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'{{Short description|Tribe of birds}} {{Other uses}} {{Automatic taxobox | name = Grouse | fossil_range = [[Early Miocene]] to recent | image = SageGrouse21.jpg | image_caption = Male [[sage grouse]]<br />''Centrocercus urophasianus'' | taxon = Tetraonini | authority = [[William Elford Leach|Leach]], 1820 | subdivision_ranks = [[Genera]] | subdivision = ''[[Bonasa]]''<br /> ''[[Tetrastes]]''<br /> ''[[Centrocercus]]''<br /> ''[[Dendragapus]]''<br /> ''[[Tympanuchus]]''<br /> ''[[Lagopus]]''<br /> ''[[Falcipennis]]''<br /> ''[[Canachites]]''<br /> ''[[Tetrao]]''<br /> ''[[Lyrurus]]''<br /> and see text | synonyms = Tetraonidae <small>[[Nicholas Aylward Vigors|Vigors]], 1825</small><br /> Tetraoninae <small>Vigors, 1825</small> }} '''Grouse''' {{IPAc-en|ɡ|r|aʊ|s}} are a group of birds from the [[order (biology)|order]] [[Galliformes]], in the [[family (biology)|family]] [[Phasianidae]]. Grouse are presently assigned to the [[Tribe (biology)|tribe]] '''Tetraonini''' (formerly the subfamily '''Tetraoninae''' and the family '''Tetraonidae'''), a classification supported by [[mitochondrial DNA]] sequence studies,<ref name=Gutierrez/> and applied by the [[American Ornithologists' Union]],<ref name=AOUlist/> [[ITIS]],<ref name=ITIS/> [[International Ornithologists' Union|International Ornithological Congress]],<ref>{{Cite web|title=Taxonomic Updates – IOC World Bird List|url=https://www.worldbirdnames.org/new/updates/taxonomy/|access-date=2021-08-01|language=en-US}}</ref> and others.<ref name=Boyd/><ref name=":0">{{Cite journal|date=2021-05-01|title=A phylogenomic supermatrix of Galliformes (Landfowl) reveals biased branch lengths|url=https://www.sciencedirect.com/science/article/abs/pii/S1055790321000245|journal=Molecular Phylogenetics and Evolution|language=en|volume=158|pages=107091|doi=10.1016/j.ympev.2021.107091|issn=1055-7903|last1=Kimball |first1=Rebecca T. |last2=Hosner |first2=Peter A. |last3=Braun |first3=Edward L. |pmid=33545275 |s2cid=231963063 }}</ref> Grouse inhabit [[temperate]] and [[subarctic]] regions of the [[Northern Hemisphere]], from [[pine trees|pine forests]] to [[moorland]] and [[mountain]]side,<ref name=EoB/> from [[83rd parallel north|83°N]] ([[rock ptarmigan]] in northern [[Greenland]]) to [[28th parallel north|28°N]] ([[Attwater's prairie chicken]] in [[Texas]]).<ref name=Firefly/> The [[Turkey (bird)|turkeys]] are [[koklass pheasant]] are also closely allied with them.<ref name=":0" /> ==Description== Grouse are heavily built like other Galliformes, such as [[chicken]]s. They range in length from {{convert|31|to|95|cm|in|frac=2|abbr=on}}, and in weight from {{convert|0.3|to|6.5|kg|lb|frac=4|abbr=on}}. Males are larger than females—twice as heavy in the [[western capercaillie]], the largest member of the family. Grouse have feathered nostrils. Their legs are covered in feathers down to the toes, and in winter the toes, too, have feathers or small scales on the sides, an adaptation for walking on snow and burrowing into it for shelter. Unlike other Galliformes, they have no [[Spur (zoology)|spurs]].<ref name=Firefly/> ==Feeding and habits== These birds feed mainly on vegetation—buds, [[catkin]]s, leaves, and twigs—which typically accounts for over 95% of adults' food by weight. Thus, their diets vary greatly with the seasons. Hatchlings eat mostly insects and other [[invertebrate]]s, gradually reducing their proportion of animal food to adult levels. Several of the forest-living species are notable for eating large quantities of [[conifer]] needles, which most other [[vertebrate]]s refuse. To digest vegetable food, grouse have big [[crop (anatomy)|crops]] and [[gizzard]]s, eat grit to break up food, and have long [[intestine]]s with well-developed [[caecum|caeca]] in which [[symbiosis|symbiotic]] bacteria digest [[cellulose]].<ref name=Firefly/> [[Forest]] species flock only in autumn and winter, though individuals tolerate each other when they meet. [[Prairie]] species are more social, and [[tundra]] species (ptarmigans, ''[[Lagopus]]'') are the most social, forming flocks of up to 100 in winter. All grouse spend most of their time on the ground, though when alarmed, they may take off in a flurry and go into a long glide.<ref name=Firefly/> Most species stay within their breeding range all year, but make short seasonal movements; many individuals of the [[rock ptarmigan|ptarmigan]] (called rock ptarmigan in the US) and [[willow grouse]] (called willow ptarmigan in the US) [[bird migration|migrate]] hundreds of kilometers.<ref name=Firefly/> ==Reproduction== In all but one species (the [[willow ptarmigan]]), males are [[Animal sexuality#Polygamy|polygamous]]. Many species have elaborate [[courtship display]]s on the ground at dawn and dusk, which in some are given in [[Lek (biology)|leks]]. The displays feature males' brightly colored [[comb (anatomy)|combs]] and in some species, brightly colored inflatable sacs on the sides of their necks. The males display their [[plumage]], give vocalizations that vary widely between species, and may engage in other activities, such as drumming or fluttering their wings, rattling their tails, and making display flights. Occasionally, males fight.<ref name=Firefly/> The nest is a shallow depression or [[Bird nest#Scrape|scrape]] on the ground—often in cover—with a scanty lining of plant material. The female lays one [[Clutch (eggs)|clutch]], but may replace it if the eggs are lost. She begins to lay about a week after mating and lays one egg every day or two; the clutch comprises five to 12 eggs. The eggs have the shape of hen's eggs and are pale yellow, sparsely spotted with brown. On laying the second-last or last egg, the female starts 21 to 28 days of incubation. Chicks hatch in dense, yellow-brown down and leave the nest immediately. They soon develop feathers and can fly shortly before they are two weeks old. The female (and the male in the willow grouse) stays with them and protects them until their first autumn, when they reach their mature weights (except in the male [[capercaillie]]s). They are sexually mature the following spring, but often do not mate until later years.<ref name=Firefly/> ==Populations== [[File:Bonasa-umbellus-001edit1.jpg|thumbnail|right|A ruffed grouse in Canada]] Grouse make up a considerable part of the vertebrate [[Biomass (ecology)|biomass]] in the Arctic and Subarctic. Their numbers may fall sharply in years of bad weather or high predator populations—significant grouse populations are a major food source for [[lynx]], [[fox]]es, [[marten]]s, and [[bird of prey|birds of prey]]. However, because of their large clutches, they can recover quickly.{{citation needed|date=October 2020}} The three tundra species have maintained their former numbers. The prairie and forest species have declined greatly because of habitat loss, though popular game birds such as the [[red grouse]] and the [[ruffed grouse]] have benefited from habitat management. Most grouse species are listed by the [[IUCN]] as "least concern" or "near threatened", but the [[greater prairie chicken|greater]] and [[lesser prairie chicken]] are listed as "vulnerable" and the [[Gunnison grouse]] is listed as "endangered". Some subspecies, such as Attwater's prairie chicken and the [[Cantabrian capercaillie]], and some national and regional populations are also in danger.<ref name=Firefly/> The wild turkey suffered a precipitous decline in the past, but has since become abundant, even in developed areas. ==Sexual size dimorphism== === Male size selection === The phenotypic difference between males and females is called [[sexual dimorphism]].<ref name=ssd /> Male grouse tend to be larger than female grouse,<ref name=ssd /> which seems to hold true across all the species of grouse, with some difference within each species in terms of how drastic the size difference is.<ref name=ssd /> The hypothesis with the most supporting evidence for the evolution of sexual dimorphism in grouse is sexual selection.<ref name=ssd /> Sexual selection favors large males; stronger selection for larger size in males leads to greater size dimorphism.<ref name=ssd /> Female size will increase correspondingly as male size increases, and this is due to heredity (but not to the extent of the male size).<ref name=ssd /> This is because females that are smaller will still be able to reproduce without a substantial disadvantage, but this is not the case with males.<ref name=ssd /> The largest among the male grouse (commonly dubbed 'Biggrouse') attract the greatest numbers of females during their mating seasons.{{citation needed|date=October 2020}} === Mating behavior selection === {{Main|Lek mating}} Male grouse display [[Lek mating|lekking]] behavior, which is when many males come together in one area and put on displays to attract females.<ref name=Fiske>{{cite journal |last1=Fiske |first1=Peder| last2=Rintamaki |first2=Pekka |last3=Karvonen |first3=Eevi|title=Mating success in lekking males: a meta-analysis |journal=Behavioral Ecology |date=1998 |volume=9 |issue=4 |pages=328–338 |doi=10.1093/beheco/9.4.328 |doi-access=free}}</ref> Females selectively choose among the males present for traits they find more appealing.<ref name="Fiske"/> Male grouse exhibit two types: typical lekking and exploded lekking.<ref name=ssd /> In typical lekking, males display in small areas defending a limited territory, and in exploded lekking, displaying males are covered over an expansive land area and share larger territories.<ref name=ssd /> Male grouse can also compete with one another for access to female grouse through territoriality, in which a male defends a territory which has resources that females need, like food and nest sites.<ref name=ssd /> These differences in male behavior in mating systems account for the evolution of body size in grouse.<ref name=ssd /> Males of territorial species were smaller than those of exploded lekking species, and males of typical lekking species were the largest overall.<ref name=ssd /> The male birds that exhibit lekking behavior, and have to compete with other males for females to choose them, have greater sexual dimorphism in size.<ref name=Soulsbury>{{cite journal|last1=Soulsbury|first1=Carl D|last2=Kervinen|first2=Matti|last3=Lebigre|first3=Christophe|title=Sexual size dimorphism and the strength of sexual selection in mammals and birds|journal=Evolutionary Ecology Research|date=2014|volume=16|pages=63–76}}</ref> This suggests the hypothesis of sexual selection affecting male body size and also gives an explanation for why some species of grouse have a more drastic difference between male and female body size than others.{{citation needed|date=October 2020}} === Contrast with other bird species === Sexual size dimorphism can manifest itself differently between grouse and other birds. In some cases, the female is dominant over the male in breeding behavior, which can result in females that are larger than the males.<ref name="ReferenceA">{{cite journal| last1= Mueller |first1= H. C.|title= The Evolution of Reversed Sexual Dimorphism in Owls: An Empirical Analysis of Possible Selective Factors |journal= [[The Wilson Bulletin]] |volume=98|issue=3|pages=387–406}}</ref> ==In culture== [[File:Enontekiö.vaakuna.svg|thumb|upright|A [[willow grouse]] in the coat of arms of [[Enontekiö]]]] Grouse are [[game (food)|game]], and hunters kill millions each year for food, sport, and other uses. In the United Kingdom, this takes the form of [[driven grouse shooting]]. The male [[black grouse]]'s tail feathers are a traditional ornament for hats in areas such as [[Scotland]] and the [[Alps]]. Folk dances from the Alps to the North American prairies imitate the displays of lekking males.<ref name=Firefly/> ==Species== === Extant genera === {| class="wikitable" |- ! Image !! Genus !! Extant Species |- |[[File:Ruffed Grouse (32036698415).jpg|frameless|262x262px]] |''[[Ruffed grouse|Bonasa]]'' – ruffed grouse | * [[Ruffed grouse]], ''Bonasa umbellus'' |- |[[File:Spruce Grouse (Falcipennis canadenis) RWD.jpg|frameless|175x175px]]||''[[Spruce grouse|Canachites]]'' – spruce grouse || * [[Spruce grouse]], ''Canachites canadensis'' ** [[Franklin's grouse]], ''C. c. franklinii'' |- |[[File:Breeding male dusky grouse (30749272762).jpg|border|frameless|175x175px]] |''[[Dendragapus]]'' – blue grouse | * [[Sooty grouse]], ''Dendragapus fuliginosus'' * [[Dusky grouse]], ''Dendragapus obscurus'' |- |[[File:Greater sage-grouse (Centrocercus urophasianus).jpg|frameless|175x175px]] |''[[Centrocercus]]'' – sage grouse | * [[Gunnison grouse]], ''Centrocercus minimus'' * [[Greater sage-grouse]], ''Centrocercus urophasianus'' |- |[[File:Prairie Chicken.jpg|175x175px]] |''[[Tympanuchus]]'' – prairie grouse | * [[Greater prairie chicken]], ''Tympanuchus cupido'' ** [[Attwater's prairie chicken]], ''T. c. attwateri'' **{{extinct}}[[Heath hen]], ''T. c. cupido'' * [[Lesser prairie chicken]] ''Tympanuchus pallidicinctus'' * [[Sharp-tailed grouse]], ''Tympanuchus phasianellus'' ** [[Columbian sharp-tailed grouse]], ''T. p. columbianus'' |- |[[File:Haselhuhn-01.jpg|frameless|175x175px]] |''[[Tetrastes]]'' – hazel grouse | * [[Hazel grouse]], ''Tetrastes bonasia'' * [[Chinese grouse]], ''Tetrastes sewerzowi'' |- |[[File:Willow Ptarmigan- George C. Wood resize (16273373035).jpg|frameless|175x175px]] |''[[Lagopus]]'' – ptarmigans | * [[Willow ptarmigan]], ''Lagopus lagopus'' ** [[Red grouse]], ''L. l. scotica'' * [[White-tailed ptarmigan]], ''Lagopus leucura'' * [[Rock ptarmigan]], ''Lagopus muta'' |- |[[File:Falcipennis falcipennis displaying.jpg|frameless|175x175px]]||''[[Falcipennis]] –'' Siberian grouse || * [[Siberian grouse]], ''Falcipennis falcipennis'' |- |[[File:Capercaillie (8751340764).jpg|175x175px]]|| ''[[Tetrao]]'' – capercaillies || * [[Black-billed capercaillie]], ''Tetrao urogalloides'' * [[Western capercaillie]], ''Tetrao urogallus'' ** [[Cantabrian capercaillie]], ''T. u. cantabricus'' |- |[[File:Birkhahn (cropped).jpg|frameless|175x175px]]||''[[Lyrurus]]'' – black grouse || * [[Caucasian grouse]], ''Lyrurus mlokosiewiczi'' * [[Black grouse]], ''Lyrurus tetrix'' |- |} === Extinct genera === * Genus ''[[Proagriocharis]]'' † ** ''[[Proagriocharis kimballensis]]'' †<ref>Note: extinct species, late [[Miocene]] and early [[Pliocene]].</ref><ref>Larry D. Martin; James Tate, Jr. (1970). "A new turkey from the Pliocene of Nebraska"</ref> * Genus ''[[Rhegminornis]]'' † ** ''[[Rhegminornis|Rhegminornis calobates]]'' †<ref>Note: extinct species, early [[Miocene]].</ref><ref>Storrs L. Olson; John Farrand, Jr. (1974). "Rhegminornis restudies: a tiny Miocene turkey"</ref><ref>{{Cite web |title=BioLib: Biological library |url=https://www.biolib.cz/en/taxon/id469956/}}</ref> ==References== ===Footnotes=== {{Reflist|refs= <ref name=Gutierrez>{{cite journal |author1=Gutiérrez, R. J. |author2=Barrowclough, G. F. |author3=Groth, J. G. |title=A classification of the grouse (Aves: Tetroninae) based on mitochondrial DNA sequences |journal=Wildlife Biology |volume=6 |issue=4 |year=2000 |pages=205–212 |url=http://fwcb.cfans.umn.edu/research/owls/lit%20folder/gutierrez%20et%20al.%202000.pdf |url-status=dead |archive-url=https://web.archive.org/web/20150327211714/http://fwcb.cfans.umn.edu/research/owls/lit%20folder/gutierrez%20et%20al.%202000.pdf |archive-date=2015-03-27 |doi=10.2981/wlb.2000.017 |s2cid=85807603 }}</ref> <ref name=AOUlist>{{cite web|url=http://checklist.aou.org/taxa/ |title=AOU Checklist of North and Middle American Birds |publisher=American Ornithologists' Union|access-date=2014-03-13}}</ref> <ref name=ITIS>{{ITIS|id=553485|taxon=''Tetraoninae''|access-date=2013-03-13}}</ref> <ref name=Boyd>{{cite web|url=http://jboyd.net/Taxo/List2.html#phasianidae|title=Phasianidae: Turkeys, Grouse, Pheasants, Partridges |work=Aves – A taxonomy in flux|author=Boyd, John|access-date=2014-03-13}}</ref> <ref name=Firefly>{{cite book | last = Storch | first = Ilse | author2 = Bendell, J. F. | year = 2003 | chapter = Grouse | editor = Perrins, Christopher | editor-link = Chris Perrins | title = The Firefly Encyclopedia of Birds | publisher = Firefly Books | pages = [https://archive.org/details/fireflyencyclope0000unse/page/184 184–187] | isbn = 978-1-55297-777-4 | chapter-url-access = registration | chapter-url = https://archive.org/details/fireflyencyclope0000unse | url = https://archive.org/details/fireflyencyclope0000unse/page/184 }}</ref> <ref name=EoB>{{cite book |editor=Forshaw, Joseph|author= Rands, Michael R.W.|year=1991|title=Encyclopaedia of Animals: Birds|publisher= Merehurst Press|location=London|pages= 91|isbn= 978-1-85391-186-6}}</ref> <ref name=ssd>{{cite journal|last1=Drovetski|first1=S. V.|last2=Rohwer|first2=S.|last3=Mode|first3=N. A.|title=Role of sexual and natural selection in evolution of body size and shape: a phylogenetic study of morphological radiation in grouse|journal=Journal of Evolutionary Biology|date=2006|volume=19|issue=4|pages=1083–1091|doi=10.1111/j.1420-9101.2006.01097.x|pmid=16780509|s2cid=6144117 }}</ref> }} ===General=== * {{cite book |author=De Juana, E. |year=1994 |chapter=Family Tetraonidae (Grouse) |pages=[https://archive.org/details/handbookofbirdso0001unse/page/376 376–411] |editor1=del Hoyo, J. |editor2=Elliott, A. |editor3=Sargatal, J. |title=Handbook of the Birds of the World, Vol. 2. New World Vultures to Guineafowl |publisher=[[Lynx Edicions]] |location=Barcelona |isbn=978-84-87334-15-3 |chapter-url=https://archive.org/details/handbookofbirdso0001unse/page/376 }} * {{cite web|url=https://theuplandhunter.com/ruffed-grouse-eat-spring-fall-winter/ |title=What Do Grouse Eat? |publisher=The Upland Hunter |date=1 September 2017}} ==External links== <!-- Auk113:811. MolPhylEvol20:149. SystBiol51:930. WildlBiol6:205. --> {{Commons|Tetraoninae}} * [http://ibc.lynxeds.com/family/grouse-tetraonidae Grouse videos] on the Internet Bird Collection * {{cite journal | last1 = Johnsgard | first1 = P. | year = 1982 | title = Etho-Ecological Apects of Hybridization in the Tetraonidae | url = http://digitalcommons.unl.edu/johnsgard/16/ | journal = World Pheasant Association Journal | volume = VII | pages = 42–57 }} * {{Cite NIE|wstitle=Grouse|short=x}} {{Birds}} {{Phasianidae|state=all}} {{Meat}} {{Taxonbar|from=Q201240}} {{Authority control}} [[Category:Grouse| ]] [[Category:Tetraonini|*]] [[Category:Bird common names]] [[Category:Taxa named by William Elford Leach]]'