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The Relationships of the Starlings (Sturnidae: Sturnini) and the Mockingbirds (Sturnidae: Mimini)

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The Relationships of the Starlings (Sturnidae: Sturnini) and the Mockingbirds (Sturnidae: Mimini) THE RELATIONSHIPS OF THE STARLINGS (STURNIDAE: STURNINI) AND THE MOCKINGBIRDS (STURNIDAE: MIMINI) CHARLESG. SIBLEYAND JON E. AHLQUIST Departmentof Biologyand PeabodyMuseum of Natural History,Yale University, New Haven, Connecticut 06511 USA ABSTRACT.--OldWorld starlingshave been thought to be related to crowsand their allies, to weaverbirds, or to New World troupials. New World mockingbirdsand thrashershave usually been placed near the thrushesand/or wrens. DNA-DNA hybridization data indi- cated that starlingsand mockingbirdsare more closelyrelated to each other than either is to any other living taxon. Some avian systematistsdoubted this conclusion.Therefore, a more extensiveDNA hybridizationstudy was conducted,and a successfulsearch was made for other evidence of the relationshipbetween starlingsand mockingbirds.The resultssup- port our original conclusionthat the two groupsdiverged from a commonancestor in the late Oligoceneor early Miocene, about 23-28 million yearsago, and that their relationship may be expressedin our passerineclassification, based on DNA comparisons,by placing them as sistertribes in the Family Sturnidae,Superfamily Turdoidea, Parvorder Muscicapae, Suborder Passeres.Their next nearest relatives are the members of the Turdidae, including the typical thrushes,erithacine chats,and muscicapineflycatchers. Received 15 March 1983, acceptedI November1983. STARLINGS are confined to the Old World, dine thrushesinclude Turdus,Catharus, Hylocich- mockingbirdsand thrashersto the New World. la, Zootheraand Myadestes.d) Cinclusis closerto Starlingsusually have been thought to be re- the thrushes,flycatchers, starlings and mocking- lated to crows and other members of the cot- birds than to the wrens." void assemblage,or to New World troupials A Distance Wagner tree (Farris 1972) based (Icterini) and/or Old World weaverbirds (Plo- on the DNA hybridization data showed Mimus ceinae).Mockingbirds and thrashershave been and Sturnusto be more closely related to each viewed as relatives of thrushes (Turdidae) and/ other than to any other taxon, and they clus- or wrens (Troglodytidae). tered with the thrushes,muscicapine flycatch- Evidenceof a relationship between starlings ers, erithacine chats, and dippers. and mockingbirds had been reported at least The same pattern emerged from a more ex- twice (Beecher 1953, Stallcup 1961) before we tensivestudy of the passetines(Sibley and Ahl- published the results of a comparisonof their quist in pressa) in which the mimine-sturnine DNAs (Sibley and Ahlquist 1980). The 1980 relationship was based on the same data as in study was based on 153 DNA-DNA hybrids the 1980 paper, but many additional data per- among 18 oscinegenera used as radio-labeled taining to other passefinegroups were includ- single-copy"tracers." Among our conclusions ed. In the classificationderived from this study, was the following statement.(DNAs of the cit- we treated the tribes Sturnini and Mimini as ed genera were used in the 1980 study). sister groups in the family Sturnidae. The fol- "The thrushes,muscicapine flycatchers, mock- lowing is a partial outline of this classification ingbirds, starlings,and dippers are membersof to indicate the relative positions and the names a monophyleticassemblage. a) Within this group we have assignedto the taxa mentioned in the the muscicapineflycatchers (Muscicapa, Melae- present paper. Details for most groups have nornis,Niltava, Rhinomyias)are closely related to been omitted. the chat-like thrushes (Erithacus,Erythropygia, Order Passeriformes Phoenicurus,Luscinia, Cossypha, Pogonocichla, Myr- mecocichla,Copsychus). b) The starlings (Sturnus, Suborder Oligomyodi, the suboscines Onychognathus,Spreo, Ampeliceps, Aplonis) are Suborder Passeres, the oscines closestto the mockingbirdsand thrashers(Mi- Parvorder Corvi mus,Dumetella, Toxostoma, Oreoscoptes. c) The tur- SuperfamilyCorvoidea 23O The Auk 101:230-243. April 1984 April 1984] Starling-MockingbirdRelationships 231 Family Corvidae tallica builds a pendant, globular nest with a Subfamily Corvinae, crows, jays, side entrance,and Acridotheresginginianus digs magpies, birds-of-paradise, a nest hole in an earthen bank. The color of Australian magpies, etc. starling eggsvaries from white to green or blue- Parvorder Muscicapae green; some are unmarked, and others have Superfamily Turdoidea reddish or brownish markings. Family Bombycillidae,waxwings Davisand Miller (1960)recognized 31 species Family Cinclidae, dippers in 13 genera in the "family Mimidae," includ- Family Turdidae ing Donacobiusatricapillus, which Miller (1964) Subfamily Turdinae, typical noted as having the voice and habits of a wren thrushes (Troglodytidae).Clench et al. (1982) concluded Subfamily Muscicapinae that Donacobiusactually is a wren, which is con- Tribe Muscicapini, muscicapine sistent with several DNA comparisons.In Ta- flycatchers bles ! and 3, Donacobiusclusters with the Syl- Tribe Erithacini, chats vioidea, including the wrens, not with the Family Sturnidae mimines in the Turdoidea. Tribe Sturnini, starlings Most mockingbirdsand thrashersoccur from Tribe Mimini, mockingbirds southern Canada and the United States to the and thrashers West Indies and Central America. With the ex- Superfamily Sylvioidea, Old World clusion of Donacobius,only Mirnus occurs in warblers, wrens, titmice, nut- South America. hatches, swallows, etc. All mimines build open, cup-shaped nests, Superfamily Fringilloidea, larks, sun- usually placed in bushesor trees. The Pearly- birds, weavers, fringillines, trou- eyed Thrasher (Margaropsfuscatus) places its pials, etc. "bulky, cup-shaped nest . .. in a bush or tree, cavity of a tree, or on the side of cave or cliff" Although this classificationcontains depar- and the Trembler (Cinclocerthiaruficauda) builds tures from previous arrangements,most of the its nest "in a cavity of a tree or tree fern, or at changeswere received with approval or mild the baseof a palm frond" (Bond 1971:169). Egg doubts,but the inclusion of the starlingsand colorsare blue or green with various amounts mockingbirdsin the same family elicited vig- of spotting or streaking. Thus, the eggs and orousdissent from severalcolleagues. We have nests,although variable, are somewhatsimilar therefore made a new study, using different and do not opposea relationship between the taxafor the "tracers,"several additional species, two groups. and many morecomparisons. Our original con- clusions have been confirmed by these new data, and we have alsofound severalcongruent TAXONOMIC HISTORY morphological characters.We have been un- Opinionsabout the relationshipsof starlings able to find hard evidencethat precludesa close have ranged widely, but most of the classifi- relationship between starlings and mocking- birds. cationsof the past130 yr have placedstarlings closeto crows and their allies, including birds- In this paper we review the taxonomic his- of-paradise,Old World orioles,and drongos.A tories of the two groups, note evidence from relationship to the corvoid groups was advo- other sources,and present the results of the new DNA hybridization study. catedby Sharpe(1890, 1891),Reichenow (1914), Stresemann (1934), Stonor (1938), Delacour and Vaurie (1957), and Bock (1963). A relationship DIVERSITY AND DISTRIBUTION to the corvoids and the New World troupials (Icterini) was indicated in the classificationsof Amadon (1962) recognized 111 speciesof Bonaparte(1850), Gray (1870),Sundevall (1872), starlings in 26 genera, approximately evenly Sclater (1880), the early A.O.U. Check-lists divided between Africa and Asia. Three species (1886, 1895, 1910), Shufeldt (1889), and Coues of Sturnusoccur in Europe,and Aplonisreaches (1896). Sharpe (1890: 1) declared that the star- Australia and many southwest Pacific islands. lings are "undoubtedly allied" to the Corvidae Most starlingsnest in cavities,but Aplonisme- and repeated this emphatic opinion in his ex- 232 SlBLEYAND AHLQUIST [Auk, Vol. 101 tensive review of avian classification in 1891 starlings and corvids seemsto have been based (p. 85). The corvoidsand ploceineswere viewed on the similar shape of the bill in starlings, as starling relatives by Reichenow (1882), von some corvines, and Old World orioles and the Boetticher(1931), Amadon (1943, 1956), Mayr fact that some starlings and some corvines are and Amadon (1951),Mayr and Greenway(1956, black. It appears that the most influential 1962), Storer (1971), and Voous (1977). "character" has been tradition, with most au- Wallace (1874) associated starlings with thors following the opinions of previous au- weaverbirds, woodswallows, and larks as thors. "Sturnoid Passeres."Parker (1875) reported The taxonomic history of the mockingbird- similarities between the skulls of the Celebean thrashergroup hasbeen relatively simple. Most Myna (Enodes)and that of a Song Thrush (Tur- authors have placed them with or near the dusphilomelos) but decided that the myna was thrushes; some have allied them to the wrens. more shrike-like and possibly related to the They were considered to be thrushes by Bo- birds-of-paradiseand the drongos(Dicrurus). naparte(1850), Gray (1869),Coues (1896), Ridg- Stejneger (1885) placed starlings with cor- way (1907), Beecher (1953, 1978), Morioka vids and meliphagids, which may have influ- (1967), and Gulledge (1975). Of these authors, enced Wetmore (1930, 1951, 1960), who listed only Beecher (1953) compared mockingbirds starlings near meliphagids, although far from and thrusheswith starlings, and he found sev- Corvidae. Wetmore (1960) also placed New
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