American Cliff Swallow
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The cliff swallow or American cliff swallow (''Petrochelidon pyrrhonota'') is a member of the passerine bird family
Hirundinidae The swallows, martins, and saw-wings, or Hirundinidae, are a family of passerine songbirds found around the world on all continents, including occasionally in Antarctica. Highly adapted to aerial feeding, they have a distinctive appearance. The ...
, the swallows and martins. The generic name ''Petrochelidon'' is derived from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
''petros'' meaning "rock" and ''khelidon'' "swallow", and the specific name ''pyrrhonota'' comes from ''purrhos'' meaning "flame-coloured" and ''-notos'' "-backed". Cliff swallows are extremely social songbirds that can be found in large nesting colonies reaching over 2,000
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materia ...
s. They are frequently seen flying overhead in large
flocks Flocking is the behaviour exhibited when a group of birds, called a flock, are foraging or in flight. Computer simulations and mathematical models that have been developed to emulate the flocking behaviours of birds can also generally be applie ...
during
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
, gracefully foraging over fields for flying insects or perching tightly together on a wire preening under the sun. Cliff swallows build gourd-shaped nests made from mud with small entrance holes. They build their nests tightly together, on top of one another, under
bridge A bridge is a structure built to span a physical obstacle (such as a body of water, valley, road, or rail) without blocking the way underneath. It is constructed for the purpose of providing passage over the obstacle, which is usually somethi ...
s or alongside mountain cliffs. Living in large populations, these aerial
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
s use extensive vocalizations to communicate warnings or food availability to the other individuals.


Description

The cliff swallow's average body length is , and they have short legs and small bills with relatively long pointed wings. Adult cliff swallows have an overall dark brownish
plumage Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...
covering both their back and wings, and they have a characteristic white forehead, rich red-coloured cheeks with a dark throat, basic white underparts and a buffy-coloured rump. In good lighting conditions, their crowns and mantle feathers are
iridescent Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, butterfl ...
. The Northern population is slightly larger in body size and also differs in facial markings from the Mexican population of cliff swallows, which have a chocolate-brown patch on their foreheads. The male and female have identical plumage, therefore sexing them must be done through
palpation Palpation is the process of using one's hands to check the body, especially while perceiving/diagnosing a disease or illness. Usually performed by a health care practitioner, it is the process of feeling an object in or on the body to determine ...
of the
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
. During the breeding season, the males will have a harder cloaca that is more pronounced because the
seminal vesicle The seminal vesicles (also called vesicular glands, or seminal glands) are a pair of two convoluted tubular glands that lie behind the urinary bladder of some male mammals. They secrete fluid that partly composes the semen. The vesicles are 5†...
s are swollen. In addition, during incubation females will lose feathers on their lower breast to create a warm patch for sitting on their eggs. Cliff swallows are similar in body plumage colouring to the related
barn swallow The barn swallow (''Hirundo rustica'') is the most widespread species of swallow in the world. In fact, it appears to have the largest natural distribution of any of the world's passerines, ranging over 251 million square kilometres globally. ...
species but lack the characteristic fork-shaped tail of the barn swallow prominent during flight. The cliff swallows have a square-shaped tail. Juvenile cliff swallows have an overall similar body plumage colouring to the adults, with paler tones. The juveniles lack the iridescent adult plumages, and their foreheads and throats appear speckled white. The juvenile cliff swallows’ white forehead and throat markings have high variance between unrelated individuals compared with those from the same clutch. These distinctive white facial markings disappear during maturity following their complex-basic moult pattern, because their pre-formative plumage is different from the basic plumage. The pre-formative facial plumage has been suggested as a possible way for parents nesting in large colonies to recognize their chicks.


Taxonomy

The cliff swallow belongs to the largest order and dominant avian group – ''
Passeriformes A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
''. They are the perching birds, or the passerines. All the bird species in this order have 4 toes, 3 pointing forward and one pointing backwards ( anisodactylous), that enable them to perch with ease. The sub-order that the cliff swallow belongs to is ''Oscines'' (or Passeri), for the songbirds. The family that encompasses approximately 90 species of swallows and martins, ''Hirundinidae'', includes birds that have small stream-lined bodies made for great agility and rapid flight. Furthermore, those in the family Hirundinidae have short-flat bills for their largely insectivorous diets, small feet because they spend much of their time in flight and long wings for energy-efficient flight. There are 5 subspecies of cliff swallow distinguished on the basis of plumage colour, body size and distribution – ''Petrochelidon pyrrhonota pyrrhonota'', ''P. p. melanogaster'', ''P. p. tachina'', ''P. p. hypopolia'', ''P. p. ganieri''. In addition, three core genera of ''hirundo'' were established on the basis of molecular studies: ''Hirundo sensu stricto'', containing the barn swallow; ''Cecropis'', containing the red-rumped swallow; and ''Petrochelidon'', containing the cliff swallow. The genetic tests deemed ''Petrochelidon'' and ''Cecropis'' sister to each other and both closest to ''Delichon'', the house martins. Finally, the cave swallow was identified as the nearest living relative in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
of the cliff swallow. The cave swallow has a similar plumage to the cliff swallow; however, the former has a dark cap and pale throat, and also a much smaller distribution in North America, most likely due to a decline in suitable cave sites.


Habitat and distribution

As their name suggests, throughout history the cliff swallows concentrated their nesting colonies along mountain cliffs, primarily by the western North American coast. Today, with the development of highways, concrete bridges and buildings this adaptable bird species is rapidly adjusting their common nesting sites, with populations expanding further east and building their mud nests on these concrete
infrastructure Infrastructure is the set of facilities and systems that serve a country, city, or other area, and encompasses the services and facilities necessary for its economy, households and firms to function. Infrastructure is composed of public and priv ...
s. Thus, the cliff swallow's breeding range includes large areas across
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
and the
United States of America The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territo ...
, excluding some Southern and Northern areas. The majority of nesting colonies are situated in close proximity to fields,
pond A pond is an area filled with water, either natural or artificial, that is smaller than a lake. Defining them to be less than in area, less than deep, and with less than 30% emergent vegetation helps in distinguishing their ecology from th ...
s and other
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s that would hold a large variety of flying insect populations to sustain their energy requirements during the
breeding season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
. The cliff swallows' wintering grounds have been recorded as
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
n countries, such as Southern
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
,
Uruguay Uruguay (; ), officially the Oriental Republic of Uruguay ( es, RepĂșblica Oriental del Uruguay), is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast; while bordering ...
, and parts of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, RepĂșblica Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. However, their behavior and populations have yet to be extensively studied on their wintering grounds leaving room for new information about this species. The cliff swallows are long-distance day-migrants that generally travel along the North American coast lines. The Eastern populations travel through
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
, and the Western populations through Mexico and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
down to their destinations. Flocks containing large numbers of cliff swallows have been recorded migrating together, but whether they stay together or disperse to different locations is unknown.


Swallows of Capistrano

The cliff swallow is famous for their regular migration to
San Juan Capistrano, California San Juan Capistrano (Spanish for "St. John of Capistrano") is a city in Orange County, California, located along the Orange Coast. The population was 34,593 at the 2010 census. San Juan Capistrano was founded by the Spanish in 1776, when St. ...
.


Behavior

Cliff swallows live in a colonial lifestyle during the breeding season, composed of a large number of pairs per nesting site. This group-style life can present the birds with some benefits and disadvantages; valuable information can be shared through group learning about food location and
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
preferences, but it is also much easier to transmit parasites or
disease A disease is a particular abnormal condition that negatively affects the structure or function of all or part of an organism, and that is not immediately due to any external injury. Diseases are often known to be medical conditions that a ...
s when living in close proximity. The cliff swallows have an unusually large parasite community that includes
ectoparasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
, ticks,
flea Flea, the common name for the order Siphonaptera, includes 2,500 species of small flightless insects that live as external parasites of mammals and birds. Fleas live by ingesting the blood of their hosts. Adult fleas grow to about long, a ...
s and swallow bugs, among others. These parasite
infestation Infestation is the state of being invaded or overrun by pests or parasites. It can also refer to the actual organisms living on or within a host. Terminology In general, the term "infestation" refers to parasitic diseases caused by animals su ...
s have been shown to negatively affect juvenile growth and developmental rates. Cliff swallows are socially
monogamous Monogamy ( ) is a form of Dyad (sociology), dyadic Intimate relationship, relationship in which an individual has only one Significant other, partner during their lifetime. Alternately, only one partner at any one time (Monogamy#Serial monogamy, ...
, one pair will look after each nest, but many occurrences of sexual polygamy have been noted because of the varying
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar wor ...
throughout the colony and within many individual nests. Both the female and the male cliff swallows will contribute to the colony's genetic variability by performing various actions of brood-parasitism. The cliff swallows have an “aggressive and fearless” personality in comparison to their relatives the barn swallows who were noted as being “timid and fearful”.


Diet

The cliff swallows feed on a diet consisting of
flying insects The Pterygota ( grc, Ï€Ï„Î”ÏÏ…ÎłÏ‰Ï„ÏŒÏ‚, pterugƍtĂłs, winged) are a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless (that is, insect groups whose ancestors once had wings b ...
, particularly swarming species such as: flies, bees, lacewings, mayflies, butterflies, moths, grasshoppers, and damselflies. The birds forage high (usually 50 m or higher) over fields or
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found at ...
es, and tend to rely on bodies of water like ponds during bad weather with high winds. These birds are day-hunters ( diurnal), returning to their nesting sites at
dusk Dusk occurs at the darkest stage of twilight, or at the very end of astronomical twilight after sunset and just before nightfall.''The Random House College Dictionary'', "dusk". At predusk, during early to intermediate stages of twilight, enou ...
, and are not very active during cold or rainy weather because of the low number of prey available.
Foraging Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...
behaviors related closely to their reproduction cycle; when the birds first arrive at the nesting site they will forage as far as 10 miles from the colony, in the hopes of increasing body fat reserves to prepare for cold-windy days and their energy extensive egg-laying stage. When the swallows return to the nesting site at dusk, they often fly in a tightly coordinated flock overhead, in such close
synchronization Synchronization is the coordination of events to operate a system in unison. For example, the conductor of an orchestra keeps the orchestra synchronized or ''in time''. Systems that operate with all parts in synchrony are said to be synchronou ...
that they may appear as one large organism. These large group formations are called creches. The cliff swallows' social behavior does not end with these “synchronized flying” displays; they use special vocalizations to advise other colony members of a good prey location where ample food is available. It has been thought that colony sites located close to marshes would have larger quantities of insects to support big populations; however, there are equally large nesting colonies located at a great distance from marshes.


Vocalisations

The social structure of these cliff swallow colonies has evolved a complex Bird vocalization, vocalisation system. Five vocalisations have been identified, which are used by both juveniles and adults for different reasons. These vocalisations are structurally similar across the age groups and can be described as: begging, alarm, recognition and squeak calls all with some variations. Juvenile cliff swallows are said to have established a unique call by the approximate age of 15 days, which allows the parents to identify their chick from others in the colony. The “squeak” call is of great interest to researchers because this is the special vocalisation made at a distance from the colony when a bird encounters a good foraging area. When this call is heard, large groups of their colony "neighbours" will arrive at the location. This "squeak" call is used greatly during bad weather conditions. For this specific call, the cliff swallows are one of the few known vertebrates to make a “competitively disadvantageous” cue to their peers for food availability. Alarm calls are heard at the colony while the birds are flying over-and-around the colony entrance way, and serve as a signal of danger close to the nests. When this call is heard by the other colony members, a mass fleeing of birds out and away from their nests is observed.


Reproduction

The breeding season of cliff swallows starts with the return of the birds from their wintering grounds. They usually arrive in large groups and start immediately to choose their nesting sites. The cliff swallows have been observed to skip from one to five years between breeding at the same place to avoid parasite infestations, but some pairs will return annually to the same site. Particularly for younger pairs, the size of the colony can affect their reproductive success, because they seem to rely on the valuable information that can be obtained from a large colony. Older birds are usually found in smaller colonies and exhibit earlier nesting times, avoiding the parasite manifestation that comes with the hot mid-summer season. Cliff swallows decide upon arrival at their nesting site whether they will fix a nest from the previous season or build a new nest. Building a new nest may have the benefit of lower parasite numbers, but it is very energy expensive and time-consuming. Further, taking the extra time to build a nest from scratch will mean reproducing later which could negatively affect their chicks’ survival. Nests constructed with Clay, sticky clay can last a number of years and are further supported by the cliff swallows’ tier-stacking construction strategy. Cliff swallows from the same colony socially collect mud for nest building, being seen converging at small areas together and then carrying globs of mud in their bills back to their nests. Each bird pair will have about 3–4 nestlings per brood; a clutch size of 4 has been identified as the most common and most successful. The cliff swallows brood-parasitize neighboring nests, where the females may move their eggs into or lay their eggs in other nests. The females who show intra-specific parasitism tend to have greater reproductive success than those that were brood-parasitized. The “victims” of brood-parasitism must nurture more chicks with higher energy costs and decreased Fitness (biology), fitness because they are raising young that will not pass on their own genetic material. The male cliff swallows will also take part in this gene-spread by mating with more than just one female, contributing to genetic variation throughout the colony. The nesting sites can be vulnerable to predation by other bird species, such as the house sparrow. These birds will search a number of swallow nests for the perfect place to make their own nest, destroying numerous eggs in the process. Nests, especially those at the periphery of colonies, are vulnerable to snake predation. Central nests are more coveted, have larger clutches, and are preferred for reuse in subsequent years Osborne, S. D. R. Leasure, S. Huang, R. Kannan. 2017. Central Nests Are Heavier and Have Larger Clutches Than Peripheral Nests in Cliff Swallow Colonies. Journal of the Arkansas Academy of Science 71: 206–208. Available at http://scholarworks.uark.edu/jaas/vol71/iss1/36/ Once the house sparrows pick their nest, they will bring in grass and other materials making it impossible for the cliff swallows to re-establish their place. Thus, the colonies with house sparrow predation have an overall lower success rate and fewer previous-year nests being used.


References


Books

*


External links

*
Cliff swallow – ''Petrochelidon pyrrhonota''
– USGS Patuxent Bird Identification InfoCenter

– Cornell Lab of Ornithology *
eBird Interactive Species Map- Cliff swallow

Birds of North America – Cliff swallow (''Petrochelidon pyrrhonota'')

Cliff Swallow Vocalisations – Song and Calls
*
Cliff Swallow Project
{{Taxonbar, from=Q1591775 Petrochelidon, cliff swallow Birds of North America Birds of South America Migratory birds (Western Hemisphere) Birds described in 1817, cliff swallow Taxa named by Louis Jean Pierre Vieillot, cliff swallow