Disparity in Onset Timing and Frequency of Flowering and Fruiting Events in Two Bornean Peat‐Swamp Forests

The timing and frequency of flowering and fruiting events are key tropical forest characteristics that have substantial influence on fauna. Although our understanding of geographic variation in habitat‐wide timing and frequency of flowering and fruiting is advancing, corresponding information for individual tree species is limited. Thus, we compared climate and reproductive phenology of 16 tree species over 70 mo at two Bornean tropical peat‐swamp forest sites. We found significant inter‐site correlations in rainfall and temperature, and only small absolute temperature differences. In both sites, most species exhibited within‐site synchrony in flowering and fruiting onset. Broad‐scale flowering and fruiting onset frequency classifications showed high congruence between sites. Significant correlations in flowering and fruiting onset timing between sites were found for only 19 and 17 percent of the species, respectively. This remained the case when applying 1‐ and 2‐month lag periods for both sites, with neither site consistently lagging behind. Significant differences in the exact frequency of new flowering and fruiting events were detected for 44 and 58 percent of species, respectively, and no significant relationships between the onset timing synchrony and exact frequency of new reproductive events were found for either flowers or fruit. We conclude that inter‐site climatic and ecological similarities do not necessarily lead to high inter‐site synchrony in either onset timing or exact frequency of tree reproductive events. Potential reasons for this are discussed, as are the implications for understanding tropical forest ecology and improving forest restoration project seed collections.     

Trait‐dependent distributional shifts in fruiting of common British fungi

Despite the dramatic phenological responses of fungal fruiting to recent climate warming, it is unknown whether spatial distributions of fungi have changed and to what extent such changes are influenced by fungal traits, such as ectomycorrhizal (ECM) or saprotrophic lifestyles, spore characteristics, or fruit body size. Our overall aim was to understand how climate and fungal traits determine whether and how species‐specific fungal fruit body abundances have shifted across latitudes over time, using the UK national database of fruiting records. The data employed were recorded over 45 yr (1970–2014), and include 853 278 records of Agaricales, Boletales and Russulales, though we focus only on the most common species (with more than 3000 records each). The georeferenced observations were analysed by a Bayesian inference as a Gaussian additive model with a specification following a joint species distribution model. We used an offset, random contributions and fixed effects to isolate different potential biases from the trait‐specific interactions with latitude/climate and time. Our main aim was assessed by examination of the three‐way‐interaction of trait, predictor (latitude or climate) and time. The results show a strong trait‐specific shift in latitudinal abundance through time, as ECM species have become more abundant relative to saprotrophic species in the north. Along precipitation gradients, phenology was important, in that species with shorter fruiting seasons have declined markedly in abundance in oceanic regions, whereas species with longer seasons have become relatively more common overall. These changes in fruit body distributions are correlated with temperature and rainfall, which act directly on both saprotrophic and ECM fungi, and also indirectly on ECM fungi, through altered photosynthate allocation from their hosts. If these distributional changes reflect fungal activity, there will be important consequences for the responses of forest ecosystems to changing climate, through effects on primary production and nutrient cycling.     

Cattle change plant reproductive phenology,promoting community changes in a post-fire Nothofagus forest in northern Patagonia,Argentina

Aims Variations in rates and length of flowering and fruiting not only affect the reproduction of a given plant species but also the behavior and reproduction of associated taxa. Flowering and fruiting variations may be influenced by herbivory, especially by large mammals. The aim of this study was to determine the effects of cattle browsing on the reproductive phenology of understory species in a subalpine post-fire Nothofagus forest in Patagonia.Methods The effects of herbivory on plant reproductive phenology were studied in a set of experimental exclosures (fenced plots) installed since 2001 in a post-fire N. pumilio forest, located in Nahuel Huapi National Park (NHNP), Argentina. We monitored the beginning and duration of each reproductive phenological stage: floral bud, open flower, immature fruit and mature fruit. We also counted the number of flowers, fruits, seeds and viable seeds of the dominant plants to assess whether browsing modifies temporal patterns of the flowering and fruiting periods.Important findings Cattle reduced the total number of species flowering and fruiting and changed the reproductive phenology of some species. We found that palatable species seem to be negatively affected by browsing in terms of reduced fitness due to changes in flowering and fruiting periods. In contrast, cattle benefitted the reproduction of non-palatable species and could promote the invasion of shade-intolerant exotic forbs such as Cirsium vulgare. The effects of livestock reported in this study are important to understanding how browsing could alter native species establishment and possibly alter successional trajectories during recolonization after fire.     

Are mismatches the norm? Timing of flowering,fruiting, dispersal and germination and their fitness effects in Frangula alnus (Rhamnaceae)

The close morphological and temporal links between phases of plant growth and reproduction call for integrated studies incorporating several reproductive phases from flowering to recruitment, and associated plant‐animal interactions. Phenological strategies, as well as plastic phenological response to climate change, incorporate complex interactions between developmental constraints, pollination and seed dispersal. Relationships between reproductive phenology and components of fitness were studied for two years in the north‐temperate, self‐incompatible, insect‐pollinated, and bird‐dispersed shrub Frangula alnus (Rhamnaceae). Fruit set, dispersal, germination and juvenile survival, as well as seed mass and juvenile size were measured in relation to flowering, fruiting and germination time. The results suggest that effects of flowering and fruiting time prevailed in subsequent phases, to some extent as far as to the juvenile phase, but effects of timing were complex and had partly opposing effects on different fitness components. Early flowers had higher fruit‐set and experiments indicated that synchronous peak flowering increased fruit‐set, but later flowers had higher seed mass. Peak fruiting was not associated with peak dispersal. Late fruits derived from late flowers promoted dispersal. Juvenile recruitment was enhanced by increasing seed size. We conclude that the phenology of flowering and fruiting in F. alnus comprises several features, each with different and sometimes counteracting effects on fitness components. From a general perspective, this result implies that we should not expect to find finely tuned matches in timing specifically between flowering and pollinators, and fruiting and seed dispersing birds.     

The timing of frugivore‐mediated seed dispersal effectiveness

The seed dispersal effectiveness framework allows assessing mutualistic services from frugivorous animals in terms of quantity and quality. Quantity accounts for the number of seeds dispersed and quality for the probability of recruitment of dispersed seeds. Research on this topic has largely focused on the spatial patterns of seed deposition because seed fates often vary between microhabitats due to differences in biotic and abiotic factors. However, the temporal dimension has remained completely overlooked despite these factors—and even local disperser assemblages—can change dramatically during long fruiting periods. Here, we test timing effects on seed dispersal effectiveness, using as study case a keystone shrub species dispersed by frugivorous birds and with a fruiting period of 9 months. We evaluated quantity and quality in different microhabitats of a Mediterranean forest and different periods of the fruiting phenophase. We identified the bird species responsible for seed deposition through DNA barcoding and evaluated the probability of seedling recruitment through a series of field experiments on sequential demographic processes. We found that timing matters: The disperser assemblage was temporally structured, seed viability decreased markedly during the plant's fruiting phenophase, and germination was lower for viable seeds dispersed in the fruiting peak. We show how small contributions to seed deposition by transient migratory species can result in a relevant effectiveness if they disperse seeds in a high‐quality period for seedling recruitment. This study expands our understanding of seed dispersal effectiveness, highlighting the importance of timing and infrequent interactions for population and community dynamics.     

Climate warming: a loss of variation in populations can accompany reproductive shifts

The most documented response of organisms to climate warming is a change in the average timing of seasonal activities (phenology). Although we know that these average changes can differ among species and populations, we do not know whether climate warming impacts within‐population variation in phenology. Using data from five study sites collected during a 13‐year survey, we found that the increase in spring temperatures is associated with a reproductive advance of 10 days in natural populations of common lizards (Zootoca vivipara). Interestingly, we show a correlated loss of variation in reproductive dates within populations. As illustrated by a model, this shortening of the reproductive period can have significant negative effects on population dynamics. Consequently, we encourage tests in other species to assess the generality of decreased variation in phenological responses to climate change.     

Predicting species‐specific responses of fungi to climatic variation using historical records

Although striking changes have been documented in plant and animal phenology over the past century, less is known about how the fungal kingdom's phenology has been changing. A few recent studies have documented changes in fungal fruiting in Europe in the last few decades, but the geographic and taxonomic extent of these changes, the mechanisms behind these changes, and their relationships to climate are not well understood. Here, we analyzed herbarium data of 274 species of fungi from Michigan to test the hypotheses that fruiting times of fungi depend on annual climate and that responses depend on taxonomic and functional groups. We show that the fungal community overall fruits later in warmer and drier years, which has led to a shift toward later fruiting dates for autumn‐fruiting species, consistent with existing evidence. However, we also show that these effects are highly variable among species and are partly explained by basic life‐history characteristics. Resulting differences in climate sensitivities are expected to affect community structure as climate changes. This study provides a unique picture of the climate dependence of fungal phenology in North America and an approach for quantifying how individual species and broader fungal communities will respond to ongoing climate change.     

Fruiting phenology is linked to rainfall variability in a tropical rain forest

As the influence of climate change on tropical forests becomes apparent, more studies are needed to understand how changes in climatic variables such as rainfall are likely to affect tree phenology. Using a twelve‐year dataset (2005–2016), we studied the impact of seasonal rainfall patterns on the fruiting phenology of 69 tree species in the rain forest of southeastern Madagascar. We found that average annual rainfall in this region has increased by >800 mm (23%) during this period relative to that recorded for the previous 40 years and was highly variable both within and between years. Higher monthly measures of fruiting richness and the intensity of fruiting in our sample community were associated with significantly higher levels of rainfall. We also found that less rainfall during the dry season, but not the wet season, was associated with a significant shift toward later timing of peak richness and peak intensity of fruiting in the subsequent 12 months; however, this pattern was driven primarily by an extreme drought event that occurred during the study period. Longer time scales of phenology data are needed to see whether this pattern is consistent. Madagascar is expected to experience more extremes in rainfall and drought with increasing climate change. Thus, the linkages between variable precipitation and the fruiting phenology of forest trees will have important consequences for understanding plant reproduction and the ability of Madagascar's wildlife to cope with a changing climate.     

Independent effects of warming and nitrogen addition on plant phenology in the Inner Mongolian steppe

Background and Aims

Phenology is one of most sensitive traits of plants in response to regional climate warming. Better understanding of the interactive effects between warming and other environmental change factors, such as increasing atmosphere nitrogen (N) deposition, is critical for projection of future plant phenology.

Methods

A 4-year field experiment manipulating temperature and N has been conducted in a temperate steppe in northern China. Phenology, including flowering and fruiting date as well as reproductive duration, of eight plant species was monitored and calculated from 2006 to 2009.

Key Results

Across all the species and years, warming significantly advanced flowering and fruiting time by 0·64 and 0·72 d per season, respectively, which were mainly driven by the earliest species (Potentilla acaulis). Although N addition showed no impact on phenological times across the eight species, it significantly delayed flowering time of Heteropappus altaicus and fruiting time of Agropyron cristatum. The responses of flowering and fruiting times to warming or N addition are coupled, leading to no response of reproductive duration to warming or N addition for most species. Warming shortened reproductive duration of Potentilla bifurca but extended that of Allium bidentatum, whereas N addition shortened that of A. bidentatum. No interactive effect between warming and N addition was found on any phenological event. Such additive effects could be ascribed to the species-specific responses of plant phenology to warming and N addition.

Conclusions

The results suggest that the warming response of plant phenology is larger in earlier than later flowering species in temperate grassland systems. The effects of warming and N addition on plant phenology are independent of each other. These findings can help to better understand and predict the response of plant phenology to climate warming concurrent with other global change driving factors.     

Impact of nutrient enrichment and alkalinization on periphyton communities in the New Jersey Pine Barrens

The impact of residential and agricultural development on stream periphyton communities in the New Jersey Pine Barrens was examined by comparison with communities in undeveloped areas. Watershed disturbance resulted in stream water primarily characterized by greatly elevated pH levels and nitrate concentrations. A total of 53 periphyton species were encountered in bimonthly samples over a one year period in the three disturbed and three undisturbed study streams. Species richness was significantly greater in the disturbed streams based on three criteria: the average number of species per stream on each sampling occasion (disturbed = 6.3; undisturbed = 4.9), the average number of species per stream for the entire year (disturbed = 19.3; undisturbed = 16.0), and the total number of species found in streams within a type (disturbed = 40; undisturbed = 31). Species composition also changed significantly as the result of disturbance. There appeared to be replacement of species characteristic of undisturbed Pine Barrens streams with species peripheral to the region. The expected effects of both elevated pH and nitrate were consistent with these results.     

Wood-inhabiting fungi on fallen logs of Norway spruce: relations to forest management and substrate quality

A survey of the patterns of wood decaying fungi as to occurrence of sporocarps on naturally fallen logs of Norway spruce ( Picea abies ) was undertaken in two nearby forest stands with different histories of management. One stand was an old-growth forest with few signs of logging, and the other stand was selectively logged 60–80 years ago. Altogether 118 species were found. Forest management had a negative impact on the species diversity. Newly fallen and weakly decayed logs in a natural forest had a higher species richness, more red-listed species, as well as more indicator species compared to similar logs in a managed forest. The importance of dead wood for species diversity of wood inhabiting fungi was clearly demonstrated. Presence of logs in later stages of decomposition increased the total species number in a natural forest stand with 42 (63 %), compared to a survey of only newly fallen and weakly decayed logs. Presence of logs in later stages of decomposition also increased the diversity of the species pool colonising newly fallen and weakly decayed logs. The highest number of fruiting species was found on intermediately decayed logs and on logs lying in contact with the ground. The fungal gradient as revealed in a DCA ordination was primarily related to decay. A successional pathway based on the primary decayer Fomitopsis pinicola was not detected.     

Diet and Dietary Preferences of the Southern Cassowary (Casuarius casuarius) in North Queensland, Australia

We investigated the diet of the southern cassowary (Casuarius casuarius) by identifying the seeds and fruits in fecal droppings encountered on a set of transects over 2 yr in upland rain forest in the wet tropics of North Queensland. A total of 198 droppings containing 56 plant species were found. We surveyed fleshy fruit availability over the subsequent 68 mo on transects in the same area to ascertain fruiting patterns in the study area. The number of droppings found each month did not correspond to the pattern of available fruit biomass. There was no relationship between the fruit traits of moisture content, flesh to seed mass ratio, color, or crop size to contribution of a species to the diet. During the lean fruiting season (May–July) cassowaries relied more on species that fruited continuously throughout the year as they were significantly over‐represented in droppings, while annual fruiting species were under‐represented. During months of high fruit availability (October–December), continuously fruiting species were still over‐represented in the diet but became less important while annual and biennial species became more important. Significantly more species with large fruit and large seeds appeared in the diet than expected and we confirm that the cassowary contributes to the continued dispersal of these species over long distances and in large quantities.     

Utilization of fruiting trees by monkeys as analyzed from feeding traces under fruiting trees in the tropical rain forest of Cameroon

Individual trees of the food species of monkeys were identified by placing plastic tapes with an identification number on them in the tropical rain forest of Cameroon, West Africa. In order to determine the use of the feeding trees by monkeys, the ground under each of the trees was checked at least once a week to see if there were any fallen fruits or traces of feeding on fruits. Some fruit species were not fed on by either monkeys or large arboreal squirrels. Among the food species common to both the monkeys and large squirrels, a larger proportion in terms of quantity in each species was mainly eaten by the monkeys except in the case of super-abundantly fruit producing species. The monkeys and large arboreal squirrels were well segregated in their diets. Larger proportions (more than 85% for most of the monkeys' major foods) of fruits of larger sizes were made to fall on the ground by the monkeys and squirrels. The monkeys displayed a tendency to visit fruiting trees rather evenly (even rate of visit = even frequency of visit/duration of fruiting) not ignoring any area of the home range, although a small difference in this tendency was observed between the two study periods, one an abundant season and the other a poor fruiting season. On average, one associated polyspecific group of monkeys encountered only 14 fruiting trees per day. On the other hand, fruits were available all around the year, as the fruiting periods of different tree species were widely distributed around the year, or the fruiting periods of some species were very long. Although the monkeys are able to depend heavily on fruits, the quantity of fruits is not so great. The population size of monkeys is well balanced with the available food supply in the tropical rain forest of West Africa.     

Mushroom's spore size and time of fruiting are strongly related: is moisture important?

Most basidiomycete fungi produce annual short-lived sexual fruit bodies from which billions of microscopic spores are spread into the air during a short time period. However, little is known about the selective forces that have resulted in some species fruiting early and others later in the fruiting season. This study of relationships between morphological and ecological characteristics, climate factors and time of fruiting are based upon thorough statistical analyses of 66 520 mapped records from Norway, representing 271 species of autumnal fruiting mushroom species. We found a strong relationship between spore size and time of fruiting; on average, a doubling of spore size (volume) corresponded to 3 days earlier fruiting. Small-spored species dominate in the oceanic parts of Norway, whereas large-spored species are typical of more continental parts. In separate analyses, significant relationships were observed between spore size and climate factors. We hypothesize that these relationships are owing to water balance optimization, driven by water storage in spores as a critical factor for successful germination of primary mycelia in the drier micro-environments found earlier in the fruiting season and/or in continental climates.     

Patterns of phenological changes in migratory birds

The phenology of avian migration appears to be changing in response to climate change. Seemingly contradictory differences in the timing of these annual cycles have been reported in published studies. We show that differences between studies in the choice of songbird species, as well as in the measurements of migration phenology, can explain most of the reported differences. Furthermore, while earlier spring arrival is evident across these studies, trends in timing of departure show large variation between species and according to individual timing of migration (early-arriving vs. late-departing individuals). Much of the variation in departure between species could be explained by each species’ migratory status. We present a detailed analysis of migrants recorded at a Danish migration site, and reveal that although shifts in migration timing can be demonstrated for almost all species, these shifts are either most pronounced in the early arriving/late departing individuals or the changes are similar. Thus most individuals do not seem to change their breeding-area residence time (BART). As BART is likely to reflect ecologically important factors, e.g. number of clutches, we expect that only small effects have been exerted on the breeding ecology of the studied species in the time period investigated. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Avian species richness and abundance at Lake Constance: diverging long-term trends in Passerines and Nonpasserines

In Central Europe, massive losses in species richness of breeding birds have been documented in the last decades, but the question arises whether species richness is currently still decreasing or again increasing due to conservation efforts. In this study, we investigated regional and local species richness as well as mean number of breeding pairs and mean biomass per tetrad over a period of some 20 years at Lake Constance. Three quantitative censuses of 303 tetrads (2 × 2 km²) repeated at 10-year intervals (1980–1981, 1990–1992, 2000–2002) revealed an increase in regional species richness (total number of breeding species). At the same time, however, a strong decline in local species richness (number of breeding species per tetrad), number of breeding pairs, and estimated biomass were observed. Changes of species richness differed markedly between Nonpasserine and Passerine birds. Whereas species richness of Nonpasserines remained constant from 1980 to 1990, and even increased between 1990 and 2000, that of Passerines decreased in both periods. This indicates that effects of conservation efforts apparently eclipse more general effects of climate and habitat change in Nonpasserines. The massive abundance and biomass losses observed in formerly common Passerine species are not compensated by gains in populations of Nonpasserine species. The results of the three bird censuses at Lake Constance imply that ongoing habitat degradation and human impacts as well as increasing effects of climate change are the main drivers of the observed population changes.     

Fine‐scale spatiotemporal dynamics of fungal fruiting: prevalence,amplitude, range and continuity

Despite the critical importance of fungi as symbionts with plants, resources for animals, and drivers of ecosystem function, the spatiotemporal distributions of fungi remain poorly understood. The belowground life cycle of fungi makes it difficult to assess spatial patterns and dynamic processes even with recent molecular techniques. Here we offer an explicit spatiotemporal Bayesian inference of the drivers behind spatial distributions from investigation of a Swiss inventory of fungal fruit bodies. The unique inventory includes three temperate forest sites in which a total of 73 952 fungal fruit bodies were recorded systematically in a spatially explicit design between 1992 and 2006. Our motivation is to understand how broad‐scale climate factors may influence spatiotemporal dynamics of fungal fruiting within forests, and if any such effects vary between two functional groups, ectomycorrhizal (ECM) and saprotrophic fungi. For both groups we asked: 1) how consistent are the locations of fruiting patches, the sizes of patches, the quantities of fruit bodies, and of prevalence (occupancy)? 2) Do the annual spatial characteristics of fungal fruiting change systematically over time? 3) Are spatial characteristics of fungal fruiting driven by climatic variation? We found high inter‐annual continuity in fruiting for both functional groups. The saprotrophic species were characterised by small patches with variable fruit body counts. In contrast, ECM species were present in larger, but more distinctly delimited patches. The spatial characteristics of the fungal community were only indirectly influenced by climate. However, climate variability influenced overall yields and prevalence, which again links to spatial structure of fruit bodies. Both yield and prevalence were correlated with the amplitudes of occurrence and of fruit body counts, but only prevalence influenced the spatial range. Summarizing, climatic variability affects forest‐stand fungal distributions via its influence on yield (amount) and prevalence (occupancy), whereas fungal life‐history strategies dictate fine‐scale spatial characteristics.     

Exploring the compass of potential changes induced by climate warming in plant communities

New models are required to predict the impacts of future climate change on biodiversity. A move must be made away from individual models of single species toward approaches with synergistically interacting species. The focus should be on indirect effects due to biotic interactions. Here we propose a new parsimonious approach to simulate direct and indirect effects of global warming on plant communities. The methodology consists of five steps: a) field survey of species abundances, b) quantitative assessment of species co-occurrences, c) assignment of a theorised effect of increased temperature on each species, d) creation of a community model to project community dynamics, and e) exploration of the potential range of temperature change effects on plant communities.We explored the possible climate-driven dynamics in an alpine vegetation community and gained insights into the role of biotic interactions as determinants of plant species response to climate change at local scale. The study area was the uppermost portion of Alpe delle Tre Potenze (Northern Apennines, Italy) from 1500 m up to the summit at 1940 m.Our work shows that: 1) unexpected climate-driven dynamics can emerge, 2) interactive communities with indirect effects among species can overcome direct effects induced by global warming; 3) if just one or few species react to global warming the new community configuration could be unexpected and counter-intuitive; 4) timing of species reactions to global warming is an important driver of community dynamics; 5) using simulation models with a limited amount of data in input, it is possible to explore the full range of potential changes in plant communities induced by climate warming.     

Global patterns in fruiting seasons

Aim   To identify geographical and climatic correlates of the timing of fruit production in fleshy fruited plant communities.
Location   Global.
Methods   We searched the literature for studies documenting monthly variation in the number of fleshy fruited species bearing ripe fruits in plant communities (i.e. fruit phenologies). From these data, we used circular vector algebra to characterize seasonal peaks in fruit production (mean date, as an angle) and the length of fruiting seasons (as a circular standard deviation). Generalized linear models and circular correlations were used to assess whether latitudinal patterns in fruit phenologies could be explained by variation in temperature, precipitation and actual evapotranspiration (AET).
Results   Dates of peak fruit production and the length of fruiting seasons showed consistent differences with latitude. Annual peaks in fruit production occurred 1 to 3 months after the summer solstice at high-latitude sites in both hemispheres. Fruiting seasonality increased with latitude, indicating that fruiting seasons were longer in the tropics and shorter toward the poles. AET was the best climatic predictor of fruit phenologies. Annual peaks in fruit production were positively associated with annual peaks in AET and temperature, while fruiting seasons were shorter in areas with pronounced annual variation in AET.
Main conclusions   Global patterns in fruiting seasons are associated with global variation in climate. Across the globe, fleshy fruits are produced after annual periods of elevated water–energy availability. Fruiting seasonality is also more pronounced in areas with strongly seasonal water–energy inputs. Therefore, the timing of reproduction in fleshy fruited plant communities appears to be determined, at least in part, by spatial and temporal variation in energy supplies needed to subsidise plant reproduction.     

Microvariation in the fauna of a sublittoral sand bank,Moreton Bay,Queensland

Benthic macrofauna was sampled from Naval Reserve Bank, Moreton Bay, Queensland, using an 0.05 m² van Veen grab, and data were used to investigate spatial micropatterning. Samples were taken within a 5 × 6 square sampling grid (total area 730 in2) with an intersampling distance of 6 m. Samples were taken in duplicate on each occasion at 2-inonth intervals. Particle size of sediments and depth data were also collected. Data on other abiotic factors such as temperature, salinity and currents were inferred. Sampling gave data for 131 species. Hierarchical numerical classificatory techniques were employed to obtain (a) spatial patterns (site-groups) based on biotic data, and (b) species-groups which characterize the above site groups. Species diversity (SH) species richness (S), species evenness (J’) and individual species diversity (I) indices were computed for the total data set for each site with the species summated over the sampling times. A selected number of species were used in the classification. The spatial patterns were well defined but the species-groups categorizing them were somewhat confused. Species diversity indices were found to be of little aid in the interpretation of the site patterns or in generating alternative hypotheses.