Effect of population size on viable seed output, seed rand and natural regeneration pattern of a tropical conifer Widdringtonia whytei-Rendle in Malawi

Thesis (PhD(For))--Stellenbosch University, 2013.ENGLISH ABSTRACT: Widdringtonia whytei is a tropical endemic, fire-adapted pioneer coniferous tree species within natural fire-fragmented Afromontane forest patches in a confined area on Mulanje Mountain in Malawi. Natural and anthropogenic fires within the surrounding fire-prone landscape, insect attacks, and uncontrolled harvesting of mature trees for timber threaten the survival of W. whytei. This study investigated the effects of population fragmentation on the reproductive biology of W. whytei, through four specific studies: effects of population size, tree size and crown position on viable seed output; seed rain variation among population sizes; effects of temperature and light on viability and germination of W. whytei seeds; and the influence of population size on natural regeneration patterns. The study was conducted during 2008 and 2009 on three sites using three W. whytei population sizes: small (≤10 reproductively mature cone-bearing W. whytei trees), medium (11-20 cone-bearing trees) and large (>20 cone-bearing trees). Data were collected using field and laboratory experiments and a review of secondary information. The effect of fragment size and crown position on viable seed output and seed rain pattern was tested with a nested linear mixed model. The influence of stem diameter (dbh) on viable seed output was tested using a separate data set. The variation in seed rain was explored with a generalised linear model (GLM) with a negative binomial link function. Variations in seed germination was analysed with a Weibull regression model. The influence of fragment size on seedling regeneration was tested with a GLM (binary logistic regression). Viable seed output from W. whytei cones was very low (23%) and was not affected by population size, tree diameter or crown position. W. whytei population size influenced seed rain with large fragments having higher seed rain densities. Number of dispersed seeds was generally very low and poorly dispersed. W. whytei cones are semiserotinous and depends on moderate to severe fires for heavy seedfall and wider dispersal. Such fires did not occur during the study period. Temperature is a critical factor for seed germination. Seeds germinate between 15 and 25°C with the optimum at ~ 20°C either under light or dark conditions. Regeneration density followed the seed rain pattern indicating that seed availability is a critical factor for regeneration. Regular cool fires outside the forest patches and along the edges during the hot dry season, and the thick litter layer and shady conditions inside forest patches, caused most of the seedling mortality. This study highlighted the difficulties W. whytei experiences with low output of viable seed, limited seed dispersal and low seedling regeneration in and around forest patches, i.e. factors important for conservation management of this species. It is recommended that gaps with diameter equal to canopy height and occasional spot fires would promote seed dispersal onto exposed mineral soil for rapid seedling establishment and subsequent population growth. Such natural fire disturbance events may show the true seed rain and seedling establishment patterns in this species.AFRIKAANSE OPSOMMING: Widdringtonia whytei is ‘n tropiese endemiese, vuuraangepaste pionier naaldhoutagtige boomsoort binne die natuurlike vuurgefragmenteerde Afromontane woudkolle binne ‘n beperkte area op Mulanje berg in Malawi. Natuurlike en menslike vure binne die omringende vuurgeneigde landskap, insekaanvalle, en onbeheerde kap van volwasse bome vir hout bedreig die voortbestaan van W. whytei. Hierdie studie het die effek van populasiefragmentasie op die voortplantingsbiologie van W. whytei ondersoek, deur vier spesifieke studies: die effek van populasiegrootte, boomgrootte en kroonposisie op produksie van lewenskragtige saad; saadreënvariasie tussen populasiegroottes; die effek van temperatuur en lig op lewenskragtigheid en ontkieming van W. whytei saad; en die invloed van populasiegrootte op natuurlike verjongingspatrone. Die studie is gedurende 2008 en 2009 uitgevoer op drie groeiplekke met drie W. whytei populasiegroottes: klein (≤10 keëldraende (volwasse) W. whytei bome), medium (11-20 keëldraende bome) en groot (>20 keëldraende bome). Data is versamel in veld- en laboratorium eksperimente asook 'n oorsig van sekondêre inligting. Die invloed van populasie fragmentgrootte en kroonposisie van bome op saadproduksie en die saadreënpatroon is getoets met ‘n geneste liniêre gemengde model. Die invloed van boomstamdeursnee op die produksie van kiemkragtige saad is met ‘n aparte datastel getoets. Die variasie in saadreënpatroon is met 'n algemene liniêre model (GLM) met 'n negatiewe binomiese skakelfunksie getoets. Variasies in ontkiemingspersentasie van sade is met 'n Weibull regressiemodel ontleed. Die invloed van fragmentgrootte op saailingverjonging is met 'n algemene linêre model (binêre logistiese regressie) getoets. Produksie van lewenskragtige saad in W. whytei keëls was baie lag (23%) en was nie deur populasiegrootte, boomstamdeursnee of kroonposisie beïnvloed nie. W. whytei populasiegrootte het saadreën beïnvloed en groot fragmente het groter saadreëndigthede gehad. Die aantal verspreide sade was in die algemeen baie laag en swak versprei. W. whytei keëls is semi-saadhoudend (serotinous) en is afhanklik van matige tot intense vure vir massiewe saadvrystelling en wyer saadverspreiding. Sulke vure het nie gedurende die studieperiode voorgekom nie. Temperatuur is ‘n kritiese factor vir saadontkieming. Sade ontkiem goed tussen 15 en 25°C met die optimum rondom ~ 20°C onder beide lig en donker toestande. Fragmentgrootte het nie die saailingverjonging in W. whytei beïnvloed nie. Verjongingsdigtheid het die saadreënpatroon gevolg, wat daarop dui dat saadbeskikbaarheid ‘n kritiese factor is vir saailingverjonging. Gereelde koel vure buite die woudkolle en langs die woudrand gedurende die droë warm seisoen, en die dik lae blaarval (litter) en skadutoestande binne ‘n woud, het meeste van die saailingmortaliteit veroorsaak. Hierdie studie het die probleme beklemtoon wat W. whytei ondervind met die lae produksie van kiemkragtige saad, saadverspreiding en saailingverjonging binne en rondom die woudkolle, m.a.w. faktore wat belangrik is vir bewaringsbestuur van hierdie boomsoort. Die aanbeveling word gemaak dat kroonopeninge met ‘n deursnee gelyk aan minstens die kroonhoogte, en sporadiese kolbrande binne die woudkolle kan saadverspreiding op blootgestelde minerale grond vir vinnige saailingvestiging en gevolglike populasiegroei bevorder. Sodanige vuurversteuringsgeleenthede mag die werklike patrone in saadreën en saailingvestiging in hierdie species toon

Germination response and viability of an endangered tropical conifer Widdringtonia whytei seeds to temperature and light

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Data from: Mediation of seed provisioning in the transmission of environmental maternal effects in Maritime pine (Pinus pinaster Aiton)

Although maternal environmental effects are increasingly recognized as an important source of phenotypic variation with relevant impacts in evolutionary processes, their relevance in long-lived plants such as pine trees is largely unknown. Here, we used a powerful sample size and a strong quantitative genetic approach to analyse the sources of variation of early seedling performance and to identify seed mass (SM)-dependent and -independent maternal environmental effects in Maritime pine. We measured SM of 8924 individual seeds collected from 10 genotypes clonally replicated in two environments of contrasting quality (favourable and stressful), and we measured seedling growth rate and biomass allocation to roots and shoots. SM was extremely variable (up to 14-fold) and strongly determined by the maternal environment and the genotype of the mother tree. The favourable maternal environment led to larger cones, larger seeds and reduced SM variability. The maternal environment also determined the offspring phenotype, with seedlings coming from the favourable environment being 35% larger and with greater root/shoot ratio. Transgenerational plasticity appears, thus, to be a relevant source of phenotypic variation in the early performance of this pine species. Seed provisioning explained most of the effect of the maternal environment on seedling total biomass. Environmental maternal effects on seedling biomass allocation were, however, determined through SM-independent mechanisms, suggesting that other epigenetic regulation channels may be involved