Beth Slikas

PHYLOGENY OF TITMICE (PARIDAE): II. SPECIES RELATIONSHIPS BASED ON SEQUENCES OF THE MITOCHONDRIAL CYTOCHROME-B GENE

Abstract We present a phylogenetic hypothesis for 40 species in the bird family Paridae, based on comparisons of nucleotide sequences of the mitochondrial cytochrome-b gene. Parids, including tits and chickadees, are an older group than their morphological stereotypy suggests. The longest cytochrome-b distances between species reach 12% in uncorrected divergence. With the exception of one thrasher-like terrestrial tit species of the Tibetan plateau (Pseudopodoces humilis), morphological and ecological stasis have prevailed since the initial parid radiation in the Old World during the mid-Tertiary. All trees support monophyly of the family Paridae, which includes Parus (sensu lato) and the monotypic Oriental genera Sylviparus, Melanochlora, and Pseudopodoces. Within the clade of chickadees and gray tits (Parus, subgenus Poecile), three Old World species, Parus lugubris of the eastern Mediterranean and Balkan regions, P. superciliosus of high elevations in the Himalayas of western China, and P. varius of the Orient are sisters to all other species. The Eurasian crested titmice (subgenus Lophophanes) and North American crested titmice (subgenus Baeolophus) are sister groups. Our data suggest two colonizations of the New World by parids in the late Tertiary. The ancestor of modern Baeolophus colonized North America 4 mya, and the ancestor of all North American chickadees colonized North America 3.5 mya. Phylogénie chez la mésange (Paridés): II. Relations entre les espèces basées sur des séquences du gène mitochondrial cytochrome-b

DNA-DNA HYBRIDIZATION EVIDENCE OF PHYLOGENETIC-RELATIONSHIPS AMONG MAJOR LINEAGES OF PARUS

DNA-DNA hybrids were formed among 2 outgroups and 12 taxa of titmice and chickadees to estimate the genealogical relationships of the main Parus lineages. A complete matrix of reciprocal comparisons among seven parids and the Verdin (Auriparus flaviceps) indicated that the Blue Tit (P. caeruleus) and Great Tit (P. major) together form the sister group of the rest of the genus, and that the Bridled Titmouse (P. wollweberi) is more closely related to North American titmice than to the Old World crested tits. The DNA-hybridization data complement information from allozyme and mtDNA studies of closely related parids and provide historical insight into patterns of tit behavior.

AN ENDEMIC RADIATION OF MALAGASY SONGBIRDS IS REVEALED BY MITOCHONDRIAL DNA SEQUENCE DATA

Abstract The bird fauna of Madagascar includes a high proportion of endemic species, particularly among passerine birds (Aves: Passeriformes). The endemic genera of Malagasy songbirds are not allied obviously with any African or Asiatic taxa, and their affinities have been debated since the birds first were described. We used mitochondrial sequence data to estimate the relationships of 13 species of endemic Malagasy songbirds, 17 additional songbird species, and one species of suboscine passerine. In our optimal trees, nine of the 13 Malagasy species form a clade, although these birds currently are classified in three different families. In all optimal trees, the sister to this endemic clade is a group of Old World warblers including both African and Malagasy birds. The endemic Malagasy songbird clade rivals other island radiations, including the vangas of Madagascar and the finches of the Galapagos, in ecological diversity. Corresponding Editor: S. Edwards

PHYLOGENETIC RELATIONSHIPS OF MICRONESIAN WHITE-EYES BASED ON MITOCHONDRIAL SEQUENCE DATA

Abstract Using mitochondrial sequence data, we estimated phylogenetic relationships and genetic divergence among selected species of white-eyes (Zosteropidae). We focused on taxa endemic to islands in Micronesia, specifically Zosterops conspicillatus, Z. semperi, Z. hypolais, Rukia oleaginea, and Cleptornis marchei. We also included in our data set five additional species of Zosterops from the Indo-Australian region and three species from Africa, as well as additional passerine outgroups. Our mitochondrial sequence data revealed substantial genetic divergence (5.7 to 7.3%) among Z. conspicillatus, Z. semperi, and Z. hypolais, three taxa that formerly were treated as a single species. In addition, a sequence divergence of 6.5 ± SE of 1.7% was found between the population of Z. conspicillatus from Rota and “conspecific” populations on Guam, Tinian, and Saipan. The distinctiveness of the Rota population suggests that this taxon should be recognized as a distinct species, a result that bears on the conservation of this population because it has been declining dramatically in recent years. All optimal trees based on analysis of the mitochondrial sequence data place Rukia oleaginea within the genus Zosterops. In all optimal trees, Cleptornis marchei positions as the sister taxon to a clade including all other zosteropids included in this study. The trees based on our data strongly contradict the traditional classification of Cleptornis as a honeyeater (family Meliphagidae). Our data cannot resolve with any confidence the sister relationships of the insular endemic white-eyes, although the optimal trees suggest multiple colonizations of Micronesia by more than one white-eye lineage.

Phylogeny of titmice (Paridae): I. Estimate of relationships among subgenera based on DNA-DNA hybridization

The phylogenetic relationships of titmice and chickadees (Passeriformes: Paridae) were estimated by DNA-DNA hybridization. Fifteen species of parids (representing all six subgenera) and two outgroups, Penduline Tit Remiz pendulinus and Whitebreasted Nuthatch Sitta carolinensis, were radiolabeled and compared pairwise to produce a complete reciprocal set of distance measures. The resulting phylogeny supported the traditional allocation of species to subgenera, except that Parus wollweberi clustered with the North American crested tits (subgenus Baeolophus) rather than the Old World crested tits (Lophophanes). The interrelationships of most subgenera remained unresolved. Nevertheless, a major subdivision in the family was strongly indicated: i.e., the great tits (subgenus Parus) and the blue tits (subgenus Cyanistes) formed a clade that is the sister taxon of the rest of the family. This bifurcation has important ecological, physiological, and morphological ramifications, as it correlates exactly with seed-caching and flocking habits, hippocampal development, and leg-muscle character distributions. These DNA-hybridization results corroborate or complement the phylogenetic relationships among parids supported by other molecular data sets (viz., allozyme, restriction site, and preliminary DNA hybridization analyses).

RECOGNIZING AND TESTING HOMOLOGY OF COURTSHIP DISPLAYS IN STORKS (AVES: CICONIIFORMES: CICONIIDAE)

Ethological studies in the 1940s and 1950s, most notably those of Lorenz and Tinbergen, emphasized a historical perspective. By the 1970s, the notion that behavioral traits are too plastic to retain historical information became prevalent, and evolutionary approaches in behavioral studies were largely abandoned. However, several recent studies have demonstrated that behavioral characters are remarkably consistent with phylogenies obtained from other data and not particularly prone to homoplasy. In this study, I coded descriptions of courtship display behaviors in stork species (Aves: Ciconiiformes: Ciconiidae) as a matrix of discrete characters. I mapped each behavioral character onto a phylogeny based on DNA‐DNA hybridization distances to test the homology of individual characters. Generally, displays occurring early in courtship were congruent with phylogenetic relationships and showed little homoplasy, while displays occurring late in courtship were more homoplastic. I also performed a phylogenetic analysis of the behavioral data matrix using maximum parsimony. The strict consensus of the 24 most‐parsimonious trees was congruent with the DNA‐DNA hybridization tree in all nodes having greater than 70% bootstrap support.

GENETIC VARIABILITY AND TAXONOMIC STATUS OF THE NIHOA AND LAYSAN MILLERBIRDS

Abstract The Millerbird (Acrocephalus familiaris) is an endemic Northwestern Hawaiian Islands reed warbler that existed until about 1923 on Laysan Island (A. f. familiaris) and currently occurs in a small population on Nihoa Island (A. f. kingi). The two populations are described as separate subspecies or species on the basis of size and plumage differences. We assessed genetic variation in blood samples from 15 individuals in the modern Nihoa population using approximately 3000 base pairs (bp) of mitochondrial DNA (mtDNA) sequence and 14 microsatellite loci. We also obtained up to 1028 bp of mtDNA sequence from the fragmented DNA of museum specimens of three birds collected on Nihoa in 1923 and five birds collected on Laysan in 1902 and 1911 (ancient samples). Genetic variation in both marker types was extremely low in the modern Nihoa population (nucleotide diversity [π]  =  0.00005 for mtDNA sequences; observed heterozygosity was 7.2% for the microsatellite loci). In contrast, we found three mtDNA haplotypes among the five Laysan individuals (π  =  0.0023), indicating substantially greater genetic variation. The Nihoa and Laysan taxa differed by 1.7% uncorrected mtDNA sequence divergence, a magnitude that would support designation at the subspecies, and perhaps species, level relative to other closely related Acrocephalus species pairs. However, in light of strong ecological similarity between the two taxa, and a need to have additional populations to prevent extinction from stochastic effects and catastrophes, we believe these genetic differences should not deter a potential translocation of individuals from Nihoa to Laysan.

HAWAIIAN BIRDS: LESSONS FROM A REDISCOVERED AVIFAUNA

IN THE INTRODUCTION to Hawaiian Birdlife, Andrew Berger (1972) wrote with frustration and dismay about his futile search for several species of endemic Hawaiian birds that had been described by collectors in the 1890s, but apparently were extinct by the mid-20th century. He bemoaned the fact that the ecology and behavior of those birds could never be studied. In fact, the loss of diversity and the gaps in our knowledge of the ecology and evolution of Hawaiis birds both are far greater than Berger could have imagined. During the past three decades, paleontological efforts spearheaded by Storrs Olson and Helen James have unearthed an unsuspected number and diversity of extinct birds. Bones collected from sand dunes, lava tubes, sinkholes, and former lakebeds have approximately doubled the number of endemic Hawaiian species. Among the fossil taxa collected and described by Olson and James are at least 15 new species in the spectacular honeycreeper radiation (tribe Drepanidini); several species of raptors unknown in the historical avifauna, including an eagle, a harrier, and a radiation of bird-eating owls; and many flightless birds, including rails, ibises, ducks, and geese (Olson and James 1982a, b, 1991; James and Olson 1991). Olson and James have demonstrated that the historically recorded Hawaiian avifauna, although spectacular, is a meager and biased sample of the fauna that existed prehistorically (i.e. before 1778, the year of the arrival of James Cook in the Hawaiian Islands). Radiocarbon dating of bones of extinct birds and presence of extinct birds in some archaeological sites have shown that extinctions on Hawaii occurred after humans colonized the archipelago, -1,600 years ago (Olson and James 1982a, b; James et al. 1987, Burney et al. 2001). That pattern of massive, rapid extinction of endemic birds following human colonization has been found on oceanic islands throughout the

DNA Sequence assessment of phylogenetic relationships among New World martins (Hirundinidae: Progne)

Abstract The classification of New World martins (Progne) has a convoluted history because taxonomists have relied on plumage traits that vary continuously across populations. We estimated the phylogeny of Progne by analyzing mitochondrial cytochrome b DNA sequences of 27 individuals of eight of the nine species (10 subspecies) and nuclear β-fibrinogen intron 7 sequences of 20 individuals of six species (8 subspecies). The Brown-chested Martin (P. tapera) is sister to other Progne species. The Middle American taxa—Sinaloa Martin (P. sinaloae), Cuban Martin (P. cryptoleuca), Caribbean Martin (P. dominicensis), and Central American populations of Gray-breasted Martin (P. chalybea)—form a well supported clade. This group is distinct from Purple Martin (P. subis), which has no particularly close relatives. All four Middle American taxa appear to be good species, although Cuban and Caribbean martins could be merged in view of their similar plumage and low genetic divergence (1.2%). Two of the South American taxa, the Peruvian Martin (P. murphyi) and Southern Martin (P. elegans), are also distinct species. We did not examine the Galapagos Martin (P. modesta) for lack of DNA, but it is likely to be a good species as well. An unexpected result of the study was that Gray-breasted Martin appears polyphyletic; its South American populations are closer to the Southern Martin than to its Central American populations.