Eurasian Crag Martin
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The Eurasian crag martin or just crag martin (''Ptyonoprogne rupestris'') is a small passerine bird in the swallow family. It is about long with ash-brown upperparts and paler underparts, and a short, square tail that has distinctive white patches on most of its feathers. It breeds in the mountains of southern Europe, northwestern Africa and across the
Palearctic The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sibe ...
. It can be confused with the three other species in its genus, but is larger with brighter tail spots and different plumage tone. Many European birds are resident, but some northern populations and most Asian breeders are migratory, wintering in northern Africa, the Middle East or India. The Eurasian crag martin builds a nest adherent to the rock under a cliff overhang or increasingly onto a man-made structure. It makes a neat half-cup
mud A MUD (; originally multi-user dungeon, with later variants multi-user dimension and multi-user domain) is a Multiplayer video game, multiplayer Time-keeping systems in games#Real-time, real-time virtual world, usually Text-based game, text-bas ...
nest with an inner soft lining of feathers and dry grass. Nests are often solitary, although a few pairs may breed relatively close together at good locations. Two to five brown-blotched white eggs are incubated mainly by the female, and both parents feed the chicks. This species does not form large breeding colonies, but is gregarious outside the breeding season. It feeds on a wide variety of insects that it catches with its beak while flying near to cliff faces or over streams and alpine meadows. Adults and young may be hunted and eaten by birds of prey or corvids, and this species is a host of blood-sucking mites. With its large, expanding range and large population, there are no significant conservation concerns involving the species. This bird is closely related to the other three crag martins which share its genus, and has sometimes been considered to be the same species as one or more of them, although it appears that there are areas where two species' ranges overlap without hybridisation occurring. All four ''Ptyonoprogne'' crag martins are quite similar in behaviour to other
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
swallows that build mud nests, and are sometimes subsumed into the larger genus '' Hirundo'', but this approach leads to inconsistencies in classifying several other genera, particularly the
house martin ''Delichon'' is a small genus of passerine birds that belongs to the swallow family and contains four species called house martins. These are chunky, bull-headed and short-tailed birds, blackish-blue above with a contrasting white rump, and w ...
s.


Taxonomy

The Eurasian crag martin was formally described as ''Hirundo rupestris'' by Italian naturalist Giovanni Antonio Scopoli in 1769Scopoli (1769) p. 172 and was moved to the new genus ''
Ptyonoprogne The crag martins are four species of small passerine birds in the genus ''Ptyonoprogne'' of the swallow family. They are the Eurasian crag martin (''P. rupestris''), the pale crag martin (''P. obsoleta''), the rock martin (''P.  ...
'' by German ornithologist Heinrich Gustav Reichenbach in 1850.Reichenbach (1850) plate LXXXVII figure 6 Its nearest relatives are the three other members of the genus, the
pale crag martin The pale crag martin (''Ptyonoprogne obsoleta'') is a small passerine bird in the swallow family that is resident in Northern Africa and in Southwestern Asia, east to Pakistan. It breeds mainly in the mountains, but also at lower altitudes, esp ...
, ''P. obsoleta'', the rock martin, ''P. fuligula'', and the
dusky crag martin The dusky crag martin (''Ptyonoprogne concolor'') is a small passerine bird in the swallow family. It is about long with a broad body and wings, and a short square tail that has small white patches near the tips of most of its feathers. This ma ...
, ''P. concolor''.Turner (1989) pp. 160–164 The genus name is derived from the Greek ''ptuon'' (πτύον), "a fan", referring to the shape of the opened tail, and Procne (Πρόκνη), a mythological girl who was turned into a swallow. The specific ''rupestris'' means "of rocks", from the Latin ''rupes'' "rock". There are no generally recognised subspecies. Two races, Central Asian ''P. r. centralasica'' and ''P. r. theresae'' in the
Atlas Mountains The Atlas Mountains are a mountain range in the Maghreb in North Africa. It separates the Sahara Desert from the Mediterranean Sea and the Atlantic Ocean; the name "Atlantic" is derived from the mountain range. It stretches around through Moroc ...
of Morocco, have been proposed, but the slight differences in size and colour show no consistent geographical pattern.Turner (1989) pp. 158–160 Fossils of this species have been found in
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
deposits in Bulgaria,Boev Zlatozar. (2001).
Late Pleistocene birds from the Kozarnika Cave (Montana District; NW Bulgaria)
in Delchev P., Shanov S., Benderev A. (Eds) ''Proceedings of the First national Conference on Environment and Cultural Heritage in Karst. Sofia 10–11.11.2000. Volume 1'' 113–128.
and in central France in layers dated at 242,000 to 301,000 years ago. The four ''Ptyonoprogne'' species are members of the swallow family, and are placed in the subfamily Hirundininae, which comprises all swallows and martins except the very distinctive
river martin The river martins form a distinctive subfamily Pseudochelidoninae within the swallow and martin bird family Hirundinidae. The two species are the African river martin ''Pseudochelidon eurystomina'', found in the Congo and Gabon, and the white-e ...
s. DNA studies suggest that there are three major groupings within the Hirundininae, broadly correlating with the type of nest built. The groups are the "core martins" including burrowing species like the
sand martin The sand martin (''Riparia riparia''), also known as the bank swallow (in the Americas), collared sand martin, or common sand martin, is a migratory passerine bird in the swallow family. It has a wide range in summer, embracing practically the w ...
, the "nest-adopters", which are birds like the
tree swallow The tree swallow (''Tachycineta bicolor'') is a migratory bird of the family Hirundinidae. Found in the Americas, the tree swallow was first described in 1807 by French ornithologist Louis Vieillot as ''Hirundo bicolor''. It has since been mov ...
that use natural cavities, and the "mud nest builders". The ''Ptyonoprogne'' species construct an open mud nest and therefore belong to the last group; '' Hirundo'' species also build open nests, ''
Delichon ''Delichon'' is a small genus of passerine birds that belongs to the swallow family and contains four species called house martins. These are chunky, bull-headed and short-tailed birds, blackish-blue above with a contrasting white rump, and w ...
'' house martins have a closed nest, and the '' Cecropis'' and ''
Petrochelidon ''Petrochelidon'' is a genus of birds known as cliff-nesting swallows. The genus name ''Petrochelidon'' is from the Ancient Greek words ''petros'', "rock", and ''khelidon'', "swallow". The genus includes all of the five species of birds commonly ...
'' swallows have retort-like closed nests with an entrance tunnel. ''Ptyonoprogne'' is closely related to the larger swallow genus ''Hirundo'' into which it is often subsumed, but a DNA analysis showed that an enlarged genus ''Hirundo'' should logically contain all the mud-builder genera, including the ''
Delichon ''Delichon'' is a small genus of passerine birds that belongs to the swallow family and contains four species called house martins. These are chunky, bull-headed and short-tailed birds, blackish-blue above with a contrasting white rump, and w ...
'' house martins, a practice which few authorities follow. Although the nests of the ''Ptyonoprogne'' crag martins resemble those of typical ''Hirundo'' species like the barn swallow, the research showed that if ''Delichon'', '' Cecropis'' and ''
Petrochelidon ''Petrochelidon'' is a genus of birds known as cliff-nesting swallows. The genus name ''Petrochelidon'' is from the Ancient Greek words ''petros'', "rock", and ''khelidon'', "swallow". The genus includes all of the five species of birds commonly ...
'' are split from ''Hirundo'', ''Ptyonoprogne'' should also be treated as a separate genus.


Description

The Eurasian crag martin is long with a wingspan, and weighs an average . It has ash-brown upperparts and paler underparts, and has a broader body, wings and tail than any other European swallow. The tail is short and square, with white patches near the tips of all but the central and outermost pairs of feathers. The underwing and undertail
coverts A covert feather or tectrix on a bird is one of a set of feathers, called coverts (or ''tectrices''), which, as the name implies, cover other feathers. The coverts help to smooth airflow over the wings and tail. Ear coverts The ear coverts are sm ...
are blackish, the eyes are brown, the small bill is mainly black, and the legs are brownish-pink. The sexes are alike, but juveniles have buff-brown tips to the plumage of the head, upperparts and wing coverts. This species can be distinguished from the
sand martin The sand martin (''Riparia riparia''), also known as the bank swallow (in the Americas), collared sand martin, or common sand martin, is a migratory passerine bird in the swallow family. It has a wide range in summer, embracing practically the w ...
by its larger size, the white patches on the tail, and its lack of a brown breast band. Where the range overlaps with that of another ''Ptyonoprogne'' species, the Eurasian crag martin is darker, browner and 15% larger than the rock martin,Snow & Perrins (1998) pp. 1059–1061 and larger and paler, particularly on its underparts than the dusky crag martin.Grimmett ''et al.'' (2002) p. 268 The white tail spots of the Eurasian crag martin are significantly larger than those of both its relatives. The crag martin's flight appears relatively slow for a swallow. Rapid wing beats are interspersed with flat-winged glides, and its long flexible primaries give it the agility to manoeuvre near cliff faces. The average migration flight speed has been measured at 9.9 m/s (32.5 ft/s), less than the roughly 11 m/s (36 ft/s) typical for hirundines, but the data is limited. The bird often flies high, and shows the white spots as it spreads its tail. The vocalisations include short high ''pli'', and ''piieh'' and ''tshir'' calls resembling those of the linnet and the
house martin ''Delichon'' is a small genus of passerine birds that belongs to the swallow family and contains four species called house martins. These are chunky, bull-headed and short-tailed birds, blackish-blue above with a contrasting white rump, and w ...
, respectively.Mullarney ''et al.'' (1999) p.240


Distribution and habitat

The Eurasian crag martin breeds in mountains from Iberia and northwesternmost Africa through southern Europe, the Persian Gulf and the Himalayas to southwestern and northeastern China. Northern populations are migratory, with European birds wintering in north Africa, Senegal, Ethiopia and the
Nile Valley The Nile, , Bohairic , lg, Kiira , Nobiin: Áman Dawū is a major north-flowing river in northeastern Africa. It flows into the Mediterranean Sea. The Nile is the longest river in Africa and has historically been considered the longest rive ...
, and Asian breeders going to southern China, the Indian subcontinent and the Middle East. Some European birds stay north of the Mediterranean, and, like martins in warmer areas such as India, Turkey and Cyprus, just move to lower ground after breeding. The breeding range is bounded by the July isotherm, and wintering areas need a temperature of about for enough insect food to be available. This is a rare species any distance north of its breeding areas. For example, there are only 12 records from the UK, none from Ireland, and the first record for Sweden was reported as recently as 1996. South of its normal wintering range, it has occurred as a vagrant in The Gambia. Retrieved 26 March 2010 In 2022, the first nesting of the species was recorded on the territory of the
Slovak Republic Slovakia (; sk, Slovensko ), officially the Slovak Republic ( sk, Slovenská republika, links=no ), is a landlocked country in Central Europe. It is bordered by Poland to the north, Ukraine to the east, Hungary to the south, Austria to the s ...
in the
Malá Fatra National Park The Malá Fatra National Park (Slovak: ''Národný park Malá Fatra'') is a national park in the northern part of the Malá Fatra mountains called Krivánska Malá Fatra. It has an area of 226.3 km2 (87.37 mi2) and a 232.62 km2 (89. ...
. Crag martins breed on dry, warm and sheltered cliffs in mountainous areas with crags and gorges. The typical altitude is but breeding occurs up to in Central Asia. The Eurasian crag martin's choice of nest sites is very similar to that of
Savi's pipistrelle Savi's pipistrelle (''Hypsugo savii'' sometimes classified as ''Pipistrellus savii'') is a species of vesper bat found across North West Africa, the Mediterranean region and the Middle East. It feeds at night on flying insects. In the summer it ro ...
, ''Hypsugo savii''; the bird and the bat often breed in the same locations and have almost identical ranges in Europe. Retrieved 26 March 2010 In South Asia, migrant Eurasian birds sometimes join with flocks of the dusky crag martin and roost communally on ledges of cliffs or buildings.Ali & Ripley (1986) pp. 53–54 The largest known wintering roost of Eurasian crag martins has been recorded at the
Gorham's Cave Gorham's Cave ( es, Cueva de Gorham, ) is a sea-level cave in the British overseas territory of Gibraltar. Though not a sea cave, it is often mistaken for one. Considered to be one of the last known habitations of the Neanderthals in Europe, the ...
Complex in
Gibraltar ) , anthem = " God Save the King" , song = " Gibraltar Anthem" , image_map = Gibraltar location in Europe.svg , map_alt = Location of Gibraltar in Europe , map_caption = United Kingdom shown in pale green , mapsize = , image_map2 = Gib ...
. The caves were home to a maximum of 12,000 birds during the 2020-2021 winter season; 1-2% of the entire European population of Eurasian crag martins.


Behaviour


Breeding

Crag martin pairs nest alone or in small colonies, usually containing fewer than ten nests. Nests are on average apart and each pair aggressively defends its breeding territory against other crag martins and most other bird species. Nesting takes place from May to August, and usually two broods are raised. The nest, built by both adults, is an open half cup made of mud and lined with soft material such as feathers or dry grass.Hume (1890) pp
180–183
/ref> It is constructed under an overhang on a rock cliff face, in a crevice or cave, or on a man-made structure. It takes one to three weeks to build and is re-used for the second brood and in subsequent years. The clutch is two to five eggs with an average of three. The eggs are white with brownish blotches particularly at the wide end, and average with a weight of . The eggs are incubated mainly by the female for 13–17 days to hatching, and the chicks take another 24–27 days to fledge. Both parents feed the chicks bringing food every two to five minutes, and the young are fed for 14–21 days after fledging. With such frequent feeding rates the adults mainly forage in the best hunting zones in the immediate vicinity of the nest, since the further they fly to forage the longer it would take to bring food to the chicks in the nest. In an Italian study, the hatching rate was 80.2 percent, and the average number of fledged young was 3.1. The crag martin has over the last few decades increasingly used houses and other man-made sites to nest. This greater availability of breeding sites has enabled the species to expand its range, but it is possible that this will lead to competition with other hirundines, such as the barn swallow and common house martin, which also use artificial nest sites. An Italian study showed that, as with other aerial feeders, the start of breeding was delayed by cold or wet weather, but this had no influence on the clutch size nor on the number of fledged young. Unexpectedly, it was found that once the eggs had hatched there was a negative relationship between temperature and the number of fledged young. The authors suggested that hot weather dried up the small rivers where the parents found food. Colony size did not influence the laying date, the clutch size or the number of successfully fledged young, but this species does not form large colonies anyway.


Feeding

The Eurasian crag martin feeds mainly on insects caught in its beak in flight, although it will occasionally take prey items off rocks, the ground, or a water surface. When breeding, birds often fly back and forth near to a rock face hunting for insects, feeding both inside and outside the nesting territory. At other times, they may hunt while flying above streams or alpine meadows. The insects taken depend on what is locally available and may include flies, ants, aerial spiders, and beetles. Aquatic species such as
stoneflies Plecoptera is an order of insects, commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to be one of the mo ...
, caddisflies and
pond skaters The Gerridae are a family of insects in the order Hemiptera, commonly known as water striders, water skeeters, water scooters, water bugs, pond skaters, water skippers, or water skimmers. Consistent with the classification of the Gerridae as tr ...
appear to be important in at least Spain and Italy. Unlike other hirundines, these birds feed close to their breeding sites, and may be locally vulnerable to fluctuations in insect availability. This martin is gregarious outside the breeding season, and may form sizeable flocks where food is abundant. Cliff faces generate
standing wave In physics, a standing wave, also known as a stationary wave, is a wave that oscillates in time but whose peak amplitude profile does not move in space. The peak amplitude of the wave oscillations at any point in space is constant with respect ...
s in the airflow which concentrate insects near vertical areas. The crag martin exploits the area close to the cliff when it hunts, relying on its high manoeuvrability and ability to perform tight turns.


Winter roost fidelity

Eurasian crag martins are known to form large roosts in winter, with the largest known roost being the
Gorham's Cave Gorham's Cave ( es, Cueva de Gorham, ) is a sea-level cave in the British overseas territory of Gibraltar. Though not a sea cave, it is often mistaken for one. Considered to be one of the last known habitations of the Neanderthals in Europe, the ...
Complex in Gibraltar. A study carried out at these caves and published in
Scientific Reports ''Scientific Reports'' is a peer-reviewed open-access scientific mega journal published by Nature Portfolio, covering all areas of the natural sciences. The journal was established in 2011. The journal states that their aim is to assess solely th ...
in 2021, revealed that birds showed very high fidelity towards individual caves within and between years. Mark-recapture showed there was over a 90% chance of recapturing birds at the caves where they were first caught. The condition of birds from different caves suggests differences in roost quality which correlates to the fitness of Eurasian crag martins and, ultimately, survivorship.


Predators and parasites

This species is occasionally hunted by the
peregrine falcon The peregrine falcon (''Falco peregrinus''), also known as the peregrine, and historically as the duck hawk in North America, is a Cosmopolitan distribution, cosmopolitan bird of prey (Bird of prey, raptor) in the family (biology), family Falco ...
, which shares its mountain habitat, and during its migration over the Himalayas, it is reported to be subject to predation by crows. Common kestrels,
Eurasian sparrowhawk The Eurasian sparrowhawk (''Accipiter nisus''), also known as the northern sparrowhawk or simply the sparrowhawk, is a small bird of prey in the family Accipitridae. Adult male Eurasian sparrowhawks have bluish grey upperparts and orange-barred ...
s,
Eurasian jay The Eurasian jay (''Garrulus glandarius'') is a species of passerine bird in the crow family Corvidae. It has pinkish brown plumage with a black stripe on each side of a whitish throat, a bright blue panel on the upper wing and a black tail. The ...
s and common ravens are also treated as predators and attacked by repeated dives if they approach nesting cliffs. Despite the general aggressiveness of the crag martin, it tolerates
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
common house martins, perhaps because the large numbers of that highly colonial species provide an early warning of predators. The crag martin is a host of blood-sucking mites of the genus ''
Dermanyssus Dermanyssus is a genus of mites in the family Dermanyssidae. The etymology of the word ''Dermanyssus'' is from the Greek: ''derma'', meaning skin, and ''nyssein'', meaning bite. Genera *''Dermanyssus'' **''Dermanyssus americanus'' Ewing, 192 ...
'', including '' D. chelidonis'', and of the nasal mite '' Ptilonyssus ptyonoprognes''. Retrieved 30 March 2010 Two new species of parasites were first discovered on this martin, the fly ''
Ornithomya rupes ''Ornithomya'' are genus of biting flies in the family of louse flies, Hippoboscidae. There are 29 known species. All species are parasites of birds. Distribution ''Ornithomya'' are found worldwide with the exception of Antarctica, however the l ...
'' in Gibraltar and the flea ''
Ceratophyllus nanshanensis ''Ceratophyllus nanshanensis'' is a species of flea in the family Ceratophyllidae Ceratophyllidae is a family (biology), family of fleas. Its members are Parasitism, parasites of mainly rodents and birds. It contains two subfamilies, one contai ...
'' from China.


Status

The European population of the Eurasian crag martin is estimated to be 360,000–1,110,000 individuals, including 120,000–370,000 breeding pairs. A rough estimate of the worldwide population is 500,000–5,000,000 individuals, with Europe hosting between one-quarter and one-half of the total. The population is estimated to be increasing following a northward expansion, which may be partly due to increased use of man-made structures as nest sites. Expansions of the range have been reported in Austria (where motorway bridges are used as nest sites), Switzerland, the former Yugoslavia, Romania, and Bulgaria. With its very large range and high numbers, the Eurasian crag martin is not considered to be threatened, and it is classed as Least Concern on the IUCN Red List.


Citations


Cited texts

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External links


Ageing and sexing (PDF; 2.8 MB) by Javier Blasco-Zumeta & Gerd-Michael Heinze
{{Taxonbar, from=Q213216 Ptyonoprogne Birds of Eurasia Birds of North Africa Birds described in 1769 Taxa named by Giovanni Antonio Scopoli