Tree species diversity and ecosystem function: Can tropical multi-species plantations generate greater productivity?

Results from the humid tropics of Australia demonstrate that diverse plantations can achieve greater productivity than monocultures. We found that increases in both the observed species number and the effective species richness were significantly related to increased levels of productivity as measured by stand basal area or mean individual tree basal area. Four of five plantation species were more productive in mixtures with other species than in monocultures, offering on average, a 55% increase in mean tree basal area. A general linear model suggests that species richness had a significant effect on mean individual tree basal area when environmental variables were included in the model. As monoculture plantations are currently the preferred reforestation method throughout the tropics these results suggest that significant productivity and ecological gains could be made if multi-species plantations are more broadly pursued.

Growth and species interactions of Eucalyptus pellita in a mixed and monoculture plantation in the humid tropics of north Queensland

This study investigated whether mixed-species designs can increase the growth of a tropical eucalypt when compared to monocultures. Monocultures of Eucalyptus pellita (E) and Acacia peregrina (A) and mixtures in various proportions (75E:25A, 50E:50A, 25E:75A) were planted in a replacement series design on the Atherton Tablelands of north Queensland, Australia. High mortality in the establishment phase due to repeated damage by tropical cyclones altered the trial design. Effects of experimental designs on tree growth were estimated using a linear mixed-effects model with restricted maximum likelihood analysis (REML). Volume growth of individual eucalypt trees were positively affected by the presence of acacia trees at age 5 years and this effect generally increased with time up to age 10 years. However, the stand volume and basal area increased with increasing proportions of E. pellita, due to its larger individual tree size. Conventional analysis did not offer convincing support for mixed-species designs. Preliminary individual-based modelling using a modified Hegyi competition index offered a solution and an equation that indicates acacias have positive ecological interactions (facilitation or competitive reduction) and definitely do not cause competition like a eucalypt. These results suggest that significantly increased in growth rates could be achieved with mixed-species designs. This statistical methodology could enable a better understanding of species interactions in similarly altered experiments, or undesigned mixed-species plantations.

Developing African mahogany (Khaya senegalensis) germplasm and its management for a sustainable forest plantation industry in northern Australia: Progress and needs.

The demonstrated wide adaptability, substantial yield potential and proven timber quality of African mahogany (Khaya senegalensis) from plantings of the late 1960s and early 1970s in northern Australia have led to a resurgence of interest in this high-value species. New plantations or trials have been established in several regions since the early 1990s -in four regions in north Queensland, two in the Northern Territory and one in Western Australia. Overall, more than 1500 ha had been planted by early 2007, and the national annual planting from 2007-2008 as currently planned will exceed 2400 ha. Proceedings of two workshops have summarised information available on the species in northern Australia, and suggested research and development (R&D) needs and directions. After an unsustained first phase of domestication of K. senegalensis in the late 1960s to the early 1970s, a second phase began in northern Australia in 2001 focused on conservation and tree improvement that is expected to provide improved planting stock by 2010. Work on other aspects of domestication is also described in this paper: the current estate and plans for extension; site suitability, soils and nutrition; silviculture and management; productivity; pests and diseases; and log and wood properties of a sample of superior trees from two mature plantations of unselected material near Darwin. Some constraints on sustainable plantation development in all these fields are identified and R&D needs proposed. A sustained R&D effort will require a strategic coordinated approach, cooperative implementation and extra funding. Large gains in plantation profitability can be expected to flow from such inputs.

The original native pasture ecosystems of the eastern and western Darling Downs - Can they be restored?

The original pasture ecosystems of southern inland Queensland ranged from treeless grasslands on cracking clays through grassy woodlands of varying density on a great range of soil types to those competing at the dynamic edges of forests and scrubs. Fire, both wild and aboriginal-managed, was a major factor, along with rainfall extremes, in shaping the pastures and tree:grass balance. Seedling recruitment was driven by rainfall extremes, availability of germinable seed and growing space, with seed availability and space being linked to the timing and intensity of recent fires and rain. The impact of insects, diseases, severe wind and hailstorms on recruitment should not be underestimated. The more fertile soils had denser grass growth, greater fire frequency and thinner tree cover than infertile soils, except where trees were so dense that grass growth was almost eliminated. The pastures were dominated by perennial tussock grasses of mid-height but included a wide array of minor herbaceous species whose abundance was linked to soil type and recent seasonal conditions. Many were strongly perennial with Asteraceae, Fabaceae, Malvaceae, Cyperaceae and Goodeniaceae most common in an environment, which can experience effective rainfall at any time of year. The former grassland communities that are now productive farming lands are not easily returned to their original composition. However, conservation of remnant examples of original pasture types is very achievable provided tree density is controlled, prescribed burning and grazing are used and rigorous control of invasive, exotic species is undertaken. This should be done with a clear understanding that significant short-and medium-term fluctuations in botanical composition are normal.

Performance of seven hardwood species underplanted to Pinus elliottii in south-east Queensland

Seven hardwood species were tested as underplants under Pinus elliottii plantations on the coastal lowlands of south-east Queensland. The species tested were: Flindersia brayleyana (F. Muell) (Queensland maple), F. australis (R. Br.), (crow's ash), Swietenia macrophylla (King) (American mahogany), Grevillea robusta (A. cunn) (southern silky oak), Elaeocarpus grandis (F. Muell) (silver quandong), F. ifflaiana (F. Meull) (Cairns hickory) and Ceratopetalum apetalum (D. Don) (coachwood). Most species (except E. grandis) established successfully but slowly. Underplants suffered 9-16% mortality during thinning of the overstorey. By 2004 when aged c. 38 years, four underplanted species; F. brayleyana, S. macrophylla, F. ifflaiana and E. grandis, had attained predominant heights of 20 m and mean diameter at breast height of 25 cm or better. The presence of underplants increased total site productivity by up to 23% and did not have any detrimental effect on the development of the overwood.This experiment has demonstrated that some rainforest species will survive and grow healthily as underplants in exotic pine plantations plus produce small merchantable logs within a 38 year rotation. The results also indicated the importance of correct species selection if an underplanting option is to be pursued as some species have been a complete failure (notably G. robusta).

Growth and physiological response of six Australian rainforest tree species to a light gradient

An understanding of growth and photosynthetic potential of subtropical rainforest species to variations in light environment can be useful for determining the sequence of species introductions in rainforest restoration projects and mixed species plantations. We examined the growth and physiology of six Australian subtropical rainforest tree species in a greenhouse consisting of three artificial light environments (10%, 30%, and 60% full sunlight). Morphological responses followed the typical sun-shade dichotomy, with early and late secondary species (Elaeocarpus grandis, Flindersia brayleyana, Flindersia schottiana, and Gmelina leichhardtii) displaying higher relative growth rate (RGR) compared to mature stage species (Cryptocarya erythroxyion and Heritiera trifoliolatum). Growth and photosynthetic performance of most species reached a maximum in 30-60% full sunlight. Physiological responses provided limited evidence of a distinct dichotomy between early and late successional species. E. grandis and F brayleyana, provided a clear representation of early successional species, with marked increase in Am in high light and an ability to down regulate photosynthetic machinery in low light conditions. The remaining species (F. schottiana, G. leichhardtii, and H. trifoliolatum) were better represented as failing along a shade-tolerant continuum, with limited ability to adjust physiologically to an increase or decrease in light, maintaining similar A(max) across all light environments. Results show that most species belong to a shade-tolerant constituency, with an ability to grow and persist across a wide range of light environments. The species offer a wide range of potential planting scenarios and silvicultural options, with ample potential to achieve rapid canopy closure and rainforest restoration goals.

Modelling predicts positive and negative interactions between three Australian tropical tree species in monoculture and binary mixture

Computer modelling promises to be an important tool for analysing and predicting interactions between trees within mixed species forest plantations. This study explored the use of an individual-based mechanistic model as a predictive tool for designing mixed species plantations of Australian tropical trees. The `spatially explicit individually based-forest simulator' (SeXI-FS) modelling system was used to describe the spatial interaction of individual tree crowns within a binary mixed-species experiment. The three-dimensional model was developed and verified with field data from three forest tree species grown in tropical Australia. The model predicted the interactions within monocultures and binary mixtures of Flindersia brayleyana, Eucalyptus pellita and Elaeocarpus grandis, accounting for an average of 42% of the growth variation exhibited by species in different treatments. The model requires only structural dimensions and shade tolerance as species parameters. By modelling interactions in existing tree mixtures, the model predicted both increases and reductions in the growth of mixtures (up to +/-50% of stem volume at 7 years) compared to monocultures. This modelling approach may be useful for designing mixed tree plantations.

The suitability of non-target native mangroves for the survival and development of the lantana bug Aconophora compressa, an introduced weed biological control agent

Aconophora compressa (Hemiptera: Membracidae), a biological control agent introduced against the weed Lantana camara (Verbenaceae) in Australia, has since been observed on several non-target plant species, including native mangrove Avicennia marina (Acanthaceae). In this study we evaluated the suitability of two native mangroves, A. marina and Aegiceras corniculatum (Myrsinaceae), for the survival and development of A. compressa through no-choice field cage studies. The longevity of females was significantly higher on L. camara (57.7 ± 3.8 days) than on A. marina (43.3 ± 3.3 days) and A. corniculatum (45.7 ± 3.8 days). The proportion of females laying eggs was highest on L. camara (72%) followed by A. marina (36%) and A. corniculatum (17%). More egg batches per female were laid on L. camara than on A. marina and A. corniculatum. Though more nymphs per shoot emerged on L. camara (29.9 ± 2.8) than on A. marina (13 ± 4.8) and A. corniculatum (10 ± 5.3), the number of nymphs that developed through to adults was not significantly different. The duration of nymphal development was longer on A. marina (67 ± 5.8 days) than on L. camara (48 ± 4 days) and A. corniculatum (43 ± 4.6 days). The results, which are in contrast to those from previous glasshouse and quarantine trials, provide evidence that A. compressa adults can survive, lay eggs and complete nymphal development on the two non-target native mangroves in the field under no-choice condition.

Test of the enemy release hypothesis: the native magpie moth prefers a native fireweed (Senecio pinnatifolius) to its introduced congener (S. madagascariensis).

The enemy release hypothesis predicts that native herbivores will either prefer or cause more damage to native than introduced plant species. We tested this using preference and performance experiments in the laboratory and surveys of leaf damage caused by the magpie moth Nyctemera amica on a co-occuring native and introduced species of fireweed (Senecio) in eastern Australia. In the laboratory, ovipositing females and feeding larvae preferred the native S. pinnatifolius over the introduced S. madagascariensis. Larvae performed equally well on foliage of S. pinnatifolius and S. madagascariensis: pupal weights did not differ between insects reared on the two species, but growth rates were significantly faster on S. pinnatifolius. In the field, foliage damage was significantly greater on native S. pinnatifolius than introduced S. madagascariensis. These results support the enemy release hypothesis, and suggest that the failure of native consumers to switch to introduced species contributes to their invasive success. Both plant species experienced reduced, rather than increased, levels of herbivory when growing in mixed populations, as opposed to pure stands in the field; thus, there was no evidence that apparent competition occurred.

Predicting risk and benefit a priori in biological control of invasive plant species: a systems modelling approach

While the method using specialist herbivores in managing invasive plants (classical biological control) is regarded as relatively safe and cost-effective in comparison to other methods of management, the rarity of strict monophagy among insect herbivores illustrates that, like any management option, biological control is not risk-free. The challenge for classical biological control is therefore to predict risks and benefits a priori. In this study we develop a simulation model that may aid in this process. We use this model to predict the risks and benefits of introducing the chrysomelid beetle Charidotis auroguttata to manage the invasive liana Macfadyena unguis-cati in Australia. Preliminary host-specificity testing of this herbivore indicated that there was limited feeding on a non-target plant, although the non-target was only able to sustain some transitions of the life cycle of the herbivore. The model includes herbivore, target and non-target life history and incorporates spillover dynamics of populations of this herbivore from the target to the non-target under a variety of scenarios. Data from studies of this herbivore in the native range and under quarantine were used to parameterize the model and predict the relative risks and benefits of this herbivore when the target and non-target plants co-occur. Key model outputs include population dynamics on target (apparent benefit) and non-target (apparent risk) and fitness consequences to the target (actual benefit) and non-target plant (actual risk) of herbivore damage. The model predicted that risk to the non-target became unacceptable (i.e. significant negative effects on fitness) when the ratio of target to non-target in a given patch ranged from 1:1 to 3:2. By comparing the current known distribution of the non-target and the predicted distribution of the target we were able to identify regions in Australia where the agent may be pose an unacceptable risk. By considering risk and benefit simultaneously, we highlight how such a simulation modelling approach can assist scientists and regulators in making more objective decisions a priori, on the value of releasing specialist herbivores as biological control agents.

Buffel grass in Queensland's semi-arid woodlands: response to local and landscape scale variables, and relationship with grass, forb and reptile species.

Buffel grass [Pennisetum ciliare (L.) Link] has been widely introduced in the Australian rangelands as a consequence of its value for productive grazing, but tends to competitively establish in non-target areas such as remnant vegetation. In this study, we examined the influence landscape-scale and local-scale variables had upon the distribution of buffel grass in remnant poplar box (Eucalyptus populnea F. Muell.) dominant woodland fragments in the Brigalow Bioregion, Queensland. Buffel grass and variables thought to influence its distribution in the region were measured at 60 sites, which were selected based on the amount of native woodland retained in the landscape and patch size. An information-theoretic modelling approach and hierarchical partitioning revealed that the most influential variable was the percent of retained vegetation within a 1-km spatial extent. From this, we identified a critical threshold of similar to 30% retained vegetation in the landscape, above which the model predicted buffel grass was not likely to occur in a woodland fragment. Other explanatory variables in the model were site based, and included litter cover and long-term rainfall. Given the paucity of information on the effect of buffel grass upon biodiversity values, we undertook exploratory analyses to determine whether buffel grass cover influenced the distribution of grass, forb and reptile species. We detected some trends; hierarchical partitioning revealed that buffel grass cover was the most important explanatory variable describing habitat preferences of four reptile species. However, establishing causal links - particularly between native grass and forb species and buffel grass - was problematic owing to possible confounding with grazing pressure. We conclude with a set of management recommendations aimed at reducing the spread of buffel grass into remnant woodlands.

A review of current knowledge of the weedy species Themeda quadrivalvis (grader grass)

This review of grader grass (Themeda quadrivalvis) attempts to collate current knowledge and identify knowledge gaps that may require further research. Grader grass is a tropical annual grass native to India that is now spread throughout many of the tropical regions of the world. In Australia, it has spread rapidly since its introduction in the 1930s and is now naturalised in the tropical areas of Queensland, the Northern Territory and Western Australia and extends south along the east coast to northern New South Wales. It is a vigorous grass with limited palatability, that is capable of invading native and improved pastures, cropping land and protected areas such as state and national parks. Grader grass can form dense monocultures that reduce biodiversity, decrease animal productivity and increase the fire hazard in the seasonally dry tropics. Control options are based on herbicides, grazing management and slashing, while overgrazing appears to favour grader grass. The effect of fire on grader grass is inconclusive and needs to be defined. Little is known about the biology and impacts of grader grass in agricultural and protected ecosystems in Australia. In particular, information is needed on soil seed bank longevity, seed production, germination and growth, which would allow the development of management strategies to control this weedy grass.

Potential for using through-boring to improve groundline treatment of Australian wood species: A preliminary study.

An aging electricity distribution system and reduced availability of naturally durable tropical hardwoods in Australia will combine in the next decade to produce a major shortage of poles. One approach to mitigating this shortage is to utilize lower durability species and improve the penetration of preservatives into the refractory heartwood by introducing additional pretreatment processes. A potential method for improving preservative penetration in the critical ground-line zone is through-boring. This process, in which holes are drilled through the pole perpendicular to the grain in the ground-line zone, is widely used in the western United States for treatment of Douglas-fir and may be Suitable for many Australian wood species. The potential for improving heartwood penetration in eucalypts with alkaline-copper-quaternary (ACQ) compound was assessed on heartwood specimens from four species (Eucalyptus cloeziana F.Muell., E. grandis W.Hill ex Maiden, E. obliqua L'Her. and E. pellita F.Muell.) and Lophostemon confertus (R.Br.) Peter G.Wilson & J.T.Wateril). Longitudinal ACQ penetration was extremely shallow in L. confertus and only slightly better in E. cloeziana. Longitudinal penetration was good in both E. obliqua and E. pellita, although there was some variation in treatment results with length of pressure period. The results suggest that through-boring might be a reasonable approach for achieving heartwood penetration in some Eucalyptus species, although further studies are required to assess additional treatment schedules and to determine the effects of the process oil flexural properties of the poles.

A comparison of alternative plant mixes for conservation bio-control by native beneficial arthropods in vegetable cropping systems in Queensland Australia.

Cucurbit crops host a range of serious sap-sucking insect pests, including silverleaf whitefly (SLW) and aphids, which potentially represent considerable risk to the Australian horticulture industry. These pests are extremely polyphagous with a wide host range. Chemical control is made difficult due to resistance and pollution, and other side-effects are associated with insecticide use. Consequently, there is much interest in maximising the role of biological control in the management of these sap-sucking insect pests. This study aimed to evaluate companion cropping alongside cucurbit crops in a tropical setting as a means to increase the populations of beneficial insects and spiders so as to control the major sap-sucking insect pests. The Population of beneficial and harmful insects, with a focus on SLW and aphids, and other invertebrates were sampled weekly oil four different crops which could be used for habitat manipulation: Goodbug Mix (GBM; a proprietary seed Mixture including self-sowing annual and perennial herbaceous flower species); lablab (Lablab purpureus L. Sweet); lucerne (Medicago sativa L.); and niger (Guizotia abyssinica (L.f.) Cass.). Lablab hosted the highest numbers of beneficial insects (larvae and adults of lacewing (Mallada signata (Schneider)), ladybird beetles (Coccinella transversalis Fabricius) and spiders) while GBM hosted the highest numbers of European bees (Apis mellifera Linnaeus) and spiders. Lucerne and niger showed little promise in hosting beneficial insects, but lucerne hosted significantly more spiders (double the numbers) than niger. Lucerne hosted significantly more of the harmful insect species of aphids (Aphis gossypii (Glover)) and Myzus persicae (Sulzer)) and heliothis (Heliothis armigera Hubner). Niger hosted significantly more vegetable weevils (Listroderes difficillis (Germar)) than the other three species. Therefore, lablab and GBM appear to be viable options to grow within cucurbits or as field boundary crops to attract and increase beneficial insects and spiders for the control of sap-sucking insect pests. Use of these bio-control strategies affords the opportunity to minimise pesticide usage and the risks associated with pollution.

Variation in ecophysiology and carbon economy of invasive and native woody vines of riparian zones in south-eastern Queensland

Exotic and invasive woody vines are major environmental weeds of riparian areas, rainforest communities and remnant natural vegetation in coastal eastern Australia, where they smother standing vegetation, including large trees, and cause canopy collapse. We investigated, through glasshouse resource manipulative experiments, the ecophysiological traits that might facilitate faster growth, better resource acquisition and/or utilization and thus dominance of four exotic and invasive vines of South East Queensland, Australia, compared with their native counterparts. Relative growth rate was not significantly different between the two groups but water use efficiency (WUE) was higher in the native species while the converse was observed for light use efficiency (quantum efficiency, AQE) and maximum photosynthesis on a mass basis (Amax mass). The invasive species, as a group, also exhibited higher respiration load, higher light compensation point and higher specific leaf area. There were stronger correlations of leaf traits and greater structural (but not physiological) plasticity in invasive species than in their native counterparts. The scaling coefficients of resource use efficiencies (WUE, AQE and respiration efficiency) as well as those of fitness (biomass accumulated) versus many of the performance traits examined did not differ between the two species-origin groups, but there were indications of significant shifts in elevation (intercept values) and shifts along common slopes in many of these relationships – signalling differences in carbon economy (revenue returned per unit energy invested) and/or resource usage. Using ordination and based on 14 ecophysiological attributes, a fair level of separation between the two groups was achieved (51.5% explanatory power), with AQE, light compensation point, respiration load, WUE, specific leaf area and leaf area ratio, in decreasing order, being the main drivers. This study suggests similarity in trait plasticity, especially for physiological traits, but there appear to be fundamental differences in carbon economy and resource conservation between native and invasive vine species.