Vegetation response to removal of non-native feral pigs from Hawaiian tropical montane wet forest

Globally, non-native ungulates threaten native biodiversity, alter biotic and abiotic factors regulating ecological processes, and incur significant economic costs via herbivory, rooting, and trampling. Removal of non-native ungulates is an increasingly common and crucial first step in conserving and restoring native forests. However, removal is often controversial and there is currently little information on plant community responses to this management action. Here, we examine the response of native and non-native understory vegetation in paired sites inside and outside of exclosures across a 6.5–18.5 year chronosequence of feral pig (Sus scrofa) removal from canopy-intact Hawaiian tropical montane wet forest. Stem density and cover of native plants, species richness of ground-rooted native woody plants, and abundance of native plants of conservation interest were all significantly higher where feral pigs had been removed. Similarly, the area of exposed soil was substantially lower and cover of litter and bryophytes was greater with feral pig removal. Spatial patterns of recruitment were also strongly affected. Whereas epiphytic establishment was similar between treatments, the density of ground-rooted woody plants was four times higher with feral pig removal. Abundance of invasive non-native plants also increased at sites where they had established prior to feral pig removal. We found no patterns in any of the measured variables with time, suggesting that commonly occurring species recover within 6.5 years of feral pig removal. Recovery of species of conservation interest, however, was highly site specific and limited to areas that possessed remnant populations at the time of removal, indicating that some species take much longer (>18.5 years) to recover. Feral pig removal is the first and most crucial step for conservation of native forests in this area, but subsequent management should also include control of non-native invasive plants and outplanting native species of conservation interest that fail to recruit naturally.     

Ecological impacts of feral pigs in the Hawaiian Islands

The foraging habits of exotic ungulate species can directly and indirectly affect native plant and animal distribution and abundance patterns. Most of the studies on feral pig interactions with other biota in the Hawaiian Islands have been published as difficult to access reports to governmental and nongovernmental organizations, graduate student theses, and a few in peer reviewed journals. In this paper we discuss the origins of pig introductions to Hawaii, their feralization process, population expansion, and interactions with native and non-native biota. We also consider the environmental effects triggered by pigs on local ecosystems and biotic communities. Feral pig activities can reduce the abundance of native plant species, enhance conditions for the establishment of invasive non-indigenous plants, and perhaps indirectly negatively impact native forest bird species. Pig foraging and traveling patterns may also lead to physical alteration of ecosystems by increasing soil erosion that may lead to watershed degradation. However, much remains to be learned about the strength and significance of aforementioned interactions and their long-term effects on Hawaiian biota and ecosystems due to some confounding events. Elucidating the dynamics and long-term ecological effects generated by pigs is a crucial step towards increasing our understanding of and more effectively managing biotic interactions.     

Invasive aphids attack native Hawaiian plants

Invasive species have had devastating impacts on the fauna and flora of the Hawaiian Islands. While the negative effects of some invasive species are obvious, other species are less visible, though no less important. Aphids (Homoptera: Aphididae) are not native to Hawai’i but have thoroughly invaded the Island chain, largely as a result of anthropogenic influences. As aphids cause both direct plant feeding damage and transmit numerous pathogenic viruses, it is important to document aphid distributions and ranges throughout the archipelago. On the basis of an extensive survey of aphid diversity on the five largest Hawaiian Islands (Hawai’i, Kaua’i, O’ahu, Maui, and Moloka’i), we provide the first evidence that invasive aphids feed not just on agricultural crops, but also on native Hawaiian plants. To date, aphids have been observed feeding and reproducing on 64 native Hawaiian plants (16 indigenous species and 48 endemic species) in 32 families. As the majority of these plants are endangered, invasive aphids may have profound impacts on the island flora. To help protect unique island ecosystems, we propose that border vigilance be enhanced to prevent the incursion of new aphids, and that biological control efforts be renewed to mitigate the impact of existing species.     

Invasive plant suppresses the growth of native tree seedlings by disrupting belowground mutualisms

The impact of exotic species on native organisms is widely acknowledged, but poorly understood. Very few studies have empirically investigated how invading plants may alter delicate ecological interactions among resident species in the invaded range. We present novel evidence that antifungal phytochemistry of the invasive plant, Alliaria petiolata, a European invader of North American forests, suppresses native plant growth by disrupting mutualistic associations between native canopy tree seedlings and belowground arbuscular mycorrhizal fungi. Our results elucidate an indirect mechanism by which invasive plants can impact native flora, and may help explain how this plant successfully invades relatively undisturbed forest habitat.     

Influence of woody invader control methods and seed availability on native and invasive species establishment in a Hawaiian forest

When invasive woody plants become dominant, they present an extreme challenge for restoration of native plant communities. Invasive Morella faya (fire tree) forms extensive, nearly monospecific stands in wet and mesic forests on the Island of Hawai’i. We used logging, girdling, and selective girdling over time (incremental girdling) to kill stands of M. faya at different rates, with the objective of identifying a method that best promotes native forest re-establishment. We hypothesized that rapid canopy opening by logging would lead to establishment of fast-growing, non-native invaders, but that slower death of M. faya by girdling or incremental girdling would increase the establishment by native plants adapted to partial shade conditions. After applying the M. faya treatments, seed banks, seed rain, and plant recruitment were monitored over 3 years. Different plant communities developed in response to the treatments. Increased light and nitrogen availability in the logged treatment were associated with invasion by non-native species. Native species, including the dominant native forest tree, (Metrosideros polymorpha) and tree fern (Cibotium glaucum), established most frequently in the girdle and incremental girdle treatments, but short-lived non-native species were more abundant than native species. A diverse native forest is unlikely to develop following any of the treatments due to seed limitation for many native species, but girdling and incremental girdling promoted natural establishment of major components of native Hawaiian forest. Girdling may be an effective general strategy for reestablishing native vegetation in areas dominated by woody plant invaders.     

Non-timber forest product harvesting in alien-dominated forests: effects of frond-harvest and rainfall on the demography of two native Hawaiian ferns

Non-timber forest products (NTFP) represent culturally and economically important resources for millions of people worldwide. Although many NTFP are harvested from disturbed habitats and therefore subject to multiple pressures, few quantitative studies have addressed this issue. Similarly few NTFP studies have assessed seasonal variation in demographic rates even though this can confound harvest effects. In Hawaiȁ8i, the wild-gathered ferns, Microlepia strigosa and Sphenomeris chinensis, represent highly important cultural resources but declining populations have led to conservation concerns. Both ferns are harvested from disturbed, alien-dominated forests and contemporary Hawaiian gathering practices often consist of harvest and concurrent weeding of alien invasive species. We assessed the effects of concurrent frond-harvest and alien species weeding on frond structure, density, and rates of production by comparing experimentally harvested vs. control plots, and documented relationships between frond demographic patterns and precipitation. Gathering practices had no impact on frond density of either species or on most other demographic parameters over the short term. Exceptions included a significant decrease in the density of the longest S. chinensis fronds and a significant decrease in M. strigosa frond production when fronds were gathered without alien weeding. However, seasonal and annual changes in frond density and production occurred across all plots of both species and were significantly correlated with precipitation. The relatively low harvest effects for both species are likely due to several factors including short frond longevity and the strict criteria used by gatherers to select harvestable fronds. The potential for sustainable harvest in the context of alien-dominated forests is discussed.     

Response of native Hawaiian woody species to lava-ignited wildfires in tropical forests and shrublands

Wildfires are rare in the disturbance history of Hawaiian forests but may increase in prevalence due to invasive species and global climate change. We documented survival rates and adaptations facilitating persistence of native woody species following 2002–2003 wildfires in Hawaii Volcanoes National Park, Hawaii. Fires occurred during an El Niño drought and were ignited by lava flows. They burned across an environmental gradient occupied by two drier shrub-dominated communities and three mesic/wet Metrosideros forest communities. All the 19 native tree, shrub, and tree fern species demonstrated some capacity of postfire persistence. While greater than 95% of the dominant Metrosideros trees were top-killed, more than half survived fires via basal sprouting. Metrosideros trees with diameters >20 cm sprouted in lower percentages than smaller trees. At least 17 of 29 native woody species colonized the postfire environment via seedling establishment. Although the native biota possess adaptations facilitating persistence following wildfire, the presence of highly competitive invasive plants and ungulates will likely alter postfire succession.     

Architecture of remnant trees influences native woody plant recruitment in abandoned Hawaiian pastures

Plant Ecology - Abandoned tropical pastures offer opportunities for passive and active restoration of native forest communities. Tree architecture of remnant canopy trees may be one important...     

Invasive insects impact forest carbon dynamics

Invasive insects can impact ecosystem functioning by altering carbon, nutrient, and hydrologic cycles. In this study, we used eddy covariance to measure net CO₂ exchange with the atmosphere (NEE), and biometric measurements to characterize net ecosystem productivity (NEP) in oak‐ and pine‐dominated forests that were defoliated by Gypsy moth (Lymantria dispar L.) in the New Jersey Pine Barrens. Three years of data were used to compare C dynamics; 2005 with minimal defoliation, 2006 with partial defoliation of the canopy and understory in a mixed stand, and 2007 with complete defoliation of an oak‐dominated stand, and partial defoliation of the mixed and pine‐dominated stands. Previous to defoliation in 2005, annual net CO₂ exchange (NEE_yr) was estimated at ?187, ?137 and ?204 g C m^?2 yr^?1 at the oak‐, mixed‐, and pine‐dominated stands, respectively. Annual NEP estimated from biometric measurements was 108%, 100%, and 98% of NEE_yr in 2005 for the oak‐, mixed‐, and pine‐dominated stands, respectively. Gypsy moth defoliation strongly reduced fluxes in 2006 and 2007 compared with 2005; NEE_yr was ?122, +103, and ?161 g C m^?2 yr^?1 in 2006, and +293, +129, and ?17 g C m^?2 yr^?1 in 2007 at the oak‐, mixed‐, and pine‐dominated stands, respectively. At the landscape scale, Gypsy moths defoliated 20.2% of upland forests in 2007. We calculated that defoliation in these upland forests reduced NEE_yr by 41%, with a 55% reduction in the heavily impacted oak‐dominated stands. ‘Transient’ disturbances such as insect defoliation, nonstand replacing wildfires, and prescribed burns are major factors controlling NEE across this landscape, and when integrated over time, may explain much of the patterning of aboveground biomass and forest floor mass in these upland forests.     

Photosynthesis, photoinhibition, and nitrogen use efficiency in native and invasive tree ferns in Hawaii

Photosynthetic gas exchange, chlorophyll fluorescence, nitrogen use efficiency, and related leaf traits of native Hawaiian tree ferns in the genus Cibotium were compared with those of the invasive Australian tree fern Sphaeropteris cooperi in an attempt to explain the higher growth rates of S. cooperi in Hawaii. Comparisons were made between mature sporophytes growing in the sun (gap or forest edge) and in shady understories at four sites at three different elevations. The invasive tree fern had 12-13 cm greater height increase per year and approximately 5 times larger total leaf surface area per plant compared to the native tree ferns. The maximum rates of photosynthesis of S. cooperi in the sun and shade were significantly higher than those of the native Cibotium spp (for example, 11.2 and 7.1 µmol m^-2 s^-1, and 5.8 and 3.6 µmol m^-2 s^-1 respectively for the invasive and natives at low elevation). The instantaneous photosynthetic nitrogen use efficiency of the invasive tree fern was significantly higher than that of the native tree ferns, but when integrated over the life span of the frond the differences were not significant. The fronds of the invasive tree fern species had a significantly shorter life span than the native tree ferns (approximately 6 months and 12 months, respectively), and significantly higher nitrogen content per unit leaf mass. The native tree ferns growing in both sun and shade exhibited greater photoinhibition than the invasive tree fern after being experimentally subjected to high light levels. The native tree ferns recovered only 78% of their dark-acclimated quantum yield (F_v/F_m), while the invasive tree fern recovered 90% and 86% of its dark-acclimated F_v/F_m when growing in sun and shade, respectively. Overall, the invasive tree fern appears to be more efficient at capturing and utilizing light than the native Cibotium species, particularly in high-light environments such as those associated with high levels of disturbance.     

Broadcast application of a placebo rodenticide bait in a native Hawaiian forest

This study consisted of three replicates of controlled field trials using a pelletized placebo (Ramik® Green formulated without diphacinone) bait treated with a biological marker and broadcast at three application rates — 11.25, 22.5 and 33.75 kg/ha. We determined that Polynesian (Rattus exulans) and roof rats (Rattus rattus) consumed this bait when broadcast on the ground and assessed the optimal sowage rate to result in maximum exposure of bait to the rats while minimizing bait usage. All Polynesian rats captured in all application rates had eaten the bait. The percentage of roof rats that had eaten the bait increased with application rate, however, 22.5 kg/ha was clearly the optimal application rate. Bait degradation and invertebrate activity was documented and assessed.     

Interspecific interactions between feral pigs and native birds reveal both positive and negative effects

In tropical Australian rainforests, predators and scavengers aggregate beneath emergent trees that house large colonies of metallic starlings (Aplonis metallica), feeding in the nutrient‐rich open areas below. Analysis of camera‐trap records shows that the presence of feral pigs (Sus scrofa) is associated with an absence of birds (cockatoos and brush turkeys), presumably reflecting behavioural avoidance (pigs pose a direct danger to birds). However, bird numbers increase as soon as pigs depart, then fall if pigs are absent for long periods. Feral pigs thus displace native birds from these resource hotspots; but by turning over the soil and enhancing the birds' access to food, the pigs also have a positive impact on food availability for the avifauna. Thus, although invasive species have caused irreparable environmental damages worldwide, they may also provide positive benefits for certain species. The net benefit of such interspecific interactions will depend on the outcome of both positive and negative effects.     

Evaluation of feral pig removal in Hawaiian preserves

The Nature Conservancy of Hawaii (TNC) recently embarked on an ambitious ungulate control programme throughout their preserves on the islands of Maui and Molokai. The aim of the programme was local eradication of feral pigs (Sus scrofa) and they wanted some way of evaluating their progress. Catch-effort models have previously been applied to cumulative pig dispatches during an island eradication programme (Ramsey et al. 2009; Conservation Biology 23: 449-459). These models simultaneously estimate the parameters describing the initial population size and the probability of detecting an individual per unit of hunting effort, which can then be used to evaluate the likelihood of eradication. However these models rely on a number of assumptions including that the system is closed except for removals and that the relationship between hunting effort and the probability of detection is constant throughout the experiment. As the TNC control programme progressed it became clear the both these assumptions were violated and more pigs were often caught per unit of effort on the later compared with the earlier hunts. There was ongoing immigration into the preserves through breaks in the fence and via unfenced boundaries. Also, later hunts seemed to be more successful per unit of effort than earlier ones, presumably because hunters learnt the best way to cover the area and where the sites most likely to contain pigs were. We described how we incorporated this learning process into a catch-effort model using Bayesian updating in order to evaluate the efficacy of the control programme.     

Temporal and spatial partitioning of water resources among eight woody species in a Hawaiian dry forest

Lowland dry forests are unique in Hawaii for their high diversity of tree species compared with wet forests. We characterized spatial and temporal partitioning of soil water resources among seven indigenous and one invasive dry forest species to determine whether the degree of partitioning was consistent with the relatively high species richness in these forests. Patterns of water utilization were inferred from stable hydrogen isotope ratios (δD) of soil and xylem water, zones of soil water depletion, plant water status, leaf phenology, and spatial patterns of species distribution. Soil water δD values ranged from –20‰ near the surface to –48‰ at 130 cm depth. Metrosideros polymorpha, an evergreen species, and Reynoldsia sandwicensis, a drought-deciduous species, had xylem sap δD values of about –52‰, and appeared to obtain their water largely from deeper soil layers. The remaining six species had xylem δD values ranging from –33 to –42‰, and apparently obtained water from shallower soil layers. Xylem water δD values were negatively correlated with minimum annual leaf water potential and positively correlated with leaf solute content, an integrated measure of leaf water deficit. Seasonal patterns of leaf production ranged from dry season deciduous at one extreme to evergreen with near constant leaf expansion rates at the other. Species tapping water more actively from deeper soil layers tended to exhibit larger seasonality of leaf production than species relying on shallower soil water sources. Individuals of Myoporum sandwicense were more spatially isolated than would be expected by chance. Even though this species apparently extracted water primarily from shallow soil layers, as indicated by its xylem δD values, its nearly constant growth rates across all seasons may have been the result of a larger volume of soil water available per individual. The two dominant species, Diospyros sandwicensis and Nestegis sandwicensis, exhibited low leaf water potentials during the dry season and apparently drew water mostly from the upper portion of the soil profile, which may have allowed them to exploit light precipitation events more effectively than the more deeply rooted species. Character displacement in spatial and temporal patterns of soil water uptake was consistent with the relatively high diversity of woody species in Hawaiian dry forests. Received: 20 May 1999 / Accepted: 2 March 2000     

Persistence of an endangered native duck,feral mallards,and multiple hybrid swarms across the main Hawaiian Islands

Interspecific hybridization is recognized as an important process in the evolutionary dynamics of both speciation and the reversal of speciation. However, our understanding of the spatial and temporal patterns of hybridization that erode versus promote species boundaries is incomplete. The endangered, endemic koloa maoli (or Hawaiian duck, Anas wyvilliana) is thought to be threatened with genetic extinction through ongoing hybridization with an introduced congener, the feral mallard (A. platyrhynchos). We investigated spatial and temporal variation in hybrid prevalence in populations throughout the main Hawaiian Islands, using genomic data to characterize population structure of koloa, quantify the extent of hybridization, and compare hybrid proportions over time. To accomplish this, we genotyped 3,308 double‐digest restriction‐site‐associated DNA (ddRAD) loci in 425 putative koloa, mallards, and hybrids from populations across the main Hawaiian Islands. We found that despite a population decline in the last century, koloa genetic diversity is high. There were few hybrids on the island of Kaua?i, home to the largest population of koloa. By contrast, we report that sampled populations outside of Kaua?i can now be characterized as hybrid swarms, in that all individuals sampled were of mixed koloa × mallard ancestry. Further, there is some evidence that these swarms are stable over time. These findings demonstrate spatial variation in the extent and consequences of interspecific hybridization, and highlight how islands or island‐like systems with small population sizes may be especially prone to genetic extinction when met with a congener that is not reproductively isolated.     

Bryophytes display allelopathic interactions with tree species in native forest ecosystems

Bryophytes are widespread in terrestrial ecosystems but little is known about their influence on vascular species. Water‐soluble leachates (0%, 1%, 5%, 10% concentration) derived from 18 species of bryophytes (mosses 11 species; liverworts 7 species) were tested on the germination and seedling growth of Lactuca sativa and two common trees Melicytus ramiflorus (Violaceae) and Fuchsia excorticata (Onagraceae) in southern New Zealand forests. Bryophyte water soluble extracts (BWSE) have minor impact on seed germination of Lactuca, stimulatory effects on radical growth at low (1%) concentrations and inhibitory effects at higher concentrations (5–10%). For Melicytus the BWSE had variable effects, with evidence of strong stimulatory (Dendrohypopterygium filiculiforme) and inhibitory (Lepidozia concinna) effects on germination, but generally inhibited radical growth. BWSE at all test concentrations consistently inhibit both germination and seedling radicle growth in Fuchsia. The toxicity effect of water‐soluble leachates varies significantly between bryophyte species but not consistently between mosses and liverworts. Bryophyte species exhibiting strongest inhibition effects under control conditions were associated with significantly reduced densities of broadleaved tree seedlings in forest ecosystems. Our results demonstrate that some bryophyte species via allelopathic interactions can inhibit seedling establishment and growth of forest trees. This mechanism provides an additional factor constraining the spatial distribution of the regeneration niche in forest communities.     

Response of forest tree seedlings to simulated litterfall damage.

Litterfall is an important cause of seedling mortality in many forests ranging from wet tropical to boreal. However, there is a lack of studies that investigate differences between species in seedling resilience to litterfall damage. We selected seedling pairs of seven tree species and simulated litterfall damage by pinning one of each pair to the ground. The mortality and growth rates varied significantly between species for pinned individuals, but were similar for unpinned seedlings. The mortality of pinned Nestegis cunninghamii and Prumnopitys ferruginea was significantly greater than that of unpinned individuals (P < 0.05). However, contrary to expectations, the growth rates of pinned Hedycarya arborea and Nothofagus menziesii were much greater than for those left unpinned (P < 0.05). In general, seedling resilience to the bending damage differed substantially between species. N. cunninghamii and P. ferruginea suffered high mortalities and did not increase growth rates in response to damage, whereas, H. arborea and N. menziesii suffered few mortalities and regained height quickly. Other study species demonstrated intermediate resilience. This study demonstrates that some species are more likely to survive in high-risk litterfall regimes than are others. Given that litterfall risk can vary greatly between microsites, these results suggest that litterfall can contribute to regeneration niche differentiation. This revised version was published online in August 2006 with corrections to the Cover Date.