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Effect of Light on the Germination of Forest Trees in Ghana

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Effect of Light on the Germination of Forest Trees in Ghana Journal of Eect of light on the germination of forest trees in Ecology 1999, 87, 772±783 Ghana B. KYEREH*, M. D. SWAINE{ and J. THOMPSON{{ *Institute of Renewable Natural Resources, University of Science and Technology, Kumasi, Ghana; {Department of Plant and Soil Science, University of Aberdeen, Aberdeen AB24 3UU, UK; and {Forestry Research Institute of Ghana, University PO Box 63, Kumasi, Ghana Summary 1 Seed germination in light and dark, and responses to irradiance and light quality, were tested in shadehouse experiments for 19 West African tropical forest tree spe- cies representing a wide range of ecological types. Germination in forest gaps of dierent size was tested for 11 species. 2 Percentage germination was reduced in the dark only for three small-seeded spe- cies that are common in forest soil seed banks: Musanga cecropioides, Nauclea diderrichii and Milicia excelsa. Percentage germination of the other 16 species, including four widely regarded as `pioneers', was unaected. 3 Eects of dierent irradiances in shadehouses, where the seeds were watered, were signi®cant for some species, but there was no consistent pattern. Irradiance eects in forest gaps, where the seeds received only natural wet season rainfall, were more widespread and substantial, and were most commonly shown as a depression of percentage germination at high irradiance. 4 Eects of light quality (neutral vs. green shade; red : far-red = 0.43) were insignif- icant at 5% irradiance in shadehouses for all species except Nauclea diderrichii.In growth chamber experiments, the low energy response was only evident at 1.0 mmol m±2 s±1 (< 1% of unshaded forest irradiance) in Musanga and Nauclea. 5 The speed of germination was aected by irradiance in many species, but the eect was small compared with dierences between species, in which time to com- plete germination varied between 3 weeks and over 6 months. 6 Seeds of Ceiba pentandra and Pericopsis elata planted in deep forest shade (2% irradiance) and in a small gap (30% irradiance) germinated well in both sites, showed exponential biomass growth in the gap but a linear decline in mean seed- ling biomass and subsequent death in deep shade. 7 Light-mediated germination is relatively rare among these forest trees, even among pioneers, so that the working de®nition of a pioneer should be seen to depend more on a species' ability to survive in forest shade. The eects of canopy opening on seed germination are small except in the largest openings, which severely depress germination in a number of species, including some species with strongly light-demanding seedlings. Key-words: irradiance, photoblastic seeds, pioneers, red : far-red ratio Journal of Ecology (1999) 87, 772±783 Introduction butions have been made by Va zquez-Ya nes and co- workers in Mexico (Va zquez-Yanes 1977, 1980; Environmental in¯uences on the germination of tro- Va zquez-Yanes & Orozco-Segovia 1982a, b, 1984, pical forest species have received much less attention 1990, 1993; Va zquez-Yanes et al. 1990). This may, in than temperate species, although signi®cant contri- part, be due to the perception that most timber tree species from tropical rain forest have recalcitrant # 1999 British Correspondence: M.D. Swaine (fax 01224 272703; e-mail seeds (sensu Roberts 1973; Corbineau & Comb Ecological Society [email protected]). 1989), which have no dormancy, rapid germination 773 and short viability. With typically high seed moist- able from ®eld experience in any tropical forest B. Kyereh, ure content, recalcitrant seeds are dicult to store, region, but the de®ning characteristics have not M.D. Swaine & but if collected fresh they are easy to germinate and been widely tested by experiment. Raich & Gong J. Thompson establish in nurseries and germinate abundantly in (1990) planted seeds of 43 Malaysian tree species in natural forest. The contrasting seed type, known as forest shade, as well as in small and large canopy orthodox because their seeds can be readily stored gaps, and recorded percentage germination. They in arti®cial conditions, have low seed moisture and recognized three species groups: those that showed generally show various degrees of dormancy. higher germination in gaps, those with higher germi- Recalcitrant seeds are characteristic of non-pioneer nation in forest shade and those that germinated (climax) species, while orthodox seeds are associated well both in shade and in gaps. There were, how- with pioneer species (sensu Swaine & Whitmore ever, no clear discontinuities between the three cate- 1988), which include relatively few timber species gories, and in some cases there were marked (Whitmore 1983). dierences in response to canopy cover between dif- This dichotomy among tropical forest tree species ferent seed collections of the same species. is a useful generalization, but conceals considerable However, there are several reports of pioneers diversity of species' germination response. Garwood germinating in deep forest shade (e.g. Endospermum (1983), for example, has shown considerable varia- peltatum in Sabah; Kennedy & Swaine 1992). Ceiba tion both between and within species in the period pentandra was observed by M. Dike and D. U. U. of seed dormancy following dispersal, and suggested Okali (personal communication) to germinate in for- that this often allows the seed to germinate at the est shade in Nigeria; and Cecropia spp. were the most favourable season for seedling establishment. most abundant germinants in the understorey of a The speed of germination is also very variable: in Costa Rican rain forest (Li et al. 1996). Such obser- 335 Malaysian tree species, Ng (1980) showed that vations cast doubt on the gap-dependence of pio- the time taken to complete germination varied neers for germination, and thus on the naturalness between 1 week and more than 1 year. of the dichotomy between them and non-pioneers. If Swaine & Whitmore (1988) proposed that tropical this widely recognized and frequently applied forest tree species could be divided into two guilds dichotomy is to be preserved, it becomes more (functional groups; sensu Gitay & Noble 1997), the dependent on the second of the Swaine & Whitmore pioneers and non-pioneers, distinguished by the (1988) criteria for pioneers, the seedling's require- dependence of pioneers on canopy gaps for seed ger- ment for high irradiance. Thus, although pioneers mination and seedling establishment. The implica- may germinate in deep shade, they will usually be tion was that pioneers had seed dormancy that was overlooked and very small seedlings die swiftly in broken by gap conditions (increased irradiance and low irradiance (Kennedy & Swaine 1992). red : far-red ratio, elevated or ¯uctuating tempera- The environmental conditions that break seed tures), associated with a requirement of their seed- dormancy in tropical forest tree seeds may be a high lings for high irradiance, so that the guilds were red : far-red ratio (Va zquez-Ya nes 1977, 1980; naturally distinct. The hypothesis was based on the Orozco-Segovia & Va zquez-Ya nes 1989; Valio & circumstantial evidence that pioneer species were Joly 1979), high maximum temperature or tempera- never found as young seedlings in forest shade, and ture range (Va zquez-Ya nes & Orozco-Segovia only occurred as young plants in canopy gaps or 1982a), ¯uctuating soil moisture (e.g. Terminalia sp.; other disturbed areas such as roadsides and farms. Taylor 1960) or some combination of these. All of Various studies have shown that some pioneer spe- these changes in microclimate can occur following cies require light for germination (photoblasticity) canopy opening, and because they are substantial (Valio & Joly 1979; Va zquez-Yanes & Smith 1982; may also reduce seed germination in non-dormant Orozco-Segovia & Va zquez-Yanes 1989; Va zquez- (non-pioneer) species by causing desiccation or heat- Yanes et al. 1990), which lends support for pioneer ing (insolation). Such environmental eects have gap-dependence determined by seed germination. implications for forest management: exploitation of Hawthorne (1993, 1995) provided a subdivision of tropical forest for timber commonly creates large the more speciose non-pioneer guild on the basis of gaps that appear to favour pioneer species over non- seedling ability to survive and grow in deep forest pioneers, because of the dierential eect on seed shade. In such conditions, seedlings of non-pioneer germination and on seedling growth and survival. light demanders (NPLD) have higher mortality, The present study set out to test if West African most dying before exceeding a metre or so in height, pioneer trees have gap-dependent germination as while non-pioneer shade bearers (NPSH) have lower proposed by Swaine & Whitmore (1988), or if their mortality and continue to grow in deep shade, even- failure to establish is due to negative carbon balance # 1999 British tually becoming established trees. in young seedlings. Experimental testing is necessary Ecological Society Journal of Ecology, The pioneer/non-pioneer dichotomy has the because existing allocation to the pioneer guild has 87, 772±783 advantage that the groups are readily distinguish- been based on the subjectively perceived distribution 774 of young plants in forest habitats. We used an has a large and persistent soil seed bank (Hall & Light and experimental approach in shadehouses, growth Swaine 1980) of very small seeds and is an abundant germination of chambers and forest gaps to determine germination colonizer of abandoned farmland in wet forest. forest trees responses to light in West African tree species, cho- Terminalia ivorensis and Terminalia superba are also sen to include both pioneers and a range of non-pio- fast-growing pioneers but achieve larger size and neers as previously categorized by Hall & Swaine their larger seeds (in winged fruits) are uncommon (1981) and Hawthorne (1995). Our strategy was to in soil seed banks. Like Musanga, their seedlings are ®nd out ®rst if the species were photoblastic, in intolerant of shade.
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