Ornithol. Sci. 2: 89–96 (2003)

INVITED ARTICLE Translocations in avian conservation: reintroduction biology of the (Philesturnus carunculatus carunculatus)

Johanna P. PIERRE#

Marine Conservation Unit, Department of Conservation, P. O. Box 10–420, Wellington,

Abstract Translocation is a commonly used tool in conservation management. ORNITHOLOGICAL However, because post-release monitoring has been infrequent in the past, reasons for SCIENCE the outcomes of translocations have often been unknown. Here, I review the reintro- © The Ornithological Society duction biology (including dispersal patterns, social organisation, survival, habitat use of Japan 2003 and foraging patterns) of a population of 26 South Island Saddlebacks (Philesturnus carunculatus carunculatus), on Motuara Island, New Zealand. After release on Motu- ara Island, South Island Saddlebacks dispersed widely through forest areas. During their first post-release breeding season, saddlebacks established territories of 1.9 ha- 8.8 ha (X¯ 4.21 ha, SD2.42) in size, and territorial confrontations were very rare. Saddlebacks bearing both adult and subadult plumage held territories and attempted to breed, and successful breeding produced approximately 10 fledglings. Saddlebacks foraged on a variety of plant species, dead wood and the ground. Except for five-fin- ger (Pseudopanax arboreus), a preferred foraging substrate, foraged in all plant material in proportion to its availability. Saddlebacks preferred to forage in the lower levels of the forest. Although vegetation composition differed significantly between territories, all territories contained forest areas, and birds appeared to prefer foraging in larger sized trees. Large territory sizes, breeding attempts by young birds and rarity of territorial confrontations are most likely products of low population density. As density increases, birds are expected to occupy smaller territories, forage more effi- ciently within these smaller areas, start breeding at older ages, and possibly colonise scrub habitats. The translocated group sustained no more than 50% mortality at 8-10 months after release. In the past, translocations of 15–59 South Island Saddlebacks have been successful, suggesting that the relatively small founder group does not threaten the success of the transfer to Motuara Island. Saddlebacks are flexible in their habitat use, appear to readily adapt to ‘new’ environments and have high repro- ductive potential, increasing the likelihood of success of translocations of this species.

Key words New Zealand, Philesturnus carunculatus carunculatus, Reintroduction, , Translocation

Translocation is becoming increasingly commonly 1973; Merton 1973, 1975; Bell 1978; Butler & Mer- used for conservation, and can be defined as the in- ton 1992). In New Zealand, translocations have often tentional release of plants or to the wild been incorporated into species management programs to establish, re-establish or augment a population in the past, and have been successful in saving some (IUCN 1987; Griffith et al. 1989). The technique has New Zealand birds from extinction. Since the 1880s, been applied to the conservation of mammals (e.g. almost 400 translocations of 50 taxa (42 birds, five Dufty et al. 1994; Short et al. 1994), invertebrates reptiles and three invertebrates) have been conducted (e.g. Sherley 1994) and birds (e.g. Atkinson & Bell in New Zealand, often in emergency situations, such as in 1962 when ship rats (Rattus rattus) arrived at (Received 4 March 2003; Accepted 23 September 2003) the last stronghold of the South Island Saddleback # E-mail: [email protected] (Philesturnus carunculatus carunculatus) (Atkinson Proceeding of the Symposium on Behavioral Studies and Conserva- tion Biology. 15 September 2002, Tokyo; Organized by Go Fujita & Bell 1973; Merton 1973, 1975; Bell 1978). and Hisashi Nagata Despite the increasingly common application of

89 J. P. PIERRE translocations in conservation management, and wide shore islands, it is unable to coexist with introduced recognition of the value of post-release monitoring predators, and consequently, was virtually extinct by (e.g. Scott & Carpenter 1987; Griffith et al. 1989; 1900 (Oliver 1955; Roberts 1991). The total popula- Sarrazin & Barbault 1996), such monitoring does not tion of South Island Saddlebacks is currently about always occur. In fact, Wolf et al. (1996) reported that 650, with birds occurring on 11 islands, all of which only 45% of 336 and mammal translocation pro- are free of introduced predators. grams included tagging of released individuals or Following the South Island Saddleback Recovery post-release telemetry, and in about 30% of transloca- Plan (Roberts 1991) administered by the New tions, the causes of 90–100% of losses were Zealand Department of Conservation, 26 South Is- never identified. In New Zealand, translocation con- land Saddlebacks were translocated to Motuara Is- tinues to be widely used as a conservation tool. Post- land in the Marlborough Sounds from the Titi Islands release monitoring of translocated animals may be near Stewart Island, New Zealand, in March 1994. becoming more common (e.g. Nelson et al. 2002), (See Figure 1 for the locations of islands mentioned but such monitoring seldom occurred in the past (De- in the text). Seven adult males, 11 adult females, 5 partment of Conservation 1994; Lovegrove & Veitch subadult males, one subadult female and one adult 1994; Armstrong & McLean 1995). and subadult of unknown sex were included in the Similar to post-release monitoring of survival, translocated founder population. Motuara Island (59 monitoring the habitat use of newly translocated or- ha) was selected as the release site for translocated ganisms is important, yet seldom occurs. Globally, South Island Saddlebacks for three main reasons. translocations are deemed most likely to succeed First, introduced predators (Rattus exulans) were ex- where animals are released into areas of high habitat tirpated from the island in 1993. Second, vegetation quality and/or quantity (Wolf et al. 1996). This em- on Motuara Island has been regenerating since farm- phasizes the importance of thoroughly investigating ing was abandoned in 1926 (W.F. Cash pers. comm.), the habitats at potential release sites before conduct- and finally, the bird was historically resident in the ing translocations and monitoring the habitat use of area. Conservation managers considered Motuara Is- translocated animals after they are released. Post-re- land to be sufficiently large and with appropriate lease monitoring of habitat use can confirm suitability of the new location, and produce valuable informa- tion relevant to future translocations, thereby facili- tating adaptive wildlife management (Sarrazin & Barbault 1996). For example, mobile animals at low densities may occupy habitats on the basis of prefer- ence rather than requirement, and may colonise less ‘preferred’ areas as their population density increases (Jenkins 1976; Armstrong & McLean 1995). There- fore, monitoring spatiotemporal patterns of habitat use together with population growth can be informa- tive in determining habitat requirements and prefer- ences, and the ability of translocated organisms to colonise ‘novel’ environments. The South Island Saddleback is a forest endemic to the islands of New Zealand. It is an en- dangered subspecies of the near threatened New Zealand Saddleback (IUCN 2000). Diurnal, territorial and largely insectivorous, the South Island Saddle- back forages on foliage, live and dead wood, and the ground. It uses cavities for roosting and is monoga- mous with apparently flexible nest site requirements (Guthrie-Smith 1925; Pierre 1995). Although the South Island Saddleback was formerly widespread Fig. 1. Locations of islands mentioned in the text. (From over New Zealand’s South Island and southern off- Pierre 1999).

90 South Island Saddleback reintroduction habitat to support more South Island Saddlebacks data using paired t tests and analysis of variance than currently existed on any other island (W.F. Cash (ANOVA) conducted in SPSS 8.0 (SPSS Inc. 1997). I pers. comm.). used Games-Howell post-hoc tests (Day & Quinn In this paper, I review the reintroduction biology of 1989) to identify homogeneous subgroups after sig- the South Island Saddleback. I discuss post-release nificant ANOVAs, and Bonferroni a adjustments dispersal, social organisation, survival, habitat use when appropriate, including in G tests described and foraging patterns of the newly released popula- below (Miller 1981). See Pierre (2001) for a discus- tion on Motuara Island, New Zealand. Before the sion of how pseudoreplication and problems of inde- work described in this review was carried out, the pendence were minimized. South Island Saddleback had not been studied in de- I characterised vegetation using a transect-quadrat tail. Also, future translocations were planned to in- sampling method, and sampled both saddleback terri- crease the total population of this bird (Roberts tories and one area uninhabited by saddlebacks 1991). For these reasons, post-release monitoring of (Pierre 2001). I defined plant availability as the cross- the translocated population was critical. sectional area of each plant species at breast height, and the availability of dead wood as the proportion of METHODS total quadrat area it covered on each transect (Pierre 2001). Vegetation characteristics were compared be- Saddlebacks were weighed, measured and their tween sites using G tests (Zar 1996). plumage examined on capture to determine sex and age (W.F. Cash pers. comm.; Nillson 1978; Jenkins & RESULTS Veitch 1991). Numbered metal and unique combina- tions of coloured plastic legbands were used to iden- After release, South Island Saddlebacks ranged tify individuals. After release on Motuara Island, widely through the forested areas of Motuara Island. birds were relocated using a recording of South Is- By the start of the first post-release breeding season, land Saddleback male territorial song, and via their both adult and subadult birds appeared to have settled vocal responses and attraction to disturbances (e.g. on territories, which varied in size from 1.9–8.8 ha logs breaking). The work reviewed here results from (X¯ 4.21 ha, SD2.42, N6, Figure 2, Pierre 1999). post-release monitoring involving intensively search- Adult and subadult saddlebacks announced their ing for birds for four days, six months after they were presence vocally from a range of vertical locations in released, and searching for and monitoring birds for their territories, especially in the first two hours after 56 days, from eight months after release, during their dawn, and for approximately one hour just before first breeding season on Motuara Island. dusk. However, confrontations between neighbours Territory boundaries were identified during the appeared very rare, and I observed only one during breeding season by conducting repeated searches for the study. This low frequency was despite saddle- birds and recording the locations of song posts. I backs venturing into each other’s territories, which I monitored birds for 90 minutes during these tracking observed on five occasions during my research, and episodes; tracking beyond five 90 minute sessions did which at least sometimes appeared related to searches not alter the perceived location of territory bound- for water (Pierre 1999). aries (Pierre 1999). As well as recording song posts, I The vegetation composition of saddleback territo- monitored habitat use by birds, including substrates ries on Motuara Island differed significantly, and was used for foraging and other activities, and heights of different again in areas uninhabited by saddlebacks the forest that birds occupied (Pierre 2001). I also (G tests, vegetation composition in all territories recorded prey identity when possible, and where prey compared to all other territories and a vegetated area were caught (Pierre 2000). Saddleback nests were lo- not occupied by saddlebacks: G8 2117.36, P 0.005, cated by following birds holding nesting materials, Pierre 2001). However, all territories were in forest and by checking nest boxes. To avoid disturbing nest- areas and included plants of similar size distributions ing birds, I did not check nests frequently after locat- (G tests: plant sizes in all territories compared to all ing them. Instead, I monitored breeding activities in- other territories, G5 14, P 0.005, NS). In contrast, directly by assessing the behaviour of adult birds saddlebacks did not inhabit a locality covered by (Pierre 1999). smaller diameter plants, in this case scrub (G tests: all I conducted analyses of habitat use and foraging territories compared to an area uninhabited by saddle-

91 J. P. PIERRE

backs, all G5 26, P 0.005, Pierre 2001). At least five, and possibly six pairs of saddlebacks were present during the first post-release breeding season on Motuara Island. Pairs were not knowingly transferred together, and after release at least some birds interacted with more than one potential mate before settling on a territory with what appeared to be a stable partner (Pierre 1999). Both adult birds and those in subadult plumage formed pairs and at- tempted breeding during the first post-release breed- ing season. Pairs were able to form and then initiate breeding with what appeared to be minimal delay; one pair built a nest and reared two offspring no more than 35 days after pairing. This pair then appeared to renest (Pierre 1999). The nests of two other pairs were found in a tree hole and a nest box, and held one and two eggs, respectively. However, these nests were deserted for unknown reasons. In all, an esti- mated 10 fledglings were hatched the first breeding season after saddlebacks were released onto Motuara Island (Pierre 1999; W.F. Cash pers. comm.). Translocated saddlebacks utilised a range of forag- ing substrates, including various plant species, dead wood and the ground. Foraging patterns differed be- tween males and females, with males spending the most foraging time on the ground, whereas females Fig. 2. South Island Saddleback territory boundaries during preferred to forage in five-finger (Pseudopanax 11 Nov 94–13 Jan 95, the first breeding season after release. arboreus) (ANOVA: F 4.59, P0.003, Figure 3, Letters denote legband combinations: Bblue, Ggreen, 5,29 Mmetal, Rred, Wwhite, Yyellow. (From Pierre 1999). Pierre 2001). Overall, birds apportioned their forag- ing time differently among different foraging sub- strates, and spent the most time foraging on the

Fig. 3. Percent time spent on foraging sites (X¯ 1 SE) by male (open bars) and female (hatched bars) South Is- land Saddlebacks on Motuara Island. BrarepBrachyglottis repanda, CoplucCoprosma lucida, CorlaeCoryno- carpus laevigatus, Deaddead wood, HedarbHedycarya arborea, KuneriKunzea ericoides, Macexc Macropiper excelsum, MelramMelicytus ramifloris, PhocooPhormium cookianum, PsearbPseudopanax ar- boreus. Numbers above bars represent the number of birds contributing to means. (From Pierre 2001).

92 South Island Saddleback reintroduction

land had strong effects on the area of territories. The ground and five-finger (ANOVA: F5,29 13.08, P0.001, Figure 3, Pierre 2001). However, birds size of territories has been used most plant species (and dead wood) in propor- recorded to change temporally, with territories being largest during breeding (O’Callaghan 1980). tion to their availability (t tests: t1–6 1.14–2.87, P0.21–0.99), except for five-finger (used more than South Island Saddlebacks on Motuara Island were much less vocal than North Island Saddlebacks oc- expected, t test: t6 2.74, P 0.03). Flax (Phormium cookianum) may also have been used more than ex- curring in higher density populations. However, tem- pected, although small sample size rendered this im- poral singing patterns appear similar for both sub- possible to test statistically (Pierre 2001). The num- species (pers. obs.; Jenkins 1976; Murphy 1989). Ter- ber of prey items South Island Saddlebacks obtained ritorial confrontations occurred rarely on Motuara Is- from foraging substrates was roughly in proportion to land, relative to North Island Saddleback populations the amount of foraging time spent, although dead (Jenkins 1976; O’Callaghan 1980). This is probably wood was a particularly rich foraging substrate. Sad- because saddleback territories on Motuara Island dlebacks were largely insectivorous, but also fed on were sufficiently large that neighbours would seldom nectar and honeydew (Pierre 2000). meet, and may have been out of earshot of each other Saddlebacks were recorded significantly more fre- often, if not most of the time. The utility and impor- quently in the lowest two metres of the forest than at tance of song in maintaining territorial integrity might be expected to increase with increasing popula- any other level (ANOVA: F4,30 41.31, P 0.001, Pierre 2001). This was the case for both males and fe- tion densities, however Murphy (1989) showed that males, whose vertical use of the forest did not appear the number of neighbours around North Island Sad- dleback territories did not affect singing rates. The to differ (ANOVA: F4,30 0.20, P 0.94, however 1- b0.16 at a0.1, Pierre 2001). frequency of territorial displays, however, is known Saddlebacks were observed drinking water fre- to increase with population densities in North Island quently, both from natural sources and troughs con- Saddlebacks (Jenkins 1976; O’Callaghan 1980). structed on the island before birds were released. South Island Saddleback territories on Motuara They also bathed in these human-made water troughs Island were not areas of completely exclusive use frequently (Pierre 2001). by territory ‘owners’. This may not be unusual for Mortality of South Island Saddlebacks was 35% the species. In a relatively high density population, (9/26 birds dead) at six months after release. Maxi- O’Callaghan (1980) documented areas of overlap mum mortality was 50% 8–10 months after release. between North Island Saddleback territories, as Of the surviving 13/26 translocated saddlebacks, well as subadults and non-territorial adults moving there were 3/7 adult males, 4/11 adult females and through the territories of others. 6/7 subadults remaining (Pierre 1999). Vegetation composition of all saddleback territo- ries differed on Motuara Island. Further, saddlebacks seemed to prefer areas with larger trees; all territories DISCUSSION were at least partially forested, compared to uninhab- After release on Motuara Island, South Island Sad- ited areas in which the dominant vegetation type was dlebacks traversed forested areas widely, before set- scrub. The species composition of forest however, did tling on territories after about eight months. Translo- not seem important. Habitat characteristics, e.g. the cations of North Island Saddlebacks (P. c. rufusater) availability of food, affect the size of North Island also result in wide post-release dispersal, suggesting Saddleback territories (Blackburn 1964; O’Callaghan that this is the norm for the species (Jenkins 1976; 1980). Despite this, South Island Saddleback territo- Armstrong & Craig 1995). South Island Saddlebacks ries were probably much larger than the area required settled on territories that were unusually large and to support a breeding pair (also see above) and as variable in size. There is little data on territory size in previously mentioned, the large size is almost cer- saddlebacks, but on Cuvier Island (170 ha), two pairs tainly a product of low population density. of North Island Saddlebacks held territories of 0.89 In low density North Island Saddleback popula- ha and 1.22 ha in November/December, the height of tions, birds also preferentially occupy forest habitat. the breeding season. The smallest territory on Motu- However, with increasing density, birds will colonise ara Island was about 1.5–2 times this size, suggesting scrub areas, and can breed very successfully in this that the low density of saddlebacks on Motuara Is- habitat type (Jenkins 1976; Craig 1994; B. Walter

93 J. P. PIERRE pers. comm.). Thus, it is expected that as the South bly flax were used more than expected. Foraging se- Island Saddleback population density increases on lectivity of high or low density North Island Saddle- Motuara Island, birds will colonise scrub areas. On back populations cannot be compared, due to lack of , the suitability of scrub may research. However, I expect that as population den- have been increased by the erection of roost and nest sity increases, South Island Saddlebacks may use a boxes (B. Walter pers. comm.). Thus, the addition of wider spectrum of plants to increase the efficiency of roost and nest boxes may enhance the suitability of foraging in smaller territories. Published data report- scrub areas for saddlebacks on Motuara Island. In any ing relative richness of locations of prey capture are case, the colonisation of scrub demonstrates the be- nonexistent for North Island Saddleback populations. havioural plasticity of saddlebacks. Both for saddle- However, as saddlebacks appear flexible with respect backs and other organisms, behavioural plasticity can to foraging strategies and diet, these are expected to be an important factor increasing the success of vary with habitat type as well as season, as are the translocations. importance of different foraging substrates and prey After release on Motuara Island, but before settling types (Atkinson 1964, 1966; Blackburn 1964, 1967; on a territory in a stable pair, some South Island Sad- Lovegrove 1980, 1992; O’Callaghan 1980; Pierre dlebacks were observed with more than one bird of 1995). the opposite sex. This has also been reported in South Island Saddlebacks on Motuara Island for- newly translocated North Island Saddleback popula- aged mostly in lower levels of the forest, like North tions (Armstrong & Craig 1995). Further, pairs of Island Saddlebacks at high density (Lovegrove 1980; North Island Saddlebacks translocated together did O’Callaghan 1980). However, high density popula- not maintain their pair bond after release (Armstrong tions of North Island Saddlebacks also display verti- & Craig 1995). Also, similar to my results for South cal stratification within pairs when foraging (Love- Island Saddlebacks, North Island Saddlebacks in low grove 1980; O’Callaghan 1980). This may function density populations are recorded breeding at one year to reduce intersexual competition within pairs, and old (Craig 1994). However, in high density popula- improve the efficiency of resource use in territories. tions, North Island Saddlebacks breed at two or more As increasing population density causes territory years of age (Lovegrove 1980). Further, South Island sizes to decrease on Motuara Island, birds may de- Saddlebacks were able to breed successfully with one velop vertically stratified foraging behaviour. Coinci- pair taking a maximum of only 35 days between pair- dent with this speculation, an increased degree of ver- ing and nest building. This pair fledged two chicks tical stratification in North Island Saddleback forag- and probably renested in the first breeding season ing behaviour has been related to decreases in terri- after translocation. Although there are no published tory size (Lovegrove 1980). records of the length of time taken from pairing to Although the diet of saddlebacks contains some nesting, newly released North Island Saddlebacks water, most of Motuara Island is very dry, and birds have also been reported to raise more than one brood used both natural and human-constructed water per breeding season (Jenkins 1976; Craig 1994; Arm- sources frequently. That saddlebacks used water strong & Craig 1995). sources outside their own territories suggests a very Like South Island Saddlebacks on Motuara Island, strong need for water, and as predicted prior to the North Island Saddlebacks appear to be flexible in translocation, the instalment of water troughs may terms of foraging substrates they can use (Atkinson have increased the quality of saddleback habitat on 1964, 1966; Lovegrove 1980). Male and female Motuara Island, possibly increasing the likelihood of South Island Saddlebacks had slightly different forag- success of the translocation. ing strategies in terms of substrate used and vertical Mortality rate is one of the most important factors location. Similarly, albeit at high density, North Is- determining the size of founder groups, making it a land Saddleback males spent more time foraging on key consideration when planning translocations. On the ground than their female counterparts (Blackburn Motuara Island, maximum mortality was 50% 8–10 1964; Lovegrove 1980; O’Callaghan 1980), however, months after saddlebacks were released. Published whether there are intersexual differences in their for- records of North Island Saddleback mortality range aging patterns at low density is unknown. Out of all from 8% at six months to 52% at two years after re- plant species (including dead wood) that saddlebacks lease (Jenkins 1976; Armstrong & Craig 1995). Natu- on Motuara Island foraged on, five-finger and possi- rally, mortality will vary due to site-specific charac-

94 South Island Saddleback reintroduction teristics as well as the ability of birds to deal with the nections to the world outside Motuara. I would like to stresses of capture and translocation, and their flexi- thank my field assistants for their hard work, especially bility in adapting to new environments. Weather con- Sally Truman and Maureen and Richard Pierre for help ditions after release and the abundance of natural both in the field and beyond. Figures were reproduced predators are other important considerations. with permission from Elsevier Science. In general, for a translocation into excellent quality habitat to have a 60% chance of success, the recom- REFERENCES mended size of the founder group is fifty individuals (Griffith et al. 1989). Only 26 South Island Saddle- Armstrong DP & Craig JL (1995) Effects of familiarity backs were released onto Motuara Island, but in the on the outcome of translocations, I. A test using sad- past, translocations of 15–59 South Island Saddle- dlebacks Philesturnus carunculatus rufusater. Biol backs have been successful (Nillson 1978; Roberts Conserv 71: 133–141. 1991). Thus, from the outset, the success of the trans- Armstrong DP & McLean IG (1995) New Zealand fer to Motuara Island may not have been jeopardized translocations: theory and practice. Pacific Conserv Biol 2: 39–54. by the relatively small founder group. In combination Atkinson IAE (1964) Feeding stations and food of with the predator-free environment of Motuara Is- North Island saddleback in August. Notornis 11: 93– land, the saddlebacks’ flexible habitat requirements 97. and foraging strategies, ability to readily adapt to Atkinson IAE (1966) Feeding stations and food of the ‘new’ habitats, and potential for high reproductive North Island saddleback in May. Notornis 13: 7–11. output increased the likelihood that translocations to Atkinson IAE & Bell BD (1973) Offshore and outlying this island would be successful. islands. In: Williams GR (ed) The Natural History of In the eight years since translocation, estimates of New Zealand. pp 372–392. AH & AW Reed, Welling- the number of saddlebacks Motuara Island can sup- ton. port have ranged from about 70 (in years of cold, wet Bell BD (1978) The Big South Cape Islands rat irrup- climatic conditions, when breeding success is low) to tion. In: Dingwall PR, Atkinson IAE & Hay C (eds) 150–200 (in the first years after release, with high The Ecology and Control of Rodents in New Zealand reproductive output, an abundance of prey and Nature Reserves. pp 33–40. Department of Lands and favourable climate) (Pierre 1995; W. F. Cash pers. Survey, Wellington. comm.). Social organisation, survival, foraging ecol- Blackburn A (1964) Some observations on behaviour of ogy and diet have not been investigated in detail the North Island saddleback in August. Notornis 11: since my work was completed. However, due to the 87–92. success of this translocation over the first eight years Blackburn A (1967) Feeding stations and food of the at least, the Motuara Island population is now being North Island saddleback in November. Notornis 14: used as a source population for other South Island 67–70. Saddleback translocations. Butler D & Merton D (1992) The Black Robin: Saving the Worlds’ Most Endangered Bird. Oxford Univer- sity Press, Auckland. ACKNOWLEDGMENTS Craig JL (1994) Meta-populations: is management as flexible as nature? In: Olney PJS, Mace GM & Feist- I wish to thank the Ornithological Society of Japan ner ATC (eds) Creative Conservation: Interactive for the invitation to present a symposium paper at their Management of Wild and Captive Animals. pp 50–66. 2002 Annual Meeting. I also extend thanks to the Royal Chapman and Hall, London. Forest and Bird Protection Society Waikato Branch, the Day RW & Quinn GP (1989) Comparisons of treat- Pacific Development and Conservation Trust, the Or- ments after an analysis of variance in ecology. Ecol nithological Society of New Zealand and the University Monogr 59: 433–463. of Canterbury for funding my study, and Dr. I. McLean Department of Conservation (1994) New Zealand trans- who proposed work on Motuara Island initially. The De- fer guidelines for indigenous terrestrial flora and partment of Conservation Permission granted me per- fauna. In: Serena M (ed) Reintroduction Biology of mission to stay on the island, and staff of the Depart- Australian and New Zealand Fauna. pp 253–255. ment of Conservation (especially Bill Cash), Cougar Surrey Beatty and Sons, Chipping Norton. Line, Dolphin Watch Marlborough, and the Sandpiper Dufty AC, Seebeck JH, McKay J & Watson AJ (1994) were a great help with field work, as well as valued con- Reintroduction of the eastern barred bandicoot Per-

95 J. P. PIERRE

ameles gunnii at Gellibrand Hill Park, Victoria. In: (Aves). M.Sc. Thesis, University of Auckland, Auck- Serena M (ed) Reintroduction Biology of Australian land. and New Zealand Fauna. pp 219–225. Surrey Beatty Oliver WRB (1955) New Zealand Birds. 2nd ed. AH & and Sons, Chipping Norton. AW Reed, Wellington. Griffith B, Scott JM, Carpenter JW & Reed C (1989) Pierre JP (1995) Behaviour, ecology and reintroduction Translocation as a species conservation tool: status biology of the South Island saddleback Philesturnus and strategy. Science 245: 477–480. carunculatus carunculatus. B Sc (Hons) Thesis, Uni- Guthrie-Smith H (1925) Bird Life on Island and Shore. versity of Canterbury, Christchurch. William Blackwood and Sons, London. Pierre JP (2000) Foraging behaviour and diet of a rein- IUCN (1987) The IUCN position statement on translo- troduced population of the South Island Saddleback cation of living organisms: Introductions, re-intro- (Philesturnus carunculatus carunculatus). Notornis ductions and re-stocking. IUCN, Gland. 47: 7–12. IUCN (2000) 2000 IUCN Red List of Threatened Pierre JP (1999) Reintroduction of the South Island sad- Species. IUCN, Gland and Cambridge. dleback (Philesturnus carunculatus carunculatus): Jenkins PF (1976) Social organisation and vocal behav- dispersal, social organisation and survival. Biol. Con- iour of the saddleback Philesturnus carunculatus ru- serv. 89: 153–159. fusater (Aves). Ph D Thesis, University of Auckland, Pierre JP (2001) Reintroduction of the South Island sad- Auckland. dleback (Philesturnus carunculatus carunculatus): Jenkins PF & Veitch CR (1991) Sexual dimorphism and habitat use including foraging patterns. Notornis 48: age determination in the North Island saddleback 63–71. (Philesturnus carunculatus rufusater). N Z J Zool 18: Roberts A (1991) A Recovery Plan for the South Island 445–450. Saddleback. Department of Conservation, Southland. Lovegrove TG (1980) The saddleback pair bond. M Sc Sarrazin F & Barbault R (1996) Reintroduction: chal- Thesis, University of Auckland, Auckland. lenges and lessons for basic ecology. TREE 11: 474– Lovegrove TG (1992) The effects of introduced preda- 478. tors on the saddleback (Philesturnus carunculatus), Scott JM & Carpenter JW (1987) Release of captive- and implications for management. Ph D Thesis, Uni- reared or translocated endangered birds: What do we versity of Auckland, Auckland. need to know? Auk 104: 544–545. Lovegrove TG & Veitch CR (1994) Translocating wild Sherley G (1994) Translocations of the Mahoenui giant forest birds. Ecological Management 2: 23–35. weta Deinacrida n. sp. and Placostylus land snails in Merton D (1973) Conservation of the saddleback. New Zealand: What have we learnt? In: Serena M Wildlife: A Review 4: 13–23. (ed) Reintroduction Biology of Australian and New Merton D (1975) The saddleback: its status and conser- Zealand Fauna. pp 57–63. Surrey Beatty and Sons, vation. In: Martin RD (ed) Breeding Endangered Chipping Norton. Species in Captivity. pp 61–74. Academic Press, Lon- Short J, Turner B, Parker S & Twiss J (1994) Reintro- don. duction of endangered mammals to mainland Shark Miller RG (1981) Simultaneous statistical inference. Bay: a progress report. In: Serena M (ed) Reintroduc- McGraw Hill, New York. tion Biology of Australian and New Zealand Fauna. Murphy SM (1989) The interaction of social organisa- pp 183–188. Surrey Beatty and Sons, Chipping Nor- tion and parentage with vocalisation in the North Is- ton. land saddleback. M Sc Thesis, University of Auck- SPSS Inc. (1997) SPSS for Windows. SPSS Inc., land, Auckland. Chicago. Nelson NJ, Keall SN, Brown D & Daugherty CH (2002) Wolf CM, Griffith B, Reed C & Temple SA (1996) Establishing a new wild population of tuatara (Sphen- Avian and mammalian translocations: update and re- odon guntheri). Conserv Biol 16: 887–894. analysis of 1987 survey data. Conserv Biol 10: 1142– Nillson RJ (1978) The South Island saddleback. 1154. Wildlife: A Review 9: 32–36. Zar JH (1996) Biostatistical Analysis. 3rd ed Prentice O’Callaghan AP (1980) Use of space by the North Is- Hall, Upper Saddle River. land saddleback Philesturnus carunculatus rufusater

96