Multiple Reversals of Bill Length over 1.7 Million Years in a Hawaiian Bird Lineage.

Evolutionary change has been documented over geological time, but reversals in morphology, from an ancestral state to a derived state and back again, tend to be rare. Multiple reversals along the same lineage are even rarer. We use the chronology of the Hawaiian Islands and an avian example, the Hawaiian honeycreeper 'amakihi (Hemignathus spp.) lineage, which originated on the oldest main island of Kaua'i 1.7 million years ago, to examine the process of sequential reversals in bill length. We document three single and two multiple reversals of bill length on six main islands from oldest to youngest, consistent with the phylogeny of the lineage. Longer bills occur on islands with endemic species, including phylogenetically relevant outgroups, that may compete with or dominate the 'amakihi. On islands without those species, the 'amakihi had shorter bills of similar length. Both types of reversals in morphology in this lineage integrate microevolutionary processes with macroevolution in the adaptive radiation of Hawaiian honeycreepers.

Vector movement underlies avian malaria at upper elevation in Hawaii: implications for transmission of human malaria.

With climate warming, malaria in humans and birds at upper elevations is an emerging infectious disease because development of the parasite in the mosquito vector and vector life history are both temperature dependent. An enhanced-mosquito-movement model from climate warming predicts increased transmission of malaria at upper elevation sites that are too cool for parasite development in the mosquito vector. We evaluate this model with avian malaria (Plasmodium relictum) at 1,900-m elevation on the Island of Hawaii, with air temperatures too low for sporogony in the vector (Culex quinquefasciatus). On a well-defined site over a 14-year period, 10 of 14 species of native and introduced birds became infected, several epizootics occurred, and the increase in prevalence was driven more by resident species than by mobile species that could have acquired their infections at lower elevations. Greater movement of infectious mosquitoes from lower elevations now permits avian malaria to spread at 1,900 m in Hawaii, in advance of climate warming at that elevation. The increase in malaria at upper elevations due to dispersal of infectious mosquitoes is a real alternative to temperature for the increased incidence of human malaria in tropical highlands.

Females lead population collapse of the endangered Hawaii creeper.

Population collapses result from drastic environmental changes, but the sexes may differ in vulnerability. Collapse of the endangered Hawaii creeper (Oreomystis mana) at Hakalau Forest National Wildlife Refuge resulted from food limitation associated with increased numbers of an introduced bird (Japanese white-eye, Zosterops japonicus), which competes with the creeper for food. Both creeper sexes had stunted bill growth and the greatest change in molt of native species in the community. With a surge in numbers of white-eyes, a recent cohort of adult females had very low survival after breeding, while adult males from the same cohort, and older females and males, continued to have high survival. Lower female survival resulted in a significantly more male-biased adult sex ratio. Recent low female survival was based on a great cost of reproduction, indicated by molt-breeding overlap that was previously avoided, and lower fat during the lengthy fledgling period. The difference in female survival between cohorts was associated with stunted bills from being reared in and then breeding in an increasingly poor food environment. Trend analysis of survey data indicate that the bird is declining throughout the refuge, with males being 72-80% of adults left six years after the white-eye increased. Competition over time was consistent with that previously documented over space on the Island of Hawaii. Adaptive management to recover the bird in this protected area needs to focus on improving both adult female survival and the adult sex ratio.

Changes in timing, duration, and symmetry of molt of Hawaiian forest birds.

Food limitation greatly affects bird breeding performance, but the effect of nutritive stress on molt has barely been investigated outside of laboratory settings. Here we show changes in molting patterns for an entire native Hawaiian bird community at 1650-1900 m elevation on the Island of Hawaii between 1989-1999 and 2000-2006, associated with severe food limitation throughout the year beginning in 2000. Young birds and adults of all species took longer to complete their molt, including months never or rarely used during the 1989-1999 decade. These included the cold winter months and even the early months of the following breeding season. In addition, more adults of most species initiated their molt one to two months earlier, during the breeding season. Suspended molt, indicated by birds temporarily not molting primary flight feathers during the months of peak primary molt, increased in prevalence. Food limitation reached the point where individuals of all species had asymmetric molt, with different primary flight feathers molted on each wing. These multiple changes in molt, unprecedented in birds, had survival consequences. Adult birds captured during January to March, 2000-2004, had lower survival in four of five species with little effect of extended molt. Extended molt may be adaptive for a nutrient stressed bird to survive warm temperatures but not cool winter temperatures that may obliterate the energy savings. The changing molt of Hawaiian birds has many implications for conservation and for understanding life history aspects of molt of tropical birds.

Negative effects of an introduced bird species on growth and survival in a native bird community.

Exploitative competition is a major determinant of community structure in natural assemblages [1, 2], but, introduced species are rarely competitors that lead to extinction of native species [3, 4]. Here we document strong community-wide competition from the Japanese white-eye (Zosterops japonicus) on native Hawaiian passerine birds. Introduced in 1929 [5], white-eye successfully invaded old-growth forest and coexisted with eight native species [6], overlapping multiple foraging substrates with each but evidencing no agonistic interactions [7]. The endangered Hawaii akepa (Loxops coccineus coccineus) was viable during 1987-1999 but became nonviable during 2000-2006 in association with an abrupt increase in white-eyes [7]. We show that after 2000, juveniles of every native bird species measured had lower mass and shorter bills and tarsi. For most species, lower mass led to decreased juvenile survival, and shorter bills to decreased survival of second-year and older adults. Lower survival of smaller birds represents normalizing selection that is restoring previous size means to future generations [8]. Birds at a nearby site with fewer white-eyes had normal size. White-eye had less stunting of bills and did not suffer the survival consequences of native species. Exploitative competition for food between native birds and an introduced species requires intensive management to prevent further declines.

Dispersal ghosts in Oceania.

Anthropological genetics helps expand our understanding of human phenotypes in the Pacific, in part because of its focus on gene genealogies to infer past episodes of dispersal and to differentiate these events from adaptations due to long-duration directional selection. Sewall Wright's 1949 seminal paper on population structure emphasized that there were two strong forces that exerted systematic and therefore determinant pressure on the gene pool: recurrent immigration and gene flow. These are important topics to all discussions of human dispersal in any region of the world. Furthermore, Wright listed five unique kinds of events that produced indeterminate or unpredictable changes that could lead to phenotypic and genotypic effects. In this category, he placed unique selective incidents, unique hybridization events, unique reductions in number, swamping by mass immigration, and mutational drive due to an allele always being favored since its origin or introduction. This discussion of human dispersal in the Pacific will touch on these topics, since they provide a second level of complexity in knowing who moved about a region of the world found already settled when rediscovered by colonial explorers during the 16-18th centuries.