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Maintenance of Key Tree Species

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Maintenance of Key Tree Species Maintenance of key tree species SCIENCE FOR CONSERVATION 190 Robert B. Allen, Geoff M. Rogers and Glenn H. Stewart Published by Department of Conservation P.O. Box 10-420 Wellington, New Zealand Science for Conservation is a scientific monograph series presenting research funded by New Zealand Department of Conservation (DOC). Manuscripts are internally and externally peer-reviewed; resulting publications are considered part of the formal international scientific literature. Titles are listed in the DOC Science Publishing catalogue on the departmental website http:// www.doc.govt.nz and printed copies can be purchased from [email protected] © Copyright March 2002, New Zealand Department of Conservation ISSN 11732946 ISBN 0478222335 This report was prepared for publication by DOC Science Publishing, Science & Research Unit; editing and layout by Lynette Clelland. Publication was approved by the Manager, Science & Research Unit, Science Technology and Information Services, Department of Conservation, Wellington. CONTENTS Abstract 5 1. Background 6 2. Objective 8 3. Case studies of selected species 8 3.1 Kamahi (Weinmannia racemosa)8 3.2 Southern rata (Metrosideros umbellata)11 3.3 Northern rata (Metrosideros robusta)14 3.4 Hinau (Elaeocarpus dentatus)18 3.5 Red beech (Nothofagus fusca)20 3.6 Pahautea (Libocedrus bidwillii)23 3.7 Halls totara (Podocarpus hallii)28 4. Critical stages of life history in indigenous trees 31 4.1 Flowering and pollination 31 4.2 Seed productionasynchrony across species 31 4.3 Seed predation 32 4.4 Dispersal 33 4.5 Seed predation following dispersal 33 4.6 Seedling establishment 34 4.7 Canopy defoliation 34 4.8 Stage of stand development 35 4.9 Modifications to disturbance regimes and successional development 35 5. Establishing priorities among species 36 5.1 Using demographic parameters to show that there is reason for concern about the maintenance of a tree species 38 5.2 Understanding the causes of any imbalances in the demographics of a tree species 41 5.2.1 Measure the correct parameters in the most appropriate way 41 5.2.2 Develop an encompassing framework for the demographics of tree species 43 5.3 Determining the significance of maintaining a tree species within a context of its contribution to ecosystem dynamics 44 5.4 Showing that the population trajectory of a tree species can be changed to something considered more desirable 47 5.4.1 The traditional option of minimising introduced animal impacts 47 5.4.2 Is there a need for more direct management? 48 5.5 Comparing the vulnerability of tree species 49 5.5.1 Vulnerability of tree species 49 5.5.2 A systematic approach to scoring species 50 6. Recommendations 52 7. Acknowledgements 52 10. References 53 4 Allen et al.Maintenance of key tree species Maintenance of key tree species Robert B. Allen1, Geoff M. Rogers2, Glenn H. Stewart3 1Landcare Research, PO Box 69, Lincoln 8152 2Department of Conservation, PO Box 5244, Dunedin 3Lincoln University, PO Box 84, Lincoln 8152 ABSTRACT There is a widespread perception that certain dominant (key) tree species are declining on conservation lands in New Zealand. This concern is usually based on observations suggesting undesirable shifts in the demographic processes regulating the distribution, size and viability of tree populations. Our project was initiated to define a process to understand and compare the vulnerability to population decline of key tree species at a range of scales and to set priorities for conservation management. We first consider in detail observations and possible causes of population decline of seven selected tree species, then consider more generally the critical stages in tree life cycles likely to affect population viability of indigenous species and, finally, suggest an approach to establishing priorities among tree species. Prioritising tree species should involve: showing there is reason for concern about the maintenance of a tree species in terms of a population imbalance; understanding the causes of any imbalances in the demographics of a tree species; determining the significance of maintaining a tree species within a context of its contribution to ecosystem functions; and showing that the trajectory of a tree species population can be changed to something judged more desirable. In all of these we assess the adequacy of knowledge, suggest research questions in need of answers, and consider the implications for management. Keywords: Forest, tree species, vulnerability, risk assessment, monitoring, forest health, indicators. © March 2002, Department of Conservation. This paper may be cited as: Allen, R.B.; Rogers, G.M.; Stewart, G.H. 2002: Maintenance of key tree species. Science for Conservation 190. 60 p. Science for Conservation 190 5 1. Background There is a widespread perception among indigenous forest managers, ecologists, and the wider public, that certain dominant (key) tree species are declining in indigenous forests. In a general sense, this is not a new concern as numerous examples of extensive tree death can be listed from around New Zealand over the past century (e.g. Morgan 1908; Chavasse 1955; Holloway 1959; Stewart & Veblen 1982b; Rose et al. 1992). Usually this concern is based on observations suggesting undesirable shifts in the demographic processes regulating population size and viability, e.g.: Low numbers of mountain beech (Nothofagus solandri var. cliffortioides) seedlings in the Kaweka Range (Allen & Allan 1997). Extensive mortality of pahautea (Libocedrus bidwillii) trees at Hihitahi (Rogers 1997). To assess the significance of such observations, a number of related questions should be asked, such as: Is pahautea regenerating in stands where trees are dying, or in other nearby stands at Hihitahi? Is recent mortality of pahautea a feature of most forests in the Hihitahi area, or certain areas of the country, or nationally? Are there some areas where there is compensatory regeneration of pahautea? Is the imbalance of mortality and regeneration of pahautea at an area or na- tional scale of more concern than that for other tree species (e.g. mountain beech or Halls totara (Podocarpus hallii)? Addressing such a sequence of questions is essential in establishing whether there is a problem, the scale of any problem and, therefore, our ability to prioritise which species should be of most concern. This prioritisation is necessary because of the need to make informed management choices and focus research efforts. The process of prioritising which species are of most concern should also take account of what is causing any undesirable shifts in the demographic processes regulating population size and viability. The difficulty of ascribing cause has bedeviled pathologists, ecologists, and forest managers for a long time (e.g. Chavasse 1955; Wardle 1966; Veblen & Stewart 1982a; Bellingham et al. 1999a). Too often a simplistic view is presented that does not stand further scrutiny, for example: In a simple sense it could be argued, as many authors have, that defoliation by possums (Trichosurus vulpecula) kills southern rata (Metrosideros umbellata) trees. However, not all southern rata trees that die are killed by possums. For example, most tree death in natural populations is the result of competitive thinning. The important question, then, becomes: what proportion of southern rata tree death is primarily caused by possums defoliating trees?. In a general sense, this last question is difficult to answer for at least two reasons. Firstly, the structure, composition and functioning of forest ecosystems is driven by a wide range of interacting factors (Table 1), some of 6 Allen et al.Maintenance of key tree species TABLE 1. FACTORS COMMONLY INFLUENCING THE COMPOSITION OF FORESTS. SOME OF THE VARIABLES COMMONLY MEASURED TO REPRESENT THESE FACTORS, AS WELL AS SOME OF THE MECHANISMS THROUGH WHICH THESE FACTORS MAY OPERATE, ARE GIVEN (MODIFIED FROM ALLEN 2000). )$&7256 9$5,$%/(6 (;$03/(Ã2)Ã0(&+$1,60 'LVWXUEDQFH &KDQJHVÃLQÃELRPDVVÃRUÃQXPEHUÃRIÃWUHHV ,QGLYLGXDOVÃNLOOHGÃRIÃRQHÃRUÃPRUHÃVSHFLHV +HUELYRU\ /HYHOÃRIÃGHIROLDWLRQÃLQGLYLGXDOÃKHLJKWÃJURZWK 5HGXFHGÃSKRWRV\QWKHWLFÃDELOLW\ 6SHFLHVÃHIIHFWV /LWWHUÃTXDOLW\ 0RGLILHVÃWKHÃDELOLWLHVÃRIÃVHHGVÃRIÃRWKHUÃVSHFLHV WRÃJHUPLQDWHÃDQGÃJURZ &OLPDWH 7HPSHUDWXUHÃSUHFLSLWDWLRQ &KDQJHVÃSK\VLRORJLFDOÃSURFHVVHV 6RLO 7H[WXUHÃ1ÃDYDLODELOLW\ÃFDWLRQÃDYDLODELOLW\ ,QIOXHQFHVÃUHVRXUFHVÃHVVHQWLDOÃIRUÃJURZWKÃDQG GHYHORSPHQW 'LVSHUVDO 6HHGÃGLVSHUVDOÃDYDLODEOHÃUHJHQHUDWLRQÃQLFKHV 6HHGVÃGRÃQRWÃDUULYHÃDWÃDQÃRWKHUZLVHÃVXLWDEOH VLWH 7LPH 7UHHÃDJHÃUHODWLYHÃELRPDVV 6SHFLHVÃGLIIHUHQWLDOÃORQJHYLW\ which are relatively easy to measure (e.g. rainfall), others we know little about (e.g. infrequent disturbance impacts); and finally, we must add chance. Additionally, because many of these factors are correlated with each other at any particular temporal or spatial scale, it is difficult to separate the influence of an individual factor and define causal relationships. The following example shows this limitation for spatial data. A study in south Westland evaluated tree mortality in three localities along a possum invasion front; it was only in the write-up stage of this study that the invasion front was also shown to coincide with a compositional gradient usually related to soil fertility (Stewart 1992). Because we know from elsewhere that mortality in such forests apparently relates to a soil fertility gradient, it was therefore difficult to partition the effects of a possum density gradient along the Westland invasion front from those
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