Comparing Traditional and Modern Methods of Kākahi Translocation: Implications for Ecological Restoration

New Zealand Journal of Marine and Freshwater Research

ISSN: 0028-8330 (Print) 1175-8805 (Online) Journal homepage: https://www.tandfonline.com/loi/tnzm20

Comparing traditional and modern methods of kākahi translocation: implications for ecological restoration

Amber Julie McEwan, Aaria Ripeka Dobson-Waitere & Jeffrey S. Shima

To cite this article: Amber Julie McEwan, Aaria Ripeka Dobson-Waitere & Jeffrey S. Shima (2019): Comparing traditional and modern methods of kākahi translocation: implications for ecological restoration, New Zealand Journal of Marine and Freshwater Research, DOI: 10.1080/00288330.2019.1636099 To link to this article: https://doi.org/10.1080/00288330.2019.1636099

Published online: 08 Jul 2019.

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RESEARCH ARTICLE Comparing traditional and modern methods of kākahi translocation: implications for ecological restoration Amber Julie McEwana, Aaria Ripeka Dobson-Waitereb and Jeﬀrey S. Shimaa aSchool of Biological Sciences, Victoria University of Wellington, Wellington, New Zealand; bZEALANDIA Te Māra a Tāne, Wellington, New Zealand

ABSTRACT ARTICLE HISTORY Handling methods are an important determinant of translocation Received 31 January 2019 success. In Aotearoa/New Zealand, tangata whenua traditionally Accepted 21 June 2019 used insulated kete to translocate aquatic animals to new HANDLING EDITOR environments as part of ahumoana tawhito (ancient aquaculture). Joanne Clapcott In this study we investigated the influence of three transport methods (traditional [flax kete], modern [bucket], and a hybrid of KEYWORDS the two [bucket with flax support structures]) on the short-term Kākahi; freshwater mussel; performance (burrowing speed) of kākahi (freshwater mussels). translocation; burrowing; We also tested whether assisted release (planting kākahi in the Matauranga Māori substrate) resulted in enhanced burrowing speeds. Kākahi that were transported using the traditional method were slower to begin probing the substrate, but there was no difference in overall burrowing speed. We also found that assisted release resulted in faster burrowing speeds. We conclude that handling and release procedures can influence the short-term performance of translocated kākahi, and we recommend procedures for future translocation projects, including transporting animals in immersion vessels where practical, and planting them at the release site

Introduction Translocations are increasingly used for the restoration of species and ecosystems (Fischer and Lindenmayer 2000; Seddon et al. 2014 and references therein). While translocation can be a valuable tool, negative outcomessuchasincreasedmortalityandlow reproductive rates can occur (Fischer and Lindenmayer 2000; Letty et al. 2007). The use of appropriate handling procedures is an important determinant of population estab- lishment (Dickens et al. 2010). Aquatic animals are particularly vulnerable to conditions in transport vessels, where excreta buildup and thermal stress can become an issue (Lim et al. 2003). Freshwater mussels (Bivalvia Palaeoheterodonta) are important components of many aquatic ecosystems, where they act as primary consumers (Spooner et al. 2012), and as ecosystem engineers–inﬂuencing water clarity by ﬁlter-feeding (Haag 2012) and biotur- bating sediments by burrowing (Vaughn and Hakenkamp 2001). Most species have a complex life history that includes a relatively sessile adult stage and a larval stage that is

CONTACT Amber Julie McEwan [email protected] © 2019 The Royal Society of New Zealand 2 A. J. MCEWAN ET AL. parasitic on fishes, a strategy that facilitates dispersal and/or up-current migrations (Lydeard et al. 2004). Freshwater mussels are exposed to stressors throughout watersheds, including sedimentation, contamination, impoundment, river realignment (Williams et al. 2017; Naimo 1995; Brim Box and Mossa 1999), and changes to host fish populations (Watters 1996; Downing et al. 2010). Translocation has been identified as a key tool to combat freshwater mussel declines (Haag and Williams 2014). However, newly-translocated mussels are at heightened risk of displacement and predation until they have secured shelter by burrowing into the substrate (Peck et al. 2014). Aotearoa/New Zealand has three presently described species of freshwater mussel: Echyridella menziesii, E. aucklandica, and E. onekaka (Fenwick 2006; Marshall et al. 2014). The first two occur sympatrically in the Wellington region, where they are collec- tively referred to as kākahi by local mana whenua (indigenous Māori people with tribal authority in the region). Kākahi feature in many Aotearoa/New Zealand place-names, and their prevalence in whakataukī (Māori proverbs; Whaanga et al. 2018) is a strong indi- cator of the cultural importance of these animals. Kākahi are threatened by the suite of anthropogenic stressors that affect freshwater ecosystems throughout New Zealand (McDowall 2006; Allibone et al. 2010; McIntosh et al. 2010; Baskaran et al. 2009; Joy 2015; Larned et al. 2016), and the three species are classified by the New Zealand Department of Conservation as either ‘At Risk–Declining’ (E. menziesii), ‘Threatened–Nationally Vulnerable’ (E. aucklandica), or ‘Data Deficient’ (E. onekaka; Grainger et al. 2018 ). In 2018, the Zealandia wildlife sanctuary in Wellington proposed to introduce a translocated population of kākahi into a lake (called Roto Mahanga) within the sanctuary, for the purposes of conservation, education, and potential future bioremediation of the Kaiwharawhara catchment (McEwan 2018). The project is a long-term collaboration between multiple organisations, with knowledge-sharing as a specific goal. During planning discussions, transport and release methods were identified as an important factor in maximising both kākahi welfare during the translocation process, and short-term performance of kākahi at the release site. Predation of adult freshwater mussels tends to be opportunistic and in response to conditions that result in mussels being exposed (Cosgrove et al. 2007). In New Zealand, evidence of rat predation has been observed on freshwater mussel shells on the beach of a hydroelectricity reservoir (Moore et al. 2019), and wading birds have been seen feeding on kākahi in water that had been abruptly lowered as part of a farmland drainage scheme (A. McEwan Pers. Obs.). We recognised that the period immediately following translocation (i.e. before the animals have burrowed into the substrate) could be important as this is when they would be most vulnerable to predation or displacement. Experiments on freshwater mussels in India (Yusufzai et al. 2010) and the USA (Chen et al. 2001) showed that animals which were transported in wetted sacks were more stressed than those transported in immersion vessels (tissue glucose levels were used to measure stress). Chen et al. (2001) investigated 5 species, and found that the strength of stress responses varied between species. Kākahi have historically received little attention compared to other New Zealand fauna, and there is no published information available regarding appropriate translocation methods for these species. Tangata whenua have traditionally translocated aquatic animals to new environments as part of ahumoana tawhito (ancient aquaculture)–mainly to replenish traditional mahinga kai stocks and to create new food stores in other locations. Documented NEW ZEALAND JOURNAL OF MARINE AND FRESHWATER RESEARCH 3 examples of animal translocations include tuna (Anguilla spp. and kōura [Paranephrops planifrons] McDowall 2011), toheroa ([Paphies ventricosa] Futter 2011; Taikato and Ross 2018; Ross et al. in press), and kākahi (Rainforth 2008). Baskets woven from natural materials were common transport vessels for translocations. Examples of vessels include pōhā rimurapa (bull kelp baskets) and kete harakeke (flax baskets) packed with wet sphagnum moss. The benefits of incorporating such mātauranga Māori (Māori wisdom, but see Royal (2009) for a more detailed definition) in modern-day conservation management initiatives is becoming better understood in Aotearoa/New Zealand (e.g. Kusabs and Quin 2009; Awatere and Harmsworth 2014). However, it is important that intellectual property rights are respected by using appropriate procedures and obtaining appropriate permissions (Royal 2009). In this study we used laboratory mesocosm experiments to evaluate the influence of transport method: ‘traditional’ (kete stuffed with wet moss), ‘modern’ (lidded plastic bucket with water and an aerator), and ‘hybrid’ (bucket containing kete submerged in water and an aerator) on the short-term performance (burrowing speed) of kākahi. We also tested whether assisted release (planting kākahi in the substrate at the release site) resulted in enhanced burrowing speeds.

Materials and methods Raranga (weaving) The kete used in the experiment were woven from pā harakeke (ﬂax) and piritā (supple- jack) sourced from the same location as the kākahi. Materials were boiled and/or soaked prior to construction to leach out potentially harmful phytochemicals. The kete were designed using a combination of kete kono (food basket) and kete pipi (pipi basket) fea- tures, and the weaving process involved the use of karakia to acknowledge the whakapapa (genealogy) of the taonga (treasured species e.g. harakeke, kākahi) and give recognition to the diﬀerent atua (deities) involved in their creation. The raranga was carried out by people with mātauranga (wisdom/skills) and mana whenua (ancestral connection to the land) appropriate to the locations, resources, and tīkanga involved in the experiment (meaning that indigenous intellectual property rights were observed).

Transport trials We tested three transport methods: ‘modern’, ‘traditional’, and hybrid (Table 1)onkākahi (only E. menziesii was used in these experiments) from Lake Wairarapa in the lower North Island of New Zealand. Kākahi were collected by hand (with permission from mana whenua), assigned randomly to a transport method (n = 28 per transport vessel for block 1 and 32 per vessel for block 2), and driven in a vehicle for two hours (to simulate the time to travel for the planned translocation to Zealandia in Wellington). They were then immediately placed (right valve down) into a pre-prepared observation aquarium (90 × 38 × 38 cm; all treatments were observed simultaneously in a single aquarium), containing sediment and water from Lake Wairarapa (approximately 30 litres of substrate and 50 litres of water, which resulted in aquarium depths of 150 and 200 mm respectively). The sediment was composed of sand/mud collected from the source location and 4 A. J. MCEWAN ET AL.

Table 1. descriptions and photos of transport vessels used in the experiment. Method Description Photos ‘Modern’ A 20 L, lidded plastic bucket, ﬁlled with 15 L of water and with a coarse-bubble aerator attached. Kākahi are submerged in water but are crowded together at the bottom of the bucket, knocking together during transport-induced turbulence.

‘Traditional’ A kete stuﬀed with wet sphagnum moss. Kākahi are not submerged in water but are kept moist and cool with wet moss. Kākahi are packed tightly so don’t knock together with turbulence.

Hybrid Two stacked kete suspended inside a lidded bucket ﬁlled with 15 L of water and with a coarse-bubble aeratorattached. Kākahi are nestled in the kete so are submerged in water and kept separate from each other on the tiers to reduce jostling and crushing risk. Water can circulate through the tiersa.

aModelled after the ‘stacking’ described in Patterson et al. (2018). repeatedly washed with tap water. The observation aquarium was contained in a quiet, cool room with blinds drawn and no other activities being carried out. The kākahi were assigned individual numbers based on their location in the tank, and then observed con- tinuously, using a stopwatch to record the time at which each of three pre-determined burying stages (Figure 1) occurred for each kākahi. We initially ran the experiment using 28 kākahi for each treatment and observed them for 3 h (block 1). Analyses of these data revealed that 3 h was not enough time to obtain a large enough dataset–particularly regarding righting and burying, so we repeated the experiment (including collection, transport, and observation with new animals) using a further 32 kākahi per treatment and observed them for 8 h (block 2). All experimental conditions remained the same, except for temperature in the observation tank, which was 11°C during the ﬁrst block and 15° C during the second.

Figure 1. Deﬁnitions of terms used to describe burying behaviour used in the experiment. NEW ZEALAND JOURNAL OF MARINE AND FRESHWATER RESEARCH 5

Release trial Following the transport trial observation period, the 84 kākahi that were used for the first block of trials (see above) were excavated by hand into a bucket, and gently swished around to ‘shuffle’ the transport treatment groups. They were then replaced into the observation tank for the release trial (i.e. the same animals were used for one block of the transport trials and the release trial). They were placed in one of two alternating positions: side- lying or planted. Side-lying kākahi were placed right valve down on the top of the substrate, and planted kākahi were inserted (anterior end down), approximately 25% of their length into the substrate. They were then observed once per hour for 8 h, and each assigned a ‘yes’ or ‘no’ classification based on whether 60% or more of their shells were buried at that point (estimated as a level of burying sufficient to secure placement in moderate flows).

Data analysis We used Pearson’s Correlation analysis to examine the relationships between time at first probing, time at burying, and total burying time. To examine the effect of transport, we used data from both of our experimental blocks. We examined the behaviour of the mussels that did undergo burrowing using Kaplan-Meier survival analysis. We analysed time to probe using data from both blocks, then analysed time to righting, and time to bury using only the 8-hour dataset (i.e. block 2; because not enough animals completed righting and burying within 3 h to provide a large enough dataset.). Given that data was collected over two blocks with different water temperature (which has the potential to influence mussels burrowing behaviour [Waller et al. 1999]), we used Cox’s Pro- portional Hazards Model to test the effects of block on time to probe, in order to assess whether the temperature difference could be confounding our results To examine the effect of release we analysed (also using Kaplan-Meier models on the kakahi that completed burrowing) time to bury for planted and for side-lying kākahi. For both transport and release trials, we tested for differences between treatments using the log-rank test. All analyses were conducted using R version 3.2.3 (R Core Team 2015).

Results During the 8-hour observation period, 89 out of 96 (93%) of the kākahi probed, 79 out of 96 (82%) ‘got up’, and 73 out of 96 (76%) buried. The mean ‘time until probing’ (i.e. from the beginning of the observation period until first probing) was 178 ± 13 (mean ± 1 SE) minutes, the mean duration of the probing stage (i.e. the time interval between first probe and righting) was 69 ± 8 min, and the mean duration of the burrowing phase (i.e. the time between righting and being buried) was 42 ± 6 min. The mean total burying time (i.e. the time between first probe and being buried) was 105 ± 10 min. Pearson’s Cor- relation analysis showed a significant positive correlation between time at first probe and time at being buried (r = 0.75, n = 73, P < 0.0001), but no correlation between time at first probe and total burying time (r = −0.14, n = 73, P = 0.2375; Figure 2), demonstrating that kākahi that probed earlier also buried earlier, but this was due to the earlier initiation of the burying process, rather than to faster burrowing speed. 6 A. J. MCEWAN ET AL.

Figure 2. Scatterplots showing correlations between (top) time at being buried and time at ﬁrst probing and (bottom) total burying duration and time at ﬁrst probing.

During the ﬁrst two hours of observation, 19 (32%) of the traditional treatment, 20 (33%) of the modern, and 23 (38%) of the hybrid treatment probed the substrate at least once. Survival analysis showed that–of the kākahi that did probe–those in the traditional treatment were slower to probe compared to those transported in the bucket NEW ZEALAND JOURNAL OF MARINE AND FRESHWATER RESEARCH 7

Figure 3. Behaviour curves generated from survival analysis performed on kākahi that probed during the first two hours of observation. Each curve represents one of the transport treatments (bucket, kete, hybrid) and shows the times at which probing occurred in each group. and the hybrid transporter (P = 0.032; Figure 3). Median time at fist probing was 79 min for the traditional treatment, compared with 26 and 50 min for the modern and hybrid treatments respectively. When the 8-hour observation period was examined in its entirety, there were no significant differences in probing time (P = 0.92), righting time (P = 0.16), and burrowing time (P = 0.91) between transport treatments (Figure 4). Experimental

Figure 4. Behaviour curves generated from survival analysis performed on kākahi that completed (clockwise from top left) probing, righting, and burying. Each curve represents one of the transport treatments (bucket, kete, hybrid) and shows the times at which burrowing behaviour occurred in each group. 8 A. J. MCEWAN ET AL.

Figure 5. Behaviour curves generated from survival analysis performed on kakahi that completed burying during the observation period. Each curve represents one of the release treatments (planted and side-lying) and shows the times at which burying occurred in each group. block had no signiﬁcant eﬀect on survival curves (Cox proportional hazards HR = 0.7, 95% CI = 0.41–1.20, P = 0.18). During the 8-hour release trial, 35 (73%) of the planted kākahi buried, compared to 10 (21%) of the side-lying kākahi. Survival analysis of the release data showed that–of the kākahi that completed burying during the observation period–planted kākahi did so more rapidly than side-lying kākahi (P < 0.0001; Figure 5).

Discussion Transport trials Kākahi that were transported using the traditional method (kete) were slower to begin probing the substrate, compared to those that were transported using the modern (bucket) and hybrid methods. In addition, those kakahi that began probing sooner were also quicker to secure a position in the substrate. Other freshwater mussel species showed higher levels of stress (measured as increased tissue glucose levels) when transported in wetted sacks versus immersion vessels (Chen et al. 2001; Yusufzai et al. 2010). The results of this experiment indicate that the same is true for E. menziesii and that the stress of emersion-vessel transport also affects burrowing behaviour at the release site. Time to bury will influence how quickly a disturbed mussel achieves safety from flow displacement and predation (Peck et al. 2014). Therefore, increased time until probing as a result of transport method could be viewed as a behavioural cost of that method and, as such, be of interest to future mussel translocation projects. During emersion, mussels are exposed to lower oxygen levels and higher temperature fluctuations, so must keep their valves closed. This requires prolonged contraction of the adductor muscle (Wilson et al. 2011), a process which incurs an energetic cost of a magnitude related to the duration of emersion. Lower energy levels in the kete group may have NEW ZEALAND JOURNAL OF MARINE AND FRESHWATER RESEARCH 9 contributed to the observed slower probing times. Wilson et al. (2011) found that juvenile PIT-tagged freshwater mussels were slower to burrow than non-tagged animals and attrib- uted this difference to a possible energy deficit incurred by the tagging and handling process. Freshwater mussels can withstand prolonged emersion by switching to anaerobic metabolism (Aldridge et al. 1987), and the kākahi in the kete treatment may have done this, while those in immersion vessels may have been able to continue functioning ‘as usual’. This difference may have resulted in the latter groups being ready to burrow when placed in the tank, compared to the kete group, which may have needed more time to make metabolic adjustments prior to initiating burrowing. Probing onset time varied between treatments, but individual variation in this behaviour was also observed. Behavioural traits can influence post-release survival of translocated animals (May et al. 2016; Haage et al. 2017). For example, bolder animals can be less likely to survive translocation (Bremner-Harrison et al. 2004). If probing propensity is a behavioural trait that varies among individual mussels, then such a trait could influence the short-term performance of translocated mussels and would thus be an important consideration for future translocation projects. We recommend further work to determine whether individual mussel probing propensity varies along a shy-bold personality gradient as demonstrated by other invertebrates such as spiders (Riechert and Hedrick 1990), land snails (Maxime et al. 2017), and hermit crabs (Briffa et al. 2008).

Release trial Kākahi that were planted in the substrate buried more rapidly than those that were side lying on top of the substrate. The magnitude of the time difference suggests that this was not wholly attributable to the partial burying completed by us (we placed them approximately 25% of their length into the substrate), but also to the apparently more rapid foot action once the foot was enclosed in the substrate i.e. planted mussels didn’t need to initiate open water probing prior to burrowing. During the experiment, the average time until first probing was longer than the time between probing and burying (i.e. the entire burying process). Given that burrowing likely requires more energy than probing in open water (as the foot must drag the mussel through the substrate), the quicker burying time we observed is likely not a result of lower energy cost of this action. When the foot is exposed, a mussel is potentially vulnerable to sub-lethal predation, such as ‘foot-cropping’ by benthic fishes or crabs, which has been documented in some marine bivalves (Salas et al. 2001; Mouritsen and Poulin 2003). New Zealand freshwater mussels share benthic habitat with several small, car- nivorous fishes (e.g. Gobiomorphus spp.), and omnivorous freshwater crayfish (Parane- phrops spp.). Foot cropping–while not documented in freshwater bivalves–may be occurring and may have driven the evolution of behaviour which minimises foot exposure and thus cropping risk. The long average time to probe when lying on top of the substrate (which we observed across all our experiments) compared with the apparent readiness of the planted kākahi to employ foot action in the substrate may indicate that foot action when the foot is exposed is riskier than foot action when the foot is concealed within the substrate. If: (1) open water probing incurs a risk of predation, (2) open water probing is necessary to initiate burrowing, and (3) lying exposed on the substrate (i.e. not probing) also incurs risk, then presumably mussels must somehow optimise between these two activities in order to minimise risk. Therefore, assisted release could be a valuable tool in mussel 10 A. J. MCEWAN ET AL. translocations, particularly with very threatened species and/or where there is risk of predation and/or displacement. Debate exists among freshwater mussel scientists regarding the optimal way of placing mussels into the substrate, with some advocating for assisted planting (including using a tool to loosen the substrate beforehand), and others arguing that planting mussels in unsuitable habitat could incur unacceptable risk (e.g. if mussels died or had to expend energy moving to a new location; Patterson et al. 2018). Our experiment has demonstrated that–if workers have a high degree of confidence in the suitability of release-site habitat– assisted planting can result in more rapid burrowing of translocated mussels. This would in turn reduce mussel exposure to possible sub-lethal predation during open water probing, as well as lethal predation (e.g. from birds (Williams et al. 2017) and physical displacement while exposed on top of the substrate). We used the same animals from block 1 of the transport trials to carry out the release trial, and the trials were carried out back to back, meaning that the burrowing which was observed during the release trial was by animals which had already had to burrow once in the last 3 h. This may have affected their behaviour, for example they may have been fati- gued and may have buried more quickly if they had been new animals or had been pro- vided with a rest period in between trials. This may have been particularly so for the animals which were transported in the emersion vessel, although ‘shuffling’ of the transport treatment groups prior to the redeployment for the release trial would have served to distribute variability relating to transport method.

Recommendations for future mussel translocations To maximise short term performance and thus the chances of successful population estab- lishment at release sites, we recommend that freshwater mussels are transported in immersion vessels. The creation of hybrid transporters may add value because–as in our project– the resulting engagement with mana whenua would likely lead to valuable exchanges of cultural knowledge. However, in some situations the use of immersion vessels will be imprac- tical, for example if a translocation involves a large number of animals, then transporting the required amount of water would be challenging. The results of this experiment, as well as those of Chen et al. (2001) and Yusufzai et al. (2010) indicate that transport stress may be short-lived, particularly when transport time is low. Where possible, translocated mussels should be planted into the substrate at release sites, but only in situations where there is a high degree of confidence that the habitat is suitable, and only by experienced workers familiar with mussel anatomy (i.e. ‘which way is up’). This study examined only one mussel species, and Chen et al. (2001) recorded species-specific variation in transport-related stress. Also, Waller et al. (1999) recorded species-specificdifferences in burrowing behaviour following disturbance. It would be useful to collect information on the behavioural responses of other New Zealand freshwater mussel species in relation to transport and release before translocations of these animals are carried out.

Acknowledgements This project was carried out under permits 67922-RES and NFT 288 (from the Department of Con- servation and the Ministry for Primary Industry respectively). We are grateful to Vanessa Taikato NEW ZEALAND JOURNAL OF MARINE AND FRESHWATER RESEARCH 11 and Phil Ross for helpful discussion about ahumoana tawhito. Thanks to Jade Waters, Waiaria Pitau, and Te Pātukituki o Wairarapa Inc. for their weaving skills and to Ngati Kahungunu ki Wair- arapa and Rangitāne o Wairarapa for allowing access to their whenua and taonga tuku iho and for supporting the project.

Disclosure statement No potential conﬂict of interest was reported by the authors.

Funding This research was funded by Victoria University of Wellington Zealandia Centre for People and Nature PhD Scholarship, and the Greater Wellington Regional Council Science and Research Fund.

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