Comparing naturalized alien plants and recipient habitats across an east–west gradient in the Mediterranean Basin

Aim To investigate alien plant species invasion levels in different habitats and alien species traits by comparing the naturalized flora in different areas of the same biogeographical region. Location Spain, Italy, Greece and Cyprus. Methods Comparison of floristic composition, species traits and recipient habitats of naturalized alien neophytes across an east–west gradient comprising four countries in the European Mediterranean basin. Results A total of 782 naturalized neophytes were recorded; only 30 species were present in all four countries. Although floristic similarity is low, the four alien floras share the same patterns of growth form (mostly herbs), life cycle (mostly perennials) and life form (mostly therophytes, hemicryptophytes and phanerophytes). The majority of the recipient habitats were artificial. Wetlands were the natural habitats, with the highest numbers of naturalized species. Floristic similarity analyses revealed: (1) the highest floristic similarity between Italy and Spain, both of which were more similar to Greece than to Cyprus; (2) two groups of floristic similarity between habitat categories in each country (Greece–Cyprus and Italy–Spain); (3) a higher degree of homogenization in the plant assemblages in different habitats in Greece and Cyprus and a lower degree of homogenization in those in Italy and Spain; and (4) a higher degree of homogenization in artificial and natural fresh‐water habitats than in the other natural habitats. Main conclusions The floristic similarity of naturalized neophytes between the four countries is low, although the overall analysis indicates that the western group (Italy–Spain) is separated from the eastern group (Greece–Cyprus). Similar patterns emerged regarding the life‐history traits and recipient habitats. The artificial habitats and the natural wet habitats are those that are invaded most and display the greatest homogenization in all four countries. Coastal habitats display a lower degree of homogenization but a high frequency of aliens. Dry shrubs and rocky habitats display a lower degree of homogenization and a low frequency of aliens.     

Current progress and future prospects in phylofloristics

The species composition of regional plant assemblages can in large part be explained by a long history of biogeographical and evolutionary events. Traditional attempts of floristic studies typically focus on the analyses of taxonomic composition, often ignoring the rich context that evolutionary history can provide. In 2014, Swenson and Umaña introduced the term ‘phylofloristics’ to define a phylogenetically enabled analysis of the species composition of regional floras. Integrating phylogenetic information into traditional floristic analysis can provide a promising way to explore the ecological, biogeographic, and evolutionary processes that drive plant assemblies at multiple spatial scales. In this review, we summarize the current progress on the phylogenetic structure, spatial phylogenetic pattern, origin and diversification, phylogenetic regionalization of floristic assemblages, and application of phylogenetic information in biodiversity conservation. These summaries highlight the importance of incorporating phylogenetic information to improve our understanding of floristic assembly from an evolutionary perspective. The review ends with a brief outlook on future challenges for phylofloristic studies, including generating a highly resolved species-level phylogenetic tree, compiling detailed and refined information regarding the geographic distribution of all plant life, extracting trait information from publications and herbarium specimens, and developing technological and methodological approaches for big data analysis.     

Plant extinction dynamics in an insular metacommunity

Changes in the composition of local communities through time (i.e. species turnover) is a common phenomenon in insular biology. However, the mechanisms promoting variation in species turnover, both among islands and among species, are poorly understood. In an effort to better understand the causes of variation in species turnover, we evaluated the colonization and extinction dynamics of plant populations on 18 small islands off the west coast of Canada. In 1997, we quantified total population sizes of 10 woody angiosperm species. A decade later, we resampled islands to test whether: 1) species turnover occurred, 2) colonization events were offset by extinction events, 2) variation in extinction rates among islands was associated with population sizes, average plant heights, island area, island isolation or each island's exposure to ocean-born disturbances, and 3) variation in extinction rates among species was associated with plant life history traits. Results showed that extinction events outnumbered colonization events, suggesting that the metacommunity is in 'disequilibrium'. Variation in extinction rates among islands was unrelated to island area and isolation. However, extinction rates increased with exposure to ocean-born disturbances and decreased with both initial population sizes and average plant heights. Species with thicker, tougher leaves (i.e. high leaf mass per area) were less prone to extinction than species with thinner, more papery leaves. Overall results indicate that species turnover is common and that it is generated primarily by extinction. Variation in extinction rates appears to result from an interaction between among-island effects (exposure, population size and plant stature) and among-species effects (leaf toughness), suggesting that ocean-born disturbances play a key role in determining metacommunity structure.     

Metapopulation-level adaptation of insect host plant preference and extinction-colonization dynamics in heterogeneous landscapes

Species living in highly fragmented landscapes typically occur as metapopulations with frequent turnover of local populations. The turnover rate depends on population sizes and connectivities, but it may also depend on the phenotypic and genotypic composition of populations. The Glanville fritillary butterfly (Melitaea cinxia) in Finland uses two host plant species, which show variation in their relative abundances at two spatial scales: locally among individual habitat patches and regionally among networks of patches. Female butterflies in turn exhibit spatial variation in genetically determined host plant preference within and among patch networks. Emigration, immigration and establishment of new populations have all been shown to be strongly influenced by the match between the host plant composition of otherwise suitable habitat patches and the host plant preference of migrating butterflies. The evolutionary consequences of such biased migration and colonization with respect to butterfly phenotypes might differ depending on spatial configuration and plant species composition of the patches in heterogeneous patch networks. Using a spatially realistic individual-based model we show that the model-predicted evolution of host plant preference due to biased migration explains a significant amount of the observed variation in host plant use among metapopulations living in dissimilar networks. This example illustrates how the ecological extinction-colonization dynamics may be linked with the evolutionary dynamics of life history traits in metapopulations.     

Scale-dependent equilibrium in highly heterogeneous islands: plant geography of the northern Great Barrier Reef sand cays and shingle islets

In contrast to classical island biogeographical predictions, plant species richness is not monotonically related to island area or isolation for the northern Great Barrier Reef islands, and this cannot be accounted for as a ‘small island effect’. Many islands are composed of different terrain units bearing different floristic assemblages suggesting a modified‘terrain-floristic element’model for island biogeography. This requires definition of a time-scale for equilibrium considerations and emphasizes that equilibrium or first order perturbation (‘relaxation’) models may be inappropriate for some archipelagos.     

Presence, richness and extinction of birds of prey in the Mediterranean and Macaronesian islands

Aim We analyse modern patterns of richness, presence and extinction of birds of prey (Accipitriforms and Falconiforms) in the Mediterranean and Macaronesian islands, using an integrated approach involving both biogeographical and human‐induced factors. Location Forty‐three islands grouped into nine Mediterranean and Macaronesian archipelagos. Methods Information about 25 species breeding during the past century and their fate (permanence or extinction) was compiled from the literature and regional reports. Jaccard's similarity index and cluster analyses were applied to define island assemblages. In order to detect the factors driving richness, presence and extinction, generalized linear models (GLM) were applied to 32 explanatory variables, evaluating location, physiography, isolation of island, taxonomic affinities and life‐history patterns of the raptor species. Results Islands belonging to the same archipelago clustered when raptor assemblages were compared, revealing a marked biogeographical signal. Species richness was influenced by island area and accessibility from the continent (explained deviance of 51% in the GLM). Models of the probability of presence (explained deviance of 32%) revealed positive influences of migratory patterns (maximum for partial migrants), size of distribution areas and proximity to main migration routes. The model for probability of extinction explained only 8% of the deviance. It revealed that populations living on islands with a high density of human population were more prone to disappear. Also, raptors depending on human resources had more risk of extinction. Main conclusions Basic predictions of island biogeography can explain current patterns of raptor richness in the study area despite millennia of intense humanization processes. Colonization success appears to depend on life‐history traits linked to migratory and dispersal strategies, whereas body‐size constraints are not influential. Additionally, our results reveal the importance of species‐based analyses in studies of island biogeography.     

The role of dispersal ability, climate and spatial separation in shaping biogeographical patterns of phylogenetically distant plant groups in seasonally dry Andean forests of Bolivia

Aim To assess biogeographical patterns of Acanthaceae, Bromeliaceae, Cactaceae and Pteridophyta in Bolivian Andean seasonally dry forest islands and to explain current floristic differences between these islands by means of extrinsic (precipitation, elevation) and intrinsic (dispersal ability) factors. Location Ten isolated and disjunct seasonally dry forest areas in the Bolivian Andes and the adjacent seasonal forest areas of the Chiquitanía and Chaco regions. Methods We collated species data from recently updated and revised taxonomic treatments and herbarium collections for Acanthaceae, Bromeliaceae, Cactaceae and Pteridophyta, constructed floristic distance matrices to estimate beta diversity at the study sites and subjected them to Mantel correlation analyses. Multiple regressions on distance matrices allowed us to test the influence of geographical distance and environmental (elevation and precipitation) differences on floristic differentiation. Results Acanthaceae and Bromeliaceae, and to a lesser extent Cactaceae, showed coincident biogeographical patterns and suggested the presence of two seasonally dry forest groups in Andean Bolivia: one including all small isolated northern dry valleys and another including all southern valleys with connections to the lowland seasonal forests of the Chiquitanía and the Chaco. Most of the variation in the floristic distance matrices in these plant groups, with seed dispersal typically restricted to short distances, was explained by spatial separation between habitat islands. In contrast, pteridophytes showed a different biogeographical pattern. Their floristic differences between sites were determined by the environmental variables. The anemochorous and spore‐based dispersal system of this plant group seems to be a highly effective mechanism allowing pteridophytes to easily reach even the isolated dry valleys in inter‐Andean Bolivia. Main conclusions Current biogeographical patterns in dry Andean habitat islands can provide insights into the factors that control the processes of community assembly. We show that differences in community composition of phylogenetically distant plant groups in the understorey of seasonally dry forest islands can be explained by a combination of the habitat characteristics where the group is present (either precipitation, elevation or both) and, more interestingly, by group specific dispersal limitation (as inferred by geographical distance between island habitats).     

The influence of past land-use on understory plant distributions in a near-natural deciduous forest in Denmark

In regions with a long cultural history, past land-use may strongly determine floristic composition in seemingly natural vegetation. Here we examine if past land-use affects understory plant distribution in near-natural forest in a historically and environmentally heterogeneous setting, namely the 17 ha freshwater island of Borgnr (Denmark). In 107 randomly located 100 m² plots we recorded species composition and various environmental variables. The plots were classified into three groups according to past land-use: (1) Disturbed sites close to 12^th and 20^th century ruins and associated open areas; (2) formerly grazed meadow areas; and (3) forested sites without indications of strong anthropogenic influence. Past land-use, environmental variables and species traits were used to interpret floristic gradients. Unconstrained and constrained ordinations clearly separated plots with different past land-use, and showed that the main floristic gradients on Borgø correlated with hydrological-edaphic conditions and past land-use. Partial constrained ordination using the environmental parameters as covariables showed that past land-use had a unique influence on floristic composition.     

Frequent,low‐amplitude disturbances drive high tree turnover rates on a remote,cyclone‐prone Polynesian island

Aim How important are frequent, low‐intensity disturbances to tree community dynamics of a cyclone‐prone forest? We tested the following hypotheses concerning the ‘inter‐cataclysm’ period on a remote Polynesian island: (1) tree turnover would be high and recruitment rates would be significantly higher than mortality; (2) low‐intensity disturbance would result in a marginal increase in tree mortality in the short term; (3) turnover would vary among species and would be associated with plant traits linked to differences in life history; and (4) mortality and recruitment events would be spatially non‐random. Location Tutuila, a volcanic island in the Samoan Archipelago, Polynesia. Methods We censused the tree (stem diameter ≥ 10 cm) community in 3.9 ha of tropical forest three times over a 10‐year period, 1998–2008. We calculated annual mortality, recruitment and turnover rates for 36 tree species. We tested for non‐random spatial patterns and predictors of mortality, and non‐random spatial patterns of tree recruitment. A 2004 cyclone passing within 400 km allowed us to measure the effects of a non‐cataclysmic disturbance on vital rates. Results Annual turnover was 2.8% and annual recruitment was 3.6%; these are some of the highest rates in the tropics, and likely to be a response to a cyclone that passed < 50 km from Tutuila in 1991. Species turnover rates over 10 years were negatively correlated with wood specific gravity, and positively correlated with annual stem diameter increment. Mortality was spatially aggregated, and a function of site, species and an individual’s growth rate. Recruitment was highest on ground with low slope. The low‐magnitude cyclone disturbance in 2004 defoliated 29% of all trees, but killed only 1.8% of trees immediately and increased annual mortality over 5 years by 0.7%. Main conclusions The inter‐cataclysm period on Tutuila is characterized by frequent, low‐amplitude disturbances that promote high rates of tree recruitment and create a dynamic, non‐equilibrium or disturbed island disequilibrium tree community. Species with low wood density and fast growth rates have enhanced opportunities for recruitment between cataclysms, but also higher probabilities of dying. Our results suggest that increases in the frequency of cyclone activity could shift relative abundances towards disturbance‐specialist species and new forest turnover rates.     

Temporal trends in species composition and plant traits in natural grasslands of Uruguay

Abstract. We report the successional trends of the major life‐forms (graminoids and forbs) in natural grasslands of Uruguay over a 9‐yr period after the removal of domestic herbivores. For the whole community, species richness and diversity decreased over the successional period. In graminoids we observed clear temporal trajectories in floristic composition; the rate of floristic change decreased with time and was associated with a shift in plant traits. The exclusion of large herbivores promoted erect and tall grasses with narrow leaves and greater seed length, vegetative growth constrained to the cool season and increased frequency of annual species. Forbs did not show a clear temporal trend in species composition, but there was, nevertheless, a plot‐specific species turnover of this functional group that was reflected in their attributes. Species spreading by means of rhizomes, with vegetative growth restricted to the warm season. Species with larger seeds increased under grazing exclusion, as did annual and nitrogen‐fixing forbs. The floristic changes induced by cattle exclusion occurred early in the succession. This early high rate of change has practical implications for management and conservation programs of the natural grasslands of Uruguay. Additionally, the shift in plant traits may be helpful in devising simple indicators of grazing impact.     

Plant composition and diversity in a semi-natural Mediterranean island landscape: the importance of environmental factors

Abstract

Islands crucially contribute to the Mediterranean Basin’s high floristic diversity, which, however, is at risk facing climate and land-use changes. Besides the identification of highly diverse areas, the knowledge about factors favouring diversity is of great importance. We analysed plant species diversity and composition related to environmental factors over varied vegetation units on a former Italian prison island in the northwest of Sardinia. Due to a long history of land use with grazing and later abandonment the nowadays protected island features a semi-natural landscape and can serve as an example for strongly anthropogenic altered insular ecosystems. Floristic composition, soil properties, microclimate and ungulate abundance were assessed. Relationships of vegetation composition and diversity with abiotic variables were examined by Canonical Correspondence Analysis, which indicated the importance of air temperature, soil moisture, slope gradient and C/N ratio for floristic differentiation. Most important abiotic factors for plant species richness were relative air humidity and soil moisture, while floristic diversity was mainly determined by air temperature and pH. Furthermore, observation data pointed to an adverse influence of ungulate abundance for plant species diversity. Regarding nature conservation, grazing intensity thus must be critically taken into account, especially for sensitive vegetation units like the coastal garrigue.     

Vegetation Heterogeneity and Life-Strategy Diversity in the Flora of the Heterogeneous Landscape of Nizanda, Oaxaca, Mexico

We updated the floristic checklist of the Nizanda region, Isthmus of Tehuantepec (southern Mexico), characterized the occurring plant communities based on dominant species, and described the region’s flora according to life form, growth form, growth type, and growth habit spectra. Ten years of botanical exploration, along with surveys in 188 100-m² samples from different vegetation types, provided the baseline floristic information. Ordination and classification analyses were performed to examine the degree of differentiation between communities. Geographical ranges of all species were used to assess biogeographical relationships of this flora. The inventory includes 920 species (553 genera, 124 families). More than one-third of the families were represented by a single species, whereas the 10 richest families had 43% of the species richness. Dendrograms showing plot classification at three taxonomic levels (species, genus and family) revealed savannah as the most strongly differentiated community amid seven vegetation types. Regarding growth forms, forbs and trees prevailed. Phanerophytes were the most common life form category, whereas herbs and woody plants were the dominant growth types. The largest richness for all taxonomic levels was recorded in the tropical dry forest. The expanded floristic knowledge gained for the Nizanda region provided better criteria to revise the classification scheme of its vegetation. Our preliminary biogeographical analysis illustrates the role of the Isthmus of Tehuantepec as a corridor for thermophilous floras between two oceanic watersheds, and as a natural distributional limit for several Mesoamerican plant species.     

Source pools and disharmony of the world's island floras

Island disharmony refers to the biased representation of higher taxa on islands compared to their mainland source regions and represents a central concept in island biology. Here, we develop a generalizable framework for approximating these source regions and conduct the first global assessment of island disharmony and its underlying drivers. We compiled vascular plant species lists for 178 oceanic islands and 735 mainland regions. Using mainland data only, we modelled species turnover as a function of environmental and geographic distance and predicted the proportion of shared species between each island and mainland region. We then quantified the over‐ or under‐representation of families on individual islands (representational disharmony) by contrasting the observed number of species against a null model of random colonization from the mainland source pool, and analysed the effects of six family‐level functional traits on the resulting measure. Furthermore, we aggregated the values of representational disharmony per island to characterize overall taxonomic bias of a given flora (compositional disharmony), and analysed this second measure as a function of four island biogeographical variables. Our results indicate considerable variation in representational disharmony both within and among plant families. Examples of generally over‐represented families include Urticaceae, Convolvulaceae and almost all pteridophyte families. Other families such as Asteraceae and Orchidaceae were generally under‐represented, with local peaks of over‐representation in known radiation hotspots. Abiotic pollination and a lack of dispersal specialization were most strongly associated with an insular over‐representation of families, whereas other family‐level traits showed minor effects. With respect to compositional disharmony, large, high‐elevation islands tended to have the most disharmonic floras. Our results provide important insights into the taxon‐ and island‐specific drivers of disharmony. The proposed framework allows overcoming the limitations of previous approaches and provides a quantitative basis for incorporating functional and phylogenetic approaches into future studies of island disharmony.     

Are there morphological and life‐history traits under climate‐dependent differential selection in S Tunesian Diplotaxis harra (Forssk.) Boiss. (Brassicaceae) populations?

Adaptation of morphological, physiological, or life‐history traits of a plant species to heterogeneous habitats through the process of natural selection is a paramount process in evolutionary biology. We have used a population genomic approach to disentangle selection‐based and demography‐based variation in morphological and life‐history traits in the crucifer Diplotaxis harra (Forssk.) Boiss. (Brassicaceae) encountered in populations along aridity gradients in S Tunisia. We have genotyped 182 individuals from 12 populations of the species ranging from coastal to semidesert habitats using amplified fragment length polymorphism (AFLP) fingerprinting and assessed a range of morphological and life‐history traits from their progeny cultivated under common‐garden conditions. Application of three different statistical approaches for searching AFLP loci under selection allowed us to characterize candidate loci, for which their association with the traits assessed was tested for statistical significance and correlation with climate data. As a key result of this study, we find that only the shape of cauline leaves seems to be under differential selection along the aridity gradient in S Tunisian populations of Diplotaxis harra, while for all other traits studied neutral biogeographical and/or random factors could not be excluded as explanation for the variation observed. The counter‐intuitive finding that plants from populations with more arid habitats produce broader leaves under optimal conditions of cultivation than those from more mesic habitats is interpreted as being ascribable to selection for a higher plasticity in this trait under more unpredictable semidesert conditions compared to the more predictable ones in coastal habitats.     

Local and regional abundance of exotic plant species on Mediterranean islands: are species traits important?

Aim We assess the importance of three relevant and readily obtainable life‐history traits (dispersal syndrome, stem height and growth form) and biogeographical origin (European vs. non‐European) on the local and regional abundance of over 400 exotic plant species across eight Mediterranean islands. Location The Mediterranean islands of Lesbos, Rhodes, Crete, Malta, Corsica, Sardinia, Majorca and Minorca. Methods We adopt two abundance criteria for each exotic species: the proportion of islands in which the species occurs (regional abundance), and a qualitative estimate of species abundance within each of five islands (local abundance). Subsequently, we assess the relationship between local and regional abundance, as well as the role of key life‐history traits on both regional and local abundance. These analyses were undertaken separately for the European exotics and the non‐European exotics. Results Only 10.9% of the species occur on more than four islands, and only four species are present on all eight islands. Both local and regional abundances were higher for the non‐European than the European species. Local and regional abundances were positively correlated, particularly for exotics with non‐European origins. Wind‐dispersed species tended to have higher regional abundance than species dispersed by other means but this trend only occurred for local abundance on two islands — Corsica and Majorca. Neither a species’ growth form nor its stem height explained trends in regional or local abundance. Conclusions Although wind‐dispersed exotics are more widespread in the Mediterranean, plant life‐history traits appear to play a lesser role in invasion success than area of biogeographical origin. In general, exotic species of non‐European origin were more abundant at both local and regional scales. Invasion patterns should be interpreted at both local and regional scales, but the stochastic nature of biological invasions may limit deterministic interpretations of invasion patterns, especially if islands are studied in isolation.     

Determinants of alpha and beta vascular plant diversity in Mediterranean island systems: the Ionian islands,Greece

The Ionian archipelago is the second largest Greek archipelago after the Aegean, but the factors driving plant species diversity in the Ionian islands are still barely known. We used stepwise multiple regressions to investigate the factors affecting plant species diversity in 17 Ionian islands. Generalized dissimilarity modelling was applied to examine variation in the magnitude and rate of species turnover along environmental gradients, as well as to assess the relative importance of geographical and climatic factors in explaining species turnover. The values of the residuals from the ISAR log₁₀‐transfomed models of native and endemic taxa were used as a measure of island floristic diversity. Area was confirmed to be the most powerful single explanatory predictor of all diversity metrics. Mean annual precipitation and temperature, as well as shortest distance to the nearest island are also significant predictors of vascular plant diversity. The island of Kalamos constitutes an important plant diversity hotspot in the Ionian archipelago. The recent formation of the islands, the close proximity to the mainland source and the relatively low dispersal filtering of the Ionian archipelago has resulted in islands with a flora principally comprising common species and a low proportion of endemics. Small islands keep a key role in conservation of plant priority sites.     

Links to rare climates do not translate into distinct traits for island endemics

Current models of island biogeography treat endemic and non-endemic species as if they were functionally equivalent, focussing primarily on species richness. Thus, the functional composition of island biotas in relation to island biogeographical variables remains largely unknown. Using plant trait data (plant height, leaf area and flower length) for 895 native species in the Canary Islands, we related functional trait distinctiveness and climate rarity for endemic and non-endemic species and island ages. Endemics showed a link to climatically rare conditions that is consistent with island geological change through time. However, functional trait distinctiveness did not differ between endemics and non-endemics and remained constant with island age. Thus, there is no obvious link between trait distinctiveness and occupancy of rare climates, at least for the traits measured here, suggesting that treating endemic and non-endemic species as functionally equivalent in island biogeography is not fundamentally wrong.     

Synthesizing traditional biogeography with microbial ecology: the importance of dormancy

The discovery of biogeographical patterns among microbial communities has led to a focus on the empirical evaluation of the importance of dispersal limitation in microbial biota. As a result, the spatial distribution of microbial diversity has been increasingly studied while the synthesis of biogeographical theory with microbial ecology remains undeveloped. To make biogeographical theory relevant to microbial ecology, microbial traits that potentially affect the distribution of microbial diversity need to be considered. Given that many microorganisms in natural environments are in a state of dormancy and that dormancy is an important microbial fitness trait, I provide a first attempt to account for the effects of dormancy on microbial biogeography by treating dormancy as a fundamental biogeographical response. I discuss the effects of dormancy on the equilibrium theory of island biogeography and on the unified neutral theory of biodiversity and biogeography, and suggest how the equilibrium theory of island biogeography can produce predictions approaching those of the Baas‐Becking hypothesis (i.e. everything is everywhere, but the environment selects). In addition, I present a conceptual model of the unified neutral theory of biodiversity and biogeography, generalized to account for dormancy, from which a full model can be constructed for species with or without dormant life history stages.     

Patterns of species diversity on an oceanic versus a continental island mountain: a hypothesis on species diversification1

Abstract. The vegetation on a wet mountain slope on Haleakala (an oceanic island) is compared with that on Kinabalu (a continental island) to examine relationships between regional floristic richness and α- and β-diversities. The two mountains are similar in their constant tropical climate, generic and family-level floristic elements and geological age of the summit regions, but different in regional floristic richness (rich on Kinabalu vs. poor on Haleakala). α-diversity of canopy and subcanopy tree species was much higher on Kinabalu than in comparable zones on Haleakala. Average turnover rate of species (as logarithmic community similarity) on the slope was one order of magnitude greater on Kinabalu than on Haleakala (0.127 vs. 0.017 per 100 m alt.). While there were genera with wide altitudinal ranges on both mountains, a large proportion of the genera was differentiated into parapatric altitudinal congeners on Kinabalu. By contrast, most genera are altitudinally monotypic on Haleakala. The number of sympatric congeners per genus, and the frequency of multi-specific genera per plot were high on lower slopes but decreased with increasing altitude on Kinabalu, whereas the values were low across all altitudes on Haleakala. These patterns suggest that sympatric and parapatric species radiation was less on Haleakala than on Kinabalu. This may be related to Haleakala's initially poor and disharmonic flora.