Plucking or cutting Gelidium sesquipedale? A demographic simulation of harvest impact using a population projection matrix model.

A matrix model describes the annual dynamics of a commercial (harvested by plucking) Gelidium sesquipedale population off Cape Espichel, Portugal. Vital rates were measured from a frond population divided into size classes; annual transition probabilities among them were calculated. Transition probabilities under harvest by cutting are derived by assuming that all harvested fronds are cut to the first size class, and none are plucked. Simulations of the annual population dynamics for harvest by both plucking and cutting are used to assess which harvest strategy will optimize yields. Assuming the same efficiency for both strategies, cutting fronds to 7 cm (as mechanical harvesters do) results in a higher population growth rate ( = 1.08 to 1.35) than occurs with the plucking technique ( = 0.85). Simulations of population recovery show the number of fronds in each size class available the next harvest season will be higher when cut than plucked. This model can also optimize yields by predicting the more efficient season opening, and harvesting cutting height.     

Regeneration and reproduction of Mazzaella cornucopiae (Rhodophyta, Gigartinaceae) after frond harvesting

The effects that different intensities of frond harvesting have on frond regeneration and subsequent production of reproductive structures were investigated for the red intertidal alga Mazzaella cornucopiae (Postels & Ruprecht) Hommersand from British Columbia, Canada. Harvesting was done by pruning fronds in the late spring (when stand biomass is highest) of 1993 at two intensities: total and partial collection of fronds, in this second case leaving all frond biomass less than 1 cm high in place. Holdfasts were not damaged. Total percent cover of thalli, frond density, mean frond length, and stand biomass for these experimental quadrats were statistically similar to values for control quadrats in the spring of 1994. These results suggest that one total harvest of fronds per year, done in late spring without damaging holdfasts, may give the highest sustainable yield of biomass. The effects of harvesting intensity on reproduction were variable and difficult to explain. Neither the appearance nor the abundance of cystocarpic fronds were affected by frond pruning, compared with control areas, but pruning did affect the appearance and the abundance of tetrasporic fronds. Partial pruning resulted in a longer presence of tetrasporic fronds, whereas total pruning was associated with their complete absence. Results are compared with those for the few other species of the Gigartinaceae for which experimental harvesting has been done. This revised version was published online in August 2006 with corrections to the Cover Date.     

An adaptive management approach to an octocoral fishery based on the Beverton-Holt model

Coral reef species are frequently the focus of bio-prospecting, and when promising bioactive compounds are identified there is often a need for the development of responsible harvesting based on relatively limited data. The Caribbean gorgonian Pseudopterogorgia elisabethae has been harvested in the Bahamas for over a decade. Data on population age structure and growth rates in conjunction with harvest data provide an opportunity to compare fishery practices and outcomes to those suggested by a Beverton-Holt fishery model. The model suggests a minimum colony size limit of 7–9 years of age (21–28 cm height), which would allow each colony 2–4 years of reproduction prior to harvesting. The Beverton-Holt model assumes that colonies at or above the minimum size limit are completely removed. In the P. elisabethae fishery, colonies are partially clipped and can be repeatedly harvested. Linear growth of surviving colonies was up to 3 times that predicted for colonies that were not harvested and biomass increase was up to 9 times greater than that predicted for undisturbed colonies. The survival of harvested colonies and compensatory growth increases yield, and yields at sites that had previously been harvested were generally greater than predicted by the Beverton-Holt model. The model also assumes recruitment is independent of fishing intensity, but lower numbers of young colonies in the fished populations, compared to unfished populations, suggest possible negative effects of the harvest on reproduction. This suggests the need for longer intervals between harvests. Because it can be developed from data that can be collected at a single time, the Beverton-Holt model provides a rational starting point for regulating new fisheries where long-term characterizations of population dynamics are rarely available. However, an adaptive approach to the fishery requires the incorporation of reproductive data.     

Effects of size‐ and sex‐selective harvesting: An integral projection model approach

Harvesting is often size‐selective, and in species with sexual size dimorphism, it may also be sex‐selective. A powerful approach to investigate potential consequences of size‐ and/or sex‐selective harvesting is to simulate it in a demographic population model. We developed a population‐based integral projection model for a size‐ and sex‐structured species, the commonly exploited pike (Esox lucius). The model allows reproductive success to be proportional to body size and potentially limited by both sexes. We ran all harvest simulations with both lower size limits and slot limits, and to quantify the effects of selective harvesting, we calculated sex ratios and the long‐term population growth rate (λ). In addition, we quantified to what degree purely size‐selective harvesting was sex‐selective, and determined when λ shifted from being female to male limited under size‐ and sex‐selective harvesting. We found that purely size‐selective harvest can be sex‐selective, and that it depends on the harvest limits and the size distributions of the sexes. For the size‐ and sex‐selective harvest simulations, λ increased with harvest intensity up to a threshold as females limited reproduction. Beyond this threshold, males became the limiting sex, and λ decreased as more males were harvested. The peak in λ, and the corresponding sex ratio in harvest, varied with both the selectivity and the intensity of the harvest simulation. Our model represents a useful extension of size‐structured population models as it includes both sexes, relaxes the assumption of female dominance, and accounts for size‐dependent fecundity. The consequences of selective harvesting presented here are especially relevant for size‐ and sex‐structured exploited species, such as commercial fisheries. Thus, our model provides a useful contribution toward the development of more sustainable harvesting regimes.     

Non-timber forest product harvesting in alien-dominated forests: effects of frond-harvest and rainfall on the demography of two native Hawaiian ferns

Non-timber forest products (NTFP) represent culturally and economically important resources for millions of people worldwide. Although many NTFP are harvested from disturbed habitats and therefore subject to multiple pressures, few quantitative studies have addressed this issue. Similarly few NTFP studies have assessed seasonal variation in demographic rates even though this can confound harvest effects. In Hawaiȁ8i, the wild-gathered ferns, Microlepia strigosa and Sphenomeris chinensis, represent highly important cultural resources but declining populations have led to conservation concerns. Both ferns are harvested from disturbed, alien-dominated forests and contemporary Hawaiian gathering practices often consist of harvest and concurrent weeding of alien invasive species. We assessed the effects of concurrent frond-harvest and alien species weeding on frond structure, density, and rates of production by comparing experimentally harvested vs. control plots, and documented relationships between frond demographic patterns and precipitation. Gathering practices had no impact on frond density of either species or on most other demographic parameters over the short term. Exceptions included a significant decrease in the density of the longest S. chinensis fronds and a significant decrease in M. strigosa frond production when fronds were gathered without alien weeding. However, seasonal and annual changes in frond density and production occurred across all plots of both species and were significantly correlated with precipitation. The relatively low harvest effects for both species are likely due to several factors including short frond longevity and the strict criteria used by gatherers to select harvestable fronds. The potential for sustainable harvest in the context of alien-dominated forests is discussed.     

Population modelling of Gelidium sesquipedale (Rhodophyta, Gelidiales)

A matrix population model of Gelidium sesquipedale, a commercial agarophyte from the Northeast Atlantic, was developed based on demographic data obtained during two years in a commercial stand of Cape Espichel, Portugal. G. sesquipedale individuals were classified into categories such as life cycle phase, spores, juveniles and adult frond size, because the species vital rates, fecundity, fertility, survival, growth and breakage depend on them. We also exemplify the use of a user-friendly modelling software, Stella, to develop a structured-population model. This is the first time this software has been used to model the demography of seaweed populations. The Stella model developed here behaved very similarly to the matrix model, because of its particular construction, which causes the forcing functions to be discrete rather than continuous. The relative importance of spore recruitment and vegetative growth of new fronds in both population growth and population structure was investigated. Elasticity analysis suggests that vegetative recruitment is the most important demographic parameter controlling population growth together with survival and transitions between juveniles (1–6 cm fronds) and class 1 fronds (6–9 cm fronds). On the other hand, sexual reproduction may, by itself, efficiently control the relative proportion of gametophytes and tetrasporophytes in the population, even though its contribution to recruitment is extremely small. A 40% difference in the growth rates of gametophyte and tetrasporophyte submatrices resulted from natural differences in spore recruitment rates. This revised version was published online in June 2006 with corrections to the Cover Date.     

Social and Ecological Characteristics of an Expanding Natural Resource Industry: <Emphasis Type="Italic">Aloe</Emphasis> Harvesting in South Africa

Sustainable harvesting practices are important for conserving plant species and their habitats, but also the livelihoods of those that depend on them. Aloe ferox, a valuable natural resource harvested for its leaves, is the focus of a recent rural development initiative in the Eastern Cape of South Africa. This has the potential to benefit poor residents through a high-value, sustainable, export market. We characterize the social and ecological components of the system, in order to evaluate the potential for effective natural resource management. We interviewed aloe tappers to obtain information on their dependence on the A. ferox industry and harvesting practices. We assessed the harvesting pressure on A. ferox populations, sampling plants at three plots positioned along each of four transects at distances of 1.5, 3.45, and 7 km from the factory, grouping plants into two size classes: small (height <0.5 m) and large (>0.5 m). We investigated the influence of proximity to the factory and plant size class on the likelihood and intensity of harvest. The majority of aloe tappers were women, unemployed, and in receipt of government welfare grants, and the main reason for harvesting A. ferox was to generate a cash income for their daily needs. Training guidelines did not appear to be followed, with aloe tappers leaving on average 6 leaves, rather than the recommended 18–20 leaves, allowing insufficient time to pass between harvesting episodes and harvesting outside of the prescribed wetter periods. In line with training guidelines, aloe tappers were targeting larger plants; however, against recommendations, smaller plants were also regularly harvested. Harvesting pressure decreased with increasing distance from the factory. We discuss requirements to ensure A. ferox is harvested at sustainable levels in the region, particularly in light of a possible regional roll out of the program, and provide recommendations for regulating use and better training.     

Adaptive changes in harvested populations: plasticity and evolution of age and size at maturation

We investigate harvest-induced adaptive changes in age and size at maturation by modelling both plastic variation and evolutionary trajectories. Harvesting mature individuals displaces the reaction norm for age and size at maturation toward older ages and larger sizes and rotates it clockwise, whereas harvesting immature individuals has the reverse qualitative effect. If both immature and mature individuals are harvested, the net effect has approximately the same trend as when harvesting immature individuals only. This stems from the sensitivity of the evolutionary response, which depends on the maturity state of harvested individuals, but also on the type of harvest mortality (negatively or positively density dependent, density independent) and the value of three life-history parameters (natural mortality, growth rate and the trade-off between growth and reproduction). Evolutionary changes in the maturation reaction norm have strong repercussions for the mean size and the density of harvested individuals that, in most cases, result in the reduction of biomass--a response that population dynamical models would overlook. These results highlight the importance of accounting for evolutionary trends in the long-term management of exploited living resources and give qualitative insights into how to minimize the detrimental consequences of harvest-induced evolutionary changes in maturation reaction norms.     

Optimal harvesting strategies for timber and non-timber forest products in tropical ecosystems

Harvesting wild plants for non-timber forest products (NTFPs) can be ecologically sustainable–without long-term consequences to the dynamics of targeted and associated species–but it may not be economically satisfying because it fails to provide enough revenues for local people over time. In several cases, the same species can be harvested for NTFP and also logged for timber. Three decades of studies on the sustainability of NTFP harvest for local people’s livelihood have failed to successfully integrate these socio-economic and ecological factors. We apply optimal control theory to investigate optimal strategies for the combinations of non-lethal (e.g., NTFP) and lethal (e.g., timber) harvest that minimize the cost of harvesting while maximizing the benefits (revenue) that accrue to harvesters and the conservation value of harvested ecosystems. Optimal harvesting strategies include starting with non-lethal NTFP harvest and postponing lethal timber harvesting to begin after a few years. We clearly demonstrate that slow growth species have lower optimal harvesting rates, objective functional values and profits than fast growth species. However, contrary to expectation, the effect of species lifespan on optimal harvesting rates was weak suggesting that life history is a better indicator of species resilience to harvest than lifespan. Overall, lethal or nonlethal harvest rates must be <40 % to ensure optimality. This optimal rate is lower than commonly reported sustainable harvest rates for non-timber forest products.     

Cultivation of Gigartina skottsbergii (Gigartinales, Rhodophyta): Recent advances and challenges for the future

This study integrates landings statistics and biological studies of the red algaGigartina skottsbergii Setchell & Gardner. The analysis of the landings and carrageenan production in Chile suggeststhat this resource will suffer a strong harvesting pressure during the nextyears. Biological results on sporulation, germination, sporeling growth and survivorship in laboratory,indoor tanks and field conditions, indicated that cultivation of this species istechnically feasible, as spores can be seeded on ropes and other substrata. Vegetative propagation of this species through tissue fragmentationis also possible. Vegetative fragments of this carrageenophyte have 20 to30% higher growth rates than whole fronds in suspended culture systems. Protoplast production can be also explored for bypassing restrictions inspore availability. Major advantages that encourage the cultivation of G. skottsbergii include its gel quantity and quality, its pathogen-freecondition, a high reproduction potential and its regeneration capacity. Onthe other hand, the major constraints are related to its relatively slowgrowth as compared to other carrageenophytes, limited availability ofspores and high mortality during juvenile stages.     

The effects of harvesting of the South African kelp (Ecklonia maxima) on kelp population structure, growth rate and recruitment

Ecklonia maxima is an economically important kelp in South Africa. The harvested kelp is used mainly as feed for cultured Haliotis midae (abalone) on farms all along the South African South and West Coast. The effects that different harvesting methods have on the growth of sub-canopy kelps, kelp population structure and kelp recruitment were tested in a kelp bed at Bordjies Rif near Cape Town. Two 30 × 10 m sites were set up, about 100 m apart, in near monoculture stands of E. maxima. Each 30×10 m area was subdivided into three treatments. In treatment 1 (T1) the whole ‘head’ of each kelp sporophyte that reached the surface was cut off between the bulb and the primary blade (‘lethal’ method). In treatment 2 (T2) (‘non-lethal’ method), the secondary fronds of all sporophytes that reached the surface were cut 20–30 cm from their junction with the primary blade, and removed. In the control plot, the kelp plants were not treated. Harvesting treatments were done approximately every four months, at low spring tide, from 3 March 2003 to 3 November 2003 (three treatments). The effects of harvesting on the kelps depend largely on the size of plant and the time the fronds were removed; however, no seasonal pattern could be observed. The different treatments had no effect on the growth rate, population structure or recruitment of the kelp. This means that factors other than light play an important role in the growth, structure and recruitment of the kelp beds in False Bay. Results are discussed in relation to current commercial harvesting practices.     

QUANTIFYING GROWTH AND CALCIUM CARBONATE DEPOSITION OF CALLIARTHRON CHEILOSPORIOIDES (CORALLINALES,RHODOPHYTA) IN THE FIELD USING A PERSISTENT VITAL STAIN1

Growth and calcium carbonate deposition rates of the coralline alga Calliarthron cheilosporioides Manza were quantified by monitoring fronds in the intertidal zone that had been chemically labeled with the nontoxic fluorescent brightener Calcofluor white. This vital stain effectively labeled apical meristems of coralline thalli in the field: fronds exposed for only 5 min had detectable chemical marks at least 1.5 years later. By distinguishing portions of thalli that developed before and after exposure, this methodology permitted accurate measurement of growth and calcium carbonate deposition at each meristem. In Calliarthron, meristematic activity declined with increasing frond size. However, because growing fronds dichotomize, the total number of meristems and the deposition rate of new calcified tissue both increased with frond size. Growth rates reported here suggest that large fronds may not be as old as previously estimated. The Calcofluor white method may improve demographic studies of corallines by resolving growth and age of fronds in the field and may facilitate studies of climate change on calcium carbonate deposition in these ecologically important, calcifying algae.     

Population Dynamics of the Dioecious Amazonian Palm Mauritia flexuosa: Simulation Analysis of Sustainable Harvesting

The dioecious, tropical palm Mauritia flexuosa has high ecological and economic value, but currently some wild populations are harvested excessively, which is likely to increase. In this study, we investigated the population dynamics of this important palm, the effects of harvesting, and suggested sustainable harvesting regimes. Data were collected from populations in the Ecuadorian Amazon that were assumed to be stable. We used a matrix population model to calculate the density-independent asymptotic population growth rate (λ= 1.046) and to evaluate harvesting scenarios. Elasticity analysis showed that survival (particularly in the second and fifth size class) contributes more to the population growth rate, than growth and fecundity. To simulate a stable population at carrying capacity, density dependence was incorporated and applied to the seedling survival and growth parameters in the transition matrix. Harvesting scenarios were simulated with the density-dependent population models to predict sustainable harvesting regimes for the dioecious palm. We simulated the removal of only female palms and showed how both sexes are affected with harvest intensities between 15 and 75 percent and harvest intervals of 1–15 yr. By assuming a minimum female threshold, we demonstrated a continuum of sustainable harvesting schedules for various intensities and frequencies for 100 yr of harvest. Furthermore, by setting the population model's λ= 1.00, we found that a harvest of 22.5 percent on a 20 yr frequency for the M. flexuosa population in Ecuador is consistent with a sustainable, viable population over time.     

Testing sustainable management in Northern Chile: harvesting Macrocystis pyrifera (Phaeophyceae, Laminariales). A case study

Kelp harvesting in northern Chile is managed by local fishermen and is part of an organized industry. However, the lack of standardized harvesting protocols has made regulation difficult. This, in combination with the impacts of oceanographic disturbances has resulted in some kelp populations being considerably reduced during the last decade. Consequently, harvest methods that maintain kelp resources are sorely needed if harvesting is to remain a viable industry in Chile. Here, experiments were done to identify sustainable methods for harvesting Macrocystis pyrifera along the coast of northern Chile. Three methods were compared with regard to their impacts on kelp populations; one that involves extracting half of the fronds from each individual in a population, one that involves extracting all the fronds from half of the individuals in a population, and a third that involves extracting all the fronds from all of the individuals in a population (i.e., the method currently used). Following this, populations were evaluated over a 2-month period to monitor re-growth of the remaining individuals and recruitment of new individuals, as well as changes in understory algal diversity and herbivore abundance. Our results indicate that removing half of the fronds from each individual in a population was the best method for maintaining the resource for future harvest because, it (1) maintains rapid growth of new fronds on the harvested individuals, (2) promotes recruitment of new individuals, and (3) reduces herbivore densities through physical abrasion. Consequently, this method is recommended for future harvesting of M. pyrifera in Northern Chile.     

Effects of Commercial Harvesting on Population Characteristics and Rhizome Yield of Anemone altaica

Effects of Commercial Harvesting on Population Characteristics and Rhizome Yield of Anemone altaica. Commercial harvesting constitutes a direct threat to numerous non–timber forest products (NTFPs), but its ecological effects have not been well documented. Anemone altaica Fisch. ex C. A. Mey, a spring ephemeral plant found in temperate forests of Eurasia, is a traditional Chinese herb. Owing to medicinal value, its rhizomes have been harvested for commercial purposes in northwestern China for many years. This paper addresses the ecological effects of commercial harvesting on A. altaica populations under different harvest intensities. The results show that size–selective harvesting of rhizomes can increase population densities by asexual propagation. Currently, two– to three–year–old individuals derived from asexual propagation are the main targets of commercial harvesting. The increased demand in recent years has resulted in earlier and more intensive harvesting activities largely impacting the natural recovery of the harvested populations. For sustainable use of this traditional medicinal species, we recommend that a periodic harvest strategy of three to four years be adopted.     

Recurrent fruit harvesting reduces seedling density but increases the frequency of clonal reproduction in a tropical tree

Studies on the ecological impacts of non‐timber forest products (NTFP) harvest reveal that plants are often more resilient to fruit and seed harvest than to bark and root harvest. Several studies indicate that sustainable fruit harvesting limits can be set very high (>80% fruit harvesting intensity). For species with clonal and sexual reproduction, understanding how fruit harvest affects clonal reproduction can shed light on the genetic risks and sustainability of NTFP harvest. We studied 18 populations of a gallery forest tree, Pentadesma butyracea (Clusiaceae), to test the impact of fruits harvest, climate and habitat size (gallery forest width) on the frequency of sexual or clonal recruitment in Benin, West Africa. We sampled populations in two ecological regions (Sudanian and Sudano‐Guinean) and in each region, we selected sites with low, moderate and high fruit harvesting intensities. These populations were selected in gallery forests with varying width to sample the natural variation in P. butyracea habitat size. Heavily harvested populations produced significantly less seedlings but had the highest density and proportion of clonal offspring. Our study suggests that for plant species with dual reproductive strategy (via seeds and clonal), fruit harvesting and associated disturbances that come with it can lead to an increase in the proportion of clonal offspring. This raises the issue that excessive fruit harvest by increasing the proportion of clonal offspring to the detriment of seed originated offspring may lead to a reduction in genetic diversity with consequence on harvested species capability to withstand environmental stochasticity.     

Modeling Harvest Intensity of Sooty Shearwater Chicks by Rakiura Māori in New Zealand

Abstract: Cultural evidence suggests that sooty shearwater (Puffinus griseus) chicks have been harvested by Rakiura Māori on islands in southern New Zealand since prehistoric times. Concerns exist that modern harvests may be impacting sooty shearwater abundance. We modeled human-related and ecological determinants of harvest (total no. of individuals harvested) of sooty shearwater chicks on 11 islands and examined the relationship between shearwater abundance and harvesting rates (chicks/hr) and harvester behavior throughout the harvesting season. Models best explaining variation in harvest between harvesting areas (manu), for both the early and late parts of the harvesting season, included harvester-days (included in all models with change in deviance information criteria [ΔDIC], ΔDIC < 8.36 and ΔDIC < 11.5, for the early and late periods, respectively). Other harvest determinants included shearwater density, size of the manu, and number of people helping harvesters (all included in the top 5 models within ΔDIC = 2.25 for the late period). Areas harvested by several families under a common-property harvesting system had higher harvest intensity for their size (24% points higher, 95% credible interval 11–36%) than those managed as an exclusive resource for one family. The slowest harvesters spent more time harvesting but on average only harvested 36% (95% credible interval 15–65%) and 34% (95% credible interval 12–63%) of the harvest taken by the fastest harvesters during the early and late periods, respectively. Our results highlight the possibility of elevated harvest intensity as the population of harvesters increases. However, our models suggested that a corresponding reduction in harvesting rate at low prey densities during the most productive period could potentially regulate harvest intensity. Future research will integrate these results into prospective shearwater demographic models to assess the utility of a range of harvesting strategies in ensuring harvest sustainability.     

Impact of unintentional selective harvesting on the population dynamics of red grouse

1.?The effect of selective exploitation of certain age, stage or sex classes (e.g., trophy hunting) on population dynamics is relatively well studied in fisheries and sexually dimorphic mammals. 2.?Harvesting of terrestrial species with no morphological differences visible between the different age and sex classes (monomorphic species) is usually assumed to be nonselective because monomorphicity makes intentionally selective harvesting pointless and impractical. But harvesting of the red grouse (Lagopus lagopus scoticus), a monomorphic species, was recently shown to be unintentionally selective. This study uses a sex- and age-specific model to explore the previously unresearched effects of unintentional harvesting selectivity. 3.?We examine the effects of selectivity on red grouse dynamics by considering models with and without selectivity. Our models include territoriality and parasitism, two mechanisms known to be important for grouse dynamics. 4.?We show that the unintentional selectivity of harvesting that occurs in red grouse decreases population yield compared with unselective harvesting at high harvest rates. Selectivity also dramatically increases extinction risk at high harvest rates. 5.?Selective harvesting strengthens the 3- to 13-year red grouse population cycle, suggesting that the selectivity of harvesting is a previously unappreciated factor contributing to the cycle. 6.?The additional extinction risk introduced by harvesting selectivity provides a quantitative justification for typically implemented 20-40% harvest rates, which are below the maximum sustainable yield that could be taken, given the observed population growth rates of red grouse. 7.?This study shows the possible broad importance of investigating in future research whether unintentionally selective harvesting occurs on other species.     

Evolutionary rebound from selective harvesting

Size-selective harvesting favours reduced investment in growth and earlier age at reproduction. A recent study by Edeline and colleagues has shown that trait dynamics of Lake Windermere pike reflect the interplay of 50 years of harvest and natural selection. Growth and investment in early reproduction decreased under harvesting, and then recovered when fishing pressures declined. This is the first study to use contrasting temporal patterns of natural and harvest selection pressures to account for observed trait changes.     

Effects of size and growth rate on vegetative reproduction in Typha

Summary The objective of this study was to separate the effects of plant biomass and growth rate on vegetative reproduction in two species of cat-tail, Typha latifolia and T. angustifolia. Replicate clones of both species were grown under conditions of 100%, 42%, 24%, and 9% full sunlight with harvests at 41, 70, and 91 days after shading. T. angustifolia produced most of its vegetative offspring before the first harvest and increased biomass over the remainder of the experiment by increasing the size of its ramets. In contrast, T. latifolia produced vegetative offspring gradually throughout the experiment adding new ramets only after existing clones were of mature size. As a result of these differences in the cloning process, T. angustifolia showed little correlation between vegetative reproduction and clone size while T. latifolia showed a strong correlation between gegetative reproduction and clone size at the three highest light intensities. Growth rates, average clone size and vegetative reproduction were all reduced by reductions in light intensity for both species. However, no effect of growth rate on the relationship between clone size and vegetative reproduction in T. latifolia could be detected. T. latifolia showed greater survivorship and more biomass production under 9% light than T. angustifolia indicating a greater shade tolerance.