University of , 2015

Evaluating the progress of restoration planting on ,

Robert Vennell1*

1 University of Auckland, Private Bag 92019, Auckland 1142, New Zealand * Email for correspondence: [email protected]

Introduction

Ecological restoration is an intentional human activity that attempts to accelerate the recovery of an ecosystem that has become degraded, damaged or destroyed (SER, 2004). Often the impacts to ecosystems are so great that they cannot recover to their ecological state prior to the disturbance (SER, 2004; Suding et al. 2004). Therefore the key focus of most restoration efforts is to return ecosystems back to their historic trajectories (SER, 2004; Forbes & Craig, 2013) in order to enhance their overall health, integrity and sustainability.

In order to achieve this goal, a historic reference state is generally used as a framework and to provide a starting point for designing the restoration (Balauger et al. 2012). There is concern however that we currently lack sufficient knowledge of historic ecosystems to replicate their composition and function faithfully (Davis, 2000; Choi et al. 2007). In order to mitigate this problem, a modern analogue is often used – the reference ecosystem (SER, 2004). This consists of a contemporary habitat that is assumed to representative of the desired historic reference state, and can serve as a model that restoration activities attempt to emulate (White & Walker, 1997). As a result it also provides an ideal opportunity for evaluating the success of restoration activities, allowing comparisons to be made against restored ecosystems and outcomes effectively measured (e.g. Stephenson, 1999; Palmer et al. 2005; Klein et al. 2007; Matthews et al. 2009).

Motutapu Island in the of New Zealand represents an ideal opportunity for restoration; as a complex physical and geological landscape, protected from invasion by exotic pests by an ocean barrier but still close to New Zealand’s major metropolitan centre (Miller, 1994). Palynological evidence suggests the island was once covered in a mixed podocarp/broadleaf forest (Murray, unpublished data, cited in Miller et al. 1994). However extensive human cultivation has severely reduced native vegetation cover (Esler, 1980) and today the island consists primarily of introduced European pasture (Miller, 1994). The Motutapu Restoration Working Plan (Hawley, 1993) seeks to restore ‘the ecological community that was present on Motuapu after the Rangitoto eruption’ by

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revegetating up to one third of the island in native forest.

This study seeks to evaluate the success of the restoration activities on Motutapu Island, ten years after the first planting programme begun. In order to effectively evaluate whether or not the programme has been successful, a reference ecosystem - the oldest mature remnant forest stand on the island – was compared against three age classes of restoration plantings. Density, basal area and species composition of each of the forest stands were analysed to determine whether the restoration plantings are converging on the reference state in terms of plant community structure and composition.

Methods

Study Site & Data Collection Motutapu Island is situated in the Hauraki Gulf, approximately 16km away from the Auckland Isthmus – New Zealand’s most densely populated urban centre. The island has an extensive cultural history for both European and Māori (Hawley, 1993; Miller et al. 1994). Since the 19th century the island has been cleared and grazed and is largely covered in pasture, save for a few remnant forest patches that remained in the gullies and around the coastal fringe (Esler, 1980)

The point-centred-quarter (PCQ) method (Mitchell, 2015) was used to quantify density, basal area and species composition of the largest remnant of old-growth forest on the island against three age classes of restored forest (Planting A – c. 1995; Planting B – c. 2001, Planting C – c. 2010). At each forest stand, a 75 m transect was run along the edge of the forest fragment. PCQ plots were located at 15m intervals along the transect and sited 20 m apart working perpendicular from the length of the transect (25 plots total for each stand). Within each quarter of the plot, the closest woody tree with a DBH > 5cm was identified and its distance to the centre measured. Planting C contained very few individuals > 5cm DBH, so if no sufficiently large trees were identified within 20 m of the quarter, the distance and identity of the nearest smaller tree was used instead (DBH was not recorded). For Multi-stemmed individuals the diameter of all stems at breast height were recorded.

Data Analyses Density was calculated as the mean density of stems per ha, with confidence intervals calculated as per Mitchell (2015). Basal area was calculated in m2/ha by calculating mean the basal area of each stem for each site, and then multiplying by the density of stems per ha for that site. Similarity in species composition was assessed between sites using Sorenson’s index. Similarity was assessed further using multivariate analysis in PAST software (Hammer et al. 2001). First the data was

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weighted to give greater emphasis to larger individuals – by allocating the two closest individuals within a plot a value of two, with all other species recording a one. Then non-metric multi-dimension scaling (nMDs) Ordination was calculated using the Bray-Curtis dissimilarity measure to evaluate whether the forest sites could be distinguished from one another in terms of species composition. PERMANOVA was used to explore whether the sites were significantly different and SIMPER analysis was used to identify the species most responsible for driving the relationship.

Results

The reference forest contained a significantly lower density of stems than was found in the restored forest (Fig. 1). The restored plantings on the other hand were not shown to be statistically different from one another in terms of density. The reference forest also contained significantly higher mean basal area/m2/ha than the planted forest stands (Fig. 2). The planted forest stands showed a small but significant difference from one another in terms of basal area, with the oldest stand having the highest measure of basal area and progressively less in each of the younger stands.

Fig. 1. Mean density of stems per hectare for the reference forest and Fig. 2. Mean basal area per m2 per hectare for the reference three planted stands of different ages at Motutapu Island Reserve. forest and three planted stands of different ages at Motutapu Error bars = 95% CI as per Mitchell (2015). Island Reserve. Error bars = standard error.

None of the planted forests scored highly on the Sorenson’s index for measuring the similarity of species composition between the forest sites (0.25 – 0.12; Table 1). The older Plantings A and B showed a greater amount of similarity to the forest remnant than the more recent Planting C. Planting A and Planting B were the most similar in terms of species composition (0.33).

Table 1. Sorenson’s Index values comparing reference forest and three PERMANOVA provided strong evidence planted stands of different ages at Motutapu Island Reserve. Values are presented as a figure from 0 – 1, with 1 representing an identical that there were significant differences composition. between all of the forest sites examined Reference Planting A Planting B Planting C Reference . 0.25 0.25 0.12 (Bonferroni-corrected p-values <0.001) Planting A 0.25 . 0.33 0.25 however this a fairly conservative measure Planting B 0.25 0.33 . 0.17 Planting C 0.12 0.25 0.17 . and does not consider the extent of the

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difference. The nMDS ordination recorded a high stress value (0.488) suggesting that only broad generalisations can be made from the ordination. The ordination plot PA suggests that the reference forest is the most dissimilar PB from the other sites, with only a small degree of overlap RF between planting A and planting B and very little PC similarity with Planting C (Fig. 3). The planted forest sites are difficult hard to distinguish, and appear very similar in terms of composition with a large degree of overlap. SIMPER analysis identified the most influential species Fig. 3: nMDs Ordination of PCQ data at Motutapu Island reserve using Bray-Curtis Dissimilarity driving the above relationship. In particular, mānuka Index. Sites were colour coded according to their forest state (Reference – RF – Blue; Planting A – (Leptospermum scoparium), cabbage tree (Cordyline PA, Green; Planting B – PB, Yellow, Planting C – PC, Red). australis), karo (Pittosporum crassifolium) and karamu (Coprosma robusta) were highly abundant in the planted forest stands but absent in the reference forest. In contrast, kohekohe (Dysoxylum spectabile) and tōtara (Podocarpus totara) were predominantly found in reference forest and rare or absent in the restored plantings.

Discussion

To return to the central focus of the study; we aimed to evaluate whether the restoration plantings were converging on the reference state in terms of plant community structure and composition. Our results indicate that the restored forest plantings on Motutapu Island differ significantly from the reference ecosystem in terms of all the variables measured - density, basal area and species composition. However, they also suggest that there is a slight trend towards convergence in older plantings, with a small increase in average basal area and increasing similarity in species composition. Therefore in answer to the research question we suggest that currently, the restoration plantings do not resemble the reference state in any of the variables measured, however given a much longer length of time they may come to do so.

Reference sites are typically selected because they represent a well-developed, mature forest with high levels of biodiversity. However sites in the process of restoration generally represent a much earlier ecological stage (SER, 2004). The restoration plantings are dominated by species such as mānuka, karo, karamu and ngaio (Myoporum laetum) which are all commonly observed in primary successional communities in coastal areas of northern New Zealand (Esler, 1978; Bellingham, 1984; Atkinson, 2004; Towns & Atkinson, 2004). Many of the species that characterise the forest remnant however, such as kohekohe, karaka (Corynocarpus laevigatus) and taraire (Beilschmiedia tarairi)

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have shade tolerant seedlings that generally become established later in the successional trajectory (Bellingham, 1984; Atkinson, 2004; Towns & Atkinson, 2004). Where late-successional plants have been used prematurely at the start of restoration projects, survival and growth has been particularly poor (Cashmore, 1995; Forbes & Craig, 2003). Much greater success is observed when appropriate primary successional species are used to shade out grass and provide safe sites for late-successional plants to establish (Cashmore, 1995). Therefore the difference between the reference and restoration plantings should not necessarily be a cause for concern, but rather an indication that more time is needed for long-term successional processes to operate.

In closing, there are some limitations with using a single site contemporary reference state that should be briefly addressed. A single contemporary reference site can only present a snapshot of the desired historical ecosystem (White & Walker, 1997) and does not give a sense of the wide variability that may have been present in these ecosystems over different spatio-temporal scales. Longer term data is required in order to place these systems in their appropriate historical context and determine how valuable they are as a reference for current efforts (White & Walker, 1997). Another issue is that degraded ecosystems may have significantly altered biotic and abiotic processes (Suding et al. 2004) that mean restoration efforts can have unpredictable results (e.g. Van de Koppel et al. 1997; Anderson et al. 2000). Simply restoring historical conditions may not be sufficient to restore the ecosystem to a prior state, and altered feedbacks may drive it toward an alternative stable state (Suding et al. 2004). In such a case, the mature restoration planting may differ significantly from the reference ecosystem, and whether or not this constitutes “success” will depend on the goals and objectives of the particular restoration project. We recommend that future researchers evaluating the progress of the Motutapu Island restoration consider these factors when interpreting their data, and suggest collecting information from multiple forest remnants across the island to capture a wider degree of variability in the reference ecosystem. We commend the efforts of the Motutapu restoration project in incorporating palaeoecological data into the design of their restoration planting schemes (Pers comm. Bruce Burns, 2015) and encourage future researchers to take a similar long-term approach.

Word Count: 1770 wds (Excluding tables, figures)

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References

Anderson, R. C., Schwegman, J. E., & Anderson, M. R. (2000). Micro‐Scale Restoration: A 25‐Year History of a Southern Illinois Barrens. Restoration Ecology, 8(3), 296-306.

Atkinson, I. A. (2004). Successional processes induced by fires on the northern offshore islands of New Zealand. New Zealand Journal of Ecology, 28(2), 181-193.

Balaguer, L., Escudero, A., Martín-Duque, J. F., Mola, I., & Aronson, J. (2014). The historical reference in restoration ecology: Re-defining a cornerstone concept. Biological Conservation, 176, 12-20.

Bellingham, P. J. (1984). Forest regeneration on Lady Alice Island, Hen and Chickens Group. Tane, 30, 31-42.

Cashmore, P. B. (1995). Revegetation ecology on and its application to Motutapu Island. Doctoral dissertation, Environmental Science and Botany, University of Auckland.

Choi, Y. D. (2007). Restoration ecology to the future: a call for new paradigm. Restoration Ecology, 15(2), 351-353.

Davis, M. A. 2000. ‘‘Restoration’’—a misnomer? Science 287:1203.

Esler, A. E. (1978). Botanical features of islands near the west coast of the Coromandel Peninsula, New Zealand. New Zealand journal of botany, 16(1), 25-44.

Esler, A. E. (1980). Botanical features of Motutapu, Motuihe, and Motukorea, Hauraki Gulf, New Zealand. New Zealand Journal of Botany, 18(1), 15-36.

Forbes, A. R., & Craig, J. L. (2013). Assessing the role of revegetation in achieving restoration goals on Tiritiri Matangi Island. New Zealand Journal of Ecology, 37(3), 343-352.

Hammer, Ø., Harper, D.A.T., and P. D. Ryan, 2001. PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4(1): 9pp.

Hawley, J. (1993). Motutapu Restoration Working Plan. Wellington, Department of Conservation.

Klein, L. R., Clayton, S. R., Alldredge, J. R., & Goodwin, P. (2007). Long‐Term Monitoring and Evaluation of the Lower Red River Meadow Restoration Project, Idaho, USA. Restoration Ecology, 15(2), 223-239.

Matthews, J. W., Spyreas, G., & Endress, A. G. (2009). Trajectories of vegetation-based indicators used to assess wetland restoration progress.Ecological Applications, 19(8), 2093-2107.

Miller, C. J., Craig, J. L., & Mitchell, N. D. (1994). Ark 2020: A conservation vision for Rangitoto and Motutapu Islands. Journal of the Royal Society of New Zealand, 24(1), 65-90.

Mitchell, K. (2015). Quantitative analysis by the point-centered quarter method. Retrieved from http://people.hws.edu/mitchell/PCQM.pdf [Accessed October, 2015].

Palmer, M. A., Bernhardt, E. S., Allan, J. D., Lake, P. S., Alexander, G., Brooks, S., & Sudduth, E. (2005). Standards for ecologically successful river restoration. Journal of applied ecology, 42(2), 208-217.

[SER] Society for Ecological Restoration International Science & Policy Working Group (2004). The SER International Primer on Ecological Restoration. www.ser.org & Tucson: Society for Ecological Restoration International.

Stephenson, N. L. (1999). Reference conditions for giant sequoia forest restoration: structure, process, and precision. Ecological Applications, 9(4), 1253-1265.

Suding, K. N., Gross, K. L., & Houseman, G. R. (2004). Alternative states and positive feedbacks in restoration ecology. Trends in Ecology & Evolution, 19(1), 46-53.

Towns, D. R., & Atkinson, I. A. (2004). Restoration plan for Korapuki Island (Mercury Islands), New Zealand. Wellington, Department of Conservation.

Van de Koppel, J., Rietkerk, M., & Weissing, F. J. (1997). Catastrophic vegetation shifts and soil degradation in terrestrial grazing systems. Trends in Ecology & Evolution, 12(9), 352-356.

White, P. S., & Walker, J. L. (1997). Approximating nature's variation: selecting and using reference information in restoration ecology. Restoration Ecology, 5(4), 338-349.

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