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The Phylogeny of the Alcidae

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The Phylogeny of the Alcidae THE PHYLOGENY OF THE ALCIDAE J. G. STRAUCH,JR. UniversityMuseum (Zoology), Campus Box 315, Universityof Colorado, Boulder, Colorado 80309 USA ABSTRACT.--Anestimate of the phylogeny of 22 extant and 1 extinct speciesof the Alcidae was determinedfrom compatibilityanalyses of 33 cladisticcharacters of the skeleton,integ- ument, and natural history. The puffins were found to be a sister-group to all other alcids. Cerorhincawas found to be a puffin. The auklets were found to be a sister-groupto the remainingspecies. Brachyramphus was found to representa phyleticline separatefrom that including the other murrelets.Cepphus was found to be a member of the phyletic line in- cluding Endomychuraand Synthliboramphus.Alle was found to be a sister-groupof the auks. A compatibility analysisof musclecharacters of Hudson et al. (1969) yielded a phylogenetic tree in agreementwith that found usingmy data.The relationshipsamong Cepphus and the murrelets were found to need further study. A classificationbased on these results is sug- gested.It is recommendedthat the recent merging of genera by the A.O.U. (1982) be ac- cepted,that Cyclorrhynchusbe mergedwith Aethia,and that Pinguinusbe mergedwith Alca. Received25 January1984, accepted 3 December1984. THEAlcidae, a distinctgroup of marine,wing- evidencethat sandgrouseare charadriiforms. propelled diving birds, have been classified Although current evidence indicatesthat the during the past 150 years (literature reviewed compositionof the Charadriiformesis closeto by Sibley and Ahlquist 1972) with the loons that proposedby Wetmore (1930), future reso- (Gaviidae), grebes(Podicipedidae), diving pe- lution of the higher relationshipsof birds may trels (Pelecanoididae),and penguins(Sphenis- show this conclusionto be oversimplified. cidae). The modern consensusis that they are Given that the Alcidae are charadriiforms, members of the Charadriiformes (Wetmore there is still a questionof their affinitieswithin 1930, Mayr and Areadon 1951, Kitto and Wil- the order. Most authorshave suggestedthat al- son 1966,Storer 1971,Sibley and Ahlquist 1972, cids are most closely related to gulls (Storer Stegmann 1978, Strauch 1978, Cracraft 1981); 1960, Kozlova 1961). Ahlquist (1974) reported however, a few recent authors (Verheyen 1958, that the isoelectricfocusing in polyacrylamide Gysels and Rabaey 1964) have disputed this (IFPA) patterns of egg-white proteins of Uria opinion. The assumedmonophyly of the Cha- "shows an unmistakable likeness to those of radriiformes is based on their sharing a com- gulls." Evidencethat the ancestorof the alcids plex of character states (Zusi 1974, Strauch may have been more like a shorebirdhas been 1976), but it never has been tested by a phy- reportedby Stettenheim(1959), Hudson et al. logenetic analysis of the orders of birds. Fur- (1969), and Stegmann (1978). However, be- thermore, the limits of the order still are un- cause there is considerable evidence that Dro- resolved.Storer (1956) and Sibley and Ahlquist rnasis closely related to the Lari (Strauch 1978, (1972) presented evidence that the loons also Sibley and Ahlquist in press),a shorebird-like may be charadriiforms(but see Cracraft 1982). common ancestorof alcids and larids may not Olson and Feduccia (1980) assertedthat the fla- conflict with earlier ideas. Strauch (1978) and mingos (Phoenicopteridae)are closely related others(Stegmann 1978, Cracraft 1981)were un- to Charadriiformes, and Olson and Steadman able to identify the closestrelatives of the al- (1981) presented evidence that Pedionomusis a cids. On the basis of DNA-DNA hybridization charadriiform. Maclean (1967) and Fjeldsfi studies, Sibley and Ahlquist (in press) found (1976) argued that sandgrouse(Pteroclididae) that the Alcidae and Lari are sister-groups. are charadriiforms, but their evidence and con- The Alcidae are distinguished among cha- clusionshave been challengedby Olson (1970) radriiform birds by their compactform, short and Strauch (1979) and are not supported by wings, and feeding habits (Coues1868). Some the findings of Kitto and Wilson (1966). Sibley of the characteristics(sternum long and narrow and Ahlquist (in press), however, have new with long, rounded metasternum;wing bones 520 The Auk 102: 520-539. July 1985 July1985] Phylogenyofthe Alcidae 521 flattened)used to define them (Verheyen 1958, and Aethiinae (the remaining genera). Yudin Zusi 1974) may be related to their marine and (1965) thought that there probably were two diving habits; others (large supraorbital phyletic lines in the Alcidae but found insuf- grooves, basipterygoid processesabsent in ficient grounds on which to divide them. He adults,anterior toes fully webbed,hind toe ab- suspectedthat similarities in the jaw muscula- sent) are found in other charadriiform birds. ture of puffins and auklets were due to "par- Strauch (1978) found the Alcidae to be a mono- allel development."In a study of the wing and phyletic group defined by twisting of the leg musculature,Hudson et al. (1969) found that brachial tuberosity of the coracoid so that it the puffins differed markedly from other al- doesnot roof the triossealcanal and by lack of cids, were probably the most primitive living a lateral synsacralstrut. alcids,and were closestto Ptychoramphus,among Coues (1868) used nostril feathering, bill the genera studied. They found that Brachy- form, and presenceor absenceof creststo di- ramphuswas not particularlyclose to any other vide the Alcidae into three subfamilies: Alci- genus,but was closer to Uria and Alca than to nae (Pinguinus,Alca), Phaleridinae (Fratercula, Cepphus,other murrelets,auklets, or puffins;and Lunda, Cerorhinca,Ptychoramphus, Cyclorrhyn- they found Cepphusto be closestto Uria and chus,Aethia), and Urinae (Synthliboramphus,En- Synthliboramphus.Several authors (Storer 1945b, domychura,Brachyramphus, Cepphus, Alle, Uria). Sealy 1972, Binford et al. 1975, Jehl and Bond Beddard (1898) used the number of rectrices, 1975) have concluded that among the murre- relative size of the right and left liver lobes, lets Endomychuraand Synthliboramphusare quite leg muscleformula, and form of the syrinx to similar and that both are distinct from Brachy- divide the alcids into two families: Fraterculi- ramphus. dae (Cerorhinca,Lunda, Fratercula)and Uriidae It haslong been agreedthat the puffins (Cero- (all other genera). Shufeldt (1901), summariz- rhinca, Lunda, Fratercula),the auklets (Ptycho- ing a seriesof papers on the osteologyof the ramphus,Cyclorrhynchus, Aethia), and the auks alcids (1888, 1889a-d), decided that Beddard's (Uria, Alca,Pinguinus) are clustersof closelyre- two families representedtwo subfamilies.Shu- lated species. The relationships among the feldt (1889d) thought Alle closestto the auklets murrelets(Brachyramphus, Endomychura, Synthli- and Uria closestto the Laridae. Dawson (1920) boramphus),the relationshipsof Cepphusand Alle usedegg characteristicssupplemented by other to other alcids,and the relationshipsamong all characters to divide the alcids into five fami- of thesegroups have been debated,however. lies: Aethiidae (Alle, Ptychoramphus,Cyclorrhyn- I reexamined the relationships among the chus,Aethia), Cepphidae (Cepphus), Alcidae (Uria, speciesof the Alcidae using a modern, objec- Alca, Pinguinus),Fraterculidae (Cerorhinca, Lun- tive method to evaluate the characters on which da, Fratercula),and Synthliboramphidae (Syn- the estimateof phylogeny is based. thliboramphus,Brachyramphus, Endomychura). Storer (1945a) usedpelvis and leg morphology MATERIALS AND METHODS supplementedby plumage,soft-part, egg, and I examined skins and skeletons of each living breeding characters to divide the alcids into speciesof alcid. A completecomposite skeleton and sevengroups, which he later (1960) designated severalunassociated bones of Pinguinusalso were ex- as tribes:Alcini (Uria, Alca,Pinguinus), Cepphi- amined. Integument charactersfor Pinguinuswere ob- ni (Cepphus),Brachyramphini (Brachyramphus), tained from Ridgway(1919). Natural historydata were Plautini (Alle), Aethiini (Ptychoramphus,Cyclor- taken from the literature (Storer 1945a, Kozlova 1961, rhynchus,Aethia), Synthliboramphini (Endo- Sealy 1972, Simons 1980, Terres 1980). Natural his- mychura,Synthliboramphus), and Fraterculini tory data for Pinguinuswere obtainedfrom Bengtson (Cerorhinca, Lunda, Fratercula). Storer (1952) (1984). Character names follow Howard (1929), Bock suggestedthat Alle might be closestto his A1- and McEvey (1969), Zusi and Jehl (1970), and Strauch cini and that Cepphuswas closerto the ancestral (1978). A set of 33 cladistic characters was devised for the stockof the family than was Uria. Earlier, Stor- 22 extant and ! extinct speciesstudied. Primitive states er (1945b) thought Brachyramphusthe most were determined using other charadriiform birds, primitive genusof alcid. Verheyen (1958) clas- particularly the Lari, as outgroups(Strauch 1978). sifted the Alcidae into four subfamilies: Frater- Charactercompatibility analysis employing the pro- culinae (Cerorhinca,Lunda, Fratercula), Alcinae gram CLINCH 5 (written by K. L. Fiala) was used to (Pinguinus,Alca, Uria, Cepphus),Plautinae (Alle), find the largestset of mutually compatiblecharacters 522 J.G. STRAUCH,JR. [Auk, Vol. 102 T^I•I•E 1. Character-state trees for the Alcidae. Character-state No. Character State tree 1 Maxillopalatine strut P B • A Absent B Present A 2 Maxillopalatineshape A • B Hollow and cup-shaped A Broad, flat plate B 3 Ventral end of interpalatineprocess A • B Doesnot extendbeyond ventral end of palatine shelf A Extendsbeyond
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