Small Island States (SIDS) are among the nations most exposed to climate change (CC) and are characterised by a high degree of vulnerability. Their unique nature means there is a need for more studies focused on the limits to CC adaptation on such fragile nations, particularly regarding their problems and constraints. This paper addressed a perceived need for research into the limitations of adaptation on SIDS, focusing on the many unique restrictions. To this end, the study identified and described the adaptation limits they have by using a review of the literature and an analysis of case studies from a sample of five SIDS in the Caribbean and Pacific regions (Barbados, Trinidad and Tobago, Cook Islands, Fiji, Solomon Islands and Tonga). This research’s findings showed that an adaptable SIDS is characterised by awareness of various values, appreciation and understanding of a diversity of impacts and vulnerabilities, and acceptance of certain losses through change. The implications of this paper are two-fold. It explains why island nations continue to suffer from the impacts of CC and suggest some of the means via which adequate policies may support SIDS in their efforts to cope with the threats associated with a changing climate. This study concluded that, despite the technological and ecological limits (hard limits) affecting natural systems, adaptation to CC is limited by such complex forces and societal factors (soft limits) that more adequate adaptation strategies could overcome.

Island invasions may cause severe changes in biodiversity, but the factors that influence these changes are not well understood. We established 120 plots in Cycas micronesica habitats throughout Guam in 2005 following the invasion of the armored scale Aulacaspis yasumatsui, then observed plant mortality through 2020. We used transects in Yap as benchmarks, as the Yap C. micronesica population is not threatened. The initial Guam plots contained about 1600 seedlings, 1160 juveniles, and 1240 mature plants per ha. Seedling mortality was 100% by 2006, juvenile mortality was 100% by 2014, and the 2020 census revealed 96% mortality of the plant population. Localities in western Guam and isolated forest fragments exhibited the greatest mortality, with 100% extirpation from two fragmented western localities. The juvenile and mature trees in Yap were unchanged from 2010 to 2018, but the seedling count was heterogeneous among the years. Constrained recruitment from seedlings to juveniles explained these dynamics. Yap transects contained about 6120 seedlings, 3400 juveniles, and 1250 mature plants per ha. Biological control of the invasive insects remains the acute conservation action needed for the Guam population. Lessons learned may be useful in other regions where invasions of non-native pests threaten biodiversity.

Cycad plants possess uncommon morphological, chemical, and ontogenetic characteristics and they may introduce localized changes in soil traits that increase habitat heterogeneity. We used mature Cycas micronesica K.D. Hill trees growing in a range of soil types in Guam, Rota, and Yap to quantify differences between the soils beneath target trees and paired non-target soils away from cycad trees. The chronic presence of a C. micronesica tree introduced numerous localized changes in soil traits, increasing the heterogeneity of elemental stoichiometry in the community. Nitrogen, carbon:phosphorus, and nitrogen:phosphorus were increased in target soils among every soil type. Carbon increased and phosphorus decreased in most target soils. The habitats revealing the greatest number of elements with differences between target and non-target soils were the habitats with acid soils. The greatest number of metals exhibiting differences between the target and non-target soils occurred in the impoverished sandy habitat. This is the first report that indicates a cycad tree increases community spatial heterogeneity by localized changes in soil chemistry. Contemporary declines in cycad populations due to anthropogenic threats inadvertently decrease this spatial heterogeneity and its influences on primary producers in the landscape then cascading effects on the food web.

The literature containing which chemical elements are found in cycad leaves was reviewed to determine the range in values of concentrations reported for essential and beneficial elements. We found 46 of the 358 described cycad species had at least one element reported to date. The only genus that was missing from the data was Microcycas. Many of the species reports contained concentrations of one to several macronutrients and no other elements. The cycad leaves contained greater nitrogen and phosphorus concentrations than the reported means for plants throughout the world. Magnesium was identified as the macronutrient that has been least studied. Only 14 of the species were represented by data from in situ locations, with most of the data obtained from managed plants in botanic gardens. Leaf element concentrations were influenced by biotic factors such as plant size, leaf age, and leaflet position on the rachis. Leaf element concentrations were influenced by environmental factors such as incident light and soil nutrient concentrations within the root zone. These influential factors were missing from many of the reports, rendering the results ambiguous and comparisons among studies difficult. Future research should include the addition of more taxa, more in situ locations, the influence of season, and the influence of herbivory to more fully understand leaf nutrition for cycads.

The need for improved knowledge on conservation and management of cycad species has generated recent interest in compiling a database on leaf nutrient concentrations. However, the sampling protocols have not been consistent among reports and the influences of some plant and habitat traits on the plasticity of cycad leaf nutrient concentrations has not been adequately determined. We used Cycas micronesica K.D. Hill trees to determine the role of incident light level and leaf age on leaflet content of 11 essential elements. Shade leaves exhibited increased mass-based concentration for nitrogen, phosphorus, and potassium above that of sun leaves. Shade leaves exhibited decreased area-based concentration for all of the macro- and micronutrients below that of sun leaves. Mass-based concentration of nitrogen, phosphorus, and potassium decreased with leaf age, and that of calcium, magnesium, iron, manganese, and zinc increased with leaf age. These findings indicate the relative leaf age and the amount of shade or incident light at the leaf level must be recorded and reported for leaf tissue studies in cycads in order to reduce ambiguity and ensure repeatability.