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Home , Amalocichla, Anseranatidae, Apostlebird, Bornean bristlehead, Coraciimorphae, Corvida

THE AUK A QUARTERLY JOURNAL OF

VOL. 105 JULY 1988 No. 3

A CLASSIFICATION OF THE LIVING OF THE WORLD BASED ON DNA-DNA HYBRIDIZATION STUDIES

CHARLESG. SIBLEY,• JON E. AHLQUIST,2 AND BURTL. MONROE JR.3 •TiburonCenter for EnvironmentalStudies, San Francisco State University, Box855, Tiburon,California 94920 USA, 2Departmentof Zoologicaland Biomedical Sciences, Ohio University, Athens, Ohio 45701 USA, and 3Departmentof Biology,University of Louisville,Louisville, Kentucky 40292 USA

ABSTRACT.--Wepresent'a Classification of the living birdsof the world basedon the results of DNA-DNA hybridizationstudies. Several -group names are presentedformally for the first time; someof these nameshave appearedearlier in various publications.Received 21 January1988, accepted30 March 1988.

THIS classification, and the phylogeny on The dendrograms were constructedby the which it is based,were first presentedas posters method of average linkage, or UPGMA (un- at the XIX International Ornithological Con- weighted pair group method using arithmetic gressin Ottawa, Canada,in June 1986,and later averages,Sheath and Soka11973:230). The DNAs at the A.O.U. annual meetings in 1986 (Stark- of 1,058 ,of which 310 were radio-la- ville, Mississippi)and 1987(San Francisco,Cal- beled tracers,were used to reconstructthe phy- ifornia). Figures 1-5 depict the branching pat- logeny. Groupsat and abovethe ordinal level of the phylogeny above the level of were clustered from several overlapping ma- subfamilies.Dendrograms of the completeDNA- trices. For example, in the Passeriformescom- based phylogeny will be included in a forth- plete matriceswere preparedfor the suboscines, comingbook (Sibley and Ahlquist in prep.). We the and the ,and among these present the classificationat this time because matricesenough measurementswere made to parts of it have been, or will be, used in works establishthe positionsof these groups relative by other authors. to one another and to the non-.The We believe that DNA-DNA hybridization major groups were clustered on the basis of comparisonsprovide data that may be used to complete, or nearly complete, matricesconsist- reconstructthe branching pattern of the phy- ing of the radio-labeled species.At least one logeny of living birds. The delta T50H values speciesin each group in Figs. 1-5 was radio- (e.g. Sibley and Ahlquist 1981, 1983, 1986) are labeled and reciprocals between nearly all believed to be relative to time, but we no longer branches were made. The averaging effect of assumethat there is a simple relationship be- UPGMA smooths out some of the effects of dif- tween delta values and absolute time. Instead, ferent averagerates of genomic , but it seemslikely that the averagerate of nucleo- relative rate tests were used to determine the tide substitutionmay be correlatedwith factors branching of taxa. related to reproduction;for example,age at first Hennig (1966:160) proposed the age of origin breeding, generation time, number of cell di- as the ideal basisfor the ranking of taxonomic visions in the germline, and possibly other fac- categories.The application of this principle tors. This subjectis discussedby Ahlquist et al. would producea classificationin which differ- (1987), Catzeflis et al. (1987), Sibley and Ahl- ent lineageswith the sameage of origin would quist (1987d), and Sibley et al. (1987). be placed in the same categoricalrank, thus

409 The Auk 105: 409-423. July 1988 410 $IBLEY,AHLQUlST, AND MONROE JR. [Auk,Vol. 105

ORDER SUPERORDER PARVCLASS INFRACLASS

STinamir...... ormes 1Ratitae •>

Galliformes Galloansera• m -•25.9 (992•221.6 Crac{forrnesAnserlforrnes ]-- GallomorphaeAnserimorphae I O ...... Turniclformes

[.•24A[63)• ...... ] coraciae 25;[256• Cu;luf;r;Ss. Cuculimor haeColiae Trochiliformes• Apod•morphae • •)(2n .....i...... • g p Passerae

Fig. 1. Major divisions of the classAves. The numbers, e.g. 28.0 (316), indicate that the average delta T5oHvalue of 28.0 is basedon the delta Ts0Hvalues of 316 DNA-DNA hybridsbetween taxa on the two sides of the branch node. achievingthe desirablegoal of categoricalequiv- categoricalequivalence into our classification alence.Hennig (1966: 161-182) discussedthe by assigningranks on the basisof delta T50H problemsassociated with determiningthe ages values (Table 1). The delta values express de- of origin of systematicgroups and concluded greesof genomicdivergence and, sincethe av- that the availablemethods were inadequate.The eragerate of divergenceis relative to time, this presentmethods still fall shortof the ideal, but procedureprovides an approachto categorical we have tried to incorporate the principle of equivalencebased on time of origin. The fact

I I • THEFAMILIES OFNON- BIRDS Fig. 2. Families of non-Passerine birds, to Dacelonidae. July1988] ClassificationofLiving Birds 411

• 24.5(132) 19.1 (76} Eurypygidae 17.5 (27} • 14.5(6} CrotophagidaeNeornorphidae Otididae • 16.4(5) 16 9 (47} Cariarnidae • 176 (53) Coccyzidae Rhynochetidae ß P 15.0 (2} 14.• Centropodidae He]iornithidae Gruidae Psittacidae Pteroclidae

• . •95 (4} Hem procnidae Jacanidae 12.5 (5} Rostratulidae 14.1 (49) Thinocorldae -- 10.8 (8} onomidae (46} )acidae

• 13.6(7) Tytonidae Gharadriidae lO.6 (92} Bur hinidae 19.1{55) Aegothelidae 11 5 (27) Chionldidae 18.8 (26} Podargidae 10.3 {2} Batrachostornidae (87) Glareolidae • Nyctibiidae (34} Laridae

Falconidae • 169{10) 12.3 (3} Caprimulgidae Sagittariidae Ipitridae __ Podicipedidae Phaethontidae (112) 11.0(5} gidae 12.1(30) Phalacrocoracidae (323} Ardeldae Scopidae Phoenicopterldae Fig. 3. Families of non-passerinebirds, Coliidae to Columbidae. (104} Ciconiidae 10.1(82) Pelecanidae 10.9 (169} Fregatidae 10.7(50} Spheniscidae that different lineagesevolve at somewhatdif- Gaviidae 10.0 (47} ferent averagerates introduces an error of un- certain magnitude. 21.6(129) Passeriformes The Turnicidaemay presenta specialprob- Fig. 4. Familiesof non-passerinebirds, Rallidae lem. The 27 delta T50H values between Turnix to Procellariidae, and the position of the Passeri- and other taxa are large, ranging from 22.7 to formes. the pectoralsandpiper (Calidris melanoto•) to 29.8 to the ostrich (Struthiocamelus); average = 27.0. Does the Turnix problem cast doubt on the The rangeof 7.1 is unusuallylarge. The smaller positionsof all taxa in the tree? We think not, delta values are between Turnix and members becausenearly all other groupsconsist of many of the and Charadriides;the larger speciesand severallevels of branchings,but the delta values are between Turnix and the ratites, memberscluster in spite of somewhatdifferent , galliforms and anseriforms. Mem- agesat maturity. For example,members of the bers of severalother non-passerinegroups pro- Procellariidaerange in ageat first breedingfrom duced delta values between the extremes. Thus, ca. 3-4 years (Storm-Petrels)to more than !0 Turnixseems to have no close living relatives, years(large ).Differences in branch but Turnixspecies begin to breed at the age of lengths are revealedby relative rate tests,but three to five months (Bruning 1985), therefore all procellariids are closer to one another than it is possiblethat their averagerate of genomic to any outsidegroup. Similarly, the estrildine evolution is faster than the rate(s)among their finchesbegin to breed when lessthan one year nearestrelatives. If so, the branch length we of age,but they clusterwith the other members observemay be longer by an unknown factor of the Passeridae and are closest to the yiduines, and the correctposition of the branch node may as expected. be elsewhere in the tree, possibly among the Thus, it is possible that the Turnicidae are gruiformswhere the Turnicidaehave often been actually as isolatedas their position in Figs. ! assigned.The problemis not insoluble,but will and 2 suggests,but their exceptionallyearly require additional comparisonsand, perhaps, maturity, and possiblerapid DNA evolution, the developmentof a correctionfor different raise a question that must be considered, but agesat first breeding. For now we leave the cannot be resolved at this time. Turnicidae in the position indicated in Figs. ! Hennig (!966: !54-!60) proposedthat the sis- and 2, but designatethe group as incertaesedis. ter branchesat each dichotomy should receive 412 SmLE¾,AHLQUIST, AND MONROE JR. [Auk,Vol. 105

Suborder Suborder Passeri Parvorder Parvorder Corvida Passerida

•..=__='E •o.- ß

THE FAMILIES OF PASSERINE BIRDS t9.7 [89)

Fig. 5. Familiesof passefinebirds. the samecategorical rank, and that each differ- When the classification was first developed ent level of branching should be assigneda the categorieswere basedon units of 10 million different rank, arranged accordingto their rel- yearsand equatedwith delta T50Hvalues on the ative age of origin. From Figs. 1-5 it is apparent basisof delta T•0H 1.0 = 4.5 million years (e.g. that the DNA-based phylogeny contains nu- Sibleyand Ahlquist 1983, 1985a, b). It now seems merous dichotomies at many levels and that it likely that this calibration factor is not appli- would be impractical to assign a different cat- cable to all birds, but only to those with ages egoricalname to every branchnode in the phy- at first breeding of ca. 2-4 years. Speciesthat logeny. Therefore,to accommodatethe several branchesthat often occurwithin a single rank TABLE1. Categories,endings of categoricalnames, in our system,we have adoptedthe principles and delta Ts0Hranges for each categoricalrank. of "subordination and sequencing of units" proposedby Nelson (1973). If there are more Category Ending Delta TsoHrange than two branches within a , the Superclass -- 33-36 branchwith the largestdelta Ts0Hvalue is listed -- 31-33 first and is the sistergroup of the other branch- Subclass -ornithes 29-31 Infraclass -aves 27-29 es;the lineagewith the next largestdelta value Parvclass -ae 24.5-27 is listed next, etc. For dichotomousbranchings Superorder -morphae 22-24.5 the taxa are the same age and their positionsin Order -iformes 20-22 the dendrogram and classificationare inter- Suborder -i 18-20 changeable.It hasbeen necessary to be flexible Infraorder -ides 15.5-18 Parvorder -ida 13-15.5 in the assignmentof categoriesand there are Superfamily -oidea 11-13 several places where a may be adjusted Family -idae 9-11 up or down in rank. Thus, the classificationat- Subfamily -inae 7-9 tempts to reflect the branching pattern of the Tribe -ini 4.5-7 phylogeny, but does not claim to reproduceit Subtribe -ina 2.2-4.5 in detail. Congenericspp. -- 0-2.2 July1988] ClassificationofLiving Birds 413 begin to breed at one year of age evolve slightly ognizedby avian systematists.Most of the tra- faster, and those that begin to breed at ages ditional orders and ca. 90% of the families based greaterthan four years evolve slightly slower. on morphologicalsimilarities have also been The differencesin averagegenomic rates in birds delineatedby the DNA comparisons,although seem to be small comparedwith the range in their categoricalranks may have been changed. at least some other groups;for example, mam- For example, the ,honeyguides, mals, in which the (Catzeflis et al. 1987) barbets,and toucanscluster together; and are evolving ca. ten times as fast as the homi- goatsuckersare sistergroups; the diverse gru- noid primates (Sibley and Ahlquist 1987d). iforms (to our surprise)remain mostly intact, Although we have changedthe basisfor cat- and the charadriiformsare little changedexcept egoricalrank from absolutetime to delta values, for modifications in rank. cluster with we have retained the delta T50Hranges origi- , cluster with geese,motmots with nally assignedto the categories.We began by ,hawks with eagles,pigeons with acceptingthe traditional rank of Classfor the doves, with hoopoes, and Aves and of Order for the Passeriformes. The with .With few exceptionsthe DNA delta T50Hvalue for the passerinebranch is 21.6 comparisons"see" the sameclusters of species (Fig. 4), and the ordinal level becamethe range that are seen by the human eye. of values from delta T50H 20.0 to 22.2. The other The principal restructuring in the Passeri- rankswere developed above and below this level formes is the result of the division of the oscines (Table 1). The application of this systemto the (Passeri)into the parvordersCorvida and Pas- proposedclassification required four categories serida.The Corvidaapparently evolved in Aus- in addition to those traditionally used. These tralia and New Guinea; the Passerida in Eurasia, are Infraclass and Parvclass between Class and , and North America (Sibley and Ahlquist Order, and Infraorder and Parvorder between 1985b).This divisionis congruentwith the dis- Order and Family. For consistencywe have tribution of at least one morphological char- standardizedthe endings for the categoriesbe- acter,namely, the conditionof the tricipital fos- low the level of Class (Table 1). We have not sa in the head of the humerus; the Corvida have usedthe Subtribecategory. DNA hybridization a single fossathat is pneumatic,the Passerida datawere availablefor mostof the majorgroups have two fossaewhich are not pneumatic (Sib- of living birds. Those for which suchdata were ley and Ahlquist 1984a, 1987a; Olson 1987). lacking are listed as "incertaesedis" and placed These conditions are randomly distributed in in their traditionalposition, or in onesuggested relation to the Wetmore (1960) classification by data from other sources. (Bock 1962). At first glancethis classificationmay seemto Thus,the majordifferences between our clas- be a radical departurefrom the familiar pattern sification and traditional arrangementsare at of the Wetmore (1960) arrangement.For ex- the older levelswhere the higher categoriesare ample, the "higher" non-passerines (wood- linked to one another. At these levels we should peckers,rollers, etc.) have becomemembers of expect it to be more difficult to discern rela- the "lower" non-passerines,and the waterbirds tionships from morphology because conver- (exceptAnseriformes) have been moved in the gent and divergent evolution have had longer opposite direction, from the beginning to the to introduce complexitiesinto the interpreta- end of the non-passerineseries. The order "Pel- tion of morphological characters.The DNA ecaniformes"has been partially dismembered, comparisonsdetect only the degreesof nucleo- and the New World vultures are allied to the tide sequencesimilarity which, we assume,are , not to the Old World vultures. However, indices of genealogicalrelationships. from a morphologicalstudy Cottam (1957) con- A phylogeny,and a classificationbased upon cluded that the pelicansand the (Ba- it, arehypotheses about the evolution of a group laenicepsrex) are related, and the New World of organisms.Each segment of a classification vultures and storks were considered close rel- is a hypothesisthat may be tested by several ativesby Garrod (1873)and Ligon (1967) (Sibley methods, one of which is the degree of con- and Ahlquist 1985d: 125). gruencewith independentsources of evidence, There is also a high degree of congruence asnoted above.For example,Raikow (1987:42) between the DNA-based classification and the comparedthe phylogeny he derived from a boundariesof the familiesand orderslong rec- study of the hindlimb musculatureof the Old 414 SIBLEY,AI-ILQVIST, A•D MONROEJR. [Auk, Vol. 105

World suboscines"... to one suggestedby C. guarantee,and we do not claim, that the clas- G. Sibley and co-workers from their studies on sification is a true reflection of the phylogeny DNA-DNA hybridization. Although not iden- of living birds, but we believe that it represents tical, the results of the two studies are so similar progresstoward that elusive goal. that the agreement cannot be attributed to Someof the new namesfor higher categories chance. Given the unrelated nature of the data may seem inappropriate, for example, Passeri- used in these different studies, the correspon- morphae for a group that includes pigeons, dence between their results must be due to their cranes, shorebirds, , hawks, albatrosses, independent determination of an approxima- and passetines. This, and all family-group tion of the true historical phylogeny. The re- names,are basedon the first use of a generic lationshipsproposed in both investigationsdif- root as the basisof a family name. Such names fer markedly from those in earlier, more are not covered by the International Rules of traditional studies." Zoological Nomenclature, but we have used There are additional examplesof congruence namesthat would be acceptableunder the Rules. and of incongruencebetween evidence from In this casePasseridae (Vigors 1825) happens morphology and from DNA-DNA hybridiza- to be the basis for the names of higher cate- tion data, and there are many casesof disagree- goriesthat include the Passeriformes. ment between different studies based on mor- The following publications include parts of phology.Consensus has not yet been achieved, the classification,some of the DNA-DNA hy- but the molecular methods provide new tools bridization data on which it is based, or both: and conceptsthat are being incorporatedinto Ahlquist et al. (1984), Sibley and Ahlquist (1981, the proceduresand philosophy of systematics. 1982a,b; 1984a,b; 1985a-d; 1987a-d), and Sibley We hope that further comparativestudies will et al. (1982, 1984). These publications include be madeusing this classificationas the basisfor citations to others that pertain to the classifi- new questions and hypotheses. We cannot cation.

CLASSIFICATION OF LIVING BIRDS OF THE WORLD

* = New family-groupconcept, combining one or more family groups. ** = First modernusage at family or superfamilylevel in DNA-DNA hybridizationstudies. *** = New family-groupname createdin DNA-DNA hybridization studies. # = New class-groupor order-groupname createdin DNA-DNA hybridizationstudies (not coveredby International Code of ZoologicalNomenclature).

Class Aves Subclass Neornithes Infraclass Eoaves# Parrclass Ratitae Order Struthioniformes Suborder Struthioni Infraorder Struthionides Family Struthionidae,Ostrich Infraorder Rheides Family , Rheas Suborder Casuarii Family Casuariidae Tribe Casuariini, Cassowaries Tribe Dromaiini, Emus Family Apterygidae, Kiwis Order Tinamiformes Family Tinamidae, Tinamous Parvclass Galloanserae# Superorder Gallomorphae Order Craciformes Suborder Craci July1988] ClassificationofLiving Birds 415

Family ,Guans, Chachalacas, etc. Suborder Megapodii Family Megapodiidae, Order Parvorder Phasianida Superfamily Phasianoidea Family ,Grouse, , etc. Superfamily Numidoidea Family Numididae, Parvorder Odontophorida Family Odontophoridae,New World Quails Superorder Anserimorphae Order Infraorder Anhimides Superfamily Anhimoidea Family Anhimidae, SuperfamilyAnseranatoidea Family ,Magpie Infraorder Anserides Family Dendrocygnidae,Whistling-Ducks Family SubfamilyOxyurinae, Stiff-tailed Ducks Subfamily Cygninae, Subfamily Tribe Anserini, Geese Tribe Anatini, Typical Ducks

Infraclass? Order Turniciformes, inc. sedis Family Turnicidae,Buttonquails = Turnix,Ortyxelos Infraclas$ Parvclass Picae Order Infraorder Picides Family Indicatoridae,Honeyguides Family Picidae, Woodpeckers,Wrynecks Infraorder Ramphastides SuperfamilyMegalaimoidea Family Megalaimidae,** Asian Barbets Superfamily Lybioidea Family Lybiidae,*** African Barbets Superfamily Ramphastoidea Family Ramphastidae* SubfamilyCapitoninae, New World Barbets SubfamilyRamphastinae, Toucans Parvclass Coraciae SuperorderGalbulimorphae Order Galbuliformes Infraorder Galbulides Family Galbulidae,Jacamars Infraorder Bucconides Family Bucconidae,Puffbirds SuperorderBucerotimorphae Order Family Bucerotidae,Typical Hornbills Family Bucorvidae,Ground-Hornbills Order Upupiformes Infraorder Upupides Family Upupidae, Hoopoes 416 S•BLE¾,AHLQUIST, AND MONROE JR. [Auk, Vol. 105

Infraorder Phoeniculides Family Phoeniculidae,Wood-Hoopoes Family Rhinopomastidae,***Scimitarbills Superorder Coraciimorphae Order Trogoniformes Family Trogonidae Subfamily Apaloderminae,African Subfamily Trogoninae Tribe Trogonini, New World Trogons Tribe Harpactini, Asian Trogons Order Suborder Coracii Superfamily Coracioidea Family Coraciidae,Typical Rollers Family Brachypteraciidae,Ground-Rollers (inc. sedis) Superfamily Leptosomoidea Family Leptosomidae,-Rollers Suborder Alcedini Infraorder Alcedinides Parvorder Momotida Family Momotidae, Motmots Parvorder Todida Family Todidae, Todies Parrorder Alcedinida Family Alcedinidae, Alcedinid Kingfishers Parrorder Cerylida Superfamily Dacelonoidea Family Dacelonidae,Dacelonid Kingfishers Superfamily Ceryloidea Family Cerylidae,Cerylid Kingfishers Infraorder Meropides Family Meropidae, Bee-eaters Parrclass Coliae Order Coliiformes Family Coliidae Subfamily Coliinae, Typical Subfamily Urocoliinae, Long-tailed Mousebirds** Parvclass Passerae SuperorderCuculimorphae Order Cuculiformes Infraorder Cuculides Parrorder Cuculida Superfamily Cuculoidea Family Cuculidae, Old World Superfamily Centropodoidea Family Centropodidae,**Coucals Parvorder Coccyzida Family Coccyzidae,**American Cuckoos Infraorder Crotophagides Parvorder Opisthocomida Family Opisthocomidae,Hoatzin Parrorder Crotophagida Family Crotophagidae Tribe Crotophagini, Anis Tribe Guirini,*** Guira Cuckoo Parvorder Neomorphida Family Neomorphidae, Roadrunners, etc. Superorder Psittacimorphae Order Psittaciformes July1988] ClassificationofLiving Birds 417

Family Psittacidae,, Macaws, etc. Superorder Apodimorphae Order Family Apodidae, Typical Swifts Family Hemiprocnidae, CrestedSwifts Order Trochiliformes Family Trochilidae Subfamily Phaethornithinae, Hermits Subfamily Trochilinae, Typical Superorder Strigimorphae Order Musophagiformes Family Musophagidae Subfamily Musophaginae, Subfamily Criniferinae, Plaintain-eaters Order Strigiformes SuborderStrigi Parvorder Tytonida Family Tytonidae, Barn and Grassowls Parvorder Strigida Family Strigidae,Typical Owls SuborderAegotheli Family Aegothelidae,Owlet-nightjars SuborderCaprimulgi Infraorder Podargides Family Podargidae,Australian Family Batrachostomidae,***Asian Frogmouths Infraorder Caprimulgides Parvorder Steatornithida Superfamily Steatornithoidea Family Steatornithidae, Superfamily Nyctibioidea Family Nyctibiidae, Potoos Parvorder Caprimulgida Superfamily Eurostopodoidea FamilyEurostopodidae,*** Eared Nightjars SuperfamilyCaprimulgoidea Family Caprimulgidae SubfamilyChordeilinae, Nighthawks SubfamilyCaprimulginae, Nightjars Superorder Passerimorphae Order Columbiformes Family Columbidae,Pigeons, Doves Order Gruiformes Suborder Grui Infraorder Eurypygides Family Eurypygidae,Sunbittern Infraorder Otidides Family Otididae, Infraorder Gruides Parvorder Gruida Superfamily Gruoidea Family Gruidae, Cranes Family Tribe Aramini, Tribe Heliornithini, New World (Podica & Heliopaisinc. sedis) Superfamily Psophioidea Family Psophiidae, Trumpeters Parvorder Cariamida Family Cariamidae, Seriemas 418 SIBLE¾,ArtLQUIST, AND MONROE JR. [Auk, Vol. 105

Family Rhynochetidae, Suborder Ralli Family Rallidae,Rails, Gallinules, Suborder Mesitornithi, inc. sedis Family Mesitornithidae, Order Ciconiiformes Suborder Charadrii Infraorder Pteroclides Family Pteroclidae, Infraorder Charadriides Parvorder Scolopacida SuperfamilyScolopacoidea Family Thinocoridae,Seedsnipe Family Pedionomidae,Plains-wanderer Family Scolopacidae SubfamilyScolopacinae, Woodcock, Snipe SubfamilyTringinae, Sandpipers, Curlews, Phalaropes SuperfamilyJacanoidea Family Rostratulidae,Paintedsnipe Family Jacanidae,Lily-trotters, Jacanas Parvorder Charadriida Superfamily Chionidoidea Family Chionididae,Sheathbills SuperfamilyCharadrioidea Family Burhinidae,Thickknees Family Charadriidae Subfamily Recurvirostrinae Tribe Haematopodini,Oystercatchers Tribe Recurvirostrini, Avocets, Stilts SubfamilyCharadriinae, Plovers, Lapwings SuperfamilyLaroidea Family Glareolidae SubfamilyDromadinae, Crab-plover SubfamilyGlareolinae, Pratincoles Family Laridae Subfamily Larinae Tribe Stercorariini,Jaegers, Skuas Tribe Rynchopini,Skimmers Tribe Larini, Gulls Tribe Sternini, SubfamilyAlcinae, Auks, Murres, Puffins Suborder Ciconii Infraorder Falconides Parvorder Accipitrida Family Accipitridae Subfamily Pandioninae,Osprey SubfamilyAccipitrinae, Hawks, Eagles Family Sagittariidae,Secretarybird Parvorder Falconida Family Falconidae,Falcons, Caracaras Infraorder Ciconiides Parvorder Podicipedida Family Podicipedidae, Parvorder Phaethontida Family Phaethontidae, Parvorder Sulida Superfamily Suloidea Family Sulidae,Boobies, Gannets Family Anhingidae,Anhingas July1988] Classificationof Living Birds 419

Superfamily Phalacrocoracoidea Family Phalacrocoracidae, Parvorder Ciconiida Superfamily Ardeoidea Family Ardeidae, Herons, , Egrets Superfamily Scopoidea Family Scopidae,Hammerhead Superfamily Phoenicopteroidea Family Phoenicopteridae, Superfamily Threskiornithoidea Family Threskiornithidae,, Superfamily Pelecanoidea Family Pelecanidae Subfamily Balaenicipitinae,Shoebill Subfamily Pelecaninae, Superfamily Ciconioidea Family Ciconiidae Subfamily Cathartinae,New World vultures Subfamily Ciconiinae, Storks Superfamily Procellarioidea Family Fregatidae, Family Spheniscidae, Family Gaviidae, Family Procellariidae Subfamily Hydrobatinae, Storm-Petrels Subfamily Procellariinae,Shearwaters, Petrels, Diving-Petrels Subfamily Diomedeinae,Albatrosses Order Passeriformes SuborderTyranni (Suboscines) Infraorder Acanthisittides Family Acanthisittidae, New Zealand Infraorder Eurylaimides Superfamily Pittoidea Family Pittidae, Superfamily Eurylaimoidea Family ,Broadbills Family Philepittidae, (inc. sedis) Infraorder Parvorder Tyrannida Family Tyrannidae Subfamily Corythopinae,Corythopis, Mionectes, etc. Subfamily Tyranninae, Tyrant Flycatchers Subfamily Tityrinae Tribe Schiffornithini, Schiffornis Tribe Tityrini, Tityras, Becards Subfamily Cotinginae, Cotingas,Plantcutters, Sharpbills Subfamily Piprinae, Manakins Parvorder Thamnophilida Family Thamnophilidae,Typical Parvorder Furnariida Superfamily Furnarioidea Family Furnariidae Subfamily Furnariinae, Ovenbirds Subfamily Dendrocolaptinae, Superfamily Formicarioidea Family , Ground Antbirds Family Conopophagidae, Family Rhinocryptidae, Suborder Passeri (Oscines) 420 $IBLEY,AHLQUIST, AND MONROE JR. [Auk,Vol. 105

Parvorder Corvida Superfamily Menuroidea Family Climacteridae,Australo-Papuan Family Menuridae Subfamily Menurinae, Subfamily Atrichornithinae, Family Ptilonorhynchidae, SuperfamilyMeliphagoidea Family Maluridae Subfamily Malurinae Tribe Malurini, Fairywrens Tribe Stipiturini, Emuwrens SubfamilyAmytornithinae, Family Meliphagidae, ,incl. Ephthianura,Ashbyia Family Pardalotidae SubfamilyPardalotinae, Subfamily Dasyornithinae, Subfamily Acanthizinae Tribe Sericornithini, Scrubwrens Tribe Acanthizini, Thornbills, Whitefaces, etc. SuperfamilyCorvoidea Family Eopsaltriidae,Australo-Papuan robins, Family Irenidae, Fairy-, Family ,Log-runners or Family Pomatostomidae,***Australo-Papuan babblers Family Laniidae,True = ,Corvinella, Eurocephalus Family Vireonidae,, Greenlets, Peppershrikes Family SubfamilyCinclosomatinae, -thrushes, Whipbirds SubfamilyCorcoracinae, Australian Chough, Subfamily Tribe Neosittini, Tribe Mohouini, New Zealand Mohoua, Finschia Tribe Falcunculini, -tits, Oreoica,Rhagologus Tribe Pachycephalini,Whistlers, Shrike-thrushes Subfamily Corvinae Tribe Corvini, Crows,Magpies, Jays, Nutcrackers Tribe Paradisaeini,Birds of Paradise, Tribe Artamini, ,, Peltops, Pityriasis Tribe Oriolini, Orioles, Subfamily Dicrurinae Tribe Rhipidurini, Tribe Dicrurini, Tribe Monarchini, Monarchs,Magpie- Subfamily Aegithininae, Subfamily Malaconotinae Tribe Malaconotini, Bush-shrikes Tribe Prionopini, Helmet-shrikes,Batis, Platysteira, Family Callaeatidae,New Zealandwattlebirds (inc. sedis) Parvorder Passerida Superfamily Family Bombycillidae Tribe Dulini, Tribe Ptilogonatini,Silky-flycatchers Tribe Bombycillini, Family Cinclidae, Family Muscicapidae SubfamilyTurdinae, True thrushes,incl. Chlamydochaera,Brachypteryx, Alethe Subfamily Muscicapinae Tribe Muscicapini,Old World Flycatchers Tribe Saxicolini, Chats, Erithacus, etc. July1988] Classificationof Living Birds 421

Family Sturnidae Tribe Sturnini, ,Mynas Tribe Mimini, Mockingbirds, Thrashers,American Superfamily Family Sittidae Subfamily Sittinae, SubfamilyTichodromadinae, Family Certhiidae Subfamily Certhiinae Tribe Certhiini, Northern Creepers Tribe Salpornithini,Afro-Asian Creeper Subfamily Troglodytinae,Wrens SubfamilyPolioptilinae, , Verdin, Gnatwrens Family Paridae Subfamily Remizinae, Penduline-Tits Subfamily Parinae, Titmice, Chickadees Family , Long-tailed Tits, Bushtits Family Hirundinidae Subfamily Pseudochelidoninae,River-Martins (inc. sedis) Subfamily Hirundininae, , Martins Family Regulidae, Family Pycnonotidae, Family Hypocoliidae,Gray Hypocolius (inc. sedis) Family , African Family Zosteropidae,White-eyes Family SubfamilyPhylloscopinae, Leaf Warblers SubfamilyMegalurinae, Grass Warblers, "Bowdleria" = Megalurus Subfamily Garrulacinae,Garrulax Subfamily Sylviinae Tribe Timaliini, Babblers, Tribe Chamaeini, Wrentit Tribe Sylviini, Sylviine Warblers Superfamily Passeroidea Family Alaudidae, Larks Family Nectariniidae SubfamilyPromeropinae, African Subfamily Nectariniinae Tribe Dicaeini, Tribe Nectariniini,, Spiderhunters Family Tribe Melanocharitini,, Rhamphocharis Tribe Toxorhamphini,, Family Paramythiidae,Paramythia, Oreocharis Family Passeridae SubfamilyPasserinae, Sparrows, Rock-Sparrows, etc. SubfamilyMotacillinae, Wagtails and Subfamily Prunellinae, ,Dunnock SubfamilyPloceinae, Weaverbirds, incl. Amblyospiza,Bubalornis, etc. Subfamily Estrildinae Tribe Estrildini, Waxbills, Estrildines Tribe Viduini, Indigobirds,Whydahs Family Fringillidae Subfamily Peucedraminae,Olive Subfamily Fringillinae Tribe Fringillini, Chaffinches,Brambling Tribe Carduelini, Goldfinches, Crossbills, etc. Tribe Drepanidini, Hawaiian Honeycreepers Subfamily Emberizinae Tribe Emberizini, Buntings,Longspurs, Towhees Tribe Parulini, Wood Warblers, incl. Zeledonia 422 SIBLEY,AHLQUIST, AND MONROE JR. [Auk,Vol. 105

Tribe Thraupini,, -, Neotropical Honeycreepers, Plush- cap, Tanager- Tribe Cardinalini, Cardinals Tribe Icterini, Troupials,Meadowlarksß American Blackbirdsß Oropendolas

I•EW FAMILY-GROUP TAXA

Family Lybiidae, African Barbets(Sibley and Ahlquist 1985b).Type-: Lybius Hermann 1783. 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,Pared Nightjars (Sibley and Ahlquist 1985b).Type-Genus: Eurostopodus Gould 1838. Family Pomatostomidae,Australo-Papuan Babblers (Sibley and Ahlquist 1985b). Type-Genus: Pomatostomus Cabanis 1851.

ACKNOWLEDGMENTS HENNIG,W. 1966. Phylogeneticsystematics.Urbana, Univ. Illinois Press. The research on which this classification is based L•GON,J.D. 1967. Relationshipsof the cathartidvul- was done at Yale University between 1975 and 1986. tures. OccasionalPapers Univ. Michigan Mus. Support for the laboratory and field work was pro- Zool. No. 651: 1-26. vided by the William RobertsonCoe Fund of Yale NELSON,G. J. 1973. Classificationas an expression University and the U.S. National ScienceFoundation of phylogeneticrelationships. Syst. Zool. 22: 344- via grantsDEB-77-02594, 79-26746ß and BSR-83-15155 359. to C.G.S. Acknowledgmentsof the assistanceof in- OLSON,S. L. 1987. The relationships of the New dividualshave been made in someof the publications Guineanground-robins . Emu 87:247- cited below, and in other publicationspertaining to 248. this project. We thank G. A. Clark Jr., T. R. Howellß RAIKOW,R.J. 1987. Hindlimb myology and evolu- and R. B. Paynefor their constructivecomments. tion of the Old World suboscinepasserine birds (Acanthisittidae,Pittidae, Philepittidae, Eury- LITERATURE CITED laimidae).Ornithol. Monogr. No. 41. Washing- tonß D.C.ß Amer. Ornithol. Union. AHLQUIST,J. E., A. H. BLEDSOP,F. H. SHELDONß& C. SIBLEY,C. G., & J.E. AHLQUIST.1981. The phylogeny G. SIBLE¾.1987. DNA hybridization and avian and relationshipsof the ratite birds as indicated systematics.Auk 104: 556-563. by DNA-DNA hybridization.Pp. 301-335 in Evo- ß F. H. SHELDON, & C. G. SIBLEY. 1984. The lution TodayßProc. 2nd. Intl. Congr. Syst, and relationshipsof the Bornean Bristlehead(Pityri- Evol. Biol. (G. G. E. Scudder and J. L. Reveal, Eds.). asisgymnocephala) and the Black-collaredThrush PittsburghßHunt Inst. BotanicalDocumentationß (Chlamydochaerajefferyi).J. Ornithol. 125:129-140. Carnegie-Mellon Univ. BOCK,W. 1962. The pneumatic fossaof the humerus --, & --.. 1982a. The relationships of the in the Passeres. Auk 79: 425-443. Hawaiian honeycreepers(Drepaninini) as indi- BRUNING,D. 1985. Buttonquail.Pp. 76-77 in A dic- catedby DNA-DNA hybridization. Auk 99: 130- tionary of birds (B. Campbell and E. LackßEds.). 140. Calton, England,T. and A.D. Poyser,Ltd. --, & --. 1982b. The relationships of the CATZEFLIS,F. M., F. H. SHELDONßJ. E. AHLQUIST, & C. vireos (Vireoninae) as indicated by DNA-DNA G. SmLE¾.1987. DNA-DNA hybridization evi- hybridization.Wilson Bull. 94: 114-128. dence of the rapid rate of muroid DNA --, & ---. 1983. The phylogeny and classi- evolution. Mol. Biol. Evol. 4: 242-253. fication of birds based on the data of DNA-DNA COTTAM,P. 1957. The pelecaniformcharacters of the hybridization.In CurrentOrnithology 1:245-292 skeletonof the shoe-billstork, Balaeniceps rex. Zool. (R. F. JohnstonßEd.). New York and London, Ple- Bull. Brit. Mus. (Nat. Hist.) 5: 51-72. num Press. GARROD,A.H. 1873. On certainmuscles of the thigh ß & --. 1984a. The relationships of the of birds and on their value in classification,part starlings(Sturnidae: Sturnini) and the mocking- I. Proc. Zool. Soc. Londonß 1873 [1874]: 626-644. birds (Sturnidae: Mimini). Auk 101: 230-243. July1988] Classificationof Living Birds 423

ß & --. 1984b. The relationshipsof the ß & --.. 1987c. The relationships of four Papuan genusPeltops. Emu 84: 181-183. speciesof New Zealand passerinebirds. Emu 87: ß& --. 1985a.(publ. 1986). The phylog- 63-66. eny and classificationof the passerinebirdsß based ß & --. 1987d. DNA hybridization evi- on comparisonsof the geneticmaterialß DNA. Pp. dence of hominoid phylogeny: results from an 83-121 in Proc. 18th Intl. Ornithol. Congress, expanded data set. J. Mol. Evol. 26: 99-121. Moscow1982 (V. D. Ilyichevand V. M. Gavrilov, ß& --. In prep. Phylogeny and classifi- Eds.). Moscowß Nauka Publ. cationof the birds of the world: a study in mo- ß & 1985b. The phylogeny and clas- lecular evolution. New Havenß Yale Univ. Press. sificationof the Australo-Papuanpasserine birds. ß --, & F. H. Sheldon. 1987. DNA hy- Emu 85: 1-14. bridization and avian phylogenetics.Reply to ß & --.. 1985c. The phylogenyand clas- Cracraft.Pp. 97-125in Evolutionarybiology, vol. sificationof the New World suboscinepasserine 21 (M. K. Hecht, B. Wallaceß and G. T. Pranceß birds (Passeriformes:Oligomyodi: Tyranni). Pp. Eds.). New York and Londonß Plenum Press. 396-428 in Neotropicalornithology (P. A. Buck- ß R. $CHODDE, & J. E. AHLQUIST. 1984. The ley, M. S. FosterßE. S. Morton,R. S. Ridgely,and relationshipsof the Australo-Papuantreecreep- F. G. Buckley,Eds.). Washingtonß D.C. Amer. Or- ersClimacteridae as indicated by DNA-DNA hy- nithol. Union Monograph No. 36. bridization. Emu 84: 236-241. ß& . 1985d. The relationshipsof some --, G. R. WILLIAMSß& J. E. AHLQUIST. 1982. The groupsof African birds, basedon comparisons relationships of the New Zealand wrens (Acan- of the geneticmaterialß DNA. Pp. 115-161in Proc. thisittidae) as indicated by DNA-DNA hybrid- Syrup. on African Vertebrates (K.-L. Schuch- ization. Notornis 29: 113-130. mann, Ed.). BonnßMus. A. Koenig. $NEATH, P. H. A., & R. R. $OKAL. 1973. Numerical --, & 1986. Reconstructingbird phy- . San FranciscoßW. H. Freeman. logeny by comparingDNA's. Sci.Amer. 254: 82- VIeORS,N.A. 1825. Sketchesin ornithology;or, ob- 92. servationson the leading affinitiesof someof the --, & --.. 1987a. Avian phylogeny recon- more extensivegroups of birds. Arrangement of structedfrom comparisonsof the genetic mate- genera of birds. Zool. J. 2: 368-405. rial, DNA. Pp. 95-121 in Molecules and mor- WETMORE, A. 1960. A classification for the birds of phology in evolution: conflictor compromise(C. the world. Smithson. Misc. Coil. 139 (11): 1-37. PattersonßEd.). London, CambridgeUniv. Press. --, & 1987b. The LesserMelampitta is a of paradise. Emu 87: 66-68.

On 25 April 1988,the Daniel Giraud Elliot Medal wasawarded jointly to CharlesG. Sibley (SanFrancisco StateUniversity; Professor Emeritus, Yale University)and JonEdward Ahlquist (Departmentof Zoological and BiomedicalSciencesß Ohio University)."Their massive,insight-rich application of DNA hybridization techniquesto bird classificationrevolutionized taxonomy by showingat lasthow to distinguishavian evo- lutionary relationshipsfrom convergentsimilarities." Theprize was established in 1917through the Daniel Giraud Elliot Fund by a gift fromMargaret Henderson Elliot.It is awardedby the NationalAcademy of Sciencesfor meritoriouswork in zoologyor paleontology publishedover a three- to five-yearperiod.