Springs and wire plants: anachronistic defences against Madagascar's extinct elephant birds

The extinction of large vertebrates in the last few millennia has left a legacy of evolutionary anachronisms. Among these are plant structural defences that persist long after the extinction of the browsers. A peculiar, and controversial, example is a suite of traits common in divaricate (wide-angled branching) plants from New Zealand. Divaricate architecture has been interpreted as an adaptive response to cold climates or an anachronistic defence against the extinct moas. Madagascar, a larger tropical island, also had a fauna of large flightless birds, the elephant birds. If these extinct ratites selected for similar plant defences, we expected to find convergent features between New Zealand and Malagasy plants, despite their very different climates. We searched the southern thickets of Madagascar for plants with putative anti-ratite defences and scored candidate species for a number of traits common to many New Zealand divaricates. We found many Malagasy species in 25 families and 36 genera shared the same suite of traits, the 'wire plant' syndrome, as divaricates in New Zealand that resist ratite browsing. Neither ecologically, nor phylogenetically, matched species from South Africa shared these traits. Malagasy wire plants differ from many New Zealand divaricates in lacking the distinctive concentration of leaves in the interior of shrubs. We suggest that New Zealand divaricates have a unique amalgam of traits that acted as defences and also confer tolerance to cold. We conclude that many woody species in the thickets of southern Madagascar share, with New Zealand, anachronistic structural defences against large extinct bird browsers.     

Ratite and ungulate preferences for woody New Zealand plants: influence of chemical and physical traits

Ratites (ostriches Struthio camelus) and ungulates (red deer Cervus elaphus scoticus and goats Capra hircus) were presented with 14 indigenous shrubs in cafeteria-style trials. The shrubs represented the spectrum of woody plant architecture, ranging from broadleaved monopodial species through to small-leaved highly branched divaricates. Trials were stopped when almost all shoots of the plant expected to be most preferred had been consumed. There were considerable differences between the herbivores in their selection of certain plant species. Aristotelia fruticosa was avoided by deer, neutrally selected by goats, and preferred by ostriches. All herbivores strongly avoided two species, Pseudopanax crassifolius and Coprosma rugosa. Analysis of relative offtake (proportion of biomass consumed from each species, relative to total biomass consumed) showed that all three herbivores ate less of species with small leaves. Consumption by all herbivores was reduced by structural plant traits (i.e. divarication and related attributes) more than by chemical plant traits. The immediate impact of browsing on the plants, measured as the rank of proportion of shoots eaten, was broadly similar across the herbivores. The broadleaved species (e.g. Myrsine australis, Aristotelia serrata) experienced the greatest browsing, while divaricate (e.g. Coprosma rugosa) and conifer species (e.g. Podocarpus hallii) were generally least browsed. Although cafeteria-style experiments have limitations, our results for deer broadly correspond to those of field-based diet preference studies.     

Herbivory and the evolution of divaricate plants: Structural defences lost on an offshore island

Many island plants are characterized by unique morphology. For example, the high branching angles and small leaves of divaricate plants are a common feature of the New Zealand flora. The divaricate growth form may be an adaptation to deter browsing by extinct avian herbivores (moa); alternatively aspects of the insular climate may be responsible. However, our understanding of the selective pressures responsible for the high branching angles and small leaves of divaricate plants is incomplete. Here, I tested for differences in traits associated with the divaricate growth form between plants from Chatham Island and the New Zealand mainland. Moa never reached the Chatham Islands and its flora is derived from plants on mainland New Zealand. Therefore, I predicted Chatham Island plants to have lost morphological adaptations that may have deterred moa herbivory. Traits were quantified on 316 individuals in the field, allowing for 12 island‐mainland taxonomic comparisons. Chatham Island plants consistently produced smaller branching angles, larger leaves, shorter internodes and larger stems than related mainland plants. Results are therefore consistent with the hypothesis that selection for small leaves and high angled branching may be relaxed on the Chatham Islands due to an absence of moa. Smaller branching angles and larger leaves may offer a competitive advantage to Chatham Island plants.     

Is New Zealand vegetation really ‘problematic’? Dansereau's puzzles revisited

Over four decades ago, Pierre Dansereau, the noted North American ecologist, proposed six features of New Zealand vegetation as being problematic or unusual in a global context. We examine his propositions in the light of current ecological knowledge to determine whether or not these can still be considered unusual characteristics of New Zealand vegetation. (1) ‘Climatic change is still progressing’ resulting in disequilibrium between species' distributions and the present climate. New data and methods of analysis now available have removed the impression that Dansereau gained of imprecise zonation, unclear vegetation/climate relations and missing vegetation types. Communities cited as having regeneration failure can now be seen as even‐aged stands that developed after major disturbance, although there are other, also non‐climatic, explanations. However, the cause of the Westland ‘Nothofagus gap’ has become more, rather than less, controversial. (2) ‘Continuity of community composition defies classification’ and ‘Very few New Zealand associations have faithful species' are correct observations, but perhaps equally true of vegetation elsewhere. Dansereau's assertion of low species richness in New Zealand is not supported by the comparative data available. (3) ‘Lack of intolerant [i.e. mid‐seral] trees …’ is not evident with newer information. The order of species in succession, seen as unclear by Dansereau, has been determined by a range of approaches, largely confirming each other. (4) ‘Discrepancies of form and function …’ in divaricate shrubs and widespread heteroblasty are still controversial, with many more explanations. Several abiotic explanations have failed to stand up to investigation. Explanations in terms of herbivory have been well supported, although the extinction of the large avian herbivores makes certainty impossible. (5) ‘Incidence of hybridization …’ remains problematic. We do not know whether the incidence is unusually high, as Dansereau alleged, but the limited comparative data available suggest not. (6) The ‘overwhelming … competing power of exotics' is strongly context dependent. They are prominent in many non‐forest habitats. It seems that they are drivers of the vegetation change in some habitats, yet passengers after disturbance in others. Invasions can be slow, and may still be very incomplete in some ecosystem types. Whether exotics will eventually take over in most communities, or whether the native species will ‘laugh them to scorn’ as Cockayne suggested, only time will tell. In conclusion, some aspects of New Zealand's vegetation seem less unusual with increased knowledge, but others remain ‘problems’.     

Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

Secondary forest succession differs through naturalised gorse and native kānuka near Wellington and Nelson

The dominant native woody species forming early-successional vegetation on formerly forested sites in lowland New Zealand were kānuka (Kunzea ericoides) and mānuka (Leptospermum scoparium) (Myrtaceae). These have been replaced extensively by gorse (Ulex europaeus, Fabaceae), a naturalised species in New Zealand. Because gorse typically gives way to native broadleaved (angiosperm) forest in about 30 years, it is often considered desirable for facilitating native forest restoration. We tested three hypotheses, derived from the New Zealand literature, on gorse and kānuka: (1) kānuka stands have a different species composition and greater species richness than gorse stands at comparable successional stages; (2) differences between gorse and kānuka stands do not lessen over time; and (3) several native plant taxa are absent from or less common in gorse than in kānuka stands. We sampled 48 scrub or low-forest sites in two regions, Wellington and Nelson. Sites were classified into one of four predefined categories – young gorse, young kānuka, old gorse, old kānuka – based on canopy height of the succession and the dominant early-successional woody species. Few characteristics of the sites and surrounding landscapes differed significantly among site categories, and none consistently across regions. The vegetation composition of gorse and kānuka and their immediate successors differed in both regions, mainly in native woody species. Species richness was often lower in gorse and there were fewer smallleaved shrubs and orchids in gorse. Persistent differences at the older sites suggest the successional trajectories will not converge in the immediate future; gorse leads to different forest from that developed through kānuka. Gorse-dominated succession is therefore not a direct substitute for native successions. We suggest areas of early native succession should be preserved, and initiated in landscapes where successions are dominated by gorse or other naturalised shrubs.     

Modelling relationships between environment and canopy composition in secondary vegetation in central North Island, New Zealand

Relationships between composition of secondary vegetation and environment were studied in central North Island, New Zealand. A classification procedure was used to identify broad compositional groups which included forest, broadleaved scrub, shrub-fernland, sclerophyllous scrub and shrubland, and tussock-shrubland. Generalised additive models (GAMs) were used to examine relationships between species' distributions and mean annual temperature and rainfall, stand age, distance from intact forest, slope, topography, and drainage. There were marked differences in the environmental relationships of individual species. We conclude that temperature and rainfall have a dominant role in determining succession after disturbance at a regional scale, but distance from intact forest, topography, slope and solar radiation, become important at local scales. Variation unaccounted for by these environmental factors is most likely linked to historical factors such as variation in disturbance and/or grazing and browsing regimes. Intervention by managers will probably be required in the future if the current diversity of secondary vegetation in central North Island is to be maintained.     

Invasibility of native habitats by Argentine ants,

The Argentine ant, Linepithema humile, was found established in New Zealand in 1990. During summer 2001/2002 the spread of Argentine ants from urban environments into native habitats was investigated. During an initial large-scale survey around the northern cities of Auckland and Whangarei, Argentine ants were observed at 35 of 211 sites. Eight sites in Auckland were subsequently surveyed in greater detail to determine the extent of movement by Argentine ants into native habitats. The presence of Argentine ants was determined every 10 m along a total of 28 transects into native forest, scrub and mangrove habitats. Argentine ants moved up to 20 m into forest habitats. In habitats with more open canopy (mangrove and scrub), ants moved at least 30 m and 60 m, respectively. We predict that open habitats and relatively open canopy scrub environments in northern New Zealand are likely to be vulnerable to invasion, and to experience the highest densities and the greatest impacts of Argentine ants. Our preliminary data, coupled with data from other parts of the world suggests that intact indigenous forest in New Zealand will probably not be invaded. Indigenous forests are likely to have Argentine ants only at the boundary with open habitat, but in highly fragmented landscapes the impact could be significant.     

Areas of endemism of the Patagonian steppe: an approach based on insect distributional patterns using endemicity analysis

Aims  To delimit areas of endemism in the Patagonian steppe using endemicity analysis (EA), which evaluates areas of endemism by means of an endemicity index, and to compare the resulting endemic areas with those proposed for the Patagonian steppe by previous authors.
Location  The Patagonian steppe, a region of South America found approximately below parallel 36° S to the east of the Andes Mountains.
Methods  Distributional data for 159 species of insects collected in the Patagonian steppe, and consisting of 1317 georeferenced samples were used to identify areas of endemism. A data grid of presence and absence (with cells of 1° × 1°) was constructed. Initially, two different types of EA were performed, seeking areas defined by 'four or more' species. A first analysis was performed without taking into consideration those quadrats where no species had been recorded (empty quadrats), which in many cases meant a discontinuous distribution. The second analysis was performed assuming a continuous distribution for each species. A third analysis, assuming continuous distributions, was performed using 'three or more' as the number of species necessary for an area to be identified as an endemic area.
Results  In the first two analyses, EA recognized the same five areas of endemism: western Patagonia, south-western Payunia, northern Suabandean, southern Subandean and Austral Patagonia. The results of the third analysis allowed the identification of three more areas of endemism: northern Payunia, Chubutian and Santacrucian.
Main conclusions  We identified five areas of endemism for the Patagonian steppe, some of which have been defined in previous contributions. These areas are: Western Patagonia, Payunia and Subandean Patagonia (which can be divided into septentrional and meridional), Central Patagonia (Chubutence and Santacrucense) and Austral Patagonia.     

Effects of red deer on tree regeneration and growth in Aorangi Forest, Wairarapa

New Zealand forests have been substantially modified by introduced red deer over the past century. New Zealand’s indigenous forest managers need to know if regeneration of palatable tree species can be restored following control or eradication of browsing ungulates. Aorangi Forest, Wairarapa, suffered dramatic changes in forest understorey composition by the 1950s after more than seven decades of colonisation by red deer (Cervus elaphus), feral goats (Capra hircus) and pigs (Sus scrofa). Since then feral goats have been eradicated, but red deer and pigs persist throughout Aorangi Forest despite ongoing recreational hunting. This study uses data from paired fenced and unfenced plots established at seven sites in Aorangi Forest between 1981 and 1987, and re-measured in 2004, to show the effects of ungulates on tree (≥ 2 cm diameter at breast height) regeneration and growth. Our results show that browsing by red deer has prevented regeneration of kanono (Coprosma grandifolia), a highly palatable, fast-growing sub-canopy hardwood tree. Deer reduced the growth of mahoe (Melicytus ramiflorus) trees, probably by directly browsing epicormic shoots. The regeneration of other less palatable sub-canopy trees (e.g. porokaiwhiri, Hedycarya arborea), and slower-growing canopy species (e.g. hinau, Elaeocarpus dentatus and rewarewa, Knightia excelsa) appears to have been unaffected by deer browsing. Accordingly, tree species composition does not appear to have been greatly affected by browsing from deer populations in Aorangi Forest over the past two decades.     

Seasonal fluctuations in butterflies and nectar resources in a semi-natural grassland near Mt. Fuji, central Japan

We investigated seasonal fluctuation patterns in species and individuals of adult butterflies and flowering plants providing nectar in a semi-natural grassland in central Japan. We considered their interrelationships and implications for conservation. The semi-natural grassland included different vegetation structures and management regimes, including: (1) firebreaks where the grass was mowed and removed, (2) plantation areas that were mowed, (3) unpaved roads with mowed banks, (4) abandoned grassland, (5) scattered scrub forest, and (6) the surrounding forest. The sites with management (e.g., firebreaks), plantations and banks of the unpaved road sustained a larger number of butterflies and flowers than sites without management, such as the abandoned grassland, scrub forest and surrounding forest. The number of butterflies increased in the firebreak in June and at all sites in August and September. The firebreak sustained flowers in the spring, and the plantation area and banks of the unpaved road sustained flowers primarily in August and September, which was correlated with the distribution of butterflies. The different treatments such as mowing or mowing with removal of grass induced different numbers of flowers of each species affecting the habitat of adult butterflies through a season. On the other hand, the shrub tree species composing the scrub forest were host plants for the larvae of certain butterfly species. Our results suggest that heterogeneous environments with different human management or different vegetation structure or both could support habitat for various butterfly species, depending on the season and the seral stage.     

Dendroecological analysis of defoliator outbreaks on Nothofagus pumilio and their relation to climate variability in the Patagonian Andes

In the temperate forests of the southern Andes, Nothofagus pumilio, the dominant species of the most extensive forest type, experiences severe defoliation caused by caterpillars of the Ormiscodes genus (Lepidoptera: Saturniidae). This study uses tree rings to reconstruct the history of Ormiscodes outbreaks for the 1850–2005 period and examines relationships between outbreaks and climate variability. We used local climate records to compare outbreak–climate relationships in the northern Patagonian Andes (c. 41°S) and the cooler southern Patagonian Andes (c. 49°S). We also examined relationships between outbreak events and regional climate variability driven by variability in the Southern Annular Mode (SAM) and the El Niño‐Southern Oscillation. Although relationships between Ormiscodes outbreaks and climate proved to be complex, in northern Patagonia defoliation events are associated with drier and warmer than average growing seasons. Warming and drying trends in Patagonia during the latter part of the 20th century have been linked to a positive trend in SAM. During the post‐1976 period of accelerated warming in Patagonia, widespread defoliation outbreaks have occurred in both northern and southern Patagonia but the increase in frequency of events has been greater in the south. In southern Patagonia the increases in frequency of outbreaks in the late 20th century appear to be unprecedented over the c. 150 year tree‐ring record of reconstructed outbreaks. These results are consistent with the greater magnitude of recent warming in southern Patagonia, and suggest that under predicted warmer and drier climates in the 21st century, defoliator outbreaks may continue to increase in frequency. This study is the first systematic reconstruction of past insect outbreaks in South America and provides a preliminary understanding of how climate variability affects defoliator outbreaks in Patagonian Nothofagus forests.     

Evaluating tree wētā (Orthoptera: Anostostomatidae: <Emphasis Type="Italic">Hemideina</Emphasis> species) as bioindicators for New Zealand national biodiversity monitoring

Indicator taxa are increasingly being used to evaluate the natural environment because they provide both quantified and simplified information about complex phenomena and because they result in huge cost savings compared with monitoring entire biotas. In this paper, we examine the suitability of an iconic New Zealand invertebrate, the tree wētā (Orthoptera: Anostostomatidae: Hemideina species), as a bioindicator for invertebrates under a national biodiversity monitoring scheme in New Zealand. Tree wētā are common and widespread in New Zealand, comprising a distinctive component of the native invertebrate fauna, being large-bodied (up to 40 mm in length), relatively long-lived, flightless, and nocturnal. Arboreal tree wētā species are commonly monitored in conservation areas containing scrub or forest, particularly after mammal control, because they can be easily monitored using artificial roosts without harming them and they are readily identified by field workers. We evaluated whether data supported the use of tree wētā as a range of bioindicators for such monitoring and conclude that the arboreal species are good indicators for monitoring the effects of controlling the abundance of insectivorous mammals and that they are likely to be reliable population indicators of taxa sensitive to mammalian predation pressure, especially by rodents. However, it is unlikely that arboreal tree wētā are useful population indicators of habitat change (e.g. degradation and fragmentation) as they commonly survive in exotic vegetation and urban gardens throughout New Zealand. Although poorly studied for indicator value, tree wētā may not be good biodiversity indicators although there are insufficient data to establish this. We recommend further research be undertaken to develop standardised methods for monitoring so that conservation managers and researchers produce results that are consistent and comparable across different locations.     

Browsing‐mediated shrub canopy changes drive composition and species richness in forest‐tundra ecosystems

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long‐term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low‐productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming‐driven shrub expansion and hence maintains species richness.     

浙江天童国家森林公园常绿阔叶林主要组成种的种群结构及更新类型

 以浙江天童国家森林公园常绿阔叶林为对象,应用永久样方法和每木调查法调查了群落种类组成和结构,并对主要组成种的种群结构进行了分析。依据胸径级频率分布的形状,将各树种的种群结构归纳为5种类型,并结合生物学、生态学特征,分别讨论了其更新类型,同时对调查群落所处的演替阶段进行了诊断。结果表明：1）群落为6种共优势群落,出现木本植物69种。2）种群结构为单峰型的马尾松（Pinus massoniana）、枫香（Liquidambar formosana）和檫木（Sassafras tzumu）等为阳性高大乔木,无正常更新能力,更新类型是先锋群落优势种或是顶极性先锋种;间歇型的木荷（Schima superba）、黑山山矾（Symplocos heishanensis）和红楠（Machilus thunbergii）等为具有耐荫性的常绿阔叶树种,更新具有波动性和机会性,是演替系列或顶极群落优势种;逆-J字型的米槠（Castanopsis carlesii）、栲树（Castanopsis fargesii）和长叶石栎（Lithocarpus harlandii）等为强耐荫性的常绿阔叶树种,通过幼苗库和根萌生枝进行更新,是顶极群落的优势种;浙江新木姜子（Neolitsea aurata var. chekiangensis）、厚皮香（Ternstroemia gymnanthera）等为L字型;连蕊茶（Camellia fraternna）、老鼠矢（Symplocos stellaris）等为单柱型,此二型是群落亚乔木层和灌木层的主要组成种。3）调查群落已进入演替的顶极阶段,但仍处于其前期,将进一步向以米槠、栲树为主要优势种的方向发展。     

Major vegetation trends in the Tertiary of Patagonia (Argentina): A qualitative paleoclimatic approach based on palynological evidence

The patterns of Patagonian vegetation change suggest a strong relationship between the major thermal characters of the flora and the global paleoclimatic trends during Tertiary times. This conclusion was reached from the assessment of fossil pollen data from Patagonia throughout the Paleogene and Early Neogene periods and the subsequent comparison of palynological data to the global deep-sea oxygen isotope record. Four main time intervals were recognized based on the temporal distribution of selected angiosperm key taxa. (1) Paleocene to Early Eocene: presence of megatherm elements (e.g. Nypa, Pandanus), probably integrating mangrove communities in Patagonian lowlands. (2) Middle Eocene to Early Oligocene: rise to dominance of mesotherm and microtherm Nothofagus species. Megatherm taxa were well recorded at the beginning of this interval (e.g. Ilex) but were shown to disappear towards the end. (3) Late Oligocene to Middle Miocene: new increases of megatherm taxa such as palms, Cupania and Alchornea. First occurrences of mesotherm Asteraceae, represented by trailing Mutisieae, were reported. (4) Late Miocene: dispersal of meso-microtherm and arid adapted taxa (e.g. Ephedraceae and Asteraceae) across the non-Andean region of Patagonia. Microtherm Nothofagacean forests probably occurred on the higher rainfall regions of western Patagonia. The current vegetation was most likely reached during this last stage with the forest development under wetter conditions on the Andean sectors, and the steppe throughout the non-Andean region of Patagonia.     

Plant structural defences against browsing birds: a legacy of New Zealand's extinct moas

Browsing by large vertebrates has been a major force in the evolution of terrestrial plants but Holocene extinctions of the browsers have left a legacy of broken biotic partnerships. Ratite birds were the largest herbivores in several regions, such as the moas of New Zealand. Many woody plants there have a distinct form of branching, described as "divaricate", with thin, wide angled, branches intertwining to form a tangled canopy. Divaricate branching has been interpreted as a form of protection against climate extremes or as an anachronistic defense against the extinct moas. Here we report the first experimental evidence that many of these plants are defended against extant ratite browsers. In feeding experiments on two tree species with different (heteroblastic) juvenile and adult branch morphology, emus and ostriches obtained adequate feeding rates from adult shoots but sub-maintenance feeding rates from juvenile shoots with the ratite-resistant traits. Divaricate juvenile shoots suffered 30–70% less biomass removal to the birds than adult shoots. Ratites browse by a distinctive clamping and tugging action. Structural defence traits that exploit the limitations of this feeding mode include narrow, strong, elastic branches that resist being torn off, wide branching angle ("divaricate") that makes shoots difficult to swallow, and small, widely spaced leaves. This novel plant architectural defence has developed in at least 20% of the native woody flora of New Zealand, including 10 heteroblastic tree species that exhibit the ratite-resistant strategy until they reach ca 3 metres height. It is also a major axis of variation amongst homoblastic woody shrub species. The defences are useless against mammalian browsers that shear shoots, contributing to marked decreases in the abundances of ratite-resistant species in New Zealand after the introduction of mammals.     

Nitrogen concentration and mimicry in some New Zealand mistletoes

Summary The nitrogen concentration in photosynthetic organs of 41 pairs of mistletoes and their respective hosts was compared and found to be higher (in relation to that of their hosts) in cryptic mistletoes and lower in non-cryptic mistletoes. This supports the hypothesis that crypsis may have evolved as a mechanism for avoiding predation by vertebrate herbivores. There were no consistent differences between the nitrogen concentrations of hosts and mistletoes in trees and larger-leaved shrubs (whether native or introduced). The largest differences between mistletoes and their hosts were found on small-leaved divaricate shrubs (Coprosma ssp., Melicope simplex) where the nitrogen concentrations in cryptic mistletoes (Korthalsella spp.) were significantly higher than in their hosts, but significantly lower in prominent mistletoes (Loranthus micranthus, Tupeia antarctica) which apparently advertise their unpalatability. If crypsis in New Zealand mistletoes evolved as a protection against herbivory, then it must have evolved in the absence of mammalian herbivores, and the appropriate selection pressures could have been provided only by moas, extinct ratite birds. Otherwise, alternative explanations, such as differing relationships between water use and nitrogen uptake, must be sought for the observed associations of nitrogen concentration and cryptic mimicry in New Zealand mistletoes.     

Effects of Mycorrhizae and Nontarget Organisms on Restoration of a Seasonal Tropical Forest in Quintana Roo, Mexico: Factors Limiting Tree Establishment

We initiated a study of the effects of mycorrhizal fungal community composition on the restoration of tropical dry seasonal forest trees. Tree seedlings were planted in a severely burned experimental site (1995 fire) during the growing season of 1998 at the El Edén Ecological Reserve, in north Quintana Roo, Mexico. Seedlings of Leucaena leucocephala, Guazuma ulmifolia, Caesalpinia violacea, Piscidia piscipula, Gliricidia sepium, and Cochlospermum vitifolium were germinated in steam‐sterilized soil and either remained uninoculated (nonmycorrhizal at transplanting) or were inoculated with mycorrhizal fungi in soils from early‐seral (recently burned) or late‐seral (mature forest) inoculum. Inoculum from the early‐seral soil was largely Glomus spp., whereas a diverse community of arbuscular mycorrhizal fungi were reintroduced from the mature forest including species of Scutellospora, Gigaspora, Glomus, Sclerocystis, and Acaulospora. Plants grew better when associated with the mature forest inoculum, unlike a previous experiment in which plants grew taller with the early‐seral inoculum. Reasons for the different responses include a less‐intense burn resulting in more residual organic matter. In addition to mycorrhizal responses, plants were severely affected by deer browsing. One tree species, C. vitifolium found in the region but not in the reserve, was eliminated by a resident fungal facultative pathogen. Several practical conclusions for restoration can be made. The common nursery practice of soil sterilization may be detrimental because it eliminates beneficial mycorrhizal fungi; species not native to the site may not survive because they may not be adapted to the local pathogens; and herbivory can be severe depending on the landscape context of the restoration.     

The Seedling Ratio Method for determining ungulate impacts on forest understoreys: Utility in an Australian ecosystem

The Seedling Ratio Method was devised to assess the impacts of introduced ungulates on plant species richness in forest understoreys. The method has successfully assessed ungulate impacts on species richness in forests in New Zealand and Hawaii, which do not have native herbivores. We tested the three critical assumptions that underlie this method to investigate its potential for use in an Australian ecosystem. This study was conducted in the Yarra Ranges National Park (YRNP), Victoria, which had a high‐density population of an introduced deer species, Sambar (Cervus unicolor), in addition to several native herbivore species. One of the three key assumptions of the Seedling Ratio Method was not supported, whereas conventional differential exclosures showed clear and separate impacts by Sambar and native herbivores. We conclude that the Seedling Ratio Method could not provide a clear indication of browsing impact on forest understoreys in YRNP.