Phylogenetic relationships within the Alcidae (Charadriiformes: Aves) inferred from total molecular evidence

The Alcidae is a unique assemblage of Northern Hemisphere seabirds thatforage by "flying" underwater. Despite obvious affinities among thespecies, their evolutionary relationships are unclear. We analyzednucleotide sequences of 1,045 base pairs of the mitochondrial cytochrome bgene and allelic profiles for 37 allozyme loci in all 22 extant species.Trees were constructed on independent and combined data sets using maximumparsimony and distance methods that correct for superimposed changes.Alternative methods of analysis produced only minor differences inrelationships that were supported strongly by bootstrapping or standarderror tests. Combining sequence and allozyme data into a single analysisprovided the greatest number of relationships receiving strong support.Addition of published morphological and ecological data did not improvesupport for any additional relationship. All analyses grouped species intosix distinct lineages: (1) the dovekie (Alle alle) and auks, (2)guillemots, (3) brachyramphine murrelets, (4) synthliboramphine murrelets,(5) true auklets, and (6) the rhinoceros auklet (Cerorhinca monocerata) andpuffins. The two murres (genus Uria) were sister taxa, and the blackguillemot (Cepphus grylle) was basal to the other guillemots. The Asiansubspecies of the marbled murrelet (Brachyramphus marmoratus perdix) wasthe most divergent brachyramphine murrelet, and two distinct lineagesoccurred within the synthliboramphine murrelets. Cassin's auklet(Ptychoramphus aleuticus) and the rhinoceros auklet were basal to the otherauklets and puffins, respectively, and the Atlantic (Fratercula arctica)and horned (Fratercula corniculata) puffins were sister taxa. Severalrelationships among tribes, among the dovekie and auks, and among theauklets could not be resolved but resembled "star" phylogenies indicativeof adaptive radiations at different depths within the trees.