutches when incubation has only just started, due to flooding, predation etc. Birds may end up carrying tags for longer than intended. The authors “recommend that the detrimental effects of tagging may be avoided by developing attachment methods that are automatically released after one year, e.g. biodegradable materials”.
Balancing costs and benefits
The results from the Pakanen study of long-term survival suggest that requiring a small wader to carry a geolocator for several years may have an impact on survival. As in all mark-recapture studies, researchers are urged to assess the costs to the individual when seeking to understand what might be affecting the viability of a population.
Four previous WaderTales blogs have discussed tag effects:
- Are there costs of wearing a geolocator? (a multi-species analysis of leg-mounted geolocators)
- Leg flags and nest success (four small species of American shorebird)
- Green Sandpipers and Geolocators (geolocators mounted on harnesses)
- Flagging up potential problems (flag/geolocator effects in Common Sandpipers)
Details of the Dunlin tagging effects study:
Survival probability in a small shorebird decreases with the time an individual carries a tracking device.
Veli-Matti Pakanen, Nelli Rönkä, Thomson Robert Leslie, Donald Blomqvist, Kari Koivula. Journal of Avian Biology (2020): https://doi.org/10.1111/jav.02555
Four other papers relating to this Finnish Dunlin study
Grazing by cattle is an important management tool in coastal meadows. Pakanen et al studied the impact of trampling on artificial nests and concluded that even recommended stocking rates were too high for chick numbers that could deliver a sustainable population. Paper in Biodiversity and Conservation (2011). (Redshank on British estuaries are similarly vulnerable to trampling – see Big Foot and the Redshank Nest).
In a follow-up paper in 2016, Pakanen et al concluded that Dunlin populations could be sustained in grazed coastal meadows as long as there was no active grazing before 19 June. Meadows with grazing cattle attracted breeding birds but there was insufficient breeding success for sustainability. Paper in Ecology & Evolution (2016).
Dunlin are strongly philopatric, with both male and female chicks recruiting to suitable habitat close to natal sites. In a paper in Ibis (2017), Pakanen et al show that natal dispersal of Dunlin is strongly linked to the size of their natal site and how isolated the site is. They suggest that inbreeding may be avoided by creating a network of suitably sized patches (20–100 ha sites), no more than 20 km apart from each other. These may work as stepping stones for recruiting individuals. These results are corroborated by a 2021 microsatellite study in BMC Ecology and Evolution which shows genetic differentiation and isolation by distance within the Baltic Dunlin population.
Up to ten races
In western Europe we see three races of Dunlin – alpina, arctica and schinzii. Wintering birds are almost exclusively of the alpina race, which head north and east to northern Scandinavia, Russia and Siberia in spring. The other northern race is arctica, a very small Dunlin that breeds in low numbers in NE Greenland and possibly Spitzbergen. Schinzii has a very large breeding range, spanning the Baltic, southern Scandinavia, Northern Europe, the British Isles, Iceland and SE Greenland. There is huge variation in the timing of breeding of schinzii, as birds do not return to breeding sites in SE Greenland until the end of May, at the same time as schinzii Dunlin being studied by Veli-Matti Pakanen and colleagues in Finland will have their first young chicks.
Further east, centralis Dunlin replace alpina. Many of these birds use the Central Asian Flyway. Further east still, we find sakhalina that use the East Asian/Australasian Flyway (EAA). Two other subspecies have been identified breeding within the EAA Flyway, the more southerly kistchinski birds and actites, which breeds furthest south, on the Russian island of Sakhalin, on a similar latitude to the UK.
It is generally accepted that there are three Dunlin subspecies in North America. In autumn, arcticola head west from northern Alaska and NW Canada and follow the EAA Flyway, pacifica fly south along the Pacific coast from SW Alaska and hudsonia migrate from central northern Canada using the Atlantic Americas Flyway.
WaderTales blogs are written by Graham Appleton (@GrahamFAppleton) to celebrate waders and wader research. Many of the articles are based on published papers, with the aim of making shorebird science available to a broader audience.
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