The dynamic regeneration niche of a forest following a rare disturbance event

Aim: Knowledge of how climate and fire regimes affect regeneration in foundation species is critical to the conservation of entire ecosystems. Different stages of regeneration often require different ecological conditions, but dynamic constraints on regeneration are poorly known for species that regenerate only after infrequent wildfires. Focussing on a long-lived, foundation tree species (Eucalyptus regnans), we tested the hypothesis that the relative importance of fire regime variables (fire severity and time since previous fire) and environmental gradients on post-fire regeneration would shift as seedlings developed. Location: South-eastern Australia. Methods: Following a large (> 59,000 ha) summer wildfire in 2009, we sampled 131 sites (61 burnt) annually for four years (2009-2012), representing the range of environmental conditions in which E. regnans occurs. We analysed the effect of fire severity, time since fire and environmental variables on early regeneration processes critical for post-fire species distributions: seedling establishment, seedling density and growth through different height stages (10 cm, 25 cm, 50 cm and 200 cm). Results: The regeneration niche of E. regnans was defined by different factors at different stages of development. Initially, seedlings established prolifically on burnt sites, regardless of severity. Three years into the regeneration process, high-severity fire became the dominant driver of seedling persistence and growth over 25 cm. Growth over 50 cm was dependent on environmental conditions relating to elevation and precipitation. Main conclusions: Our results describe how fire occurrence, fire severity and environmental gradients affected seedling establishment, persistence and growth. The dynamic constraints on regeneration likely reflect temporal changes in the biotic and abiotic environment and variation in resource requirements during the early post-fire years. Our findings will enable more accurate forecasts of species distributions to assist forest conservation in the face of global changes in climate and fire regimes.

Epiphytes on Nothofagus cunninghamii and Eucalyptus regnans in a Victorian cool temperate rainforest

This study investigated the epiphytic communities on Myrtle Beech Nothofagus cunninghamii (Hook.) Oerst. and Mountain Ash Eucalyptus regnans F.Muell. trees in a pocket of Cool Temperate Rainforest in the Yarra Ranges National Park, Victoria, Australia. Twenty species were identified growing on N. cunninghamii, with nine species found on E. regnans. The dominant epiphytes were the moss Dicranoloma menziesii on N. cunninghamii, and the liverwort Bazzania adnexa var. adnexa on E. regnans.

The influence of time since fire and distance from fire boundary on the distribution and abundance of arboreal marsupials in Eucalyptus regnans-dominated forest in the Central Highlands of Victoria

The impact of time since fire after two consecutive wildfires 44 years apart (1939 and 1983) within the same area, and the distance from the fire boundary «100 m or 500-2000 m), were investigated in relation to the distribution and abundance of arboreal marsupials in 1994. Arboreal marsupials were censused by stagwatching and spotlighting in two relatively young age classes of mountain ash (Eucalyptus regnans) dominated forest in the Central Highlands of Victoria. Five species of arboreal marsupial were detected, but only three were detected in sufficient numbers to determine habitat preferences. Petauroides volans (greater glider) was statistically more abundant in 1939 regrowth forests, while Trichosurus caninus (mountain brushtail possum) showed no significant preference for either age class of forest. All but one record of Gymnobelideus leadbeateri (Leadbeater's possum) came from young forest, though the effect of age-class was not statistically significant. Distance from fire boundary explained little or no variation in mammal distribution or abundance. While the actual number of hollow-bearing trees was similar in both age classes of forest, the long-term lifespan of hollow-bearing trees in more recently burnt forest is predicted to be lower than in unburnt or not recently burnt forest. Post-fire salvage logging following the 1983 wildfires appears to have reduced the number of hollow-bearing trees at sites burnt in 1983.

A review of the conservation ecology of round-leaf Pomaderris Pomaderris vacciniifolia F. Muell. ex Reissek (Rhamnaceae)

Round-leaf Pomaderris Pomaderris vacciniifolia F. Muell. ex Reissek (Rhamnaceae) is a Victorian endemic shrub listed as threatened under the Flora and Fauna Guarantee Act 1988 and critically endangeredunder the Environment Protection and Biodiversity Conservation Act 1999. A review of the available literature for P. vacciniifolia indicated most information is anecdotal or found in unpublished works. Better understanding of the ecology of P. vacciniifolia may help explain why it is vulnerable, and enhance future management. Future research should focus towards better understanding of P. vacciniifolia habitat, reproductive ecology, seed dispersal mechanisms and competitive ability and how these compare with more common sympatric congeners, to determine whether any differences could explain the relative success of these species. Targeted searches for this species on public and private land are warranted to reveal additional populations and fully appreciate the distribution of this species. (The Victorian Naturalist 131 (2) 2014, 44-51).

Regulation of HbPIP2;3, a latex-abundant water transporter, is associated with latex dilution and yield in the rubber tree (Hevea brasiliensis Muell. Arg.)

Rubber tree (Hevea brasiliensis) latex, the source of natural rubber, is synthesised in the cytoplasm of laticifers. Efficient water inflow into laticifers is crucial for latex flow and production since it is the determinant of the total solid content of latex and its fluidity after tapping. As the mature laticifer vessel rings are devoid of plasmodesmata, water exchange between laticifers and surrounding cells is believed to be governed by plasma membrane intrinsic proteins (PIPs). To identify the most important PIP aquaporin in the water balance of laticifers, the transcriptional profiles of ten-latex-expressed PIPs were analysed. One of the most abundant transcripts, designated HbPIP2;3, was characterised in this study. When tested in Xenopus laevis oocytes HbPIP2;3 showed a high efficiency in increasing plasmalemma water conductance. Expression analysis indicated that the HbPIP2;3 gene was preferentially expressed in latex, and the transcripts were up-regulated by both wounding and exogenously applied Ethrel (a commonly-used ethylene releaser). Although regular tapping up-regulated the expression of HbPIP2;3 during the first few tappings of the virginal rubber trees, the transcriptional kinetics of HbPIP2;3 to Ethrel stimulation in the regularly tapped tree exhibited a similar pattern to that of the previously reported HbPIP2;1 in the virginal rubber trees. Furthermore, the mRNA level of HbPIP2;3 was associated with clonal yield potential and the Ethrel stimulation response. Together, these results have revealed the central regulatory role of HbPIP2;3 in laticifer water balance and ethylene stimulation of latex production in Hevea.

Lichen flora of the Mount Donna Buang Scenic Reserve, Victoria

This report investigated the lichen flora of the Mt Donna Buang Scenic Reserve in Victoria, There were several aims: to describe the lichens of the region, to produce a pictorial key enabling field identification and to determine any distribution patterns. A floristic survey covering approximately 50 square km was undertaken to determine lichen diversity of the region generally. Lichens were sampled along roads, tracks, walking trails and in sections of bush, taking into account forest type and, particularly, areas that were lichen rich. Seventy-five lichen species in 43 genera and 27 families were identified and described from the region. An unknown, species H, also was described. Of the 76 lichen species, 22 were crastose and the remainder macrolichens. The best represented families were: Cladoniaceae (8 species), Hypogymniaceae (6), Lobariaceae (7), Lecideaceae (6), Pannariaceae (6) and Parmeliaccae (6). This study described 12 species (17%) which previously were not known for Victoria and which are a first record for the state. These include: Cladonia sarmentosa (J.D. Hook & Taylor) Dodge, Graphis librata Knight, Parmelinopsis neodamaziana (Elix & Johnston) Elix & Hale, Pertusaria novaezelandiae Szatala, Placopsis pardlina f. microphylla Lamb, Porina leptalea AX. Sm., Pseudocyphellaria ardesiaca Galloway, Trapeliopsis congregant (Zahlbr.) Brako, Menegazzia myriotrema (Mull. Arg.) P. James, Bunodophoron scrobiculatum (Church. Bab,) Wedin, Parmelia testacea Stirton and Menegazzia purpurascens S. Louwhoff sp. nov.. The last eight species are new to the mainland and, apart from Menegazzia purpurascens, previously were known only from Tasmania. Five main elements of distribution were identified for the lichen flora of the Mt Donna Buang Scenic Reserve: cosmopolitan, austral/australasian, paleotropical, pantropical and western pacific. The majority of species (68%) had austral/australasian distributions, eleven (16%) were endemic to Australia and nine (13%) occurred only in Tasmania , Victoria and New Zealand. A pictorial, dichotomous key was constructed for the lichen flora of the Mt Donna Buang Scenic Reserve. Previously, keys to the lichen flora of Tasmanian rainforests were suggested as appropriate to similar areas in Victoria, however, the Victorian forests include a significant sclerophyll element The key presented is specific for the study site but is appropriate to similar regions in Victoria and has been tested in a number of these areas. The key was designed to be ‘user-friendly’ so that the experienced and inexperienced alike are able to use it. A more detailed investigation of the lichen flora of the Mt Donna Buang Scenic Reserve was carried out in order to determine distribution. A total of 50 quadrats, each 20m x 20m in size, were sampled. Within each, the dominant vegetation type was determined and individuals were identified and location noted. The cover abundance of each lichen species on each individual tree was estimated using a modified Braun-Blanquet scale. A total of 710 trees, representing 13 different species, were examined. Nothofagus cunninghamii (Hook.) Oerst, Eucalyptus regnans R Mull., Acacia dealbata Link, A. melanoxylon R. Br., Hedycarya angustifolia A. Cunn. and Atherosperma moschatum Labill. were the six most common tree species encountered at the study site. Nothofagus cunninghamii supported the greatest lichen diversity (39 species), although most species occurred on less than 10% of the trees. The majority of lichens occurring on N. cunninghamii A. melanoxylon, A. dealbata and H. angustifolia were foliose or crustose, those on Ł. regnans fruticose and foliose and those on A moschatum crustose. Bunodophoron australe was the only lichen species at the study site to occur on one host, Nothofagus cunninghamiL Many occurred on a number of different hosts, but were most common on one particular tree species. The distribution of lichens at the study site was analysed with a rnultivariate statistical package (PATN) which dealt with ‘pattern analysis’. The program ‘SSH’ in PATN which uses the Bray-Curtis ordination technique, was used to create scatterplots displaying the degree of dissimilarity between quadrats in terms of presence/absence of lichen species. The program ‘TWAY’ in PATN was used to construct a two way table to display which lichen species occurred in each vegetation type. The pattern analysis revealed that the lichens of the Mt Donna Buang Scenic Reserve were not restricted to any particular forest type, but particular lichens, or groups of lichens, tended to predominate in certain vegetation communities. This concurs with work done by others in Tasmanian forests. Quadrats which were situated in cool temperate rainforest were grouped more closely with each other than with quadrats in other vegetation types. These also supported the greatest number of lichen species. This was not surprising since N. cunninghamii the dominant tree species in cool temperate rainforest, supported the greatest lichen diversity.

Studies of the biology of four species of Dicranoloma

Populations of Dicranoloma billardierei (Brid) Par., D. dicarpum (Nees.) Par., D. menziesii (Tayl.) Par. and D. platycaulon (C. Muell) Dix, from two pockets of cool temperate rainforest within the Yarra Ranges National Park (Cement Creek and Myrtle Loop), were sampled for a period of two years to establish their reproductive biology. The population dynamics within quadrats of D. billardierei, D. menziesii and D. platycaulon at Cement Creek also was investigated over a two year period, through the seasonal recording of shoot loss and/or gain, The four species of Dicranoloma were dioicous and sexually dimorphic, with dwarf males epiphytic on the female plants. Antheridia were initiated before archegonia and required ca, 6 months to reach maturity, compared with 1 to 2 months for archegonia. More archegonia than antheridia occurred per inflorescence and were more variable Fertilization occurred during winter in D. billardierei and summer/ autumn in D. menziesii and D. platycaulon. The duration of the sporophyte cycle of D. menziesii was 12 months, shorter than that of D. billardierei and D. platycaulon which lasted for a period of 18 months to 2 years. In the latter two species an overlap of sporophyte generations occurred. This was particularly pronounced in D. billardierei as sporophytes remained in the swollen venter maturation stage for a period of 6 months. The duration of the sporophyte cycle could not be ascertained as few fruiting stems of D. dicarpum were found. All four species of Dicranoloma regenerated from fragments cultured in the laboratory, and only two of the species showed evidence of production of asexual propagules in the field. Dicranoloma dicarpum was found to produce gemmae, an observation which had not been recorded before, and most of the leaves on stems of D. platycaulon had detached subulas. Shoot loss was minimal in all four species, and when it did occur, (eg D. billardierei) it was attributed to disturbance by animals. Within quadrats there was an increase in shoot density which resulted from the development of innovation(s) and/or side branches rather than from the recruitment of new plants from spores or the regeneration of asexual propagules. The four species of Dicranoloma investigated were robust, perennial mosses and formed an important component of the bryophytes found within the study area. Dicranoloma menziesii was the predominant species establishing on a variety of substrata, particularly as an epiphyte on Nothofagus cunninghamii The other species were more selective in their choice of substratum. Dicranoloma platycaulon was found exclusively on the trunks of myrtle beech and D. billardierei on fallen logs and exposed roots. Dicranoloma dicarpum which was not common, grew as an epiphyte on myrtle beech and on rocks.

Ageing mallee eucalypt vegetation after fire : insights for successional trajectories in semi-arid mallee ecosystems

A critical requirement in the ecological management of fire is knowledge of the age-class distribution of the vegetation. Such knowledge is important because it underpins the distribution of ecological features important to plants and animals including retreat sites, food sources and foraging microhabitats. However, in many regions, knowledge of the age-class distribution of vegetation is severely constrained by the limited data available on fire history. Much fire-history mapping is restricted to post-1972 fires, following satellite imagery becoming widely available. To investigate fire history in the semi-arid Murray Mallee region in southern Australia, we developed regression models for six species of mallee eucalypt (Eucalyptus oleosa F.Muell. ex. Miq. subsp. oleosa, E. leptophylla F.Muell. ex. Miq., E. dumosa J. Oxley, E. costata subsp. murrayana L. A. S. Johnson & K. D. Hill, E. gracilis F.Muell. and E. socialis F.Muell. ex. Miq.) to quantify the relationship between mean stem diameter and stem age (indicated by fire-year) at sites of known time since fire. We then used these models to predict mean stem age, and thus infer fire-year, for sites where the time since fire was not known. Validation of the models with independent data revealed a highly significant correlation between the actual and predicted time since fire (r = 0.71, P < 0.001, n = 88), confirming the utility of this method for ageing stands of mallee eucalypt vegetation. Validation data suggest the models provide a conservative estimate of the age of a site (i.e. they may under-estimate the minimum age of sites >35 years since fire). Nevertheless, this approach enables examination of post-fire chronosequences in semi-arid mallee ecosystems to be extended from 35 years post-fire to over 100 years. The predicted ages identified for mallee stands imply a need for redefining what is meant by ‘old-growth’ mallee, and challenges current perceptions of an over-abundance of ‘long-unburnt’ mallee vegetation. Given the strong influence of fire on semi-arid mallee vegetation, this approach offers the potential for a better understanding of long-term successional dynamics and the status of biota in an ecosystem that encompasses more than 250 000 km² of southern Australia.

The use of topographic fire refuges by the greater glider (Petauroides volans) and the mountain brushtail possum (Trichosurus cunninghami) following a landscape-scale fire

We examined the abundance of arboreal marsupials in topographic fire refuges after a major fire in a stand-replacing crown-fire forest ecosystem. We surveyed the abundance of arboreal marsupials across 48 sites in rainforest gullies burnt to differing extents by the 2009 fires in the mountain ash (Eucalyptus regnans) forests of the Victorian Central Highlands, Australia. The greater glider (Petauroides volans) was less abundant within the extent of the 2009 fire. The mountain brushtail possum (Trichosurus cunninghami) was more abundant within the extent of the 2009 fire, particularly within unburnt peninsulas protruding into burnt areas from unburnt edges. Our results indicate that fire refuges may facilitate the persistence of some species within extensively burnt landscapes. Additional work should seek to clarify this finding and identify the demographic mechanisms underlying this response.

Dominant drivers of seedling establishment in a fire-dependent obligate seeder: climate or fire regimes?

Climate change is causing fire regime shifts in ecosystems worldwide. Plant species with regeneration strategies strongly linked to a fire regime, such as obligate seeders, may be particularly threatened by these changes. It is unclear whether changes in fire regimes or the direct effects of climate change will be the dominant threats to obligate seeders in future. We investigated the relative importance of fire-related variables (fire return interval andfire severity) and environmental factors (climate and topography) on seedling establishment in the world's tallest angiosperm, an obligate seeder, Eucalyptus regnans. Throughout its range, this species dominates the wet montane forests of south-eastern Australia and plays a keystone role in forest structure. Following major wildfires, we investigated seedling establishment in E. regnanswithin 1 year of fire as this is a critical stage in the regeneration niche of obligate seeders. Seedling presence and abundance were strongly related to the occurrence of fire but not to variation in fire severity (moderate vs. high severity). Seedling abundance increased with increasing fire return interval (range 26-300 years). First-year seedling establishment was also strongly associated with low temperatures and with high elevations, high precipitation and persistent soil water availability. Our results show that both climate and fire regimes are strong drivers of E. regnans seedling establishment. The predicted warming and drying of the climate might reduce the regeneration potential for some obligate seeders in future and these threats are likely to be compounded by changes in fire regimes, particularly increases in fire frequency.

A succession of theories : purging redundancy from disturbance theory

The topics of succession and post-disturbance ecosystem recovery have a long and convoluted history. There is extensive redundancy within this body of theory, which has resulted in confusion, and the links among theories have not been adequately drawn. This review aims to distil the unique ideas from the array of theory related to ecosystem change in response to disturbance. This will help to reduce redundancy, and improve communication and understanding between researchers. We first outline the broad range of concepts that have developed over the past century to describe community change in response to disturbance. The body of work spans overlapping succession concepts presented by Clements in 1916, Egler in 1954, and Connell and Slatyer in 1977. Other theories describing community change include state and transition models, biological legacy theory, and the application of functional traits to predict responses to disturbance. Second, we identify areas of overlap of these theories, in addition to highlighting the conceptual and taxonomic limitations of each. In aligning each of these theories with one another, the limited scope and relative inflexibility of some theories becomes apparent, and redundancy becomes explicit. We identify a set of unique concepts to describe the range of mechanisms driving ecosystem responses to disturbance. We present a schematic model of our proposed synthesis which brings together the range of unique mechanisms that were identified in our review. The model describes five main mechanisms of transition away from a post-disturbance community: (i) pulse events with rapid state shifts; (ii) stochastic community drift; (iii) facilitation; (iv) competition; and (v) the influence of the initial composition of a post-disturbance community. In addition, stabilising processes such as biological legacies, inhibition or continuing disturbance may prevent a transition between community types. Integrating these six mechanisms with the functional trait approach is likely to improve the predictive capacity of disturbance theory. Finally, we complement our discussion of theory with a case study which emphasises that many post-disturbance theories apply simultaneously to the same ecosystem. Using the well-studied mountain ash (Eucalyptus regnans) forests of south-eastern Australia, we illustrate phenomena that align with six of the theories described in our model of rationalised disturbance theory. We encourage further work to improve our schematic model, increase coverage of disturbance-related theory, and to show how the model may link to, or integrate with, other domains of ecological theory.