Germination of drupelets in multi-seeded drupes of the shrub Leptecophylla tameiameiae (Ericaceae) from Hawaii: A case for deep physiological dormancy broken by high temperatures

This study addressed the difficulty of germinating drupelets (hereafter seeds) in the multi-seeded stony dispersal units (drupes) of Leptecophylla tameiameiae (Ericaceae). Embryos in fresh seeds were 77% the length of the endosperm, and seeds inside the intact drupes imbibed water. We monitored germination at 15/6, 20/10 and 25/15°C for 162 weeks, after which each drupe was cut open and ungerminated seeds counted. Drupes contained 1-6 seeds, and the total number of seeds in all treatments and controls was 1977, with 20, 29, 25, 18, 7 and < 1% of them occurring in one-, two-, three-, four-, five- and six-seeded drupes, respectively. The percentage of seeds germinating in one-, two-, three-, four-, five-and six-seeded drupes was 74, 66, 65, 72, 56 and 0, respectively. Neither warm nor cold stratification for 6 or 12 weeks significantly increased germination percentages, compared to controls incubated continuously at 25/15°C for 162 weeks, where 72% of the seeds in the drupes germinated. At 25/15°C, 24-49 weeks were required for 20% of the seeds to germinate. Warm followed by cold stratification did not promote germination, and there was no widening of the temperature range for germination. Like seeds of other species known to have deep physiological dormancy (PD), those of L. tameiameiae required extended periods of time (16 to ≥ 162 weeks) to come out of dormancy and germinate, gibberellic acid (GA₃) did not promote germination and excised embryos failed to grow. Thus, we conclude that seeds of L. tameiameiae have deep PD. However, unlike seeds of other species with deep PD, those of L. tameiameiae required an extensive period of warm rather than of cold stratification to come out of dormancy. It is suggested that a subtype a (seeds require a long period of cold stratification to come out of dormancy) and a subtype b (seeds require a long period of exposure to warm stratification to come out of dormancy) of deep PD be recognized in the Nikolaeva formula system for classifying seed dormancy.