Acrocephalus Arundinaceus
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The great reed warbler (''Acrocephalus arundinaceus'') is a
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
n
passerine 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 ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
'' Acrocephalus''. The genus name ''Acrocephalus'' is 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 ...
''akros'', "highest", and ''kephale'', "head". It is possible that
Naumann Naumann is a Central German variation of the surname Neumann. Notable people with the surname include: * Albert Naumann (1875–?), German Olympic fencer * Alexander Naumann (1837–1922), German chemist * Christian August Naumann (1705–?), Germa ...
and
Naumann Naumann is a Central German variation of the surname Neumann. Notable people with the surname include: * Albert Naumann (1875–?), German Olympic fencer * Alexander Naumann (1837–1922), German chemist * Christian August Naumann (1705–?), Germa ...
thought ''akros'' meant "sharp-pointed". The specific ''arundinaceus'' is from
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
and means "like a reed", from ''arundo, arundinis'', "reed". It used to be placed in the
Old World warbler Old World warblers are a large group of birds formerly grouped together in the bird family Sylviidae. The family held over 400 species in over 70 genera, and were the source of much taxonomic confusion. Two families were split out initially, the ci ...
assemblage, but is now recognized as part of the marsh and tree-warbler
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
(
Acrocephalidae The Acrocephalidae (the reed warblers, marsh- and tree-warblers, or acrocephalid warblers) are a family of oscine passerine birds, in the superfamily Sylvioidea. The species in this family are usually rather large "warblers". Most are rather pla ...
). Great reed warblers are medium-sized birds and are the largest of the European warblers. They breed throughout mainland
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and the west
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 ...
and migrate to
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
in the winter. Great reed warblers favour reed beds as their habitat during breeding months, while living in reed beds, bush thickets, rice fields, and forest clearings during the winter. Great reed warblers exhibit relatively low
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, and both genders of the species are similar in appearance. This species mates both
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
ly and
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, ...
ly.


Description

The thrush-sized
warbler Various Passeriformes (perching birds) are commonly referred to as warblers. They are not necessarily closely related to one another, but share some characteristics, such as being fairly small, vocal, and insectivorous. Sylvioid warblers Th ...
is one of the largest species of the former family, the
Old World warbler Old World warblers are a large group of birds formerly grouped together in the bird family Sylviidae. The family held over 400 species in over 70 genera, and were the source of much taxonomic confusion. Two families were split out initially, the ci ...
s and it stands as one of the largest species in the family
Acrocephalidae The Acrocephalidae (the reed warblers, marsh- and tree-warblers, or acrocephalid warblers) are a family of oscine passerine birds, in the superfamily Sylvioidea. The species in this family are usually rather large "warblers". Most are rather pla ...
. It measures in length, in wingspan and weighs . The adult has unstreaked brown upperparts and dull buffish-white chin and underparts. The forehead is flattened, and the bill is strong and pointed. It looks very much like a giant
Eurasian reed warbler The common reed warbler (''Acrocephalus scirpaceus'') is an Old World warbler in the genus '' Acrocephalus''. It breeds across Europe into the temperate western Palaearctic where it is migratory, wintering in sub- Saharan Africa. It is also a re ...
(''A. scirpaceus''), but with a stronger
supercilium The supercilium is a plumage feature found on the heads of some bird species. It is a stripe which runs from the base of the bird's beak above its eye, finishing somewhere towards the rear of the bird's head.Dunn and Alderfer (2006), p. 10 Also ...
. The sexes are identical, as with most old world warblers, but young birds are richer buff below. The warbler's song is very loud and far-carrying. The song's main phrase is a chattering and creaking ''carr-carr-cree-cree-cree-jet-jet'', to which the whistles and vocal mimicry typical of
marsh warbler The marsh warbler (''Acrocephalus palustris'') is an Old World warbler currently classified in the family Acrocephalidae. It breeds in temperate Europe and the western Palearctic and winters mainly in southeast Africa. It is notable for incorpora ...
s are added.


Distribution and ecology

The great reed warbler breeds in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and the west
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 does not breed in
Great Britain Great Britain is an island in the North Atlantic Ocean off the northwest coast of continental Europe. With an area of , it is the largest of the British Isles, the largest European island and the ninth-largest island in the world. It is ...
, but is an irregular visitor. Its population has in recent decades increased around the eastern
Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden and the North and Central European Plain. The sea stretches from 53°N to 66°N latitude and from ...
, while it has become rarer at the western end of its range. It is a migratory bird, wintering in
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
. This bird migrates north at a rather late date, and some birds remain in their winter quarters until the end of April. While there are no
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
of this bird,
mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
haplotype A haplotype ( haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material ( DNA) which is inherited from two parents. Normally these organisms have their DNA or ...
data indicate that during the last glacial period there were two
allopatric Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
populations of great reed warbler. The great reed warblers in southwestern and southeastern Europe were at that time apparently separated by the Vistulian-
Würm The Würm is a river in Bavaria, Germany, right tributary of the Amper. The length of the river is , or including the ''Steinbach'', the main feed of Lake Starnberg. It drains the overflow from Lake Starnberg and flows swiftly through the villag ...
ice sheet In glaciology, an ice sheet, also known as a continental glacier, is a mass of glacial ice that covers surrounding terrain and is greater than . The only current ice sheets are in Antarctica and Greenland; during the Last Glacial Period at Las ...
s and the surrounding barren lands. Though the data are insufficient to robustly
infer Inferences are steps in reasoning, moving from premises to logical consequences; etymologically, the word '' infer'' means to "carry forward". Inference is theoretically traditionally divided into deduction and induction, a distinction that in ...
a date for this separation, it suggests the populations became separated around 80,000 years ago – coincident with the first major advance of the ice sheets. The populations must have expanded their range again at the start of the
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
about 13,000 years ago, but even today the western birds winter in the west and the eastern birds in the east of tropical Africa. This
passerine 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 ...
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
is found in large
reed bed A reedbed or reed bed is a natural habitat found in floodplains, waterlogged depressions and estuaries. Reedbeds are part of a succession from young reeds colonising open water or wet ground through a gradation of increasingly dry ground. As ...
s, often with some bushes. On their breeding grounds, they are
territorial A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
. In their winter quarters, they are frequently found in large groups, and may occupy a reed bed to the exclusion of other birds. Like most warblers, it is
insectivorous 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 were ...
, but it will take other prey items of small size, including
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s such as
tadpole A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found i ...
s. The great reed warbler undergoes marked long-term population fluctuations, and it is able to expand its range quickly when new
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 ...
becomes available. This common and widespread bird is considered a species of
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. Population size can be calculated with a suitability model, but direct counts of territorial males in suitable habitat and sampling the population sex-ratio can be a proper alternative to inference-rich predictive modeling based on imperfect habitat-extrapolation of densities of reed warblers at large spatial scales.


Behaviour


Diet

''A. arundinaceus'' has a primarily
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
diet. Observation of prey collection specifically during breeding season has shown the retrieval of insect larvae, moths, dragonflies, damselflies, beetles, spiders, small fish, and frogs. ''A. arundinaceus'' has also been reported to eat fruit during non-breeding seasons. Nestlings typically feed on
diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
and arachnids, though this may not be their preferred food.


Communication and courtship

Male great reed warblers have been observed to communicate via two basic song types: short songs about one second in length with few syllables, and long songs of about four seconds that have more syllables and are louder than the short variety. It has been observed that long songs are primarily used by males to attract females; long songs are only given spontaneously by unpaired males, and cease with the arrival of a female. Short songs, however, are primarily used in territorial encounters with rival males. During experimental observation, male great reed warblers showed reluctance to approach recordings of short songs, and when lured in by long songs, would retreat when playback was switched to short songs. Traditionally, monogamous species of genus ''Acrocephalus'' use long, variable, and complex songs to attract mates, whereas polygynous varieties use short, simple, stereotypical songs for territorial defence. There is evidence that long songs have been evolved through intersexual selection, whereas short songs have been evolved through intrasexual selection. The great reed warbler is a notable example of these selective pressures, as it is a partial polygynist and has evolved variable song structure (both long and short) through evolutionary compromise. In addition to communication, the great reed warbler's song size has been implicated in organism fitness and reproductive success. Though no direct relationship has been found between song size and either territory size or beneficial male qualities, such as wing length, weight, or age, strong correlation has been observed between repertoire size and territory quality. Furthermore, partial correlation analysis has shown that territory quality has significant effect on the number of females obtained, while repertoire length is linked to the number of young produced.


Mating system and sexual behavior

Great reed warbler females lay 3–6
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s in an open cup-nest in reeds. Some pairs of warblers are
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, ...
, but others are not, and unpaired, so-called "satellite" males still father some young. Great reed warblers defend their nests using graded alarm calls, directed towards a wide range of enemies, although these alarm calls might reveal the whereabouts of the nest to brood parasites. A long-term study of the factors that contribute to male fitness examined the characteristics of males and territories in relation to annual and lifetime breeding success. It showed that the arrival order of the male was the most significant factor for predicting pairing success, fledgling success, and number of offspring that survive. It also found that arrival order was closely correlated with territory attractiveness rank. Females seem to prefer early arriving males that occupy more attractive territories. These females also tend to gain direct benefits through the increased production of fledglings and
offspring In biology, offspring are the young creation of living organisms, produced either by a single organism or, in the case of sexual reproduction, two organisms. Collective offspring may be known as a brood or progeny in a more general way. This ca ...
that become adults. In addition, male song repertoire length is positively correlated to annual harem size and overall lifetime production of offspring that survive. Song repertoire size alone is able to predict male lifetime number of surviving offspring. Females tend to be attracted to males with longer song repertoires since they tend to sire offspring with improved viability. In doing so, they gain indirect benefits for their own young. Great reed warblers have a short,
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
breeding cycle in which the male contributes little to parental care. They defend large territories in reed beds where there is reduced visibility, which may allow males to practice deception by moving and attracting a second female. This second female may not realize that the male has already mated. Polygyny of the great reed warbler was assessed in another study that showed the importance of
female choice Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior.Bateson, Paul Patrick Gordon. "Mate Choice." Mate Choic ...
. The differences in territory characteristics seemed to be more important. However, there is also a strong correlation between males and their territory characteristics. Models based on the polygyny threshold and sexy son hypotheses predict that females should gain evolutionary advantage in either short-term or long-term in this mating system, yet the study did not support this. The data showed that secondary females had greatly reduced breeding success.


References


External links


Ageing and sexing (PDF; 1.9 MB) by Javier Blasco-Zumeta & Gerd-Michael Heinze
* Great reed warbler
Species text in The Atlas of Southern African Birds
{{Taxonbar, from=Q180835
great reed warbler The great reed warbler (''Acrocephalus arundinaceus'') is a Eurasian passerine in the genus '' Acrocephalus''. The genus name ''Acrocephalus'' is from Ancient Greek ''akros'', "highest", and ''kephale'', "head". It is possible that Naumann and ...
Birds of Eurasia Birds described in 1758 Taxa named by Carl Linnaeus Articles containing video clips