New Zealand pohutukawa (Metrosideros excelsa), a member of the Myrtaceae, is a large, mass-flowering tree endemic to northern New Zealand coastlines. Mainland populations have been reduced to fragmented stands, and the original suite of bird pollinators has been largely replaced by introduced species. The native pollinator fauna on several offshore islands is largely intact and includes three species of the New Zealand honeyeaters (Meliphagidae) and native, solitary bees. We estimated multilocus outcrossing rates for three mainland and two island populations and found that they were among the lowest in the Myrtaceae (t(m) = 0.22-0.53). The shift in pollinators had no measurable effect on the mating system. Mass-flowering facilitates geitonogamous selfing, and inbreeding depression in seedling height was detectable at 6 mo of growth. F(s) [Wright's (1965) Fixation Index] was consistently higher than F(m) in all populations, indicating that selection may eliminate selfed offspring from populations prior to achieving reproductive maturity. Results suggest that increased selfing in mainland populations due to pollinator changes is not responsible for current patterns of poor regeneration of this species.

The floral biology and breeding system of pohutukawa (Metrosideros excelsa, Myrtaceae), a mass-flowering tree of northern New Zealand coastlines, were examined. Trees flower over a peak period of 2 weeks, and compound inflorescences contain an average of 14.3 showy, hermaphrodite, red brush flowers that remain open for 7 days. A brief female flower stage (mean duration 1.3 d) is followed by the main hermaphrodite phase that lasts for 4 days. Neither dichogamy nor herkogamy is important in preventing pollen and stigma interference. Pollen is highly viable (93.6%), and stigma receptivity extends for at least 9 days, as indicated by peroxidase activity, pollen germination, pollen tube length 24 h after pollination, and seed set. Stigmatic exudate production appears to increase up to 5 days post-anthesis. On average, flowers produce 46 p1 nectar per day, containing 18% (w/v) sucrose. Floral design and display of pohutukawa are consistent with high levels of autogamous and geitonogamous self-pollination. Controlled pollination experiments were used to assess the effect of self-and crosspollen and a pollen mixture from five unrelated parents on capsule and seed production, and on pollen tube growth in seven trees. Three trees in the experiment were self-incompatible, as quantified by the index of self-incompatibility (ISI), indicating that natural populations may consist of a mosaic of selfcompatible and incompatible individuals. Self-in- B99002Received 22 January 1999; accepted 28 July 1999 compatibility is late-acting as pollen tubes from selfs and crosses reached the ovary simultaneously at 10-15 d after pollination. In common with other Myrtaceae, the seed/ovule ratio in pohutukawa is low and this is likely to be genetically determined rather than limited by stigmatic pollen load. Germination of fertile seeds from all pollination treatments was equally high (98.4%), indicating that no inbreeding depression is acting at this stage of the life cycle. The pollen/ovule ratio of 462.5 (s.e. ±43.3) places the breeding system of pohutukawa between facultative selfing and facultative outcrossing.

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Exclusion experiments were used to assess the effect of different pollinator groups on outcrossing and seed production in Metrosideros excelsa. The main study site was Little Barrier Island, New Zealand where indigenous bird and native solitary bees are the main flower visitors. Our results showed that native birds were more important pollinators of M. excelsa than native bees. Seed production was much higher in open pollination than in two exclusion experiments where either birds were excluded and native bees only had access to flowers, or where all pollinators had been excluded. The number of fertile seeds per capsule was 45% higher after open pollination than in treatments with bee visitation only and 28% higher than in treatments where all flower visitors were excluded. Estimated outcrossing rates were significantly higher (tm = 0.71) for open pollination in the upper canopy (>4 m above-ground level) where bird visitation is presumed to be more frequent than for a treatment with native bee access only (tm = 0.40). Our results also suggest that a large proportion of seeds (66%) arise from autonomous self-pollination when all pollinators are excluded. In four trees of a modified mainland population with predominantly introduced birds and a mixture of introduced and native bees there was no decrease in seed production for the treatment allowing bee access only, indicating that -in contrast to native bees -honeybees may be more efficient pollinators of M. excelsa. Observation of the foraging behaviour of both groups of bees showed that native bees contact the stigma of flowers less frequently than honeybees.This is likely to be a consequence of their smaller body size relative to honeybees.