Rachael V. Gallagher

Phenological trends among Australian alpine species: using herbarium records to identify climate-change indicators

Globaltemperatures areincreasingat anunprecedentedrateandthe analysisof long-termphenological records hasprovidedsomeofthemostcompellingevidencefortheeffectofthesechangesonspecies.Inregionswheresystematically collected data on the timing of life-cycle events is scarce, such as Australia, researchers must seek alternative sources of information from which climate-change signals can be identified. In the present paper, we explore the limitations and strengthsofusingherbariumspecimenstodetectchangesin floweringphenology,toselectpotentialindicatorspecies,andto pinpoint locations for potential monitoring schemes of native plants in Australias subalpine and alpine zone. We selected 20 species on the basis of a range of selection criteria, including a flowering duration of 3 months or less and the number of herbarium records available in the areas above 1500m. By the use of gridded temperature data within the study region, we identifiedanincreaseinmeanannualtemperatureof0.74 � Cbetween1950and2007.Wethenmatchedthespatiallocationsof the herbarium specimens to these temperature data and, by using linear regression models, identified five species whose flowering response may be sensitive to temperature. Higher mean annual temperatures at the point of collection were negativelyassociatedwithearlier floweringineachofthesespecies(a=0.05).Wealsofoundasignificant(P=0.02)negative relationship between year and flowering observation for Alpine groundsel, Senecio pectinatus var. major. This species is potentially a suitable candidate for monitoring responses of species to future climate change, owing to the accessibility of populations and its conspicuous flowers. It is also likely that with ongoing warming the other four species identified (Colobanthus affinis, Ewartia nubigena, Prasophyllum tadgellianum and Wahlenbergia ceracea) in the present study may show the same response.

Global climatic drivers of leaf size

Leaf size, climate, and energy balance Why does plant leaf size increase at lower latitudes, as exemplified by the evolutionary success of species with very large leaves in the tropics? Wright et al. analyzed leaf data for 7670 plant species, along with climatic data, from 682 sites worldwide. Their findings reveal consistent patterns and explain why earlier predictions from energy balance theory had only limited success. The authors provide a fully quantitative explanation for the latitudinal gradient in leaf size, with implications for plant ecology and physiology, vegetation modeling, and paleobotany. Science, this issue p. 917 Day- and nighttime leaf-to-air temperature differences drive global gradients in leaf size. Leaf size varies by over a 100,000-fold among species worldwide. Although 19th-century plant geographers noted that the wet tropics harbor plants with exceptionally large leaves, the latitudinal gradient of leaf size has not been well quantified nor the key climatic drivers convincingly identified. Here, we characterize worldwide patterns in leaf size. Large-leaved species predominate in wet, hot, sunny environments; small-leaved species typify hot, sunny environments only in arid conditions; small leaves are also found in high latitudes and elevations. By modeling the balance of leaf energy inputs and outputs, we show that daytime and nighttime leaf-to-air temperature differences are key to geographic gradients in leaf size. This knowledge can enrich “next-generation” vegetation models in which leaf temperature and water use during photosynthesis play key roles.

Trait differences between naturalized and invasive plant species independent of residence time and phylogeny

The ability to predict which alien plants will transition from naturalized to invasive prior to their introduction to novel regions is a key goal for conservation and has the potential to increase the efficacy of weed risk assessment (WRA). However, multiple factors contribute to plant invasion success (e.g., functional traits, range characteristics, residence time, phylogeny), and they all must be taken into account simultaneously in order to identify meaningful correlates of invasion success. We compiled 146 pairs of phylogenetically paired (congeneric) naturalized and invasive plant species in Australia with similar minimum residence times (i.e., time since introduction in years). These pairs were used to test for differences in 5 functional traits (flowering duration, leaf size, maximum height, specific leaf area [SLA], seed mass) and 3 characteristics of species’ native ranges (biome occupancy, mean annual temperature, and rainfall breadth) between naturalized and invasive species. Invasive species, on average, had larger SLA, longer flowering periods, and were taller than their congeneric naturalized relatives. Invaders also exhibited greater tolerance for different environmental conditions in the native range, where they occupied more biomes and a wider breadth of rainfall and temperature conditions than naturalized congeners. However, neither seed mass nor leaf size differed between pairs of naturalized and invasive species. A key finding was the role of SLA in distinguishing between naturalized and invasive pairs. Species with high SLA values were typically associated with faster growth rates, more rapid turnover of leaf material, and shorter lifespans than those species with low SLA. This suite of characteristics may contribute to the ability of a species to transition from naturalized to invasive across a wide range of environmental contexts and disturbance regimes. Our findings will help in the refinement of WRA protocols, and we advocate the inclusion of quantitative traits, in particular SLA, into the WRA schemes. Diferencia de Características entre Especies de Plantas Naturalizadas e Invasoras Independientes del Tiempo de Residencia y de la Filogenia Resumen La habilidad para predecir cuáles plantas exóticas harán la transición de naturalizadas a invasoras antes de su introducción a regiones nuevas es un objetivo clave para la conservación y tiene el potencial de incrementar la eficiencia de la evaluación de riesgo de hierbas (ERH). Sin embargo, múltiples factores contribuyen al éxito invasor de las plantas (p. ej.: características funcionales, características de cobertura, tiempo de residencia, filogenia) y todos deben considerarse simultáneamente para poder identificar correlaciones significativas del éxito invasor. Recopilamos en Australia 146 parejas de especies de plantas invasoras y naturalizadas emparejadas filogenéticamente (congéneres) y con tiempos de residencia mínima similares (es decir, el tiempo transcurrido desde su introducción en años). Estas parejas se usaron para probar diferencias en cinco características funcionales (duración de la floración, tamaño de la hoja, altura máxima, área específica de la hoja [AEH], masa de la semilla) y en tres características de cobertura nativa de las especies (ocupación de bioma, temperatura media anual y amplitud de pluviosidad) entre especies invasoras y naturalizadas. Las especies invasoras, en promedio, tuvieron una mayor AEH, periodos de floración más largos y fueron más altas que sus parientes congéneres naturalizadas. Las invasoras también exhibieron una mayor tolerancia a diferentes condiciones ambientales en su cobertura nativa, donde ocuparon más biomas y una mayor amplitud de pluviosidad y condiciones de temperatura que sus congéneres naturalizadas. Sin embargo, ni la masa de la semilla ni el tamaño de hoja difirieron entre las parejas de especies naturalizadas e invasoras. Un hallazgo relevante fue el papel de la AEH en la distinción entre las parejas naturalizadas e invasoras. Las especies con valores altos de AEH estuvieron asociadas típicamente con tasas mayores de crecimiento, pérdida rápida de volumen de material de hojas y periodos de vida más cortos que aquellas especies con AEH baja. Este conjunto de características puede contribuir a la habilidad de las especies para llevar a cabo la transición de naturalizada a invasora a lo largo de una amplia cobertura de contextos ambientales y regímenes de perturbación. Nuestros hallazgos ayudarán en la mejora de los protocolos de ERH, y abogamos por la inclusión de las características cuantitativas, en particular la AEH, en los esquemas de ERH.

Next-Generation Invaders? Hotspots for Naturalised Sleeper Weeds in Australia under Future Climates

Naturalised, but not yet invasive plants, pose a nascent threat to biodiversity. As climate regimes continue to change, it is likely that a new suite of invaders will emerge from the established pool of naturalised plants. Pre-emptive management of locations that may be most suitable for a large number of potentially invasive plants will help to target monitoring, and is vital for effective control. We used species distribution models (SDM) and invasion-hotspot analysis to determine where in Australia suitable habitat may occur for 292 naturalised plants. SDMs were built in MaxEnt using both climate and soil variables for current baseline conditions. Modelled relationships were projected onto two Representative Concentration Pathways for future climates (RCP 4.5 and 8.5), based on seven global climate models, for two time periods (2035, 2065). Model outputs for each of the 292 species were then aggregated into single ‘hotspot’ maps at two scales: continental, and for each of Australia’s 37 ecoregions. Across Australia, areas in the south-east and south-west corners of the continent were identified as potential hotspots for naturalised plants under current and future climates. These regions provided suitable habitat for 288 and 239 species respectively under baseline climates. The areal extent of the continental hotspot was projected to decrease by 8.8% under climates for 2035, and by a further 5.2% by 2065. A similar pattern of hotspot contraction under future climates was seen for the majority of ecoregions examined. However, two ecoregions - Tasmanian temperate forests and Australian Alps montane grasslands - showed increases in the areal extent of hotspots of >45% under climate scenarios for 2065. The alpine ecoregion also had an increase in the number of naturalised plant species with abiotically suitable habitat under future climate scenarios, indicating that this area may be particularly vulnerable to future incursions by naturalised plants.

Predicted impact of exotic vines on an endangered ecological community under future climate change

Potential interactions between climate change and exotic plant invasions may affect areas of high conservation value, such as land set aside for the protection of endangered species or ecological communities. We investigated this issue in eastern Australia using species distribution models for five exotic vines under climate regimes for 2020 and 2050. We examined how projected changes in the distribution of climatically suitable habitat may coincide with the remaining remnants of an endangered ecological community—littoral rainforests—in this region. The number of known infestations of each weed in tropical, subtropical and temperate areas was used to assess the likelihood of further expansion into areas projected to provide suitable habitat under future conditions. Littoral rainforest reserves were consistently predicted to provide bioclimatically suitable habitat for the five vines examined under both current and future climate scenarios. We explore the consequences and potential strategies for managing exotic plant invasions in these protected areas in the coming decades.

The grass may not always be greener: projected reductions in climatic suitability for exotic grasses under future climates in Australia.

Climate change presents a new challenge for the management of invasive exotic species that threaten both biodiversity and agricultural productivity. The invasion of exotic perennial grasses throughout the globe is particularly problematic given their impacts on a broad range of native plant communities and livelihoods. As the climate continues to change, pre-emptive long-term management strategies for exotic grasses will become increasingly important. Using species distribution modelling we investigated potential changes to the location of climatically suitable habitat for some exotic perennial grass species currently in Australia, under a range of future climate scenarios for the decade centred around 2050. We focus on eleven species shortlisted or declared as the Weeds of National Significance or Alert List species in Australia, which have also become successful invaders in other parts of the world. Our results indicate that the extent of climatically suitable habitat available for all of the exotic grasses modelled is projected to decrease under climate scenarios for 2050. This reduction is most severe for the three species of Needle Grass (genus Nassella) that currently have infestations in the south-east of the continent. Combined with information on other aspects of establishment risk (e.g. demographic rates, human-use, propagule pressure), predictions of reduced climatic suitability provide justification for re-assessing which weeds are prioritised for intensive management as the climate changes.

A tool to assess potential for alien plant establishment and expansion under climate change

Predicting the influence of climate change on the potential distribution of naturalised alien plant species is an important and challenging task. While prioritisation of management actions for alien plants under current climatic conditions has been widely adopted, very few systems explicitly incorporate the potential of future changes in climate conditions to influence the distribution of alien plant species. Here, we develop an Australia-wide screening tool to assess the potential of naturalised alien plants to establish and spread under both current and future climatic conditions. The screening tool developed uses five spatially explicit criteria to establish the likelihood of alien plant population establishment and expansion under baseline climate conditions and future climates for the decades 2035 and 2065. Alien plants are then given a threat rating according to current and future threat to enable natural resource managers to focus on those species that pose the largest potential threat now and in the future. To demonstrate the screening tool, we present results for a representative sample of approximately 10% (n = 292) of Australias known, naturalised alien plant species. Overall, most alien plant species showed decreases in area of habitat suitability under future conditions compared to current conditions and therefore the threat rating of most alien plant species declined between current and future conditions. Use of the screening tool is intended to assist natural resource managers in assessing the threat of alien plant establishment and spread under current and future conditions and thus prioritise detailed weed risk assessments for those species that pose the greatest threat. The screening tool is associated with a searchable database for all 292 alien plant species across a range of spatial scales, available through an interactive web-based portal at http://weedfutures.net/.

Effects of Growth Form and Functional Traits on Response of Woody Plants to Clearing and Fragmentation of Subtropical Rainforest

The conservation implications of large-scale rainforest clearing and fragmentation on the persistence of functional and taxonomic diversity remain poorly understood. If traits represent adaptive strategies of plant species to particular circumstances, the expectation is that the effect of forest clearing and fragmentation will be affected by species functional traits, particularly those related to dispersal. We used species occurrence data for woody plants in 46 rainforest patches across 75,000 ha largely cleared of forest by the early 1900s to determine the combined effects of area reduction, fragmentation, and patch size on the taxonomic structure and functional diversity of subtropical rainforest. We compiled species trait values for leaf area, seed dry mass, wood density, and maximum height and calculated species niche breadths. Taxonomic structure, trait values (means, ranges), and the functional diversity of assemblages of climbing and free-standing plants in remnant patches were quantified. Larger rainforest patches had higher species richness. Species in smaller patches were taxonomically less related than species in larger patches. Free-standing plants had a high percentage of frugivore dispersed seeds; climbers had a high proportion of small wind-dispersed seeds. Connections between the patchy spatial distribution of free-standing species, larger seed sizes, and dispersal syndrome were weak. Assemblages of free-standing plants in patches showed more taxonomic and spatial structuring than climbing plants. Smaller isolated patches retained relatively high functional diversity and similar taxonomic structure to larger tracts of forest despite lower species richness. The response of woody plants to clearing and fragmentation of subtropical rainforest differed between climbers and slow-growing mature-phase forest trees but not between climbers and pioneer trees. Quantifying taxonomic structure and functional diversity provides an improved basis for conservation planning and management by elucidating the effects of forest-area reduction and fragmentation. Efectos de la Forma de Crecimiento y Atributos Funcionales en la Respuesta de Plantas Leñosas al Desmonte y Fragmentación de Bosque Lluvioso Subtropical.

Characterizing opportunistic breeding at a continental scale using all available sources of phenological data: An assessment of 337 species across the Australian continent

ABSTRACT Research from the intensively studied northern temperate and boreal regions dominates avian reproductive phenology studies. However, in most other areas, long-term, high-quality phenological datasets are not available, limiting our ability to predict how reproductive timing may respond to rapid climate change. Here, we provide novel methods for combining conventional and nonconventional observations to understand phenological patterns in birds across a southern continent. Observations from egg collections, bird banding, nest record schemes, and citizen science were combined to determine egg-laying phenology for ∼50% of Australias mainland breeding species. We investigated start, peak, and length of avian egg-laying periods (1) derived from different data sources, (2) across tropical, subtropical, desert, grassland, and temperate biomes, and (3) comparing 2 representative temperate regions of the northern and southern hemispheres. We found that start and peak egg-laying dates calculated from single-visit observations of young or eggs resulted in similar dates as those from more accurate multi-visit nest observations. This demonstration suggests that future studies aimed at assessing changes in the timing of breeding in response to climate change can utilize such observational data. This will significantly increase sample sizes, rather than restricting such analyses to just intensively tracked nests, for which accurate laying dates are available. We found that egg-laying phenology varies between biomes (tropical, subtropical, desert, grassland, temperate), with birds in the desert biome having the earliest peaks of egg-laying. Finally, the length of the egg-laying period differs significantly between hemispheres. The southern temperate zone species have extensive egg-laying periods and many species breed year-round in marked contrast to the highly predictable, springtime breeding in the north. Therefore, avian phenological patterns and documented responses to climate change from the well-sampled, but highly seasonal, northern hemisphere may not be transferrable across the globe.

Variation in avian egg shape and nest structure is explained by climatic conditions

Why are avian eggs ovoid, while the eggs of most other vertebrates are symmetrical? The interaction between an egg and its environment likely drives selection that will shape eggs across evolutionary time. For example, eggs incubated in hot, arid regions face acute exposure to harsh climatic conditions relative to those in temperate zones, and this exposure will differ across nest types, with eggs in open nests being more exposed to direct solar radiation than those in enclosed nests. We examined the idea that the geographical distribution of both egg shapes and nest types should reflect selective pressures of key environmental parameters, such as ambient temperature and the drying capacity of air. We took a comparative approach, using 310 passerine species from Australia, many of which are found in some of the most extreme climates on earth. We found that, across the continent, egg elongation decreases and the proportion of species with domed nests with roofs increases in hotter and drier areas with sparse plant canopies. Eggs are most spherical in open nests in the hottest environments, and most elongate in domed nests in wetter, shadier environments. Our findings suggest that climatic conditions played a key role in the evolution of passerine egg shape.