bowerbird

File:Regent Bowerbird arranging bower items.jpg arranging items in its bower.]]

The Ailuroedus catbirds are monogamous, with males raising chicks with their partners, but all other bowerbirds are polygynous, with the female building the nest and raising the young alone. These latter species are commonly dimorphic, with the female being drabber in color. Female bowerbirds build a nest by laying soft materials, such as leaves, ferns, and vine tendrils, on top of a loose foundation of sticks.

All Papuan bowerbirds lay one egg, while Australian species lay one to three with laying intervals of two days.Higgins, P.J. and J.M. Peter (editors); Handbook of Australian, New Zealand and Antarctic Birds, Volume 6: Pardalotes to Shrike-thrushes {{ISBN|0-19-553762-9}}Frith, Clifford B.; [http://www.publish.csiro.au/?act=view_file&file_id=MU9940060.pdf "Egg laying at long intervals in bowerbirds (Ptilonorhynchidae)"] Bowerbird eggs are around twice the weight of those of most passerines of similar sizeBased on the formula of egg mass = 0.258m^0.73, where m is body mass. From Rahn, H., Sotherland, P. and Paganelli, C. V., 1985. "Interrelationships between egg mass and adult body mass and metabolism among passerine birds" in Journal für Ornithologie 126:263–271.{{cite journal|author=Hoyt, Donald F. |journal=Auk |volume=96 |issue=1 |pages=73–77 |url=http://sora.unm.edu/node/23238 |title=Practical Methods of estimating the Volume and Fresh Weight of Bird Eggs |year=1979}} – for instance eggs of the satin bowerbird weigh around {{convert|19|g|oz|abbr=on}} as against a calculated {{convert|10|g|oz|abbr=on}} for a passerine weighing {{convert|150|g|oz|abbr=on}}. Eggs hatch after 19 to 24 days, depending on the species{{cite book |editor=Forshaw, Joseph|author= Frith, Clifford B.|year=1991|title=Encyclopedia of Animals: Birds|publisher= Merehurst Press|location=London|pages= 228–331|isbn= 978-1-85391-186-6}} and are a plain cream color for catbirds and the tooth-billed bowerbird, but in other species possess brownish wavy lines similar to eggs of Australo-Papuan babblers. In accordance with their lengthy incubation periods, bowerbirds that lay more than one egg have asynchronous hatching, but siblicide has never been observed.

Bowerbirds as a group have the longest life expectancy of any passerine family with significant banding studies. The two most studied species, the green catbird and satin bowerbird, have life expectancies of around eight to ten years[http://www.environment.gov.au/cgi-bin/biodiversity/abbbs/abbbs-search.pl Species search] by Australian Bird and Bat Banding Scheme and one satin bowerbird has been known to live for twenty-six years.{{cite web|url=http://www.environment.gov.au/cgi-bin/biodiversity/abbbs/abbbs-search.pl?taxon_id=378 |title=Satin Bowerbird |publisher=Environment.gov.au |date=2007-04-13 |access-date=2013-03-27}} For comparison, the common raven, the heaviest passerine species with significant banding records, has not been known to live longer than 21 years.Wasser, D. E. and Sherman, P.W.; "Avian longevities and their interpretation under evolutionary theories of senescence" in Journal of Zoology 2 November 2009

The most notable characteristic of bowerbirds is their extraordinarily complex courtship and mating behaviour, where males build a bower to attract mates. There are two main types of bowers. Prionodura, Amblyornis, Scenopoeetes and Archiboldia bowerbirds build so-called maypole bowers, which are constructed by placing sticks around a sapling; in the former two species these bowers have a hut-like roof.{{cite journal |last1=Kusmierski |first1=Rab |title=Molecular information on bowerbird phylogeny and the evolution of exaggerated male characteristics |journal=Journal of Evolutionary Biology |date=September 1993 |volume=6 |issue=5 |pages=737–752 |doi=10.1046/j.1420-9101.1993.6050737.x |s2cid=84721791 }} Chlamydera, Sericulus and Ptilonorhynchus bowerbirds build an avenue-type bower made of two walls of vertically placed sticks.{{cite journal |last1=Kusmierski |first1=Rab |title=Labile evolution of display traits in bowerbirds indicates reduced effects of phylogenetic constraint. |journal=Proc. Biol. Sci. |date=1997-03-22 |volume=264 |issue=1380 |pages=307–13 |pmid=9107048 |pmc=1688255 |doi=10.1098/rspb.1997.0044 }} Ailuroedus catbirds are the only species which do not construct either bowers or display courts.{{cite journal |last1=Kusmierski |first1=Rab |title=Labile evolution of display traits in bowerbirds indicates reduced effects of phylogenetic constraint |journal=Proc Biol Sci |date=1997-03-22 |volume=264 |issue=1380 |pages=307–313 |pmid=9107048 |doi=10.1098/rspb.1997.0044 |pmc=1688255 }} In and around the bower, the male places a variety of brightly colored objects he has collected. These objects — usually different among each species — may include hundreds of shells, leaves, flowers, feathers, stones, berries, and even discarded plastic items, coins, nails, rifle shells, or pieces of glass. The males spend hours arranging this collection. Bowers within a species share a general form but do show significant variation, and the collection of objects reflects the biases of males of each species and its ability to procure items from the habitat, often stealing them from neighboring bowers. Several studies of different species have shown that colors of decorations males use on their bowers match the preferences of females.

File:Vogelkop Bowerbird Bower 1099.jpg

In addition to the bower construction and ornamentation, male birds perform involved courtship displays to attract the female. Research suggests the male adjusts his performance based on success and female response.{{Cite journal|last1=Patricelli|first1=Gail L.|last2=Coleman|first2=Seth W.|last3=Borgia|first3=Gerald|date=January 2006|title=Male satin bowerbirds, Ptilonorhynchus violaceus, adjust their display intensity in response to female startling: an experiment with robotic females|journal=Animal Behaviour|volume=71|issue=1|pages=49–59|doi=10.1016/j.anbehav.2005.03.029|s2cid=6096593|issn=0003-3472}}{{Cite journal|last1=Coleman|first1=Seth W.|last2=Patricelli|first2=Gail L.|last3=Borgia|first3=Gerald|date=April 2004|title=Variable female preferences drive complex male displays|journal=Nature|volume=428|issue=6984|pages=742–745|doi=10.1038/nature02419|pmid=15085130|bibcode=2004Natur.428..742C |s2cid=2744790|issn=0028-0836}}

File:Satin bowerbird.jpg]]

Mate-searching females commonly visit multiple bowers, often returning to preferred bowers several times, and watching males' elaborate courtship displays and inspecting the quality of the bower. Through this process the female reduces the set of potential mates.{{Cite journal|last1=Uy|first1=J. Albert C.|last2=Patricelli|first2=Gail L.|last3=Borgia|first3=Gerald|date=November 2001|title=Complex Mate Searching in the Satin Bowerbird Ptilonorhynchus violaceus|journal=The American Naturalist|volume=158|issue=5|pages=530–542|doi=10.1086/323118|pmid=18707307|s2cid=2506815 |issn=0003-0147}} Many females end up selecting the same male, and many under-performing males are left without copulations. Females mated with top-mating males tend to return to the male the next year and search less.{{Cite journal|last1=Uy|first1=J. Albert C.|last2=Patricelli|first2=Gail L.|last3=Borgia|first3=Gerald|date=2000-02-07|title=Dynamic mate-searching tactic allows female satin bowerbirds Ptilonorhynchus violaceus to reduce searching |journal=Proceedings of the Royal Society of London B: Biological Sciences|volume=267|issue=1440|pages=251–256|doi=10.1098/rspb.2000.0994 |pmc=1690527|pmid=10714879}}

File:20110707221304!Chlamydera nuchalis bower - Mount Carbine.jpg]]

It has been suggested that there is an inverse relationship between bower complexity and the brightness of plumage. There may be an evolutionary "transfer" of ornamentation in some species, from their plumage to their bowers, in order to reduce the visibility of the male and thereby its vulnerability to predation.{{cite journal|last=Gilliard|first=Ernest Thomas|title=Bower ornamentation versus plumage characters in bower-birds.|journal=Auk|date=July 1956|volume=73|issue=3|pages=450–451|doi=10.2307/4082011|jstor=4082011|doi-access=free}} This hypothesis is not well supported because species with vastly different bower types have similar plumage. Others have suggested that the bower functioned initially as a device that benefits females by protecting them from forced copulations and thus giving them enhanced opportunity to choose males and benefits males by enhancing female willingness to visit the bower.{{cite journal|last=Kusmierski|first=Rab |author2=G. Borgia |author3=A. Uy |author4=R.H. Crozier|title=Labile evolution of display traits in bowerbirds indicates reduced effects of phylogenetic constraint|journal=Proceedings of the Royal Society of London B|year=1997|volume=264|pages=307–313|pmc=1688255|doi=10.1098/rspb.1997.0044|pmid=9107048|issue=1380}} Evidence supporting this hypothesis comes from observations of Archbold's bowerbirds that have no true bower and have greatly modified their courtship so that the male is limited in his ability to mount the female without her cooperation. In tooth-billed bowerbirds that have no bowers, males may capture females out of the air and forcibly copulate with them. Once this initial function was established, bowers were then co-opted by females for other functions such as use in assessing males based on the quality of bower construction. Recent studies with robot female bowerbirds have shown that males react to female signals of discomfort during courtship by reducing the intensity of their potentially threatening courtship. Young females tend to be more easily threatened by intense male courtship, and these females tend to choose males based on traits not dependent on male courtship intensity.

The high degree of effort directed at mate choice by females and the large skews in mating success directed at males with quality displays suggests that females gain important benefits from mate choice. Since males have no role in parental care and give nothing to females except sperm, it is suggested that females gain genetic benefits from their mate choice. However this has not been established, in part because of the difficulty of following offspring performance since males take seven years to reach sexual maturity. One hypothesis for the evolutionary causation of the bower building display is Hamilton and Zuk's "bright bird" hypothesis, which states that sexual ornaments are indicators of general health and heritable disease resistance.{{cite journal|last=Hamilton|first=William D|author2=M. Zuk|title=Heritable true fitness and bright birds: a role for parasites?|journal=Science|year=1982|volume=218|issue=4570|pages=384–387|doi=10.1126/science.7123238|pmid=7123238|jstor=1688879|bibcode=1982Sci...218..384H }} Doucet and Montgomerie{{cite journal|last=Doucet|first=Stephanie M.|author2=R. Montgomerie|title=Multiple sexual ornaments in satin bowerbirds: ultraviolet plumage and bowers signal different aspects of male quality|journal=Behavioral Ecology|year=2003|volume=14|issue=4|pages=503–509|doi=10.1093/beheco/arg035|doi-access=free}}{{cite journal|last=Doucet|first=Stephanie M.|author2=R. Montgomerie|title=Structural plumage colour and parasites in satin bowerbirds Ptilonorhynchus violaceus: implications for sexual selection|journal=Journal of Avian Biology|year=2003|volume=34|issue=3|pages=237–242|doi=10.1034/j.1600-048x.2003.03113.x|doi-access=free}} determined that the male bowerbird's plumage reflectance indicates internal parasitic infection, whereas the bower quality is a measure of external parasitic infection. This would suggest that the bowerbird mating display evolved due to parasite-mediated sexual selection, although there is some controversy surrounding this conclusion.{{cite journal|last=Borgia|first=Gerald|author2=K. Collins|title=Female choice for parasite-free male satin bowerbirds and the evolution of bright male plumage|journal=Behavioral Ecology and Sociobiology|year=1989|volume=25|issue=6|pages=445–454|doi=10.1007/bf00300191|bibcode=1989BEcoS..25..445B |s2cid=29986069}}

This complex mating behaviour, with its highly valued types and colors of decorations, has led some researchers{{cite book | last=Choe | first=J.C. | title=Encyclopedia of Animal Behavior | publisher=Elsevier Science | year=2019 | isbn=978-0-12-813252-4 | url=https://books.google.com/books?id=O5lnDwAAQBAJ&pg=RA2-PA38 | access-date=2019-10-19}} to regard the bowerbirds as among the most behaviorally complex species of bird. It also provides some of the most compelling evidence that the extended phenotype of a species can play a role in sexual selection and indeed, act as a powerful mechanism to shape its evolution, as seems to be the case for humans. Inspired by their seemingly extreme courtship rituals, Charles Darwin discussed both bowerbirds and birds-of-paradise in his writings.{{Cite journal | doi=10.1146/annurev.es.17.110186.000313|title = Biology of Birds of Paradise and Bowerbirds| journal=Annual Review of Ecology and Systematics| volume=17| pages=17–37|year = 1986|last1 = Diamond|first1 = J.}}

In addition, many species of bowerbird are superb vocal mimics. MacGregor's bowerbird, for example, has been observed imitating pigs, waterfalls, and human chatter. Satin bowerbirds commonly mimic other local species as part of their courtship display.

Bowerbirds have also been observed creating optical illusions in their bowers to appeal to mates. They arrange objects in the bower's court area from smallest to largest, creating a forced perspective which holds the attention of the female for longer. Males with objects arranged in a way that have a strong optical illusion are likely to have higher mating success.{{cite journal|author=Maxmen, Amy |date=9 September 2010 |doi=10.1038/news.2010.458 |title=Bowerbirds trick mates with optical illusions |journal=Nature News}}

File:Masked Bowerbird.jpg, illustrated by John Gould (1804–1881)]]

Though bowerbirds have traditionally been regarded as closely related to the birds of paradise, recent molecular studies suggest that while both families are part of the great corvid radiation that took place in or near Sahul (Australia-New Guinea), the bowerbirds are more distant from the birds of paradise than was once thought. DNA–DNA hybridization studies placed them close to the lyrebirds;{{cite journal|last=Sibley|first=CG|author2=Schodde, R|author3=Ahlquist, JE|title=The relationship of the Australo-Papuan Treecreepers Climacteridae as indicated by DNA-DNA hybridization|journal=Emu|date=1 January 1984|volume=84|issue=4|pages=236–241|doi=10.1071/MU9840236|bibcode=1984EmuAO..84..236S }} however, anatomical evidence appears to contradict this placement,{{cite journal | title=Systematic affinities of the lyrebirds (Passeriformes: Menura), with a novel classification of the major groups of passerine birds | journal=Molecular Phylogenetics and Evolution | volume=25 | issue=1 | date=2002-10-01 | issn=1055-7903 | doi=10.1016/S1055-7903(02)00215-4 | pages=53–62 | ref={{sfnref | Molecular Phylogenetics and Evolution | 2002}} | last1=Ericson | first1=Per G.P | last2=Christidis | first2=Les | last3=Irestedt | first3=Martin | last4=Norman | first4=Janette A. | pmid=12383750 | bibcode=2002MolPE..25...53E }} and the true relationship remained unresolved for long. Cladistic analyses in the mid-2010s usually allied bowerbirds with the Australasian treecreepers (Climacteridae), another Sahul endemic family which is highly adapted to a woodpecker-like lifestyle (woodpeckers being absent from Sahul). This putative superfamily forms part of a large basal radiation of ancient songbirds, with the lyrebirds being part of a more ancestral branch than the bowerbirds and their DNA-DNA hybridization similarities being due to the phenetic methodology which (unlike cladistic analysis) merely assesses overall similarity without accounting for convergent evolution.

Many bowerbirds (in particular New Guinean species) are little known and even less studied. But the hypothesized relationships of 3 roughly equally distinct groups and one peculiar species inferred from courtship behaviour and external appearance are by and large confirmed by molecular phylogenetics, . Some insights from the more recent studies, however, were less expected: The Tooth-billed catbird, with its unique "stagemaker" courtship, was long suspected not to be a true catbird (genus Ailuroedus). As it turned out, this is not only correct, but in fact the Tooth-billed catbird is robustly resolved by the mtDNA data as more closely related to the "maypole"-type bower builders than to Ailuroedus, and certainly warrants separation in genus Scenopoeetes. Also, the enigmatic "maypole-builder" genus Archboldia seems to be merely a Amblyornis with unusually heavy melanin pigmentation as is often found in tropical rainforest birds. On the other hand, the "avenue-builders" also have a hypermelanic lineage, the satin bowerbird, but this seems well separable as a monotypic genus Ptilonorhynchus, as is the "maypole-building" golden bowerbird (as Prionodura). Interestingly, the widely divergent "avenue-builders" may represent the oldest living lineage, with the monogamous true catbirds, which do not build a bower and were traditionally held to be "primitive", as the most derived group among living bowerbirds – the last common ancestor of the living bowerbirds is hypothesized to have been polygynous, with sexually dimorphic plumage – cryptic greenish in the females, and probably dark with a yellow belly in the males. But overall relationships between the true catbirds, the "maypole-builders" and the "avenue-builders" were not definitely resolvable, with only a small outgroup being used and outgroup effects on intra-family relationships not being tested. Even so, it is precisely this uncertainty about inter-group relationships that strongly suggests that the "maypole"/"avenue" bowers are not one ancestral and one derived type, but evolved independent of one another, perhaps from a "clean stage"-type courtship arena which is commonly established by all bower-building species at the start of bower construction, and persists in little-altered form (just adding some remarkable leaves strewn about as decoration) in Scenopoeetes which almost certainly is the most ancient living lineage of the "maypole-builders". Among the catbirds, the white-cheeked group (A.buccoides/geislerorum/stonii) is very likely the most ancient one, which is also in line with the hypothesis that bowerbirds have become more and more drab and inconspicuous as their evolution progressed.

The cladogram below showing the relationships between the genera is based on a molecular phylogenetic study by Per Ericson and collaborators that was published in 2020. The genus Archboldia was found to be embedded in the genus Amblyornis.{{Cite journal | last1=Ericson | first1=P.G.P. | last2=Irestedt | first2=M. | last3=Nylander | first3=J.A.A. | last4=Christidis | first4=L. | last5=Joseph | first5=L. | last6=Qu | first6=Y. | date=2020 | title=Parallel evolution of bower-building behavior in two groups of bowerbirds suggested by phylogenomics | journal=Systematic Biology | volume=69 | issue=5 | pages=820-829 | doi=10.1093/sysbio/syaa040 | doi-access=free }}

{{Clade | style=font-size:100%;line-height:100%

|label1=Ptilonorhynchidae

|1={{clade

|1={{clade

|1=Sericulus – 4 species

|2={{clade

|1=Ptilonorhynchus – satin bowerbird

|2=Chlamydera – 5 species

}}

}}

|2={{clade

|1=Ailuroedus – 10 species: catbirds

|2={{clade

|1=Scenopoeetes – tooth-billed bowerbird

|2={{clade

|1=Prionodura – golden bowerbird

|2=Amblyornis – 5 species + Archboldia – Archbold's bowerbird

}}

}}

}}

}}

}}

True catbirds

Genus Ailuroedus

• Ochre-breasted catbird, Ailuroedus stonii
• White-eared catbird, Ailuroedus buccoides
• Tan-capped catbird, Ailuroedus geislerorum
• Green catbird, Ailuroedus crassirostris
• Spotted catbird, Ailuroedus maculosus
• Huon catbird, Ailuroedus astigmaticus
• Black-capped catbird, Ailuroedus melanocephalus
• Northern catbird, Ailuroedus jobiensis
• Arfak catbird, Ailuroedus arfakianus
• Black-eared catbird, Ailuroedus melanotis

Maypole-builders (including Tooth-billed catbird)

Genus Scenopoeetes

• Tooth-billed catbird, Scenopoeetes dentirostris

Genus Archboldia

• Archbold's bowerbird, Archboldia papuensis

Genus Amblyornis

• Vogelkop bowerbird, Amblyornis inornata
• MacGregor's bowerbird, Amblyornis macgregoriae
• Huon bowerbird, Amblyornis germanus
• Streaked bowerbird, Amblyornis subalaris
• Golden-fronted bowerbird, Amblyornis flavifrons

Genus Prionodura

• Golden bowerbird, Prionodura newtoniana

Avenue-builders

Genus Sericulus

• Flame bowerbird, Sericulus ardens
• Masked bowerbird, Sericulus aureus
• Fire-maned bowerbird, Sericulus bakeri
• Regent bowerbird, Sericulus chrysocephalus

Genus Ptilonorhynchus

• Satin bowerbird, Ptilonorhynchus violaceus

Genus Chlamydera

• Western bowerbird, Chlamydera guttata
• Spotted bowerbird, Chlamydera maculata
• Great bowerbird, Chlamydera nuchalis
• Eastern great bowerbird, Chlamydera (nuchalis) orientalis (possibly a distinct species)
• Yellow-breasted bowerbird, Chlamydera lauterbachi
• Fawn-breasted bowerbird, Chlamydera cerviniventris

Fossil record

Bowerbirds have a scant fossil record that nonetheless extends to the Chattian (latest Oligocene), with the fossil species Sericuloides marynguyenae dated to 26 to 23 million years ago. It was found in Faunal Zone A deposits of the White Hunter Site at D-site Plateau of the Riversleigh World Heritage Area. S. marynguyenae was a tiny member of its family, about the same size as the golden bowerbird. It is known from the proximal end of a right carpometacarpus and the proximal end of a left tarsometatarsus. The material, though fragmentary, preserves much detail, and is overall more similar to the "avenue-builders" – in particular Chlamydera – than to the other two main groups. However, the splits between the three main groups of living bowerbirds are presumed to have occurred only in the Miocene, some time after Sericuloides lived. Thus, the fossil species may have belonged to a more basal and now entirely extinct lineage, and/or it may be considered to support the hypothesis that the "avenue-builders" are the most ancient group of bowerbirds and retain many "primitive" features in their anatomy.{{cite journal | title= The earliest record of bowerbirds (Passeriformes, Ptilonorhynchidae) from the Oligo-Miocene of northern Australia | journal=Alcheringa | volume=47 | date=2023-03-15 | doi=10.1080/03115518.2023.2180537 | pages=475–483 | ref={{sfnref | Alcheringa | 2023}} | last1=Nguyen | first1=Jacqueline M.T.| issue=4 | s2cid=257578512 | doi-access=free | bibcode=2023Alch...47..475N }}

Other than S. marynguyenae, as of 2023 only one other prehistoric bowerbird species is known. This has not been named, as it is only known from the distal left ulna piece QM F57970 (AR19857), also found on the D-site Plateau of Riversleigh WHA, but in interval 3 of Faunal Zone B at the Ross Scott-Orr site, in late early Miocene (Burdigalian) sediments dated to 16.55 mya. Even though this piece of fossil bone is merely some 16 mm long, it is excellently preserved, and its features are characteristic of a smallish bowerbird the size of a black-eared catbird. Bowerbird ulnae – to the extent they have been studied – differ little between genera and species, but the Miocene fossil is unlike all living members of the family in one detail or another. If anything, it resembles the presumably more advanced groups ("maypole-builders" and true catbirds) more than the "avenue-builders" and given its age it may well have been one of the earliest members of either of the former two groups.

{{Reflist}}

• [http://ibc.lynxeds.com/family/bowerbirds-ptilonorhynchidae Bowerbird videos] on the Internet Bird Collection
• [https://web.archive.org/web/20150418032704/http://www.viralforest.com/bower-bird/ Bower bird nest building]

{{Bowerbird|state=expanded}}

{{Taxonbar|from=Q215603}}

{{Authority control}}

Category:Birds of Oceania

Category:Taxa named by George Robert Gray

Category:Extant Chattian first appearances