Effects of Drought,Pest Pressure and Light Availability on Seedling Establishment and Growth: Their Role for Distribution of Tree Species across a Tropical Rainfall Gradient

Tree species distributions associated with rainfall are among the most prominent patterns in tropical forests. Understanding the mechanisms shaping these patterns is important to project impacts of global climate change on tree distributions and diversity in the tropics. Beside direct effects of water availability, additional factors co-varying with rainfall have been hypothesized to play an important role, including pest pressure and light availability. While low water availability is expected to exclude drought-intolerant wet forest species from drier forests (physiological tolerance hypothesis), high pest pressure or low light availability are hypothesized to exclude dry forest species from wetter forests (pest pressure gradient and light availability hypothesis, respectively). To test these hypotheses at the seed-to-seedling transition, the potentially most critical stage for species discrimination, we conducted a reciprocal transplant experiment combined with a pest exclosure treatment at a wet and a dry forest site in Panama with seeds of 26 species with contrasting origin. Establishment success after one year did not reflect species distribution patterns. However, in the wet forest, wet origin species had a home advantage over dry forest species through higher growth rates. At the same time, drought limited survival of wet origin species in the dry forest, supporting the physiological tolerance hypothesis. Together these processes sort species over longer time frames, and exclude species outside their respective home range. Although we found pronounced effects of pests and some effects of light availability on the seedlings, they did not corroborate the pest pressure nor light availability hypotheses at the seed-to-seedling transition. Our results underline that changes in water availability due to climate change will have direct consequences on tree regeneration and distributions along tropical rainfall gradients, while indirect effects of light and pests are less important.     

Seed bank,seed size and dispersal in moisture gradients of temporary pools in Southern France

Plants reduce risk of extinction due to unpredictable rainfall by soil seed banks, dispersal or large seeds. However, seed size also increases independently in dry habitats, and since seed size is in a trade-off with seed number, size of seed banks is expected to increase in moister habitats. Therefore, we wanted to test if seed abundance in soil increases in wet habitats, if seed size increases in dry habitats, and if spread of seeds along the gradient is higher for plants of intermediate habitats in local moisture gradients.We studied 15 temporary pools in three biogeographically separated wetlands in Southern France. For each pool we studied five different moisture levels, totalling 75 local plant communities. We quantified soil seed bank by the seedling emergence method, seed size and an index of spatial spread of seeds in the soil for every species. We also quantified water levels for each plot.We found increasing abundance of seeds in the soil with increasing water levels but lower seed size and higher spread at intermediate water levels. When we controlled for niche position, we found no trade-off between seed size, spread and abundance in the soil seed bank.Type and importance of risk reduction strategies thus appeared to be strongly driven by the plant species’ moisture niche and the spatial arrangement of water levels.     

Rainfall seasonality and drought performance shape the distribution of tropical tree species in Ghana

Tree species distribution in lowland tropical forests is strongly associated with rainfall amount and distribution. Not only plant water availability, but also irradiance, soil fertility, and pest pressure covary along rainfall gradients. To assess the role of water availability in shaping species distribution, we carried out a reciprocal transplanting experiment in gaps in a dry and a wet forest site in Ghana, using 2,670 seedlings of 23 tree species belonging to three contrasting rainfall distributions groups (dry species, ubiquitous species, and wet species). We evaluated seasonal patterns in climatic conditions, seedling physiology and performance (survival and growth) over a 2‐year period and related seedling performance to species distribution along Ghana's rainfall gradient. The dry forest site had, compared to the wet forest, higher irradiance, and soil nutrient availability and experienced stronger atmospheric drought (2.0 vs. 0.6 kPa vapor pressure deficit) and reduced soil water potential (?5.0 vs. ?0.6 MPa soil water potential) during the dry season. In both forests, dry species showed significantly higher stomatal conductance and lower leaf water potential, than wet species, and in the dry forest, dry species also realized higher drought survival and growth rate than wet species. Dry species are therefore more drought tolerant, and unlike the wet forest species, they achieve a home advantage. Species drought performance in the dry forest relative to the wet forest significantly predicted species position on the rainfall gradient in Ghana, indicating that the ability to grow and survive better in dry forests and during dry seasons may allow species to occur in low rainfall areas. Drought is therefore an important environmental filter that influences forest composition and dynamics. Currently, many tropical forests experience increase in frequency and intensity of droughts, and our results suggest that this may lead to reduction in tree productivity and shifts in species distribution.     

Peatland distibution along a north-south transect in the Mackenzie River Basin in relation to climatic and environmental gradients

Climate is a major factor affecting the development and form of peatlands, as well as the distribution of individual bryophyte species. This paper examines the climatic and ecological gradients affecting the distribution of peatland types along a north-south gradient in the Mackenzie River Basin. Based on a TWINSPAN analysis of bryophyte abundance from 82 peatlands in the Mackenzie River Basin, seven peatland types, two with southerly geographical distributions are recognized. In the Mackenzie River Basin, such local factors as surface water chemistry, pH, and solute concentration as well as height above the water table play a significant role in determinining bryophyte species distributions. Climate is secondary. Amongst the climatic variables, precipitation, length of the growing season, and annual temperature are the most signifcant. The seven peatland groups are: widespread poor fens; peat plateaus with thermokarst pools, low-Boreal bogs; bogs and peat plateus without thermokarst pools; low-Boreal dry poor fens; wet moderate-rich fens; and wet extreme-rich fens.     

Can community structure track sea‐level rise? Stress and competitive controls in tidal wetlands

Climate change impacts, such as accelerated sea‐level rise, will affect stress gradients, yet impacts on competition/stress tolerance trade‐offs and shifts in distributions are unclear. Ecosystems with strong stress gradients, such as estuaries, allow for space‐for‐time substitutions of stress factors and can give insight into future climate‐related shifts in both resource and nonresource stresses. We tested the stress gradient hypothesis and examined the effect of increased inundation stress and biotic interactions on growth and survival of two congeneric wetland sedges, Schoenoplectus acutus and Schoenoplectus americanus. We simulated sea‐level rise across existing marsh elevations and those not currently found to reflect potential future sea‐level rise conditions in two tidal wetlands differing in salinity. Plants were grown individually and together at five tidal elevations, the lowest simulating an 80‐cm increase in sea level, and harvested to assess differences in biomass after one growing season. Inundation time, salinity, sulfides, and redox potential were measured concurrently. As predicted, increasing inundation reduced biomass of the species commonly found at higher marsh elevations, with little effect on the species found along channel margins. The presence of neighbors reduced total biomass of both species, particularly at the highest elevation; facilitation did not occur at any elevation. Contrary to predictions, we documented the competitive superiority of the stress tolerator under increased inundation, which was not predicted by the stress gradient hypothesis. Multifactor manipulation experiments addressing plant response to accelerated climate change are integral to creating a more realistic, valuable, and needed assessment of potential ecosystem response. Our results point to the important and unpredicted synergies between physical stressors, which are predicted to increase in intensity with climate change, and competitive forces on biomass as stresses increase.     

Seasonal drought limits tree species across the Neotropics

Within the tropics, the species richness of tree communities is strongly and positively associated with precipitation. Previous research has suggested that this macroecological pattern is driven by the negative effect of water‐stress on the physiological processes of most tree species. This implies that the range limits of taxa are defined by their ability to occur under dry conditions, and thus in terms of species distributions predicts a nested pattern of taxa distribution from wet to dry areas. However, this ‘dry‐tolerance’ hypothesis has yet to be adequately tested at large spatial and taxonomic scales. Here, using a dataset of 531 inventory plots of closed canopy forest distributed across the western Neotropics we investigated how precipitation, evaluated both as mean annual precipitation and as the maximum climatological water deficit, influences the distribution of tropical tree species, genera and families. We find that the distributions of tree taxa are indeed nested along precipitation gradients in the western Neotropics. Taxa tolerant to seasonal drought are disproportionally widespread across the precipitation gradient, with most reaching even the wettest climates sampled; however, most taxa analysed are restricted to wet areas. Our results suggest that the ‘dry tolerance' hypothesis has broad applicability in the world's most species‐rich forests. In addition, the large number of species restricted to wetter conditions strongly indicates that an increased frequency of drought could severely threaten biodiversity in this region. Overall, this study establishes a baseline for exploring how tropical forest tree composition may change in response to current and future environmental changes in this region.     

Diverging drought-tolerance strategies explain tree species distribution along a fog-dependent moisture gradient in a temperate rain forest

The study of functional traits and physiological mechanisms determining species’ drought tolerance is important for the prediction of their responses to climatic change. Fog-dependent forest patches in semiarid regions are a good study system with which to gain an understanding of species’ responses to increasing aridity and patch fragmentation. Here we measured leaf and hydraulic traits for three dominant species with contrasting distributions within patches in relict, fog-dependent forests in semiarid Chile. In addition, we assessed pressure–volume curve parameters in trees growing at a dry leeward edge and wet patch core. We predicted species would display contrasting suites of traits according to local water availability: from one end favoring water conservation and reducing cavitation risk, and from the opposite end favoring photosynthetic and hydraulic efficiency. Consistent with our hypothesis, we identified a continuum of water use strategies explaining species distribution along a small-scale moisture gradient. Drimys winteri, a tree restricted to the humid core, showed traits allowing efficient water transport and high carbon gain; in contrast, Myrceugenia correifolia, a tree that occurs in the drier patch edges, exhibited traits promoting water conservation and lower gas exchange rates, as well low water potential at turgor loss point. The most widespread species, Aextoxicon punctatum, showed intermediate trait values. Osmotic compensatory mechanism was detected in M. correifolia, but not in A. punctatum. We show that partitioning of the pronounced soil moisture gradients from patch cores to leeward edges among tree species is driven by differential drought tolerance. Such differences indicate that trees have contrasting abilities to cope with future reductions in soil moisture.     

Impact of competition on plant performances along a flooding gradient: a multi‐species experiment

Question: 1. How do the competitive response and the importance of competition vary between species and along a flooding gradient? 2. How does the role of competition in constraining species distribution limits along the gradient vary between lower and upper limits? Location: A 1‐ha meadow within the Alzette floodplain in Luxembourg. Methods: Competitive response and importance of competition were assessed on seven meadow species differing in their tolerance to flooding. Species were cultured in monocultures and in mixtures, in three water treatments reflecting the wet, the middle and the dry end of a natural flooding gradient. We developed two models based on a multiple regression in order to express each component of competition as a function of the neighbour biomass. Results: Five species showed variations in their competitive response across water treatments; however, these species achieved either their highest or their worst competitive response in their optimal water treatment (i.e. the treatment in which the species had the highest biomass in monoculture). Competition was more important for the flood‐tolerant species in the dry treatment than for the flood‐intolerant species in the wet treatment. Conclusions: 1. Variations in species competitive responses along flooding gradients may be the result of either an amplified effect between competition and hydrological stresses, or a hierarchical effect of stress over competition. 2. The role of competition is more important in constraining the upper distribution limits of the flood‐tolerant species than the lower limits of the flood‐intolerant species along flooding gradients.     

Seasonal Dependent Effects of Flooding on Plant Species Survival and Zonation: a Comparative Study of 10 Terrestrial Grassland Species

Past research has provided compelling evidence that variation in flooding duration is the predominant factor underlying plant species distribution along elevation gradients in river floodplains. The role of seasonal variation in flooding, however, is far from clear. We addressed this seasonal effect for 10 grassland species by testing the hypothesis that all species can survive longer when flooded in winter than when flooded in summer. We carried out an inundation experiment under simulated conditions of summer and winter flooding in the greenhouse. The results showed that all species survived longer under winter floods than under summer floods. However, responses upon flooding were species-specific. All summer flood-tolerant species had high tolerance for winter floods as well, but summer flood sensitive species survived either a little longer, or dramatically longer when flooded under simulated winter conditions. Next, we examined whether winter or summer survival best predicted the lower distribution limits of the species as measured in a natural flooding gradient after an extremely long winter flood. We found a strong significant relationship between the lower distribution limits of species in the field and their tolerance to summer floods, although we measured the lower limits 14 years after the latest major summer flood. In contrast, no such significant relationship existed with species tolerance to winter floods. Some relatively intolerant species occurred at much higher floodplain elevations as was expected from their tolerance to winter inundation in the experiments. This suggests that zonation patterns as created by occasional summer floods may be maintained for a long time, probably due to the limited ability of species to re-colonise lower positions in the floodplain.     

Within-community vegetation structure in the conifer woodlands of the Siskiyou Mountains,Oregon

The conifer woodlands found on serpentine-derived soils in southwest Oregon, USA, are characterized by a debse but discontinuous shrub stratum and a species-rich herbaceous stratum. Quantitative analysis of shrub and herb distributions within 10 sites showed that small-scale patterns within the conifer woodland sites are as important as landscape-scale patterns in community organization. Gradient analysis was used to describe the distribution of herbaceous species with respect to gradients of shrub influence (shading, decreased soil temperature, increased soil moisture, increased litter depth) within sites and topographic moisture among sites. Regression analysis of the resulting species distributions and calculations of alpha and beta diversity showed that (a) the shrub-influence microgradient significantly affected distributions for 15 out of 20 major herbaceous species, and the topographic-moisture gradient influenced 13 out of the 20 species, (b) species richness was higher under intermediate conditions along both gradients, and (c) beta diversity within communities was 2.5 to 3.8 times the beta diversity of the site-to-site topographic-moisture gradient.     

Patterns of Richness and Abundance in a Tropical African Leaf‐litter Herpetofauna1

I compared species richness and habitat correlates of leaf‐litter herpetofaunal abundance in undisturbed and selectively logged forests, and an abandoned pine plantation in Kibale National Park, Uganda. I sampled 50 randomly located 25 m² litter plots in each area during the wet and dry seasons in 1997. Ten anuran, five lizard, and three snake species were captured in plots over the study. Assemblage composition was most similar at logged and unlogged sites. The logged forest herpetofauna had higher species richness and abundance than the unlogged forest, but diversity was greater in the unlogged forest due to greater evenness. In contrast, the pine plantation site had the highest richness, abundance, and evenness of the three study sites, but species composition was distinct from the other areas. Herpetofaunal densities were significantly lower in all three areas during the dry season than in the wet season. During the dry season, soil moisture, litter mass, topography, shrub cover, and number of fallen logs were significant positive predictors of herpetofaunal presence in litter plots, but only soil moisture was significant in the wet season. The interaction of moisture and topography appears to be important in determining seasonal patterns of litter herpetofaunal distribution. Comparison of litter herpetofaunal studies across the tropics have shown that mid‐elevation faunas generally support fewer species than lowland faunas. Compared with other tropical mid‐elevation litter faunas, Kibale supports an intermediate number of species, but at lower densities than observed at any other mid‐elevation site reported in the literature.     

Rainfall seasonality and pest pressure as determinants of tropical tree species' distributions

Drought and pests are primary abiotic and biotic factors proposed as selective filters acting on species distributions along rainfall gradients in tropical forests and may contribute importantly to species distributional limits, performance, and diversity gradients. Recent research demonstrates linkages between species distributions along rainfall gradients and physiological drought tolerance; corresponding experimental examinations of the contribution of pest pressure to distributional limits and potential interactions between drought and herbivory are limited. This study aims to quantitate differential performance and herbivory as a function of species range limits across a climatic and floristic transition in Southeast Asia. Khao Chong Botanical Garden, Thailand and Pasoh Forest Reserve, Malaysia straddle the Kangar‐Pattani Line. A reciprocal transplantation across a seasonality gradient was established using two groups of species (“widespread” taxa whose distributions include seasonally dry forests and “aseasonal” taxa whose distributions are limited to aseasonal forests). Growth, biomass allocation, survival, and herbivory were monitored for 19 months. Systematic differences in performance were a function of species distribution in relation to rainfall seasonality. In aseasonal Pasoh, aseasonal species had both greater growth and survivorship than widespread species. These differences were not a function of differential herbivory as widespread and aseasonal species experienced similar damage in the aseasonal forest. In seasonally dry Khao Chong, widespread species showed higher survivorship than aseasonal species, but these differences were only apparent during drought. We link this differential performance to physiological mechanisms as well as differential tolerance of biotic pressure during drought stress. Systematic decreases in seedling survival in aseasonal taxa during drought corresponded with previously documented physiological differences and may be exacerbated by herbivore damage. These results have important implications for tropical diversity and community composition in light of predicted increases in the frequency and severity of drought in hyperdiverse tropical forests.     

The effect of soil on the growth performance of tropical species with contrasting distributions

Within the tropics, a marked gradient in rainfall between dry and wet forests correlates with a well documented turnover of plant species. While water availability along these gradients is an important determinant of species distributions, other abiotic and biotic factors correlate with rainfall and may also contribute to limit species distribution. One of these is soil fertility, which is often lower in the wetter forests. To test its possible role in species distribution along a rainfall gradient, we performed a screen‐house experiment where we measured the growth performance of seedlings of 23 species with contrasting distributions across the Isthmus of Panama. We grew seedlings in soils collected from the drier Pacific side and the wetter Atlantic side. Differences in soil fertility across the Isthmus were large enough to significantly influence the growth performance of the seedlings. However, we found no evidence of home‐soil advantage among species with contrasting distributions. Dry‐distribution species grew on average slower than wet‐distribution species suggesting a cost to drought adaptations. The response to soil differences correlated with the growth rate of the species, such that fast‐growing species responded more to changes in soil quality. We hypothesize that inherently slow growth rates of some dry‐distribution tropical species may be a more important factor limiting their colonization of wetter sites along the rainfall gradient.     

Shoreline vegetation of Lake Nubia, Sudan

The paper reports on the flora along the banks of Lake Nubia and the long-term changes that have taken place since the formation of the Aswan High Dam Lake 30 years ago. The study also aims to determine the main factors that govern plant distribution in this area. Shoreline vegetation was studied from Debeira at the Sudanese–Egyptian border to the ‘Dal’ Cataract at the south end of the lake, in Sudan. The shoreline was classified into four moisture gradient zones depending on the period of inundation, namely: wet zone (frequently inundated and recently exposed); moist zone (periodically inundated); semi-dry zone (rarely inundated); and dry zone (never inundated). Plants characterising each zone were identified. Soil texture and thickness of the deposits were measured for each zone at each site. According to Grime’s strategy, the study revealed that species of the wet and the moist zones are mainly disturbance-tolerant; species observed in the ‘semi-dry’ zone are competitive-stress-tolerant; and those of the ‘dry’ zone are stress-tolerant. The present study indicates that elevation above mean sea level, and period of inundation and texture of deposits play an important role in governing the distribution of the shoreline vegetation.     

陆地植物群落物种多样性的梯度变化特征

研究陆地植物落物种多样性随环境因子及群落演替梯度的变化特征是揭示生物多样性与生态因子相互关系的重要方面,根据近期国内外的文献,综述了这方面的研究进展。随纬度的降低,通常物咱多样性随中,随不分梯度的变化,物种多样性的变化有６种趋势;随海拔高的变化,物处多样性有５种模式;随土壤养分梯度的变化,表现出不同的规律;演替过程中物种多样生的变化趋势相似。关于植物群落物种多样性梯度格局的机制有多种假说,但仍需进     

Competition Is a Strong Driving Factor in Wetlands,Peaking during Drying Out Periods

The aim of the study is to investigate the relative importance of plant-plant interactions with regard to flooding and drought effect on perennial plant performances in wetlands. Flooding is expected to be the major driver and, accordingly, the importance of drought is hardly if ever taken into account. Focusing on five widespread species, the growth, the survival and the competitive ability of plants were monitored on permanent plots spread along two elevation gradients. Flooding duration and drought intensity were found to vary substantially along the ~ 0.5 meter range elevation gradient. Flooding and drought alternate over the hydrological year and the pin-point surveys were thus conducted over the course of one year. The data were modeled taking into account survival, recruitment and competitive growth throughout flooding and drying out periods. Flooding and drought both directly impacted the plant performances and their competitive effect, with the effect of drought being much more general among species and of higher magnitude than flooding. The importance of competition was found to be high for all species, particularly during the drying out period. It varied more along the flooding gradient than along the drought gradient. The higher flooding tolerance shown by the studied species compared to drought may be related to species specific growth timing together with efficient response traits. These results offer new insights into the filters operating over the species pools. This suggests that the drying out period and drought conditions may be even more important for species’ relative success and the importance of competition than the flooding pattern. The general applicability of this result, obtained in mild Atlantic climate and fertile wetlands, remains to be studied.     

Ecology of a steppe-tundra gradient in interior Alaska

Abstract. Subarctic steppe is currently restricted in interior Alaska and the Yukon Territory to steep, south-facing river bluffs. Paleoecological and biogeographic evidence suggests that some steppe taxa may have been more widespread during the Full-Glacial. We examined factors controlling the distribution of steppe taxa on an elevation gradient across a steppetundra ecotone; such analyses may help define potential Full-Glacial distributions of these taxa. Multivariate analyses suggest that species can be divided into four spatially distinct groups, but individualistic species distributions create considerable overlap among these groups. The steppe-tundra ecotone comprises a broad zone of mixing between steppe taxa and drought-tolerant alpine tundra taxa, followed by an abrupt shift to alpine shrub tundra. The transition from low steppe to tundra vegetation is primarily associated with a gradient of decreasing soil temperature. The more abrupt transition from mixed steppe-tundra to alpine shrub tundra vegetation is primarily associated with changes in soil depth and soil moisture. Variation in vegetation within steppe is associated with gradients in soil phosphorus and moisture. Greenhouse experiments on drought tolerance of two steppe and two tundra taxa suggest that the individualistic distribution of species along the ecotone is partly a function of physiological differences among species. Our analyses of vegetation-environment relationships support the hypothesis that some components of the steppe community could have been more widespread during the colder Full-Glacial.     

Fern species richness along a central Himalayan elevational gradient, Nepal

Aim The study explores fern species richness patterns along a central Himalayan elevational gradient (100–4800 m a.s.l.) and evaluates factors influencing the spatial increase and decrease of fern richness. Location The Himalayas stretch from west to east by 20°, i.e. 75–95° east, and Nepal is located from 80 to 88° east in this range. Methods We used published data of the distribution of ferns and fern allies to interpolate species elevational ranges. Defining species presence between upper and lower elevation limit is the basis for richness estimates. The richness pattern was regressed against the total number of rainy days, and gradients that are linearly related to elevation, such as length of the growing season, potential evapotranspiration (PET, energy), and a moisture index (MI = PET/mean annual rainfall). The regressions were performed by generalized linear models. Results A unimodal relationship between species richness and elevation was observed, with maximum species richness at 2000 m. Fern richness has a unimodal response along the energy gradients, and a linear response with moisture gradients. Main conclusions The study confirms the importance of moisture on fern distributions as the peak coincides spatially with climatic factors that enhance moisture levels; the maximum number of rainy days and the cloud zone. Energy‐related variables probably control species richness directly at higher elevations but at the lower end the effect is more probably related to moisture.     

Off-channel temporary pools contribute to native riparian plant species diversity in a regulated river floodplain

Off-channel temporary pools on riverbanks have characteristic seasonal wet–dry cycles resulting from direct inflow of river water or hyporheic flow seepage. This study addressed two questions regarding the role of temporary pools in supporting diversity of riparian vegetation in a regulated river in Japan: (1) do temporary pools maintain high native riparian plant species diversity? And, (2) how do physical environmental factors affect the pattern of plant species distribution? The study was conducted on a 1-km section of the Hayade River alluvial fan in Niigata Prefecture, central Japan. Two to five transect belts consisting of 20 contiguous plots (1 m × 1 m) were laid out in two off-channel temporary pools and six other sites classified by physiognomy. Vascular plant presence, relative elevation, and substratum type were recorded in all plots. Species richness, Simpson’s reciprocal index (1/D), and Shannon–Wiener function (H′) were calculated by life form to estimate plant species diversity. Two conclusions could be drawn from the results. First, the temporary pools maintained high native riparian plant species diversity in this regulated river floodplain. Second, several environmental factors (seasonal wet–dry cycle, low elevation, complex micro-topography, and fine substrata) created spatial and seasonal heterogeneity of moisture conditions in the temporary pools, supporting plant species diversity. For sustainable maintenance of riparian plant species diversity, progressive river-management should restore original riverine dynamics to generate a shifting mosaic of diverse geomorphology.     

Biogeographical gradients in galling species richness

Summary Five hypotheses were invoked to account for variation in galling species number per location on plants of different structural complexity, namely herbs, shrubs, and trees, both in Brazil and USA. The hypotheses were: 1) the altitudinal/latitudinal gradient hypothesis; 2) the harsh environment hypothesis; 3) the plant species richness hypothesis; 4) the host plant area hypothesis; 5) the plant structural complexity hypothesis. The altitudinal and the harsh environment hypotheses were correlated and supported with sample data in both localities, with increasing gall species number as altitude/latitude declined and as sites became hotter and drier. The two hypotheses were separated by studying riparian sites and dry hillside sites at the same elevation in Arizona. Galling species frequency was higher in dry sites than in riparian sites, supporting the harsh environment hypothesis. Of the five hypotheses tested only the harsh environment hypothesis predicted that galling insect species number should vary in response to environmental variables such as moisture and temperature. Temperate shrubs supported more galling species than did other plant types, both in dry and mesic sites. The overall difference between galling species richness for tropical and temperate latitudes was not statistically significant. Free-feeding insect herbivore species exhibited the opposite pattern of species richness to gallers, being more speciose in riparian sites. The present study corroborates the hypothesis that the gall forming habit is an adaptation to harsh or stressful environments, and we describe for the first time broad scale geographical patterns in galling insect species richness.