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The mistletoebird (''Dicaeum hirundinaceum''), also known as the mistletoe flowerpecker, is a species of
flowerpecker The flowerpeckers are a family, Dicaeidae, of passerine birds. The family comprises two genera, ''Prionochilus'' and ''Dicaeum'', with 50 species in total. The family has sometimes been included in an enlarged sunbird family Nectariniidae. The b ...
native to most of
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
(though absent from
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
and the driest desert areas) and also to the eastern
Maluku Islands The Maluku Islands (; Indonesian: ''Kepulauan Maluku'') or the Moluccas () are an archipelago in the east of Indonesia. Tectonically they are located on the Halmahera Plate within the Molucca Sea Collision Zone. Geographically they are located eas ...
of
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
in the
Arafura Sea The Arafura Sea (or Arafuru Sea) lies west of the Pacific Ocean, overlying the continental shelf between Australia and Western New Guinea (also called Papua), which is the Indonesian part of the Island of New Guinea. Geography The Arafura Sea is ...
between Australia and
New Guinea New Guinea (; Hiri Motu Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea). It is a simplified version of ...
. The mistletoebird eats mainly the berries of the
parasitic 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 c ...
mistletoe Mistletoe is the common name for obligate hemiparasitic plants in the order Santalales. They are attached to their host tree or shrub by a structure called the haustorium, through which they extract water and nutrients from the host plant. ...
and is a vector for the spread of the mistletoe's seeds through its digestive system.del Hoyo, J. et al., eds. (2008). ''
Handbook of the Birds of the World The ''Handbook of the Birds of the World'' (HBW) is a multi-volume series produced by the Spanish publishing house Lynx Edicions in partnership with BirdLife International. It is the first handbook to cover every known living species of bird. T ...
'' 13: 388. .


Taxonomy and evolution

The mistletoebird is one of 44 species of the flowerpecker family
Dicaeidae The flowerpeckers are a family, Dicaeidae, of passerine birds. The family comprises two genera, ''Prionochilus'' and ''Dicaeum'', with 50 species in total. The family has sometimes been included in an enlarged sunbird family Nectariniidae. The b ...
. The flowerpeckers are considered to be nearest in avian evolutionary relationship to the
sunbird Sunbirds and spiderhunters make up the family Nectariniidae of passerine birds. They are small, slender passerines from the Old World, usually with downward-curved bills. Many are brightly coloured, often with iridescent feathers, particularly i ...
family Nectariniidae. Both the flowerpeckers and sunbirds are thought to be early offshoots of the early passeroid radiation that occurred 20-30 million years ago. The sunbirds are found mainly in Africa and Asia and the flowerpeckers throughout Asia. The mistletoebird is a geologically recent arrival into Australia from
South-East Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...
. It is thought to have started colonizing Australia from about two million years ago. The mistletoebird is a mistletoe-feeding specialist and mistletoe-feeding specialists have evolved independently in eight of the world's
avian Avian may refer to: *Birds or Aves, winged animals *Avian (given name) (russian: Авиа́н, link=no), a male forename Aviation *Avro Avian, a series of light aircraft made by Avro in the 1920s and 1930s *Avian Limited, a hang glider manufacture ...
families. This extreme dietary specialization has evolved in non-passerine species, as well as
sub-oscine The Tyranni (suboscines) are a suborder of passerine birds that includes more than 1,000 species, the large majority of which are South American. It is named after the type genus ''Tyrannus''. These have a different anatomy of the syrinx muscul ...
and
oscine A songbird is a bird belonging to the suborder Passeri of the perching birds (Passeriformes). Another name that is sometimes seen as the scientific or vernacular name is Oscines, from Latin ''oscen'', "songbird". The Passeriformes contains 500 ...
passerines. Early scientific research had the mistletoe and mistletoebird mutually co-evolving with a high dependence on each other through their evolutionary development. As the mistletoe has been in Australia for a long time and mistletoebirds for a relatively short time, the mistletoe seed was distributed originally by non-specialized frugivore birds like the honeyeater. Even though the mistletoebird has evolved into a very efficient local distributor of mistletoe seeds, the mistletoebird needs the mistletoe but the mistletoe does not need the mistletoebird. Molecular-based scientific methods have been used recently to evaluate generic taxonomy within the flowerpecker family. The flowerpecker genetic subdivision is based on a single morphological character — the length of the outermost primary wing feather. Most flowerpeckers are sexually dichromatic, have stouter bills than sunbirds and display a broad variety of tongue structure. Genetic analysis of mitochondrial DNA of 70% of flowerpecker species showed that the mistletoebird and the red-capped flowerpecker (''D. geelvinkianum'') to be each other's closest relative. The specific epithet ''hirundinaceum'' derives from the name of the
swallow 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 ...
(''Hirundo'') and refers to its long and pointed swallow-like wings.


Subspecies

There are four recognised
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 ...
, which differ in geographical location and plumage details, primarily of the males: *''Dicaeum hirundinaceum hirundinaceum'' Shaw & Nodder, 1792 — Australia (extensive red on throat and chest; flanks grey) *''Dicaeum hirundinaceum keiense'' Salvadori, 1874
Watubela archipelago Watubela is an archipelago in the Maluku Islands, east of Ceram and north of Kai Islands, southeast of the Gorong archipelago, and southwest of the Bomberai Peninsula of Papua, Indonesia. It includes the islands of Watubela itself, Kesui (also ...
, Indonesia (red on chest limited) *''Dicaeum hirundinaceum ignicolle'' G. R. Gray, 1858
Aru Islands The Aru Islands Regency ( id, Kabupaten Kepulauan Aru) is a group of about 95 low-lying islands in the Maluku Islands of eastern Indonesia. It also forms a regency of Maluku Province, with a land area of . At the 2011 Census the Regency had a po ...
, Indonesia (flanks yellow-buff) *''Dicaeum hirundinaceum fulgidum'' P. L. Sclater, 1883
Tanimbar Islands The Tanimbar Islands, also called ''Timur Laut'', are a group of about 65 islands in the Maluku (province), Maluku Provinces of Indonesia, province of Indonesia. The largest and most central of the islands is Yamdena; others include Selaru to t ...
, Indonesia (flanks buff, red on chest limited, and throat pale buff)


Description

The mistletoebird is small, long and weight. The male is glossy blue-black above, with a red chest and a slight red under-tail, and a black center stripe running down its white belly. The female is dark grey above, with a white throat, light grey underparts, and just a touch of pinkish-red under the tail. The eyes, bill, and legs are black; the bill is just over a centimeter long, slender, slightly down-curved and sharply pointed. Immature birds are similar to the female, but have an orange-pink bill instead of black. There is a variance in size and color across their distribution. The adult mistletoebird is smaller in the north of its distribution and females in the north have lighter colored underparts compared to darker ones in the south. The mistletoebird has long pointed wings and a short square tail with a slight notch at the tip. They usually occur singularly or in twos but occasionally in small family groups or flocks and very occasionally in mixed-species flocks when food is in abundance. Their flight is swift and they are usually seen flying above or high in the tree canopy on distinctively pointed wings. Their posture is usually upright when perched but adopt a more horizontal, swaying posture when alarmed. They have a variety of vocalizations but their most familiar is a short, sharp, high-pitched or whistled mainly on the wing. Songs while perched include a whistled ''wissweet wissweet'' uttered repeatedly. The male mistletoe bird can be mistaken for a
scarlet robin The scarlet robin (''Petroica boodang'') is a common red-breasted Australasian robin in the passerine bird genus ''Petroica''. The species is found on continental Australia and its offshore islands, including Tasmania. The species was originall ...
(''Petroica boodang'') or
flame robin The flame robin (''Petroica phoenicea'') is a small passerine bird native to Australia. It is a moderately common resident of the coolest parts of south-eastern Australia, including Tasmania. Like the other two red-breasted ''Petroica'' robins&md ...
(''Petroica phoenicea)'', but it has no spot on the forehead and has a black streak on the white belly. Due to the combination of size, shape, behaviour, and plumage the mistletoebird is unlikely to be mistaken for any other Australian passerine; however, the
pardalote Pardalotes or peep-wrens are a family, Pardalotidae, of very small, brightly coloured birds native to Australia, with short tails, strong legs, and stubby blunt beaks. This family is composed of four species in one genus, ''Pardalotus'', and seve ...
and thornbill are the closest in size and shape.


Behaviour


Habitat

Mistletoebirds are nomadic where movement is generally local and is associated with the fruiting of mistletoe. Mistletoebirds mostly occur in forests and woodlands dominated by any species of
Eucalyptus ''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as euca ...
from the dry interior to the coastal rain forests, but they do not frequent high altitude areas in winter as they adopt a torpid state when cold. Mistletoebirds are more likely to occur in mature stands where trees are larger and more likely to have become infested with mistletoe, rather than in regenerated areas. There are over 1300 species of mistletoe around the world and about 100 in Australia, where a common variety in the drier climates is the grey mistletoe (''Amyema quandang''). All mistletoes share a common growth form where water and nutrients are obtained from their host tree via a specialized vascular attachment, but as mistletoe manufacture their own
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
s using
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
, they are referred to as 'half parasitic'. Although common on the mainland of Australia, there are no mistletoes in
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
. Less specialized fruit-eating species, such as the honeyeater, source much of their protein from
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s. To source the arthropods, the honeyeaters venture far and, therefore, are a conduit for long distance mistletoe seed dispersal, while the mistletoebirds generally concentrate on and limit their feeding to local mistletoe infestations. Recent ecological studies have found the mistletoe producing a rich
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') l ...
in areas where it proliferates. The litter-fall under mistletoe infestation, where there is a high turnover of rich mistletoe leaves, has a diverse animal and plant life. Also, the denseness of the mistletoe bush and the characteristics of its leaves makes it a cool and safe place for birds to rest, hide, and nest. In a recent study in southern Australia, 217 species of Australian
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
birds were reported nesting in mistletoe, including the mistletoebird.


Feeding

The mistletoebird is a specialist
frugivore A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance an ...
that primarily eats the berries of mistletoes. Foraging time takes up 25% of the mistletoebird's daytime activities in summer and 29% in winter. The increased fruit consumption in winter could be related to the bird's increased thermo-regulatory costs during the cold of winter. Mistletoe fruit dominates the mistletoebird's diet accounting for 85% of foraging observations. Arthropods formed 13% and the remainder was from nectar, insects, and other berries. The only other known frugivore with a similar degree of dietary specialization is
Pesquet's parrot Pesquet's parrot (''Psittrichas fulgidus''), also known as the Dracula parrot or as the vulturine parrot (leading to easy confusion with '' Pyrilia vulturina'' from Brazil), is the only member of its genus. It is endemic to hill and montane rai ...
(or vulturine parrot) of New Guinea that feeds almost exclusively on strangler figs. In becoming frugivore specialists, both these birds live off the fruit of parasites that germinate in the canopy of host trees, where the fruit of the parasite has specific seed placement requirements on host branches and resists consumption by frugivore nonspecialist birds. The mistletoe plant minimizes opportunistic consumption of their berries by less specialized dispersal birds by producing few, inconspicuous fruits over a long period. Specialist frugivorous birds, by eating mainly fruit, have a diet rich in carbohydrates and poor in protein. To increase the protein level, which is particularly important for reproduction and molting, the mistletoe bird has to eat large quantities of the mistletoe fruit.


Feeding adaptations

The mistletoebird has a small muscularized stomach and short
alimentary canal The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
, where the amount of mechanical grinding and chemical digestion is minimized. This enables a clear passage and quick exit of the mistletoe fruit seeds through the mistletoebird's digestive system. In comparison, the non-specialized fruit-eating birds that they compete with for the mistletoe berry, such as the
spiny-cheeked honeyeater The spiny-cheeked honeyeater (''Acanthagenys rufogularis'') is the only species in the genus ''Acanthagenys''. It is large for a honeyeater, ranging from tall and weighing around 52 grams."Spiny-cheeked Honeyeater." Featherdal Wildlife ...
(''Acanthagenys rufogularis)'', have a more complex digestive system that takes longer for the fruit seeds to pass, and in the process the seeds are subjected to more mechanical and chemical action in the gut. It takes between 4–25 minutes for the mistletoe seed to pass through the mistletoebird. In a scientific study in South Australia with the grey mistletoe (''Amyema quandang)'', it took 2.3 times longer for the mistletoe seeds to pass through the digestive system of a
spiny-cheeked honeyeater The spiny-cheeked honeyeater (''Acanthagenys rufogularis'') is the only species in the genus ''Acanthagenys''. It is large for a honeyeater, ranging from tall and weighing around 52 grams."Spiny-cheeked Honeyeater." Featherdal Wildlife ...
(''A. rufogularis'') than it did through the mistletoebird and five months after seed deposit, 43% more seedlings had established from seeds defecated by mistletoebirds than from seeds passed by the honeyeater. The mistletoebird's modified
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organ (biology), organs of the digestive syste ...
is in the same plane. This facilitates rapid passage of the large number of berries consumed. Comparing the frugivore mistletoebird with a similarly sized
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 ...
the
inland thornbill The inland thornbill (''Acanthiza apicalis'') was originally described by English ornithologist John Gould in '' The Birds of Australia.'' Inland thornbills are within the order passerines. The inland thornbill belongs to the genus '' Acanthiza ...
(''Acanthiza inornata),'' where both their body masses are approximately . The mean length of the gizzard in the mistletoebird is and in the thornbill. The mean length of the small intestine is in the mistletoebird and in the thornbill. The mistletoebird has a large
gape The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for food ...
relative to the size of its head and its total body size. This allows grasping and swallowing the mistletoe fruit, which are about long by in diameter. These are large berries relative to the diminutive mistletoebird. Efficiency of mistletoe fruit dispersal by the mistletoebird is increased by the way the bird perches on a branch to
defecate Defecation (or defaecation) follows digestion, and is a necessary process by which organisms eliminate a solid, semisolid, or liquid waste material known as feces from the digestive tract via the anus. The act has a variety of names ranging fro ...
. They sometimes stand facing along the host tree branch depositing seeds in a string on the perch. Honeyeater-dispersed seeds, in contrast, fall randomly and less efficiently onto substrates below the defecating bird. The texture of the defecated seed with the mistletoebird is stickier than other bird's
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
to facilitate a strong bond with the host tree branch.


Breeding

Mistletoebirds conduct elaborate courtship and nuptial displays where, with a horizontal body, the male sways from side to side fluttering its wings, flicking its tail and displaying flashes of red. The female flies to a nearby branch where she flutters her wings. The male flies to her, mates and immediately they depart together. Mistletoebird pairs nest solitarily and raise up to 3 broods per season. The breeding season is from August to April depending where in Australia, but usually coincides with the fruiting of mistletoe. The nest can be found in many different plants but especially in eucalyptus, mistletoe, and acacia trees where it is suspended from an outer twig or thin horizontal branch and concealed among foliage. The nest is pear shaped with a slit side entrance and is usually made of plant down, wool, spider webs and spider egg sacs. The outside of the nest is coated with dry plant material like leaves, bark and lichen. The nest is built by the female, but accompanied by the male who assists very occasionally. Incubation is by the female but it has been recorded that males do sit the nest while the female is away. She will lay a clutch of three or four white eggs, each measuring , oval in shape. Incubation period is between 10–12 days and the young remain in the nest for approximately 15 days. The fledglings are fed by both parents but mainly by the female. Newly hatched birds are initially fed arthropods, after which mistletoe berry pulp is then gradually introduced into their diet. After about two weeks, when they are ready to
fledge Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight. This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
, the mistletoe berry pulp comprises 80% of food intake of the young mistletoebird. Both parents remove the fecal sacs left by the young in the nest.


Other characteristics

The longevity of a mistletoebird has been recorded in southern
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
, where a banded adult male was recaptured near the banding site after 9 years. Mistletoebirds vocally
mimic MIMIC, known in capitalized form only, is a former simulation computer language developed 1964 by H. E. Petersen, F. J. Sansom and L. M. Warshawsky of Systems Engineering Group within the Air Force Materiel Command at the Wright-Patterson AFB in ...
other birds. Heard in all seasons and given by both sexes, they have been recorded mimicking the mulga parrot ( ''Psephotus varius)'', as well as more than 25 different species of passerines. Predators known to have taken mistletoebird nestlings are the grey shrike-thrush (''Colluricincla harmonica)'', pied butcherbird (''Cracticus nigrogularis)'',
pied currawong The pied currawong (''Strepera graculina'') is a black passerine bird native to eastern Australia and Lord Howe Island. One of three currawong species in the genus ''Strepera'', it is closely related to the butcherbirds and Australian magpie of ...
(''Strepera graculina)'', and the
Australian raven The Australian raven (''Corvus coronoides'') is a passerine bird in the genus ''Corvus'' native to much of southern and northeastern Australia. Measuring in length, it has all-black plumage, beak and mouth, as well as strong grey-black legs and ...
(''Corvus coronoides)''. Nests have also been parasitized by a number of
cuckoo Cuckoos are birds in the Cuculidae family, the sole taxon in the order Cuculiformes . The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals and anis. The coucals and anis are sometimes separ ...
species, including Horsfield's bronze-cuckoo (''Chrysococcyx basalis)'' and the
fan-tailed cuckoo The fan-tailed cuckoo (''Cacomantis flabelliformis'') is a species of cuckoo in the family Cuculidae. It is found in Australia, Fiji, New Caledonia, New Zealand, New Guinea, Solomon Islands, and Vanuatu. Taxonomy Six subspecies have been reco ...
(''Cacomantis flabelliformis)''.


Threats and human interaction

There was an ecological balance between the parasitic mistletoe and their host trees before this balance was disturbed with excessive forest clearing. Land clearing to accommodate farming practices and population growth created an imbalance that resulted in excessive infestations of mistletoe in some areas leading to
forest degradation Forest degradation is a process in which the biological wealth of a forest area is permanently diminished by some factor or by a combination of factors. "This does not involve a reduction of the forest area, but rather a quality decrease in its c ...
, and a total lack of mistletoe in other areas. Mistletoe density in Eucalyptus forests in Australia ranges from 0 to over 100 plants per
hectare The hectare (; SI symbol: ha) is a non-SI metric unit of area equal to a square with 100-metre sides (1 hm2), or 10,000 m2, and is primarily used in the measurement of land. There are 100 hectares in one square kilometre. An acre is a ...
, where this high value is from highly damaged and perturbed habitat. From natural intact forests there are typically less than 10 plants per hectare. Avian richness increases with mistletoe density until a threshold of approximately 20 plants per hectare is reached after which heavy grazing and degradation sets in. Areas of excessive infestation of the parasite mistletoe badly damages or kills the host trees. The reaction to this was to try and eradicate the bird seen as spreading the parasite, the mistletoebird. One farmer in Australia in the 1930s claimed to have destroyed over 1200 mistletoebirds in a 6-year period, but they are no longer actively persecuted.


Gallery

Image:Mistletoebird0696e.jpg, Female mistletoebird feeding young in a nest in southern Western Australia. Image:Mistletoebirds2819e.jpg, Three young mistletoebirds in their nest waiting to be fed. Image:Mistletoebird3208e.jpg, Fledgling leaving its nest for the first time.


References

{{Taxonbar, from=Q538383 Dicaeum Birds of Australia Birds of the Maluku Islands Birds described in 1792