Effects of Habitat Structure
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D.R. TOWNS1 and G.P. ELLIOTT2 191 1 Science and Research Division, Department of Conservation, Private Bag 68-908, Newton, Auckland, New Zealand 2 549 Rocks Road, Nelson, New Zealand EFFECTS OF HABITAT STRUCTURE ON DISTRIBUTION AND ABUNDANCE OF LIZARDS AT PUKERUA BAY, WELLINGTON, NEW ZEALAND __________________________________________________________________________________________________________________________________ Summary: The distribution and abundance of lizards relative to habitat structure were studied at Pukerua Bay, Wellington between December 1982 and March 1988 in order to identify options for management of the habitat of the five species of lizards present. One species, Whitaker’s skink (Cyclodina whitakeri), is a threatened species with only one known mainland population. Pitfall traps were set for 23 667 trap-days and yielded 2897 lizard captures. Highest capture rate was for common skinks (Oligosoma nigriplantare polychroma) and lowest rate was for C. whitakeri. Of the five lizard species at Pukerua Bay, C. whitakeri had the narrowest habitat range and was most sensitive to temperature extremes. These features, combined with predation, and habitat disturbance and degradation, have resulted in critically low numbers of C. whitakeri. Assessment of management options to lessen the threats to C. whitakeri concluded that risks from disturbance, predation and fire could best be minimised through a managed revegetation programme. __________________________________________________________________________________________________________________________________ Keywords: skinks; Cyclodina aenea; Cyclodina whitakeri; Oligosoma nigriplantare polychroma; Oligosoma zelandicum; geckos; Hoplodactylus maculatus; vegetation; substrate; temperature; livestock; revegetation; predators; fire. Introduction (Cyclodina whitakeri). Much of the Pukerua Bay site has recently been designated a Department of The New Zealand archipelago supports an unusually Conservation Scientific Reserve with the long term diverse fauna of lizards when compared with intention of restoring the area to coastal forest. temperate environments elsewhere (Daugherty et al., However, because of differences in habitat use, not 1990c). Before humans arrived about 1000 years ago, all lizard species present at Pukerua Bay will the highest local diversities were probably in warm necessarily benefit from reforestation (East, East and lowland forests, especially those near the coast. Daugherty, 1995). Also, other recent environmental Recent palaeoecological analyses have revealed that changes (such as the presence of introduced assemblages of up to 17 sympatric species of lizards, predators) could complicate the effects of restoration including at least six species of Cyclodina1, were on the lizards (Towns, 1991, 1994, in press). once present in Northland (Towns and Daugherty, In this paper we describe the Pukerua Bay site 1994). As introduced predatory mammals spread and its lizard assemblage. We provide estimates of following human colonisation, these large primeval the population densities of each species, investigate lizard assemblages disappeared from mainland New the relationship between lizard density and Zealand; highest species diversities are now found on environmental characteristics, discuss how reserve offshore islands (Towns and Atkinson, 1991; Towns management could reflect the habitat needs of the and Daugherty, 1994). lizard species, and identify how habitat management Part of a coastal forest lizard assemblage has could benefit Cyclodina whitakeri, the rarest species persisted at Pukerua Bay near Wellington despite in the assemblage (Towns, 1985; Molloy and Davis, extreme habitat modification. The assemblage is 1994). unique because it includes the only known remaining mainland population of Whitaker’s skink Study area ______________________________________________________________ The study site was between Pukerua Bay township 1 Nomenclature for reptiles follows Robb and Rowlands and Wairaka Point, on steep coastal hill sides formed (1977), Hardy (1977), Daugherty et al. (1990a) and from an escarpment that faces directly north and Patterson and Daugherty (1995). rises steeply to 160 m a.s.l. (Fig. 1). The escarpment New Zealand Journal of Ecology (1996) 20(2): 191-206 ©New Zealand Ecological Society 192 NEW ZEALAND JOURNAL OF ECOLOGY, VOL. 20, NO. 2, 1996 includes greywacke bluffs, screes and talus slopes of Lizards present rocks and boulders. At the foot of some slopes are narrow talus areas (10-20 m wide) and rocks and Five species of lizard persist at Pukerua Bay: large boulders are scattered along old beach terraces. 1. Common gecko (Hoplodactylus maculatus): a The coastline is also rocky, with narrow beaches of nocturnal species that is geographically and rocks, stones and gravel. ecologically ubiquitous and may comprise By 1982, the study area had been farmed at low several cryptic species (Daugherty, Patterson and intensity for at least 100 years and sheep (Ovis aries)2 Hitchmough, 1994). The Pukerua Bay taxon is had access throughout. There was also evidence of common throughout the North Island and repeated fires on the northeastern cliffs. Vegetation northern South Island (R. Hitchmough, pers. on the escarpment and along the shoreline therefore comm.; Victoria University of Wellington, N.Z.). formed a mosaic which in northeast - southwest 2. Common skink (Oligosoma nigriplantare sequence was (C. Ogle, pers. comm.; Department of polychroma): a widespread diurnal species that Conservation, Wanganui, N.Z.): is part of a complex of closely related species 1. Montpellier broom (Teline monspessulana)3 - revised by Patterson and Daugherty (1990). The Cape ivy (Senecio angulatus)-boneseed complex shows a high degree of morphological, (Chrysanthemoides monilifera) shrubland and ecological and genetic diversity (Daugherty et mountain flax (Phormium cookianum) on steep al., 1990b), but genetic analyses indicate no faces and burn scars. evidence of more than one taxon at Pukerua Bay 2. Small pockets and isolated plants of puka (C.H. Daugherty, pers. comm.; Victoria (Griselinia lucida) and karaka (Corynocarpus University of Wellington, N.Z.). laevigatus) at the foot of bluffs (sometimes with 3. Brown skink (Oligosoma zelandicum): a diurnal Olearia paniculata), and scattered along the species present only in the southern third of the lower slopes. North Island and in the northern South Island 3. A remnant stand of karaka trees with the (Pickard and Towns, 1988). In the Wellington understorey eaten out by stock. and Manawatu areas O. zelandicum is 4. A patch of coastal broadleaf forest dominated frequently sympatric with O. n. polychroma, but by kohekohe (Dysoxylum spectabile) on a ridge O. zelandicum has the narrower habitat range crest with the understorey eaten out by stock. (Gill, 1976). 5. Kanuka (Kunzia ericoides) shrubland mixed 4. Copper skink (Cyclodina aenea): like all species with flax on southwestern bluffs. of Cyclodina, confined to the North Island 6. Coprosma propinqua shrubland intermixed with where it is the most widespread species in the pohuehue (Muehlenbeckia complexa), reverting genus (Pickard and Towns, 1988). It is one of grassland, scattered tree nettle (Urtica ferox), New Zealand’s smallest lizards and is regarded and occasional kawakawa (Macropiper as both crepuscular and diurnal (Porter, 1987). excelsum) and kaikomako (Pennantia 5. Whitaker’s skink (Cyclodina whitakeri): apart corymbosa) on stable slopes and along the old from the Pukerua Bay population, was known beach terrace. only from three offshore islands that lack When this study began the northeastern portion introduced mammals (Towns and Robb, 1986), of the study area was a Recreation Reserve but is now being reintroduced to islands from administered by Porirua City Council. A further 12.3 which predators have been removed (Towns, ha of escarpment adjacent to the Recreation Reserve 1994). C. whitakeri is the largest species at was purchased from its owners by the Crown in Pukerua Bay (up to 20 cm total length), is 1984 and a sheep-proof fence was completed by nocturnal, prone to cutaneous water loss (A. March 1987. Wandering goats (Capra hircus) Cree and C.H. Daugherty, pers. comm.; Victoria entered the study area for the first time in 1986 and University of Wellington, N.Z.), and uses warm are now periodically culled. Few stock have entered moist environments, such as rocky areas and the Crown area or the Recreation Reserve since seabird burrows, as daytime retreats (Macredie, March 1987. 1984; Towns, 1994). ______________________________________________________________ Methods 2 Nomenclature for mammals follows King (1990). 3 Nomenclature for plants follows Moore and Edgar Study sites (1970), Lambrechsten (1981), Allan (1982), Connor and Edgar (1987) and Webb, Sykes and Garnock-Jones The main study grid was in the Crown reserve where (1988). exploratory 20 m transect lines (every 20 m along TOWNS and ELLIOTT: DISTRIBUTION AND ABUNDANCE OF LIZARDS 193 Figure 1: The Pukerua Bay study site (A); localities mentioned in the text (B), with the greater Wellington urban areas shaded; and location of the reserves (C). 194 NEW ZEALAND JOURNAL OF ECOLOGY, VOL. 20, NO. 2, 1996 260 m of lower slope and beach platform) revealed The following data were obtained so that MNA the highest frequency of C. whitakeri captures could be calculated for lizards at Pukerua Bay in (December 1982 - May 1983). Regular sampling on year one: the study grid began in December 1983 and 1. Mean