Imperfect replacement of native species by non-native species as pollinators of endemic Hawaiian plants.

Native plant species that have lost their mutualist partners may require non-native pollinators or seed dispersers to maintain reproduction. When natives are highly specialized, however, it appears doubtful that introduced generalists will partner effectively with them. We used visitation observations and pollination treatments (experimental manipulations of pollen transfer) to examine relationships between the introduced, generalist Japanese White-eye (Zosterops japonicus) and 3 endemic Hawaiian plant species (Clermontia parviflora, C. montis-loa, and C. hawaiiensis). These plants are characterized by curved, tubular flowers, apparently adapted for pollination by curve-billed Hawaiian honeycreepers. Z. japonicus were responsible for over 80% of visits to flowers of the small-flowered C. parviflora and the midsize-flowered C. montis-loa. Z. japonicus-visited flowers set significantly more seed than did bagged flowers. Z. japonicus also demonstrated the potential to act as an occasional Clermontia seed disperser, although ground-based frugivory by non-native mammals likely dominates seed dispersal. The large-flowered C. hawaiiensis received no visitation by any birds during observations. Unmanipulated and bagged C. hawaiiensis flowers set similar numbers of seeds. Direct examination of Z. japonicus and Clermontia morphologies suggests a mismatch between Z. japonicus bill morphology and C. hawaiiensis flower morphology. In combination, our results suggest that Z. japonicus has established an effective pollination relationship with C. parviflora and C. montis-loa and that the large flowers of C. hawaiiensis preclude effective visitation by Z. japonicus.

Climate-associated population declines reverse recovery and threaten future of an iconic high-elevation plant.

Although climate change is predicted to place mountain-top and other narrowly endemic species at severe risk of extinction, the ecological processes involved in such extinctions are still poorly resolved. In addition, much of this biodiversity loss will likely go unobserved, and therefore largely unappreciated. The Haleakala silversword is restricted to a single volcano summit in Hawai'i, but is a highly charismatic giant rosette plant that is viewed by 1-2 million visitors annually. We link detailed local climate data to a lengthy demographic record, and combine both with a population-wide assessment of recent plant mortality and recruitment, to show that after decades of strong recovery following successful management, this iconic species has entered a period of substantial climate-associated decline. Mortality has been highest at the lower end of the distributional range, where most silverswords occur, and the strong association of annual population growth rates with patterns of precipitation suggests an increasing frequency of lethal water stress. Local climate data confirm trends toward warmer and drier conditions on the mountain, and signify a bleak outlook for silverswords if these trends continue. The silversword example foreshadows trouble for diversity in other biological hotspots, and illustrates how even well-protected and relatively abundant species may succumb to climate-induced stresses.

Patterns of nucleotide variation in homoeologous regulatory genes in the allotetraploid Hawaiian silversword alliance (Asteraceae).

Genome-wide duplication (polyploidization) is prevalent in a large number of eukaryotic organisms and is particularly widespread in flowering plants. Polyploid species appear to vary from their diploid progenitors in a variety of ecologically important traits, suggesting that genome duplications provide a mechanism for ecological diversification. Studies of nucleotide variation at duplicate genes that arise via polyploidization allow us to infer the evolutionary forces that act on these polyploid loci. In an effort to examine the evolutionary dynamics of homoeologous loci, molecular population genetic analyses were undertaken for duplicate regulatory genes in the allopolyploid Hawaiian silversword alliance, a premier example of adaptive radiation. The levels and patterns of nucleotide variation for the floral homeotic genes ASAPETALA1 (ASAP1) and ASAPETALA3/TM6 (ASAP3/TM6) were studied in two species representing different lineages within the Hawaiian silversword alliance: Argyroxiphium sandwicense ssp. macrocephalum and Dubautia ciliolata ssp. glutinosa. Homoeologueous copies of ASAP1 and ASAP3/TM6 show differing levels and patterns of nucleotide polymorphism. Duplicate ASAP1 copies have similar levels of nucleotide diversity and haplotype structure in both species; by contrast, duplicate ASAP3/TM6 genes display different levels and patterns of variation in D. ciliolata ssp. glutinosa. Additionally, D. ciliolata ssp. glutinosa appears to be segregating for a moderate frequency null allele in one ASAP3/TM6 homoeologue. These results suggest that differing evolutionary forces can affect duplicate loci arising from allopolyploidization.

Tissue elastic properties of eight Hawaiian Dubautia species that differ in habitat and diploid chromosome number.

Eight Hawaiian Dubautia species grow in habitats as varied as exposed lava, dry scrub, mesic forest, wet forest, and bog. These species also differ in diploid chromosome number, with four species having 13 pairs of chromosomes and four species having 14 pairs. This ecological and chromosomal variation is paralleled by significant interspecific variation in tissue elastic properties. The four 13-paired species from dry habitats exhibit significantly lower tissue elastic moduli near full hydration (E i) than the four 14-paired species from mesic to wet habitats. Values of E i range from 2 to 4 MPa among the former species and from 9 to 18 MPa among the latter species. The turgor dependence of the elastic modulus also differs markedly between the two groups of species. As a result of these differences in tissue elastic properties, the capacity for maintaining high turgor pressures as tissue water content decreases is much greater in the 13-paired species from dry habitats than in the 14-paired species from mesic to wet habitats. These results indicate that the evolutionary diversification of the Dubautia species has been accompanied by a significant degree of change at the physiological level.

Variation in the tissue water relations of two sympatric Hawaiian Dubautia species and their natural hybrid.

At one site of sympatry on the Island of Hawaii, Dubautia ciliolata and D. scabra are restricted to different lava flows, even though individuals of the two species may be found growing within a few meters of one another. Associated with this habitat difference is a difference in the tissue water deficits experienced by these two species. Midday water potentials in D. ciliolata are typically 0.4-0.5 MPa lower than in D. scabra.These two species also exhibit significant differences in their tissue osmotic and elastic properties. Dubautia ciliolata exhibits a lower tissue osmotic potential at full hydration and a lower tissue elastic modulus near full hydration than D. scabra. As a result, high and positive tissue turgor pressures are maintained to significantly lower tissue water contents and water potentials in D. ciliolata than in D. scabra. These differences in tissue osmotic and elastic properties appear to have a marked influence on diurnal turgor maintenance. Thus, while diurnal water potentials in D. ciliolata are significantly lower than in D. scabra, the diurnal turgor pressures exhibited by these two species are very similar.The natural hybrid between D. ciliolata and D. scabra exhibits intermediate tissue osmotic and elastic properties. This is evident, in particular, for the turgor dependence of the elastic modulus.The degree of phenotypic variation in the tissue osmotic and elastic properties of D. ciliolata appears to be relatively limited. As a result, plants of D. ciliolata growing under both well-watered conditions in the glasshouse and under natural conditions in the field exhibit a large capacity for maintaining high turgor pressures as tissue water content decreases.

Environmental characteristics, field water relations, and photosynthetic responses of C4 Hawaiian Euphorbia species from contrasting habitats.

Four endemic Hawaiian Euphorbia species range in habitat from open arid coastal strand to shaded mesic forest and in growth-form from small prostrate shrubs to trees. As shown in the present study, these large differences in habitat and growth-form are paralleled by equally large differences in maximal photosynthetic rate (13.7 to 37.1 µmol CO2 m-2s-1), dark respiration rate (0.7 to 4.1 µmol CO2 m-2s-1), light level for saturation of photosynthesis (0.9 to over 2.0 mmol m-2s-1), light compensation point (0.01 to 0.11 mmol m-2s-1), leaf conductance to CO2 (1.7 to 4.9 mm s-1), and mesophyll conductance to CO2 (3.7 to 8.5 mm s-1). A principal consequence of this differentiation is that the capacity for photosynthesis at high light levels is higher in open site species, such as E. celastroides and E. degeneri, and at low light levels is higher in shade species, such as E. forbesii. E. hillebrandii, a species from intermediate semiopen habitats, exhibits an intermediate photosynthetic capacity at both high and low light levels. Despite this remarkable diversity, all four species exhibit the distinguishing physiological features of C4 photosynthesis.

Photosynthetic responses of C3 and C4 species from cool shaded habitats in Hawaii.

The C4 species, Euphorbia forbesii, and the C3 species, Claoxylon sandwicense, occupy cool, shaded habitats in Hawaii. Both of these species exhibit the photosynthetic characteristics of typical shade plants: low light-saturated photosynthetic rates, low dark respiration rates, low light levels for saturation of photosynthesis, and low light compensation points. In addition, the quantum yields of the two species are similar at leaf temperatures near 22°C, reflecting a significant increase in the quantum yield of E. forbesii over that of C4 species from open habitats. C. sandwicense has a lower dark respiration rate than E. forbesii. Hence, since the quantum yields of the two species are similar at cool temperatures, C. sandwicense has a higher photosynthetic rate than E. forbesii at low incident photon flux densities. As a consequence, C. sandwicense should have a greater carbon gain than E. forbesii under the diffuse radiation conditions of their native habitat. However, since E. forbesii has a higher light-saturated photosynthetic rate than C. sandwicense, E. forbesii may have a greater carbon gain than C. sandwicense during sunflecks.