a classification of the living birds of the world based on dna-dna hybridization studies

7/29/2019 A Classification of the Living Birds of the World Based on DNA-DNA Hybridization Studies

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410 SIBLEY, AHLQUIST, AND MONROE JR. [Auk, Vol. 105MAJOR DIVIS IONS OF CLASS AVES ORDER SUPERQADER PARVClASS INFRAClASS

=". . . " , . " . . . 8truthioniformes

-1'5,9 ' 9 G ~ 9 ) ~ 2 1 ~ , 8 ~ ' ~ 7 4 ~ ) ~ = = = = = = = = = = = = T i n a m i f o r m e sraciformes21.6 (48) Galliformes22.9 (89) Anseri forrnes

28.0 (316)_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _Turniclformes27.0(27)

] Gallomorphae- Anserimorphae

] Ratitae ]] Galloanserae

r'::-:6C":,3C":'::-:'6::-:';--)---------------- Piciforrnes - - - - - - - - - - - - - - - - - - - - - - PicaeGalbUI,", ~ r m e s _ Galbul imorphae ]~ 4 ~ 4 ~ ' ~ 6 3 ~ ) ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ B " " o l ' f "m , , ] ,20,8 (1S) Upupiformes Bucerot lmorphae Coraciae23.4 (82) Trogomtormes ] Coraci imorphae22.1 (28) Coracl i tormes

25.0 (256) ______________________ Coliae_ Cuculimorphae_ Psittacimorphae

Columblformes ]Gruiformes .Ciconii.formes Pass8nmorphaePassertformes

" 1 g ~ : ; : ; : : = = = = = = = = = = = = = = C O l i i f o r m e s~ 2 4 ' 5 (132) CUGuliformes23.7 (268) Psit taciformes23.1 (163)~ ~ E ~ = = = = = = = = = = = A P o d i f o r m e s ]1.3 (47) Trochiliformes Apodimorphae~ 2 ' 1 . 9 (97) M u ~ o ~ h a g i f o r m e S l Strigimorphae. L...{20.4 (21) Sirtgiformes J22.5 (276)

~2 0 . 8 ( 1 1 7 )20.1 (503)

21.6 (129)

Passerae

lD"'""

Fig. 1. Major divisions of the class Aves. The numbers, e.g. 28.0 (316), indicate that the average deltaT50H value of 28.0 is based on the delta T50H values of 316 DNA-DNA hybrids between taxa on the two sidesof the branch node.

achieving the desirable goal of categorical equiv-alence. Hennig (1966: 161-182) discussed theproblems associated with determining the agesof origin of systematic groups and concludedthat the available methods were inadequate. Thepresent methods still fall short of the ideal, butwe have t ried to incorporate th e principle of

categorical equivalence into our classificationby assigning ranks on the basis of del ta T50Hvalues (Table 1). The del ta values express de-grees of genomic divergence and, since th e av-erage rate of divergence is relat ive to t ime, thisprocedure provides an approach to categoricalequivalence based on time of origin. The fact

"'"UE"...

"'" "'"0:;U. ou"" ,mel

"DU""'"....

"DE lDOU_ lD" 0....JO

00ci"' NNm N ci0"' N

"' "'"'..:

I '"''"I N'";2 N ;;:'"'

THE FAMILIES OF NON-PASSERINE BIRDSFig. 2. Families of non-Passerine birds, Struthionidae to Dacelonidae.


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July 1988] Classification of Living Birds 411

'"',-,.'-,'-C'-32'-' Coliidae

~ ~ ~ ~ ~ ~ ~ ~ ~ ~ C r o t o P h a g i d a e4 .5 ( 6) Neomorphidae1 6. 4 ( 5) Opisthocomidae. - -c : - : - :=- - i17 .6 (53) Coccyzidae23.7 (268) 14.5 ( fa ) Centropodidae

13.0 Cuculidae

"'''"3.''-', - " " '63 : : - ) - - - - - - - - - - - Psittacldae~ ~ ~ = = = = ~ ~ ~ = = = = A P O d i d a e2 1 , 3 (47) 9.5 (4) ~ : o ~ i ~ i ~ i ~ ~ ~ d a e

,( Musophagidae~ i J ~'Or'_C2_"_-liTI:ill::=====." Strlgldae13.6 (7) Tytonidae!" 19.1 (55) AegothelidaefJ1 l 18 . 8 (26) 10.3 (2J Podargidae, e 47.9 (20) Batrachostomidae~ ~ ~ ~ ~ ~ ~ ~ ~ ~ Nyctibildae5 .8 (2) Steatornithidae16.9 (10) Eurostopodidae

12.3 (3J Caprimulgidae==",- Columbidae

I"""Fig. 3. Families of non-passerine birds, Coliidae

to Columbidae.

that different lineages evolve at somewhat different average rates introduces an error of uncertain magnitude.The Turnicidae may present a special prob

lem. The 27 delta TsoH values between Turnixand other taxa are large, ranging from 22.7 tothe pectoral sandpiper (Calidris melanotos) to 29.8to the ostrich (Struthio camelus); average = 27.0.The range of 7.1 is unusually large. The smallerdelta values are between Turnix and membersof th e Gruiformes and Charadriides; the largerdelta values are between Turnix and the ratites,tinamous, galliforms and anser iforms. Members of several other non-passerine groups produced delta values between the extremes. Thus,Turnix seems to have no close living relatives,but Turnix species begin to breed at th e age ofthree to five months (Bruning 1985), thereforeit is possible that their average rate of genomicevolution is faster than the rate(s) among theirnearest relatives. I f so, the branch length weobserve may be longer by an unknown factorand the correct position of the branch node maybe elsewhere in th e tree, possibly among thegruiforms where the Turnicidae have often beenassigned. The problem is not insoluble, but willrequire additional comparisons and, perhaps,t he development o f a correction for differentages at first breeding. For now we leave theTurnicidae in the position indicated in Figs. 1and 2, but designate the group as incertae sedis.

~ E ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ R a l l i d a eq.1 (76) Eurypygidae17.5 (27) Otldidae16.9 (47) Cariamidae:.... 15.0 (13) Rhynochetidae'" ~ 5 . 02J Psophiidae. 12.7 (13) Heliornithidae",9",.'-,"C",'9,-) Gruidae

, - Pteroclidae

g ~ ~ ~ ~ ~ ~ ~ ~ J a C a n i d a e12.5 (5) Rostratulidae-::: j{,4.1 (49) Thinocoridae10.8 (8) Pedionomidae'--' 13.3 (46) Scolopacidae- Charadriidae~ 10.8 (32) Burhinidae11,5 (27) ChionididaeL::.8::..C:::.87:::. - I T ~ ! & : : = = = Glareo !idae8.1 (34) Laridae

~ ~ ~ @ I ] C = = = = Falconidae

-,15 2 (60J Sagittariidae

[ ' r 9 ~ C ~ 9 5 ~ ) @ ' O ~ ' ~ C ~ 5 l ~ ~ ~ ~ ~ ; ; ; ~ ~ i ; ; ; t ~ ~ ~40(112) Sulidae11 0 (5J Anhingidae121 (30) Phalacrocoracidae133(323)

~ ~ l i ~ ~ ~ ~ Ardeidae124(231) Scopldae11.9 (67 Phoenicopteridae11,5 (105) Threskiornithidael' 1 1 ' 1 1 0 ~ ~ 0 ( ~ 2 J ~ ~ ~ ~ c ~ ~ ~ ~ :10.9 (169)I ~ ; z ; : ! ! ~ = = = = Fregatfdae'110.7 (50) Spheniscidae1J';;,0;,:,.4;';C7,::8'


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412 SIBLEY, AHLQUIST, AND MONROE JR.

Suborder SuborderTyranni PasseriParvorder Parvorder

.. Corvida Passerida;:i ;:i

~ i : 5 0 0~ 6-0 -w .- a.Eo E a; 9:: c c

~ " oW o :::>"''' "' 0 "'''" " "",,,,, u '": ~ ~ ~ " :c " u " """ " ",.-'" " uE '" '"'" ; g ~ ' ~ ~ : u " "'''' '" ~ ( ] )";: CD >. Uu :C,8 : ") '-.-E '- 0>'0.'0 ,,"'.c: '" 0- E O( l )Q) - oQ) '"o 0 ... 0' - _ u ~ "0 as.2 >. 0 as.- as " u'0; 0a . oc : c 0";; : 0 ' : e . ~ a ; ~ ~ ~ ; g . ~ ~ 1:=~ ' E o g E ~ ",,,,, ~ ~.. C._ ctI ... Ec.2 ai'Qiro c. ~ . ~ . ~ ~0 J :: J :: >. u ~ f 0 :=:Q)w u- O a: 1 -1 - ::;::;0.. w Oa.. . . .J>O CDU:::2:cn ClJO

[Auk, Vol. 105

I'"

. - . ~ o o ~ "1 " ~ ~17.9 (27) I

11.7 ( 2 5 1 f ' - - '2_.8...,(,70_0:....) --- ' ' '11.7 (237)

19.7 (89)

Fig. 5.THE FAMILIES OF PASSERINE BIRDS

Families of passerine birds.

th e same categorical rank, and that each different level of branching should be assigned adifferent rank, arranged according to their relative age of origin. From Figs. 1-5 i t is apparentthat the DNA-based phylogeny contains numerous dichotomies at many levels and that itwould be impractical to ass ign a different categorical name to every branch node i n the phylogeny. Therefore, to accommodate th e severalbranches that often occur within a single rankin our system, we have adopted th e principlesof "subordination and sequencing of units"proposed by Nelson (1973). I f there are morethan t;wo branches within a taxonomic rank, thebranch with th e largest delta TsoH value is listedfirst and is the sister group of the other branches; the lineage with the next largest delta valueis listed next, etc. For dichotomous branchingsthe taxa are the same age and their positions inthe dendrogram and classification a re interchangeable. I t has been necessary to be flexiblein th e assignment of categories and there areseveral places where a t axon may be adjustedup or down in rank. Thus, th e classification attempts to reflect the branching pattern of thephylogeny, but does not claim to reproduce itin detail.

When the classification was first developedthe categories were based on units of 10 millionyears and equated with delta TsoH values on thebasis of delta TsoH 1.0 = 4.5 million years (e.g.Sibley and Ahlquist 1983, 1985a, b). I t now seemslikely that this calibration factor is not applicable to al l birds, but only to those with agesat first breeding of ca. 2-4 years. Species thatTABLE 1. Categories, endings of categorical names,and delta TsoH ranges for each categorical rank.

Category Ending Delta TsoH rangeSuperclass 33-36Class 31-33Subclass -ornithes 29-31Infraclass -aves 27-29Parvclass -ae 24.5-27Superorder -morphae 22-24.5Order -iformes 20-22Suborder -i 18-20Infraorder -ides 15.5-18Parvorder -ida 13-15.5Superfamily -oidea 11-13Family -idae 9-11Subfamily -inae 7-9Tribe -ini 4.5-7Subtribe -ina 2.2-4.5Congeneric spp. 0-2.2


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July 1988] Classification of Living Birds 413begin to breed at one year of age evolve slightlyfaster, and those tha t beg in to breed a t agesgreater than four years evolve slightly slower.The differences in average genomic rates in birdsseem to be small compared with the range inat least some other groups; for example, mammals, in which the rodents (Catzeflis et al. 1987)are evolving ca. ten times as fast as th e hominoid primates (Sibley and Ahlquist 1987d).

Although we have changed th e basis for categorical rank from absolute time to delta values,we have retained the delta TsoH ranges originally assigned to th e categories. We began byaccepting the traditional rank of Class for theAves and of Order for the Passeriformes. Thedelta TsoH value for the passerine branch is 21.6(Fig. 4), and the ordinal level became th e rangeof values from delta TsoH 20.0 to 22.2. The otherranks were developed above and below this level(Table 1). The application of this system to theproposed classification required four categoriesin addition to those traditionally used. Theseare Infraclass and Parvclass between Class andOrder, and Infraorder and Parvorder betweenOrder and Family. For consistency we havestandardized the endings for the categories below the level of Class (Table 1). We have notused the Subtribe category. DNA hybridizationdata were available for most of the major groupsof living birds. Those for which such data werelacking are l isted as "incertae sedis" and placedin their traditional position, or in one suggestedby data from other sources.

At first glance this classification may seem tobe a radical departure from th e familiar patternof the Wetmore (1960) arrangement. For example, the "higher" non-passerines (woodpeckers, rollers, etc.) have become members ofthe "lower" non-passerines, and th e waterbirds(except Anseriformes) have been moved in theopposite direction, from the beginning to theend of th e non-passerine series. The order "Pelecaniformes" has been partially dismembered,and the New World vultures are allied to thestorks, not to the Old World vultures. However,from a morphological study Cottam (1957) concluded tha t the pelicans and the Shoebill (Ba-laeniceps rex) are related, and the New Worldvultures and storks were considered close relatives by Garrod (1873) and Ligon (1967) (Sibleyand Ahlquist 1985d: 125).

There is also a high degree of congruencebetween th e DNA-based classification and theboundaries of th e families and orders long rec-

ognized by avian systematists. Most of the traditional orders and ca. 90% of th e families basedon morphological similarities have also beendelineated by the DNA comparisons, althoughtheir categorical ranks may have been changed.For example, the woodpeckers, honeyguides,barbets, and toucans cluster together; owls andgoatsuckers are sister groups; the diverse gruiforms (to our surprise) remain mostly intact,and the charadriiforms are little changed exceptfor modifications in rank. Grouse cluster withquails, ducks cluster with geese, motmots withkingfishers, hawks with eagles, pigeons withdoves, hornbills with hoopoes, and cotingaswith manakins. With few exceptions the DNAcomparisons "see" the same clusters of speciesthat are seen by the human eye.

The principal restructuring in the Passeriformes is the result of the division of th e oscines(Passeri) into th e parvorders Corvida and Passerida. The Corvida apparently evolved in Australia and New Guinea; th e Passerida in Eurasia,Africa, and North America (Sibley and Ahlquist1985b). This division is congruent with the dist ribution of at least one morphological character, namely, th e condition of the tricipital fossa in the head of the humerus; the Corvida havea single fossa that is pneumatic, th e Passeridahave two fossae which are not pneumatic (Sibley and Ahlquist 1984a, 1987a; Olson 1987).These conditions are randomly distributed inrelation to the Wetmore (1960) classification(Bock 1962).

Thus, th e major differences between our classification and traditional arrangements are atth e older levels where the higher categories arelinked to one another. At these levels we shouldexpect it to be more difficult to discern relationships from morphology because convergent and divergent evolut ion have had longerto introduce complexities into th e interpretati on of morphological characters. The DNAcomparisons detect only the degrees of nucleotide sequence similarity which, we assume, areindices of genealogical relationships.A phylogeny, and a classification based uponit , are hypotheses about th e evolution of a groupof organisms. Each segment of a classificationis a hypothesis that may be tested by severalmethods, one of which is th e deg ree of congruence with independent sources of evidence,as noted above. For example, Raikow (1987: 42)compared th e phylogeny he derived from astudy of the hindlimb musculature of the Old


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414 SIBLEY, AHLQUIST, AND MONROE JR. [Auk, Vol. 105World suboscines " . . . to one suggested by C.G. Sibley and co-workers from their studies onDNA-DNA hybridization. Although not identical, th e results of th e two studies are so similarthat the agreement cannot be attributed tochance. Given the unrelated nature of th e dataused in these different studies, the correspondence between their results must be due to theirindependent determination of an approximation of th e true historical phylogeny. The re-lationships proposed in both investigations differ markedly from those in e arlie r, moretraditional studies."There are additional examples of congruence

and of incongruence between evidence frommorphology and f rom DNA-DNA hybridization data, and there are many cases of disagreement between different studies based on morphology. Consensus has not yet been achieved,but the molecular methods provide new toolsand concepts that are being incorporated intoth e procedures and philosophy of systematics.We hope that further comparative studieswill

be made using this classification as th e basis fornew questions and hypotheses. We cannot

guarantee, and we do not claim, t ha t the clas-sif icat ion is a true reflection of the phylogenyof living birds, but we believe that it representsprogress toward that elusive goal.Some of the new names for higher categories

may seem inappropriate, for example, Passeri-morphae for a group tha t includes pigeons,cranes, shorebirds, herons, hawks, albatrosses,and passerines. This, and all family-groupnames, are based on the first use of a genericroot as the basis of a family name. Such namesare not covered by the International Rules ofZoological Nomenclature, but we have usednames that would be acceptable under the Rules.In this case Passeridae (Vigors 1825) happensto be the basis for the names of higher cate-gories that include the Passeriformes.The following publications include parts ofthe classification, some of th e DNA-DNA hybridization data on which it is based, or both:Ahlquist et al. (1984), Sibley and Ahlquist (1981,1982a, b; 1984a, b; 1985a-d; 1987a-d),and Sibleyet al. (1982, 1984). These publications includecitat ions to others that pertain to the classifi-cation.

CLASSIFICATION OF LIVING BIRDS OF THE WORLD* = New family-group concept, combining one or more family groups.** = First modern usage at family or superfamily level in DNA-DNA hybridization studies.

*** = New family-group name created in DNA-DNA hybridization studies.# = New class-group or order-group name created in DNA-DNA hybridization studies (not

covered by International Code of Zoological Nomenclature).Class AvesSubclass Neornithes

Infraclass Eoaves#Parvclass RatitaeOrder StruthioniformesSuborder Struthioni

Infraorder StruthionidesFamily Struthionidae, Ostrich

Infraorder RheidesFamily Rheidae, RheasSuborder CasuariiFamily CasuariidaeTribe Casuariini, CassowariesTribe Dromaiini, Emus

Family Apterygidae, KiwisOrder Tinamiformes

Family Tinamidae, TinamousParvclass Galloanserae#Superorder GallomorphaeOrder CraciformesSuborder Craci


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July 1988] Classification of Living BirdsFamily Cracidae, Guans, Chachalacas, etc.

Suborder MegapodiiFamily Megapodiidae, Megapodes

Order GalliformesParvorder Phasianida

Superfamily PhasianoideaFamily Phasianidae, Grouse, Pheasants, etc.Superfamily Numidoidea

Family Numididae, GuineafowlsParvorder Odontophorida

Family Odontophoridae, New World QuailsSuperorder Anserimorphae

Order AnseriformesInfraorder Anhimides

Superfamily AnhimoideaFamily Anhimidae, Screamers

Superfamily AnseranatoideaFamily Anseranatidae, Magpie GooseInfraorder AnseridesFamily Dendrocygnidae, Whistling-DucksFamily Anatidae

Subfamily Oxyurinae, Stiff-tailed DucksSubfamily Cygninae, SwansSubfamily Anatinae

Tribe Anserini, GeeseTribe Anatini, Typical Ducks

415

Infraclass?Order Turniciformes, inc. sedisFamily Turnicidae, Buttonquails = Turnix, Ortyxelos

Infraclass NeoavesParvclass Picae

Order PiciformesInfraorder Picides

Family Indicatoridae, HoneyguidesFamily Picidae, Woodpeckers, Wrynecks

Infraorder RamphastidesSuperfamily Megalaimoidea

Family Megalaimidae:* Asian BarbetsSuperfamily Lybioidea

Family Lybiidae:** African BarbetsSuperfamily Ramphastoidea

Family Ramphastidae*Subfamily Capitoninae, New World BarbetsSubfamily Ramphastinae, Toucans

Parvclass CoraciaeSuperorder Galbulimorphae

Order GalbuliformesInfraorder Galbulides

Family Galbulidae, JacamarsInfraorder Bucconides

Family Bucconidae, PuffbirdsSuperorder Bucerotimorphae

Order BucerotiformesFamily Bucerotidae, Typical HornbillsFamily Bucorvidae, Ground-Hornbills

Order UpupiformesInfraorder Upupides

Family Upupidae, Hoopoes


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Infraorder PhoeniculidesFamily Phoeniculidae, Wood-HoopoesFamily Rhinopomastidae,*** Scimitarbills

Superorder CoraciimorphaeOrder Trogoniformes

Family TrogonidaeSubfamily Apaloderminae, African TrogonsSubfamily TrogoninaeTribe Trogonini, New World TrogonsTribe Harpactini, Asian Trogons

Order CoraciiformesSuborder Coracii

Superfamily CoracioideaFamily Coraciidae, Typical RollersFamily Brachypteraciidae, Ground-Rollers (inc. sedis)

Superfamily LeptosomoideaFamily Leptosomidae, Cuckoo-Rollers

Suborder AlcediniInfraorder AlcedinidesParvorder MomotidaFamily Momotidae, MotmotsParvorder TodidaFamily Todidae, TodiesParvorder Alcedinida

Family Alcedinidae, Alcedinid KingfishersParvorder CerylidaSuperfamily DacelonoideaFamily Dacelonidae, Dacelonid KingfishersSuperfamily Ceryloidea

Family Cerylidae, Cerylid KingfishersInfraorder Meropides

Family Meropidae, Bee-eatersParvclass Coliae

Order ColiiformesFamily ColiidaeSubfamily Coliinae, Typical MousebirdsSubfamily Urocoliinae, Long-tailed Mousebirds**

Parvclass PasseraeSuperorder CuculimorphaeOrder Cuculiformes

Infraorder CuculidesParvorder CuculidaSuperfamily CuculoideaFamily Cuculidae, Old World CuckoosSuperfamily CentropodoideaFamily Centropodidae,** Coucals

Parvorder CoccyzidaFamily Coccyzidae,** American CuckoosInfraorder CrotophagidesParvorder OpisthocomidaFamily Opisthocomidae, HoatzinParvorder CrotophagidaFamily Crotophagidae

Tribe Crotophagini, AnisTribe Guirini,*** Guira CuckooParvorder NeomorphidaFamily Neomorphidae, Roadrunners, etc.

Superorder PsittacimorphaeOrder Psittaciformes

[Auk, Vol. 105


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July 1988] Classification of Living Birds 417Family Psittacidae, Parrots, Macaws, etc.

Superorder ApodimorphaeOrder Apodiformes

Family Apodidae, Typical SwiftsFamily Hemiprocnidae, Crested Swifts

Order TrochiliformesFamily TrochilidaeSubfamily Phaethornithinae, HermitsSubfamily Trochilinae, Typical Hummingbirds

Superorder StrigimorphaeOrder Musophagiformes

Family MusophagidaeSubfamily Musophaginae, TuracosSubfamily Criniferinae, Plaintain-eaters

Order StrigiformesSuborder Strigi

Parvorder TytonidaFamily Tytonidae, Barn an d Grass owlsParvorder StrigidaFamily Strigidae, Typical Owls

Suborder AegotheliFamily Aegothelidae, Owlet-nightjars

Suborder CaprimulgiInfraorder Podargides

Family Podargidae, Australian FrogmouthsFamily Batrachostomidae,*** Asian Frogmouths

Infraorder CaprimulgidesParvorder Steatornithida

Superfamily SteatornithoideaFamily Steatornithidae, Oilbird

Superfamily NyctibioideaFamily Nyctibiidae, Potoos

Parvorder CaprimulgidaSuperfamily EurostopodoideaFamily Eurostopodidae:** Eared Nightjars

Superfamily CaprimulgoideaFamily Caprimulgidae

Subfamily Chordeilinae, NighthawksSubfamily Caprimulginae, Nightjars

Superorder PasserimorphaeOrder Columbiformes

Family Columbidae, Pigeons, DovesOrder Gruiformes

Suborder GruiInfraorder Eurypygides

Family Eurypygidae, SunbitternInfraorder Otidides

Family Otididae, BustardsInfraorder Gruides

Parvorder GruidaSuperfamily Gruoidea

Family Gruidae, CranesFamily Heliornithidae

Tribe Aramini, LimpkinTribe Heliornithini, New World Sungrebe (Podica & Heliopais inc. sedis)Superfamily Psophioidea

Family Psophiidae, TrumpetersParvorder Cariamida

Family Cariamidae, Seriemas


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Family Rhynochetidae, KaguSuborder RalliFamily Rallidae, Rails, Gallinules, CootsSuborder Mesitornithi, inc. sedisFamily Mesitornithidae, Mesites

Order CiconiiformesSuborder CharadriiInfraorder PteroclidesFamily Pteroclidae, SandgrouseInfraorder CharadriidesParvorder ScolopacidaSuperfamily ScolopacoideaFamily Thinocoridae, SeedsnipeFamily Pedionomidae, Plains-wandererFamily ScolopacidaeSubfamily Scolopacinae, Woodcock, SnipeSubfamily Tringinae, Sandpipers, Curlews, Phalaropes

Superfamily JacanoideaFamily Rostratulidae, PaintedsnipeFamily Jacanidae, Lily-trotters, Jacanas

Parvorder CharadriidaSuperfamily ChionidoideaFamily Chionididae, Sheathbills

Superfamily CharadrioideaFamily Burhinidae, ThickkneesFamily CharadriidaeSubfamily RecurvirostrinaeTribe Haematopodini, OystercatchersTribe Recurvirostrini, Avocets, Stilts

Subfamily Charadriinae, Plovers, LapwingsSuperfamily Laroidea

Family GlareolidaeSubfamily Dromadinae, Crab-ploverSubfamily Glareolinae, Pratincoles

Family LaridaeSubfamily LarinaeTribe Stercorariini, Jaegers, SkuasTribe Rynchopini, SkimmersTribe Larini, GullsTribe Sternini, Terns

Subfamily Alcinae, Auks, Murres, PuffinsSuborder CiconiiInfraorder FalconidesParvorder Accipitrida

Family AccipitridaeSubfamily Pandioninae, OspreySubfamily Accipitrinae, Hawks, Eagles

Family Sagittariidae, SecretarybirdParvorder FalconidaFamily Falconidae, Falcons, Caracaras

Infraorder CiconiidesParvorder PodicipedidaFamily Podicipedidae, GrebesParvorder Phaethontida

Family Phaethontidae, TropicbirdsParvorder SulidaSuperfamily SuloideaFamily Sulidae, Boobies, GannetsFamily Anhingidae, Anhingas

[Auk, Vol. 105


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July 1988] Classification of Living Birds 419Superfamily Phalacrocoracoidea

Family Phalacrocoracidae, CormorantsParvorder CiconiidaSuperfamily Ardeoidea

Family Ardeidae, Herons, Bitterns, EgretsSuperfamily ScopoideaFamily Scopidae, HammerheadSuperfamily Phoenicopteroidea

Family Phoenicopteridae, FlamingosSuperfamily Threskiornithoidea

Family Threskiornithidae, Ibises, SpoonbillsSuperfamily Pelecanoidea

Family PelecanidaeSubfamily Balaenicipitinae, ShoebillSubfamily Pelecaninae, Pelicans

Superfamily CiconioideaFamily CiconiidaeSubfamily Cathartinae, New World vulturesSubfamily Ciconiinae, Storks

Superfamily ProcellarioideaFamily Fregatidae, FrigatebirdsFamily Spheniscidae, PenguinsFamily Gaviidae, LoonsFamily Procellariidae

Subfamily Hydrobatinae, Storm-PetrelsSubfamily Procellariinae, Shearwaters, Petrels, Diving-PetrelsSubfamily Diomedeinae, Albatrosses

Order PasseriformesSuborder Tyranni (Suboscines)

Infraorder AcanthisittidesFamily Acanthisittidae, New Zealand Wrens

Infraorder EurylaimidesSuperfamily Pittoidea

Family Pittidae, PittasSuperfamily Eurylaimoidea

Family Eurylaimidae, BroadbillsFamily Philepittidae, Asities (inc. sedis)

Infraorder TyrannidesParvorder Tyrannida

Family TyrannidaeSubfamily Corythopinae, Corythopis, Mionectes, etc.Subfamily Tyranninae, Tyrant FlycatchersSubfamily TityrinaeTribe Schiffornithini, SchiffornisTribe Tityrini, Tityras, Becards

Subfamily Cotinginae, Cotingas, Plantcutters, SharpbillsSubfamily Piprinae, Manakins

Parvorder ThamnophilidaFamily Thamnophilidae, Typical Antbirds

Parvorder FurnariidaSuperfamily Furnarioidea

Family FurnariidaeSubfamily Furnariinae, OvenbirdsSubfamily Dendrocolaptinae, Woodcreepers

Superfamily FormicarioideaFamily Formicariidae, Ground AntbirdsFamily Conopophagidae, GnateatersFamily Rhinocryptidae, Tapaculos

Suborder Passeri (Oscines)


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420 SIBLEY, AHLQUIST, AND MONROE JR. [Auk, Vol. 105Parvorder Corvida

Superfamily MenuroideaFamily Climacteridae, Australo-Papuan TreecreepersFamily Menuridae

Subfamily Menurinae, LyrebirdsSubfamily Atrichornithinae, ScrubbirdsFamily Ptilonorhynchidae, Bowerbirds

Superfamily MeliphagoideaFamily Maluridae

Subfamily MalurinaeTribe Malurini, FairywrensTribe Stipiturini, Emuwrens

Subfamily Amytornithinae, GrasswrensFamily Meliphagidae, Honeyeaters, incl. Ephthianura, AshbyiaFamily Pardalotidae

Subfamily Pardalotinae, PardalotesSubfamily Dasyornithinae, BristlebirdsSubfamily AcanthizinaeTribe Sericornithini, Scrubwrens

Tribe Acanthizini, Thornbills, Whitefaces, etc.Superfamily Corvoidea

Family Eopsaltriidae, Australo-Papuan robins, DrymodesFamily Irenidae, Fairy-bluebirds, LeafbirdsFamily Orthonychidae, Log-runners or ChowchillasFamily Pomatostomidae, *** Australo-Papuan babblersFamily Laniidae, True Shrikes = Lanius, Corvinella, EurocephalusFamily Vireonidae, Vireos, Greenlets, PeppershrikesFamily Corvidae

Subfamily Cinclosomatinae, Quail-thrushes, WhipbirdsSubfamily Corcoracinae, Australian Chough, ApostlebirdSubfamily Pachycephalinae

Tribe Neosittini, SittellasTribe Mohouini, New Zealand Mohoua, FinschiaTribe Falcunculini, Shrike-tits, Oreoica, RhagologusTribe Pachycephalini, Whistlers, Shrike-thrushes

Subfamily CorvinaeTribe Corvini, Crows, Magpies, Jays, NutcrackersTribe Paradisaeini, Birds of Paradise, MelampittaTribe Artamini, Currawongs, Woodswallows, Peltops, PityriasisTribe Oriolini, Orioles, Cuckooshrikes

Subfamily DicrurinaeTribe Rhipidurini, FantailsTribe Dicrurini, DrongosTribe Monarchini, Monarchs, Magpie-larks

Subfamily Aegithininae, IorasSubfamily Malaconotinae

Tribe Malaconotini, Bush-shrikesTribe Prionopini, Helmet-shrikes, Batis, Platysteira, Vangas

Family Callaeatidae, New Zealand wattlebirds (inc. sedis)Parvorder Passerida

Superfamily MuscicapoideaFamily Bombycillidae

Tribe Dulini, PalmchatTribe Ptilogonatini, Silky-flycatchersTribe Bombycillini, WaxwingsFamily Cinclidae, Dippers

Family MuscicapidaeSubfamily Turdinae, True thrushes, incl. Chlamydochaera, Brachypteryx, AletheSubfamily Muscicapinae

Tribe Muscicapini, Old World FlycatchersTribe Saxicolini, Chats, Erithacus, etc.


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July 1988] Classification of Living BirdsFamily Sturnidae

Tribe Sturnini, Starlings, MynasTribe Mimini, Mockingbirds, Thrashers, American Catbirds

Superfamily SylvioideaFamily SittidaeSubfamily Sittinae, NuthatchesSubfamily Tichodromadinae, Wallcreeper

Family CerthiidaeSubfamily CerthiinaeTribe Certhiini, Northern CreepersTribe Salpornithini, Afro-Asian Creeper

Subfamily Troglodytinae, WrensSubfamily Polioptilinae, Gnatcatchers, Verdin, Gnatwrens

Family ParidaeSubfamily Remizinae, Penduline-TitsSubfamily Parinae, Titmice, Chickadees

Family Aegithalidae, Long-tailed Tits, BushtitsFamily HirundinidaeSubfamily Pseudochelidoninae, River-Martins (inc. sedis)Subfamily Hirundininae, Swallows, Martins

Family Regulidae, KingletsFamily Pycnonotidae, BulbulsFamily Hypocoliidae, Gray Hypocolius (inc. sedis)Family Cisticolidae, African WarblersFamily Zosteropidae, White-eyesFamily SylviidaeSubfamily Phylloscopinae, Leaf WarblersSubfamily Megalurinae, Grass Warblers, "Bowdleria" = MegalurusSubfamily Garrulacinae, GarrulaxSubfamily SylviinaeTribe Timaliini, Babblers, PicathartesTribe Chamaeini, WrentitTribe Sylviini, Sylviine Warblers

Superfamily PasseroideaFamily Alaudidae, LarksFamily NectariniidaeSubfamily Promeropinae, African SugarbirdsSubfamily Nectariniinae

Tribe Dicaeini, FlowerpeckersTribe Nectariniini, Sunbirds, Spiderhunters

Family MelanocharitidaeTribe Melanocharitini, Melanocharis, RhamphocharisTribe Toxorhamphini, Toxorhamphus, OedistomaFamily Paramythiidae, Paramythia, Oreocharis

Family PasseridaeSubfamily Passerinae, Sparrows, Rock-Sparrows, etc.Subfamily Motacillinae, Wagtails and PipitsSubfamily Prunellinae, Accentors, DunnockSubfamily Ploceinae, Weaverbirds, inc!' Amblyospiza, Bubalornis, etc.Subfamily EstrildinaeTribe Estrildini, Waxbills, EstrildinesTribe Viduini, Indigobirds, Whydahs

Family FringillidaeSubfamily Peucedraminae, Olive WarblerSubfamily FringillinaeTribe Fringillini, Chaffinches, Brambling

Tribe Carduelini, Goldfinches, Crossbills, etc.Tribe Drepanidini, Hawaiian Honeycreepers

Subfamily EmberizinaeTribe Emberizini, Buntings, Longspurs, TowheesTribe Parulini, Wood Warblers, incl. Zeledonia

421


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422 SIBLEY, AHLQUIST, AND MONROE JR. [Auk, Vol. 105

NEW FAMILY-GROUP TAXA

Tribe Thraupini, Tanagers, Swallow-tanager, Neotropical Honeycreepers, Plushcap, Tanager-finches

Tribe Cardinalini, CardinalsTribe Icterini, Troupials, Meadowlarks, American Blackbirds, Oropendolas

Family Lybiidae, African Barbets (Sibley and Ahlquist 1985b). Type-Genus: Lybius Hermann1783.Family Rhinopomastidae, Scimitarbills (Sibley and Ahlquist 1985b). Type-Genus: Rhinopomastus

Jardine 1828.Tribe Guirini, Guira Cuckoo. Type-Genus: Guira Lesson 1830.Family Batrachostomidae, Asian Frogmouths. Type-Genus: Batrachostomus Gould 1838.Family Eurostopodidae, Eared Nightjars (Sibley and Ahlquist 1985b). Type-Genus: EurostopodusGould 1838.Family Pomatostomidae, Australo-Papuan Babblers (Sibley and Ahlquist 1985b). Type-Genus:Pomatostomus Cabanis 1851.

ACKNOWLEDGMENTSThe research on which this classification is based

was done at Yale University between 1975 and 1986.Support for the laboratory and field work was prov id ed by the William Robertson Coe Fund of YaleUniversity and the U.S. National Science Foundationvia grants DEB-77-02594, 79-26746, and BSR-83-15155to e.G.S. Acknowledgments of the assistance of individuals have been made in some of th e publicationscited below, and in other publications pertaining tothis project. We thank G. A. Clark Jr., T. R. Howell,and R. B. Payne for their constructive comments.

LITERATURE CITEDAHLQUIST, J. E., A. H. BLEDSOE, F. H. SHELDON, & e.

G. SIBLEY. 1987. DNA hybridization and aviansystematics. Auk 104: 556-563.

--- , F. H. SHELDON, & e. G. SIBLEY. 1984. Therelationships of th e Bornean Bristlehead (Pityri-asis gymnocephala) and the Black-collared Thrush(Chlamydochaera jefferyi). J. Ornithol. 125: 129-140.

BOCK, W. 1962. The pneumatic fossa of th e humerusin the Passeres. Auk 79: 425-443.

BRUNING, D. 1985. Buttonquail. Pp. 76-77 in A dict ionary of birds (B. Campbell and E. Lack, Eds.).Calton, England, T. and A. D. Poyser, Ltd.

CATZEFLIS, F. M., F. H. SHELDON, J. E. AHLQUIST, & e.G. SIBLEY. 1987. DNA-DNA hybridization evidence of the rapid rate of muroid rodent DNAevolution. Mol. BioI. Evol. 4: 242-253.

COTTAM, P. 1957. The pelecaniform characters oftheskeletonof the shoe-bill stork,Balaeniceps rex. Zool.Bull. Brit. Mus. (Nat. Hist.) 5: 51-72.

GARROD, A. H. 1873. On certain muscles of the thighof birds and on the ir value in classification, partI. Proc. Zool. Soc. London, 1873 [1874]: 626-644.

HENNIG,W. 1966. Phylogenetic systematics. Urbana,Univ. Illinois Press.

LiGON, J. D. 1967. Relationships of th e cathartid vultures. Occasional Papers Univ. Michigan Mus.Zool. No. 651: 1-26.

NELSON, G. J. 1973. Classification as an expressionof phylogenetic relationships. Syst. Zool. 22: 344359.

OLSON, S. 1. 1987. The relat ionships of the NewGuinean ground-robins Amalocichla. Emu 87: 247248.

RAIKOW, R. J. 1987. Hindlimb myology and evolution of t he O ld World suboscine passerine birds(Acanthisittidae, Pittidae, Philepittidae, Eurylaimidae). Ornithol. Monogr. No. 41. Washington, D.e., Amer. Ornithol. Union.

SIBLEY, e. G., & J. E. AHLQUIST. 1981. The phylogenyand relationships of th e ratite birds as indicatedby DNA-DNA hybridization. Pp. 301-335 in Evolution Today, Proc. 2nd. Inti. Congr. Syst. andEvol. BioI. (G. G. E. ScudderandJ. 1. Reveal, Eds.).Pittsburgh, Hunt Inst. Botanical Documentation,Carnegie-Mellon Univ.

--- , & --- . 1982a. The relat ionships of th eHawaiian honeycreepers (Drepaninini) as indicated by DNA-DNA hybridization. Auk 99: 130140.

--- , & --_. 1982b. The relat ionships of th evireos (Vireoninae) as indicated by DNA-DNAhybridization. Wilson Bull. 94: 114-128.

--- , & --- . 1983. The phylogeny and classification of birds based on the data of DNA-DNAhybridization. In CurrentOrnithology 1: 245-292(R. F. Johnston, Ed.). New York and London, Plenum Press.

--- , & --- . 1984a. The relat ionships of th estarlings (Sturnidae: Sturnini) and the mockingbirds (Sturnidae: Mimini). Auk 101: 230-243.


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July 1988] Classification of Living Birds 423--- , & --. 1984b. The relat ionships o f the

Papuan genus Peltops. Emu 84: 181-183.--, & --. 1985a. (publ. 1986). The phylog

eny and classificationof th e passerine birds, basedon comparisons of the geneticmaterial, DNA. Pp.83-121 in Proc. 18th IntI. Ornithol. Congress,Moscow 1982 (V. D. Ilyichev and V. M. Gavrilov,Eds.). Moscow, Nauka Publ.

--, & --. 1985b. The phylogeny and classification of the Australo-Papuan passerine birds.Emu 85: 1-14.

--, & --- . 1985c. The phylogeny and classification of th e New World suboscine passerinebirds (Passeriformes: Oligomyodi: Tyranni). Pp.396-428 in Neotropical ornithology (P. A. Buckley, M. S. Foster, E. S. Morton, R. S. Ridgely, andF. G. Buckley, Eds.). Washington, D.C. Amer. Ornithol. Union Monograph No. 36.--- , & - - - . 1985d. The relationships of somegroups o f African birds, based on comparisonsofthe geneticmaterial, DNA. Pp. 115-161 in Proc.Symp. on African Vertebrates (K.-L. Schuchmann, Ed.). Bonn, Mus. A. Koenig.

--, & --. 1986. Reconstructing bird phylogeny by comparing DNA's. Sci. Amer. 254: 82 92.

--- , & --- . 1987a. Avian phylogeny reconstructed from comparisons of th e genetic material, DNA. Pp. 95-121 in Molecules and morphology in evolution: conflict or compromise (C.Patterson, Ed.). London, Cambridge Univ. Press.

--- , & ---. 1987b. The Lesser Melampitta isa bird of paradise. Emu 87: 66-68.

--- , & --- . 1987c. The relationships of fourspecies of New Zealand passerine birds. Emu 87:63-66.

--- , & --- . 1987d. DNA hybridization evidence of hominoid phylogeny: resul ts from anexpanded data set. J. Mol. Evol. 26: 99-121.

--- , & - - - . I n prep. Phy logeny and classificat ion of th e birds of the world: a study in molecular evolution. New Haven, Yale Univ. Press.

--, --, & F. H. Sheldon. 1987. DNA hybridization and avian phylogenetics. Reply toCracraft. Pp. 97-125 in Evolutionary biology, vol.21 (M. K. Hecht, B. Wallace, and G. T. Prance,Eds.). New York and London, Plenum Press.

--, R. SCHODDE, & J. E. AHLQUIST. 1984. Therelationships of the Australo-Papuan treecreepers Climacteridae as indicated by DNA-DNA hybridization. Emu 84: 236-241.--, G. R. WILLIAMS, & J. E. AHLQUIST. 1982. Therelationships of the New Zealand wrens (Acanthisittidae) as indicated by DNA-DNA hybridization. Notornis 29: 113-130.

SNEATH, P. H. A., & R. R. SOKAL. 1973. Numericaltaxonomy. San Francisco, W. H. Freeman.

VIGORS, N. A. 1825. Sketches in ornithology; or, observations on the leading affinities of some of th emore extensive groups of birds. Arrangement ofgenera of birds. Zool. J. 2: 368-405.

WETMORE, A. 1960. A classification for the b irds o fthe world. Smithson. Misc. Coll. 139 (11): 1-37.

On 25 April 1988, th e Daniel Giraud Elliot Medalwas awarded jointly to Charles G. Sibley (San FranciscoState University; Professor Emeritus, Yale University) and Jon Edward Ahlquist (Department of Zoologicaland Biomedical Sciences, Ohio University). "Their massive, insight-rich application of DNA hybridizationtechniques to bird classification revolutionized taxonomy by showing at last how to distinguish avian evolutionary relationships from convergent similarities."The prize was established in 1917 through the Daniel Giraud Elliot Fund by a gift from MargaretHendersonElliot. I t is awarded by th e National Academy of Sciences for meritorious work in zoology or paleontology

published over a three- to five-year period.

a classification of the living birds of the world based on dna-dna hybridization studies

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