Reef fish assemblage structure affected by small-scale spacing and size variations of artificial patch reefs

To examine how varying the distance between patch reefs affects reef fish assemblage structure, replicate concrete reef modules (∼ 1 m³ each) were deployed on sand bottom at 8 m depth off Ft. Lauderdale, Florida, USA (26°07N, 80°05W). Modules were positioned at the apices of one of four differently sized equilateral triangles. Triangular configurations had side lengths of: 25 m, 15 m, 5 m, and 0.33 m; each treatment with two replicates. Two additional configurations: (1) a solitary module (Single) and (2) two modules side by side (Double), also with two replicates, were deployed in order to examine the interaction of reef size with fish assemblages. SCUBA divers censused fishes monthly, for 2 years, recording the species present, their abundance and sizes (TL). Fishes were assigned to one of five length categories: < 2 cm, > 2-5 cm, > 5-10 cm, > 10-20 cm, and > 20 cm. In general and excluding the smallest three-module spacing treatment (0.33 m treatment), which may have provided unique treatment-specific refuge, total fish abundance and richness were shown to increase when isolation distance increased. However, there were also species-specific and size class differences in response to isolation distance. The second part of this study indicated varying reef size, by doubling and tripling the number of reef modules, increased total fish abundance and species richness. Nevertheless, fish abundance and species richness did not change by an identical multiplier (e.g., doubling modules ≠ double abundance). These results suggest that scientists and marine managers alike should consider reef size and isolation as habitat attributes capable of altering the structure and dynamics of reef fish assemblages.     

A size analysis of planktic foraminifera from the Arabian Sea

Planktic foraminiferal faunas from different environments in the Arabian Sea were size fractionated using 14 sieves with meshes between 100 and 710 μm, to assess the effect of the sieve mesh size cut off level on the faunal composition and to determine the size frequency distribution of individual species. Nine samples from a plankton pump and a towed net, a sediment trap, a box-core and a piston core were selected, to cover living and settling flux faunas as well as fossil faunas from the sediment. In living faunas, most species show an exponential size frequency distribution, with highest numbers in the finest interval of the size spectrum. In sediment trap and core samples, individual species size frequency distributions may consist of: (1) an exponential distribution of relatively small pre-adult specimens; (2) a Gaussian-shaped distribution of larger specimens, which may be classified as adult or terminal; or (3) a combination of both. The distributions are separated using a best fit technique. The composition of the total planktic foraminiferal fauna strongly changes along the size spectrum. Dominant taxa in >355 μm fractions are Orbulina universa, Globorotalia menardii, Globorotalia tumida, Globigerinella siphonifera and Globigerinoides sacculifer, in 125–355 μm fractions Globigerina bulloides, Globigerinoides ruber, Neogloboquadrina dutertrei and Globigerinita glutinata, and in <125 μm fractions Dentigloborotalia anfracta, Tenuitella compressa, Tenuitella iota, Turborotalita quinqueloba and the immature specimens of larger species. Consequently, the choice of the sieve mesh size strongly determines the percent composition of the assemblage and in turn the paleoceanographic interpretations based on these counts. Species richness and the Shannon diversity increase with decreasing sieve mesh size, while equitability generally decreases with decreasing size. In the water column approximately 60% of the fauna (>100 μm) is present in the 100–125 μm fraction and 1–6% is larger than 250 μm. In samples representing a settling flux (sediment trap and sediment samples) 29–57% of the fauna is present in the 100–125 μm fraction, while 6–23% is larger than 250 μm. Size frequency distributions of the dextral Neogloboquadrina complex (= Neogloboquadrina dutertrei and Neogloboquadrina pachyderma + P–D intergrades) show a bimodal pattern; a smaller peak reflecting dextral Neogloboquadrina pachyderma, and a larger peak of adult Neogloboquadrina dutertrei. By applying a best fit technique to the data, the two species may be separated from each other. In size fractions larger than 150 μm most species have reached the adult stage of ontogeny and we recommend this mesh size for standard faunal analysis. In addition, sieve mesh sizes of 125 and 250 μm have to be used to obtain a reliable estimate of the abundance of small and large species, respectively.     

At what spatial scale(s) do mammals respond to urbanization?

Spatial scale is fundamental in understanding species–landscape relationships because species’ responses to landscape characteristics typically vary across scales. Nonetheless, such scales are often unidentified or unreliably predicted by theory. Many landscapes worldwide are urbanizing, yet the spatial scaling of species’ responses to urbanization is poorly understood. We investigated the spatial scaling of urbanization effects on a community of 15 mammal species using ~60 000 wildlife detections collected from a constellation of 207 camera traps across an extensive urban park system. We embedded a bivariate Gaussian kernel in hierarchical multi-species models to determine two scales of effect (a scale of maximal effect and a broader scale of cumulative landscape effect) for two biological responses (occupancy and site visit frequency) across two seasons (winter and summer) for each species. We then assessed whether scales of effect varied according to theoretical predictions associated with biological responses and species traits (body size and mobility). Scales of effect ranged from < 50 m to > 9000 m and varied among species, but not as predicted by theory. Species’ occupancy generally showed a weak response to urbanization and the scale of this effect was both highly uncertain and consistent across species. We did not detect any relationship between scales of effect and species’ body size or mobility, nor was there any evident pattern of scaling across biological response or seasons. These results imply that 1) urbanization effects on mammals manifest across a very broad spectrum of spatial scales, and 2) current theories that a priori predict the scale at which urbanization affects mammals may be of limited use within a given system. Overall, this study suggests that developing general theory regarding the scaling of species–landscape relationships requires additional empirical work conducted across multiple species, systems and timescales.     

GENOTYPIC VARIATION AND CLONAL STRUCTURE IN CORAL POPULATIONS WITH DIFFERENT DISTURBANCE HISTORIES

Genotypic diversity in six populations of the endemic Hawaiian reef coral, Porites compressa, was directly related to habitat-disturbance history. The highest diversity (lowest amount of clonal proliferation) was found in populations that had been intensely or recently disturbed. In these populations, space was not limited and mean colony size was small (< 500 cm²), suggesting early stages of recolonization. In an undisturbed, protected habitat, lower genotypic diversity was a result of a significant degree of clonal replication of established genotypes. Unoccupied substratum was rare in this habitat, and average colony size was large (> 2500 cm²). Populations in intermediately disturbed habitats showed intermediate levels of diversity and clonal structure as a result of the combined contributions of sexual and asexual reproduction. Individual clones were distributed over small areas (< 4 m²) or distances (< 1 m) in young populations, and more broadly (> 256 m²) and over longer distances (> 90 m) in the older, undisturbed population. Interpretations of life-history parameters and estimates of total genetic variability in species that have the potential to reproduce asexually are dependent upon an assessment of the overall clonal structure of populations. The power of genotypic assays to reliably detect clonal versus unique colonies, as well as the spatial scales over which clonal populations are sampled, are critical to such assessments.     

Patterns of morphological and genetic diversity of Valeriana jatamansi Jones in different habitats and altitudinal range of West Himalaya,India

Importance to know and understand diversity of Himalayan plants is increasingly recognized considering the fact that various natural and anthropogenic pressures might bring about serious influences to morphological and genetic diversity of the vegetation in the region. In this context, Valeriana jatamansi was investigated in detail, taking into account its importance in various Ayurvedic and modern medicines. Randomly selected mature plants from twenty five different populations (located between 1215 m to 2775 m asl) of V. jatamansi were analysed for their morphological attributes. Further, ISSR markers were used to detect genetic variation among 151 plants of selected 25 populations. Use of 20 primers yielded 125 reproducible polymorphic loci which were used to estimate different parameters of genetic diversity. These parameters were in turn applied to develop relationships with habitat types and altitude range. Significant variation (p < 0.05) in above ground dry weight (AGDW) and below ground dry weight (BGDW) across the populations was observed. Nei's genetic diversity index (He) ranged from 0.25 to 0.37 across the populations, with a mean of 0.31. Genetic diversity exhibited a decreasing trend with increasing altitude, and maximum diversity (He = 0.325) was observed in the range of 1201–1500 m asl. Among the different habitat conditions, highest genetic diversity (He = 0.334; Pp = 84.38) was observed in grassland habitats while minimum in mixed forest habitats (He = 0.285; Pp = 72.433). The genetic diversity (He) had significant negative relationships with AGDW, BGDW and rhizome diameter (Pearson r = −0.359, −0.424 and −0.317, respectively; p < 0.05). The genetic characterization of V. jatamansi from the western Himalaya by this study suggests influences of habitat types and the altitudinal range upon genetic diversity, and based on these proposals for conservation strategies in favour of the species are made.     

Evaluation of low impact pressure waves as a quantitative sampling method for small fish in shallow water

Low impact pressure waves (LIMP) generated by small (< 1 g) underwater ignition capsules have been used to qualitatively and semi-quantitatively, as catch per unit effort, sample young-of-the-year (YOY) and juvenile fish in shallow (0-3 m) waters. In order to evaluate this method on small fish, four categories, 0+ and 1+ year Eurasian perch, 1+ year roach and 0+ to 1+ year pike, were subjected to the LIMP at distances of 0.5-5 m from the detonation point. At a distance < 1 m all specimens were stunned and either floated to the surface or sank to the bottom, whereas almost all were unaffected at distances > 3 m. The efficiency radius of the LIMP varied between 1.5 and 3.0 m, and the effective area was 7-28 m² depending on habitat and species. The average effective area for the two extensively studied categories (0+ perch and 1+ roach) was 17.9 m². Both depth and vegetation affected the effective area. The smallest area was obtained at < 1 m depth with vegetation present, and the largest at 1-2 m depths with no vegetation present. Within each fish category, the method sampled fish independently of length suggesting that a size/year-class is sampled equally. The proportion of affected fish that sank was large and species-dependent. In addition to the netting of surfaced fish, collecting fish from the bottom is recommended for achieving an accurate quantitative estimate. The tested parameters of the LIMP, P_max and Impulse, declined as a function of distance following a negative exponential growth model with a baseline. The similar-shaped responses among the fish categories suggest that the method works at depths < 3 m also in heterogeneous environments, and that the method can be used to quantitatively estimate abundance of small fish with swim bladders. A model for quantitative abundance estimates based on the evaluation is presented.     

Spatial patterns in impacts of fishing on temperate rocky reefs: Are fish abundance and mean size related to proximity to fisher access points?

Effects of fishing on marine communities are becoming increasingly evident, yet little is known of the spatial extent of impacts, particularly for multiple impacts distributed over broad scales. We tested the common perception that commercial and recreational fishing on inshore temperate reefs generate spatial impacts that diminish with distance from fisher access points. We collected data on harvested and non-harvested reef species using underwater visual censuses at 133 shallow rocky reef sites around Tasmania and tested for relationships between assemblage and species level indices of fishing impacts and distance to the nearest boat launching ramp. Slopes of size spectra of fish communities tended to decrease with distance from the nearest boat ramp, with this relationship apparently resulting from low numbers of large fish (> 30 cm TL) and a greater number of smaller fish (< 15 cm TL) at sites closest to access points. At the species level, relationships were evident either in the abundance of legal individuals or the mean size of harvested species with distance to the nearest boat ramp, except for rock lobster. Patterns for rock lobster differed when areas in which commercial or recreational fisheries dominated were considered separately from the statewide analysis. A pattern of increasing numbers of legal lobsters with increasing distance from boat ramps was observed, but only in the areas in which the recreational fishery dominated. Observed relationships in all species were consistent with greater fishing impacts at sites closest to boat ramps, with the exception of exploited wrasses. Banded morwong, which are subject to a live export fishery, appeared to be most affected by proximity to boat ramps. Conversely, no relationships were found between the abundance or size of the most frequently occurring non-harvested species and distance to boat ramps.These results support the hypothesis that greater fishing impacts occur at more accessible sites over the entire Tasmanian coastline. The variability of results among individual species are likely, at least in part, to be related to differences in fisheries characteristics such as vessel size and range, as well as the suitability of our methods for detecting impacts. The potential of such a pattern in fishing impacts to be evident in other locations will thus likely depend on characteristics of the particular fishes and fisheries.     

Fine‐scale Spatial Genetic Structure of Ten Dipterocarp Tree Species in a Bornean Rain Forest

Fine‐scale spatial genetic structure is increasingly recognized as an important factor in the studies of tropical forest trees as it influences genetic diversity of local populations. The biologic mechanisms that generate fine‐scale spatial genetic structure are not fully understood. We studied fine‐scale spatial genetic structure in ten coexisting dipterocarp tree species in a Bornean rain forest using microsatellite markers. Six of the ten species showed statistically significant fine‐scale spatial genetic structure. Fine‐scale spatial genetic structure was stronger at smaller spatial scales (≤ 100 m) than at larger spatial scales (> 100 m) for each species. Multiple regression analysis suggested that seed dispersal distance was important at the smaller spatial scale. At the larger scale (> 100 m) and over the entire sample range (0–1000 m), pollinators and spatial distribution of adult trees were more important determinants of fine‐scale spatial genetic structure. Fine‐scale spatial genetic structure was stronger in species pollinated by less mobile small beetles than in species pollinated by the more mobile giant honeybee (Apis dorsata). It was also stronger in species where adult tree distributions were more clumped. The hypothesized mechanisms underlying the negative correlation between clump size and fine‐scale spatial genetic structure were a large overlap among seed shadows and genetic drift within clumped species.     

Length‐weight relationships of 79 marine fish species from the coastal waters of Hong Kong

Dedicated benthic fisheries‐assessment surveys with beam trawls (2 cm mesh size) were used to sample demersal fish communities at depths 7–35 m along four transects in each of the western, southern and eastern waters of Hong Kong monthly for >24 months. We report length‐weight relationships (LWRs) for 79 species (11 orders, 37 families) of fishes for which LWRs are presently absent or scarce. For 56 of these species, relationships between total length and standard length or disc width were also derived.     

6-10× pyrosequencing is a practical approach for whole prokaryote genome studies

Next generation 454 pyrosequencing technology for whole bacterial genome sequencing involves a deep sequencing strategy with at least 15-20 × in depth proposed by official protocols but usually done with over 20 × in practices. In this study, we carried out a comprehensive evaluation of quality of the de novo assemblies based on realistic pyrosequencing simulated data from 1480 prokaryote genomes and 7 runs of machine-generated data. Our results demonstrated that for most of the prokaryote genomes, 6-10 × sequencing in qualified runs with 400 bp reads could produce high quality draft assembly (> 98% genome coverage, < 100 contigs with N50 size > 100 kb, single base accuracy > 99.99, indel error rate < 0.01%, false gene loss/duplication rate < 0.5%). Our study proves the power of low depth pyrosequencing strategy, which provides a cost-effective way for sequencing whole prokaryote genomes in a short time and enables further studies in microbial population diversity and comparative genomics.     

Length–weight relationships of nine freshwater fishes from six rivers of the Western Ghats of India

Length–weight relationships (LWRs) were estimated for nine freshwater fish species collected three times using a scoop net (mesh size 0.3–0.5 cm), cast net (mesh size 1–1.5 cm) and gillnet (mesh size 2.5–4.5 cm) from six rivers of the Western Ghats of India during August 2017–October 2018. The b values for LWRs varied from 2.862 to 3.656 (R² > 0.916 and p < 0.0001 for all species).     

Distribution of bathyal meiofauna in the region of the Subtropical Front, Chatham Rise, south-west Pacific

Metazoan meiofauna was sampled along a temperate latitude transect (41-47°S 178°30′E) across the Chatham Rise, south-west Pacific, from water depths of 350-2600 m on three occasions (austral autumn 1997, spring 1997, summer 2000). Samples were collected using a multicorer and extracted on a 63-μm mesh. Meiofaunal density and biomass were negatively correlated with water depth, median grain-size and sediment calcium carbonate content, and positively correlated with sediment chlorophyll a. There were no significant relationships between meiofaunal abundance and biomass and measures of sediment organic matter and bacterial productivity and biomass, except in summer when bacterial and meiofaunal biomass were positively correlated. Vertical penetration of meiofauna into the sediment was related to sediment organic matter and sediment chloroplastic pigments. Variation in meiofaunal abundance at different spatial scales was examined at two sites: at 450 and 2300 m on the northern slope of the rise. At both sites, variation between subcores of a multicore (< 10-cm scale) was lower than that of multicores within a deployment (< 1-m scale). However, whilst the highest variance component at the shallow site was for multicores within a deployment, it was between multicore deployments (< 1-km scale) for the deep site. Meiofaunal density and biomass were generally higher on the southern slope of the rise than on the northern slope, a trend probably attributable to high productivity of the overlying Subtropical Front and the flux of more nutritious organic material to the sea floor.     

Changing the fire management regime in the renosterveld and lowland fynbos of the Bontebok National Park

This paper evaluates the history of fire management in the Bontebok National Park (3435 ha) over a period of almost four decades. A GIS database was compiled of all fires between 1972 and 2009 and the fire regime was analysed in terms of the frequency, season, size and cause of fires. Since the early 1970s, short interval burning was implemented to promote grazing for bontebok, but from 2004 the fire interval was lengthened to favour plant species diversity, an increasingly urgent conservation priority for the park. In total, 43 fires were recorded, ranging in size from 9 to 1007 ha, collectively spanning 14 013 ha. The majority of fires were large (100–500 ha), with fires of >100 ha accounting for 96% of the area burnt. The overall mean fire return period (FRP) for the park was 7.2 years, which is short judged by fynbos standards. FRPs under the old and new management regimes were 6.7 and 10.9 years respectively. Under the old regime, FRPs in renosterveld and fynbos were 5.8 and 8.0 years respectively. Large parts of the park repeatedly experienced fires at immature vegetation ages resulting in the elimination of slow-maturing seed-regenerating plant species such as Protea repens. Post-fire age distribution was highly skewed towards young vegetation, with 75% of fire-prone vegetation burning at post-fire ages of ≤7 years, and <10% of fire-prone vegetation surviving beyond 10 years of age. Prescribed and accidental fires respectively accounted for 70% and 30% of the total area burnt. Prescribed burning was mostly done in March–April, and only 8% of the total area burnt, burnt outside of the ecologically acceptable fire season. This study identified areas which have been subject to ecologically appropriate and inappropriate fire return intervals, providing a basis for informed future management and research.     

Environmental variables explaining structural and functional diversity of seagrass macrofauna in an archipelago landscape

Thirteen seagrass beds located over a 80-km range in the brackish waters of SW, Finland, northern Baltic Sea were investigated in order to determine the environmental variables important for univariate community measures and for number, composition and redundancy of functional groups of benthic macrofauna. For species assemblages, fetch and shore angle were the best explanatory variables, followed by sediment granulometry (fine gravel) and then sediment organics. Similarly, fetch, shore angle and Zostera marina shoot density were the best explanatory variables for functional group patterns. Small (< 50 m²) inner-archipelago beds were functionally and structurally equal to the most extensive (500 to > 1000 m²) seagrass beds in the study area. Community measures (density, number of species and diversity) and functional diversity (number of functional groups) equalled or exceeded levels previously recorded in deeper, non-vegetated communities in the northern Baltic Sea. In comparison with marine seagrass assemblages, the total number of species and number of species per function were low. However, species density and derived diversity measures (Shannon-Wieners index H′) equalled or exceeded those reported for other seagrass ecosystems. It is concluded that in terms of seagrass infauna, the Baltic Sea should not be regarded species poor, as is often generally stated, and that conservation initiatives and management strategies should consider both minor as well as more extensive occurrences of seagrasses in coastal waters.     

Male powerlifting performance described from the viewpoint of complex systems

This paper reflects on the factors that condition performance in powerlifting and proposes that the result-generating process is inadequately described by the allometric equations commonly used. We analysed the scores of 1812 lifters belonging to all body mass categories, and analysed the changes in the results achieved in each weight category and by each competitor. Current performance-predicting methods take into account biological variables, paying no heed to other competition features. Performance in male powerlifting (as in other strength sports) behaves as a self-organised system with non-linear interactions between its components. Thus, multiple internal and external elements must condition changes in a competitor's score, the most important being body mass, body size, the number of practitioners, and the concurrency of favourable factors in one individual. It was observed that each behaved in a specific form in the high level, according to the individuals’ circumstances, which make up the main elements of the competitive system in every category. In powerlifting, official weight categories are generally organised in three different groups: light (<52.0 to <60 kg), medium (<67.5 to <90.0 kg) and heavy (<100 to >125 kg) lifter categories, each one of them with specific allometric exponents. The exponent should be revised periodically, especially with regard to the internal dynamics of the category, and adjusted according to possible changes affecting competition.     

Seasonal variations in seminal plasma and sperm characteristics of wild-caught and cultivated Atlantic cod, Gadus morhua

The objective was to investigate changes, throughout the spawning season, in body size attributes and quantitative semen characteristics of wild-caught and cultivated Atlantic cod, Gadus morhua L. Sperm velocity increased significantly throughout the spawning season of cod from both origins. Curvilinear velocity (VCL; 30 sec post-activation) increased from 78.9 ± 6.5 to 128.2 ± 6.5 μm/sec (mean ± SEM) between the beginning and end of the spawning season, respectively, for wild-caught cod, whereas for cultivated fish, it increased from 26.6 ± 2.4 to 48.9 ± 3.1 μm/sec between January and March. Spermatocrit did not undergo a significant seasonal change in wild-caught cod but did thicken for cultivated cod (24.6 ± 4.2% in January to 40.5 ± 4.4% in April; P < 0.01). Sperm head area, perimeter, length, and width declined significantly at the end of the spawning season of cod from both origins (all P values < 0.01). Seminal plasma osmolality and Na⁺ ion concentration followed a dome-shaped function through the spawning season for both wild-caught and cultivated cod (P < 0.05). For cultivated cod, seminal plasma pH was significantly lower at the start of the spawning season (P < 0.001), whereas Ca²⁺ increased then decreased (P < 0.05). Body size attributes, spermatocrit, and seminal plasma constituents had significant relationships with sperm activity variables. These relationships varied as a function of time post-activation, month, and fish origin. Our findings may be used to (i) assess spermiation stage without killing males; (ii) optimize semen collection for hatchery production; (iii) characterize the potential impact of farming on sperm quality; and (iv) improve success of sperm cryopreservation and short-term storage.     

Effect of fragmentation on the community structure of epixylic bryophytes in Atlantic Forest remnants in the Northeast of Brazil

The effects of fragmentation and edge effects on the floristic composition, richness, diversity and abundance of epixylic bryophytes (growing on decaying wood) were investigated in ten fragments of Atlantic Forest remnants in the Northeast of Brazil. In each fragment, four perpendicular 100 m transects were demarcated. Along these transects, samples of bryophytes growing on decaying wood were collected. The forest fragments were grouped in three size classes (small: <100 ha; medium: 100–500 ha; large: >500 ha). Correlation and multivariate analysis were undertaken between bryophyte flora and fragment metrics (size, form, isolation, altitude variation, nuclear area and secondary vegetation percentage and distance from the edge). A total of 99 species of bryophytes, 52 liverworts and 47 mosses were registered. The statistical results confirming fragment size is an important factor in epixylic community structure. Therefore, composition, richness, diversity and abundance can be better explained by a junction of all studied landscape factors. Bryophyte richness, the percentage of samples with the greatest coverage of decaying wood and shade-tolerant species’ distribution, were not correlated to distance from the forest edge. This suggests that edge effects are not linear or can be detected beyond 100 m from the edge, which is very important for inclusion in future studies.     

Common spatial patterns of trees in various tropical forests: Small trees are associated with increased diversity at small spatial scales

Tropical forests are notable for their high species diversity, even on small spatial scales, and right‐skewed species and size abundance distributions. The role of individual species as drivers of the spatial organization of diversity in these forests has been explained by several hypotheses and processes, for example, stochastic dilution, negative density dependence, or gap dynamics. These processes leave a signature in spatial distribution of small trees, particularly in the vicinity of large trees, likely having stronger effects on their neighbors. We are exploring species diversity patterns within the framework of various diversity‐generating hypotheses using individual species–area relationships. We used the data from three tropical forest plots (Wanang—Papua New Guinea, Barro Colorado Island—Panama, and Sinharaja—Sri Lanka) and included also the saplings (DBH ≥ 1 cm). Resulting cross‐size patterns of species richness and evenness reflect the dynamics of saplings affected by the distribution of large trees. When all individuals with DBH ≥1 cm are included, ~50% of all tree species from the 25‐ or 50‐ha plot can be found within 35 m radius of an individual tree. For all trees, 72%–78% of species were identified as species richness accumulators, having more species present in their surroundings than expected by null models. This pattern was driven by small trees as the analysis of DBH >10 cm trees showed much lower proportion of accumulators, 14%–65% of species identified as richness repellers and had low richness of surrounding small trees. Only 11%–26% of species had lower species evenness than was expected by null models. High proportions of species richness accumulators were probably due to gap dynamics and support Janzen–Connell hypothesis driven by competition or top‐down control by pathogens and herbivores. Observed species diversity patterns show the importance of including small tree size classes in analyses of the spatial organization of diversity.