Ant diversity in an Amazonian savanna: Relationship with vegetation structure,disturbance by fire,and dominant ants

Abstract The savannas of South America support a relatively diverse ant fauna, but little is known about the factors that influence the structure and dynamics of these assemblages. In 1998 and 2002, we surveyed the ground‐dwelling ant fauna and the fauna associated with the woody vegetation (using baits and direct sampling) from an Amazonian savanna. The aim was to evaluate the influence of vegetation structure, disturbance by fire and dominant ants on patterns of ant species richness and composition. Variations in the incidence of fires among our 39 survey plots had no or only limited influence on these patterns. In contrast, spatial variations in tree cover and cover by tall grasses (mostly Trachypogon plumosus), significantly affected ant species composition. Part of the variation in species richness among the study plots correlated with variations in the incidence of a dominant species (Solenopsis substituta) at baits. Ant species richness and composition also varied through time, possibly as an indirect effect of changes in vegetation cover. In many plots, and independently of disturbance by fire, there was a major increase in cover by tall grasses, which occupied areas formerly devoid of vegetation. Temporal changes in vegetation did not directly explain the observed increase in the number of ant species per plot. However, the incidence of S. substituta at baits declined sharply in 2002, especially in plots where changes in vegetation cover were more dramatic, and that decline was correlated with an increase in the number of ground‐dwelling species, a greater turnover of bait‐recruiting species and the appearance of the little fire ant Wasmannia auropunctata. The extent to which these changes in fact resulted from the relaxation of dominance by S. substituta is not clear. However, our results strongly suggest that the ant fauna of Amazonian savannas is affected directly and indirectly by the structure of the vegetation.     

The Role of Panicum virgatum (Switch Grass) in the Revegetation of Iron-Mine Tailings in Northern New York

We investigated the role of artificially introduced Panicum virgatum (switch grass) on the sequential natural revegetation of 15- and 35-year-old tailings in the Adirondack region of northern New York. Switch grass covered approximately 48% of the 15-year-old Chaumont Tailings. Establishment of switch-grass stands improved the fertility of the site by adding organic matter, raising pH, and elevating cation exchange capacity and concentrations of major nutrients (N, P, and K). Switch-grass stands also aided the initial recruitment of such pioneer species as Populus spp. (aspens), Salix spp. (willows), and Betula spp. (birches). This facilitation of recruitment of woody species is explained as follows: (1) robust switch-grass stands physically captured the wind-disseminated seeds of these species; (2) switch grass acted as a “nurse crop” for these species, thus these species were able to increase their density vigorously through root or stem sprouting; (3) a combination of both. Switch grass decreased its cover (14%), however, as observed in the 35-year-old South Tailings. As switch grass declined, such vigorous “root-suckering” species as aspens increased in dominance, followed by the invasion of Prunus pensylvanica (pin cherry).     

Effects of jaw adductor hypertrophy on buccal expansions during feeding of air breathing catfishes (Teleostei,Clariidae)

Some species of Clariidae (air breathing catfishes) have extremely well developed (hypertrophied) jaw closing muscles that increase the maximal biting force of these species. As these enlarged jaw muscles tightly cover the suspensoria, which are firmly connected to the neurocranium, we expect diminished lateral expansions during suction for species with hypertrophied jaw muscles. In turn, this could imply a reduced suction performance for these species. Compared to Clarias gariepinus, which has relatively small jaw closers, Clariallabes longicauda shows a clear hypertrophy of the jaw adductors. A kinematic analysis of prey capture in these two species is presented here. As predicted, Clariallabes longicauda shows less lateral expansion (average abduction of the hyoids of 19.0°) than Clarias gariepinus (abduction of 31.1°). However, our data indicate that the decrease in lateral expansion capacity in the species with excessive adductor development is compensated for by a larger and faster ventral expansion of the buccal cavity by depression of the hyoid.     

Growth characteristics in co‐occurring Upper Ordovician species of the tabulate Catenipora from southern Manitoba,Canada

Four species of the tabulate coral Catenipora are present in the Selkirk Member of the Red River Formation at Garson, Manitoba. They provide an opportunity to compare the growth characteristics of multiple, co‐occurring species that produced cateniform coralla. Corallite increase, cyclomorphism and other growth features show high variability within and/or among the species. A total of five types of lateral increase and two types of axial increase are recognized. Lateral increase accounts for over 80% of all occurrences of corallite increase in each species, with the four species differing significantly in the relative frequency of the various types of lateral increase. The type of axial increase, megacorallites and agglutinated patches of corallites that developed from normal, undamaged corallites in C. foerstei are species specific. In all species, cyclic fluctuations in the tabularial area of corallites are considered to be annual, and the variable growth rates within colonies and species are attributed to differences in astogenetic stages or environmental conditions. Average annual vertical growth was positively correlated with average tabularial area in C. foerstei, C. cf. robusta and C. rubra. Catenipora cf. agglomeratiformis, however, which had the lowest average tabularial area and greatest sensitivity to sediment influx, had a high average growth rate comparable to that of C. rubra, which had the largest average tabularial area. The formation of ranks or lacunae by certain types of lateral increase seems to have been the most effective strategy for maintaining and/or expanding the colony growth surface in all four species, and was most common in C. cf. agglomeratiformis. A reptant growth pattern, characterized by creeping ranks, permitted effective recovery of damaged parts as well as quick formation of new ranks or lacunae. The growth surface of these species was situated near the sediment–water interface. □Growth characteristics, intraspecific variation, interspecific variation, palaeobiology, tabulate corals.     

Effects of urban habitat fragmentation on common small mammals: species versus communities

There is an increasing interest in understanding how species respond to the modifications of habitat attributes in urban areas. Patterns in the occurrence and abundance of small mammal communities in 15 isolated patches of remnant natural and semi-natural habitat of Porto Metropolitan Area (Portugal) were assessed against environmental characteristics (from both the patch and the surrounding matrix) of each patch using multiple regressions and canonical correspondence analysis. Four species of common small mammals were found: wood mouse (Apodemus sylvaticus), greater white-toothed shrew (Crocidura russula), Algerian mouse (Mus spretus) and house mouse (Mus musculus). Our results showed that both relative abundance and species richness were negatively affected by urbanization. The species richness also displayed a negative association with the increase of forest around the patch but relative abundance showed the opposite trend. At the species level, the relative abundance of A. sylvaticus and C. russula showed a negative association with urbanization. Our results reveal that these two species also benefit from a mosaic of habitats and pervious areas in the surrounding matrix. The relative abundance of M. spretus and M. musculus showed a negative effect of forest area around the patch. Understanding how the increase of urbanization affects small mammals will be particularly useful for the managers of urban landscapes, as these animals occupy a pivotal position in the ecosystems.     

Effects of cutting and nitrogen deposition on biodiversity in Cantabrian heathlands

Questions: Does the diversity of heathland vegetation change when subjected to experimental disturbances such as cutting and nitrogen fertilization? Do changes in the vegetation structure negatively affect the regeneration of the dominant species Calluna vulgaris? Is cutting an alternative method of conserving the diversity and maintaining the structure of heathlands in the Cantabrian Mountains? Location: Calluna vulgaris heathlands on the southern slopes of the Cantabrian Mountain range, NW Spain. Methods: A total of 60 plots were treated with different combinations of cutting and twice the estimated atmospheric deposition of nitrogen (56 kg‐N.ha^?1.yr^?1). The changes in the cover values of the species present were monitored over a five year study period. The cover values were used to calculate abundance and species richness. Results: Fertilizing with nitrogen allows biodiversity to increase over time. However, the greatest biodiversity is associated with the cutting plus fertilization treatment, since cutting allows gaps to be opened that are easily colonized by pioneer annual species, while fertilization mainly favours an increase in the mean number of perennial herbs (graminoids and forbs). Increased perennial herb richness also corresponds to a rise in their cover values. The recovery of the dominant woody species in these communities, Calluna vulgaris, is not impeded by the increase in perennial herbs species' cover values. Conclusions: In the Calluna vulgaris heathlands studied, cutting plus fertilization allowed an increase in biodiversity over time. No displacement of the dominant woody species, Calluna vulgaris, is observed due to the presence of the perennial herbs. Cutting patches of heathland is recommended as a mechanism for maintaining high vegetation diversity, when grazing is not possible.     

Differential strategies of two species of arbuscular mycorrhizal fungi in the protection of maize plants grown in chromium-contaminated soils

Chromium (Cr) is a nonessential element for plants that is extremely toxic at high concentrations. Zea mays L. is a species of plant that has developed adaptive mechanisms to increase its tolerance and absorption capacity for this metal. One effective mechanism is to form associations with arbuscular mycorrhizal fungi (AMF), which help the plant cope with stress from heavy metals such as Cr. However, it is still not clear which species of AMF are more efficient as bioremediating agents for plants of agricultural importance. Here, we evaluated the effect of Gigaspora gigantea and Rhizophagus irregularis as protective agents of maize plants in soils contaminated with Cr at concentrations of 0, 48.5, 97 and 194 mg kg^?1 under greenhouse conditions. Maize tolerance to Cr was corroborated, as well as increased absorption of this element by plants associated with both species of AMF. G. gigantea caused an increase in nitrogen content and greater translocation of Cr to the aerial part of the plant; R. irregularis registered an increase in the formation of arbuscules and vesicles with increasing metal concentration and greater retention of Cr in the roots of the plants. Based on these results, we can conclude that the analyzed species of fungi use different strategies, with similar effectiveness, to enhance the absorption capacity of Cr by the plant and influence the differential deposition of this metal in various parts of the plant.

     

Testing genome skimming for species discrimination in the large and taxonomically difficult genus Rhododendron

Standard plant DNA barcodes based on 2–3 plastid regions, and nrDNA ITS show variable levels of resolution, and fail to discriminate among species in many plant groups. Genome skimming to recover complete plastid genome sequences and nrDNA arrays has been proposed as a solution to address these resolution limitations. However, few studies have empirically tested what gains are achieved in practice. Of particular interest is whether adding substantially more plastid and nrDNA characters will lead to an increase in discriminatory power, or whether the resolution limitations of standard plant barcodes are fundamentally due to plastid genomes and nrDNA not tracking species boundaries. To address this, we used genome skimming to recover near-complete plastid genomes and nuclear ribosomal DNA from Rhododendron species and compared discrimination success with standard plant barcodes. We sampled 218 individuals representing 145 species of this species-rich and taxonomically difficult genus, focusing on the global biodiversity hotspots of the Himalaya-Hengduan Mountains. Only 33% of species were distinguished using ITS+matK+rbcL+trnH-psbA. In contrast, 55% of species were distinguished using plastid genome and nrDNA sequences. The vast majority of this increase is due to the additional plastid characters. Thus, despite previous studies showing an asymptote in discrimination success beyond 3–4 plastid regions, these results show that a demonstrable increase in discriminatory power is possible with extensive plastid genome data. However, despite these gains, many species remain unresolved, and these results also reinforce the need to access multiple unlinked nuclear loci to obtain transformative gains in species discrimination in plants.     

The effect of increased nutrient availability on leaf turnover and aboveground productivity of two evergreen ericaceous shrubs

Summary Leaf turnover and aboveground productivity in relation to nutrient availability were studied in the evergreen shrubs Erica tetralix and Calluna vulgaris. In monospecific stands of these species four levels of nutrient (NPK) availability were created during three growing seasons. Percentage survival and life expectancy of Erica leaves decreased with increasing nutrient availability. For Calluna there was no effect. Winter mortality of Erica leaves was smaller than growing season mortality. These was no difference for Calluna. The timing of leaf mortality of both species was not affected by nutrient treatment. At the end of the experimental period current year leaf biomass, total biomass and current year second year and third year biomass of both species showed a significant increase with increasing nutrient availability. The relative increase was greater for Calluna, except for second and third year biomass. Stem production and stem mortality of both species increased with increasing nutrient availability. The increased stem mortality resulted also for Calluna in an increased leaf turnover (per unit ground area) with increasing nutrient availability. Nutrient cycling in ecosystems dominated by these species will increase with increasing nutrient availability, because of increased leaf and stem turnover and productivity. This phenotypic effect is similar to the effect of the shift in dominance between different species which occurs along natural gradients of nutrient availability.     

Spatial organization of a desert rodent community: food addition and species removal

Summary From 1977 through 1983 we conducted experiments on a desert rodent community where supplemental seeds were added or certain rodent species and ants were removed from 0.25-ha fenced plots in a Chihuahuan Desert site in southeastern Arizona, USA. In this paper we examine the patterns of microhabitat use relative to vegetative cover by 11 rodent species. The results show that: i) removal of the largest seed-eating species, Dipodomys spectabilis, produced the most pervasive and dramatic shifts in microhabitat use by the remaining rodent species; ii) adding seeds or removing ants had little effect on the spatial use of microhabitats by rodents in this community; and iii) non-granivores were just as likely as granivores to shift microhabitat use when other granivores were removed. We believe these results indicate that both food and foraging microsites are limited but the relegation of subdominant species to lesspreferred microhabitats by the large Dipodomys spectabilis is the major factor underlying the spatial organization of this community. Results also demonstrate that strong interactions among species increase the probability that pathways of indirect interactions through intermediary species are important; these complex linkages may include species that overlap little in food preferences.     

Diversified chromosomal characteristics in Centropyge fishes (Pomacanthidae, Perciformes)

We studied karyotypes and other chromosomal markers such as C-banded heterochromatin and Ag-stained nucleolus organizer regions (Ag-NORs), in seven Centropyge fishes (Pomacanthidae, Perciformes). These results revealed diversified chromosomal characteristics in Centropyge species. Three species had 2n = 48 chromosomes, whereas four species had 2n = 52 chromosomes. Fundamental numbers showed a large variation from 48 to 82, particularly in the species with 2n = 52 chromosomes. In all the species, Ag-NORs were located in a single chromosome pair and C-bands were mainly distributed in the centromeric regions of most chromosomes, as commonly seen in teleostean fishes. However, these chromosomal markers showed species-specific variations and provided us with useful information that could help us in understanding chromosomal evolution. On the basis of these chromosomal characteristics, we infer the process of chromosomal evolution, which according to us involves an increase in chromosome number from 2n = 48 to 2n = 52 through centric fission or other mechanisms, and in fundamental number through pericentric inversion. In particular, karyotypic evolution involving the increase in chromosome number is an unusual event in the evolution of higher teleostean groups. Handling editor: K. Martens     

Phylogenetic analysis of eleven species of Biomphalaria Preston, 1910 (Gastropoda: Planorbidae) based on comparisons of allozymes

Freshwater snails in the genus Biomphalaria transmit Schistosoma mansoni in Africa, South America and the Caribbean region. Although considerable attention has been given to the identification of species, little is known of evolutionary relationships among the species. A phylogenetic analysis of 25 populations representing 11 species was performed on 25 enzyme loci examined using starch gel electrophoresis. A phylogenetic analysis of the individual populations produced 60 trees of equal length. The 60 trees have a consistency index value of 75.9% and a retention index value of 76.5%. The phylogenetic analysis provided strong support for the monophyly of Biomphalaria with either 14 or 15 synapomorphies uniting all of the species included and separating them from the outgroup, two species of Helisoma. Four nominal species represented by multiple populations formed monophyletic groups. Populations of B. sudanica, B. choanomphala, and B. alexandrina were interspersed. Ten arrangements were obtained for the populations of these three species. A variety of ingroup taxa were used to root the trees, and all provided support for the use of Helisoma species as an outgroup. In all of the trees obtained, the African species together formed a monophyletic group. In none of the trees obtained did the neotropical species form a monophyletic group. A constrained analysis requiring the monophyly of the neotropical species as well as the African species resulted in 90 trees just two steps longer than the shortest trees. Analysis of the species from either hemisphere alone resulted in decreased resolution, as measured by an increase in the number of trees obtained. This finding suggests that further comparisons of species from the two hemispheres will be of considerable value. Finally, two species which are resistant to infection with S. mansoni were included among the eleven studied. Neither of these species formed the sister group to all of the other species included, indicating that susceptibility is the plesiomorphic state, and that resistance is derived. Similarly, in none of the trees obtained did the two resistant species fall out as sister taxa, indicating that resistance arose independently twice.     

The effect of sub-lethal increases in temperature on the growth and population trajectories of three scleractinian corals on the southern Great Barrier Reef

To date, coral death has been the most conspicuous outcome of warming tropical seas, but as temperatures stabilize at higher values, the consequences for the corals remaining will be mediated by their demographic responses to the sub-lethal effects of temperature. To gain insight into the nature of these responses, here I develop a model to test the effect of increased temperature on populations of three pocilloporid corals at One Tree Island, near the southern extreme of the Great Barrier Reef (GBR). Using Seriatopora hystrix, S. caliendrum and Pocillopora damicornis as study species, the effects of temperature on growth were determined empirically, and the dynamics of their populations determined under natural temperatures over a 6-month period between 1999 and 2000 [defined as the study year (SY)]. The two data sets were combined in a demographic test of the possibility that the thermal regime projected for the southern GBR in the next 55–83 years—warmer by 3°C than the study year (the SY+3 regime), which is equivalent to 1.4°C warmer than the recent warm year of 1998—would alter coral population trajectories through the effects on coral growth alone; the analyses first were completed by species, then by family after pooling among species. Laboratory experiments showed that growth rates (i.e., calcification) varied significantly among species and temperatures, and displayed curvilinear thermal responses with growth maxima at ∼27.1°C. Based on these temperature-growth responses, the SY+3 regime is projected to: (1) increase annualized growth rates of all taxa by 24–39%, and defer the timing of peak growth from the summer to the autumn and spring, (2) alter the intrinsic rate of population growth (λ) for S. hystrix (λ decreases 26%) and S. caliendrum (λ increases 5%), but not for P. damicornis, and (3) have a minor effect on λ (a 0.3% increase) for the Pocilloporidae, largely because λ varies more among species than it does between temperatures. Ten-year population projections suggest that the effects of a sub-lethal increase in temperature (i.e., the SY+3 regime) are relatively small compared to the interspecific differences in population dynamics, but nevertheless will alter the population size and increase the relative abundance of large colonies at the expense of smaller colonies for all three species, as well as the Pocilloporidae. These effects may play an important role in determining the nuances of coral population structure as seawater warms, and their significance may intensity if the coral species pool is depleted of thermally sensitive species by bleaching.     

BREEDING SYSTEMS OF CENTRAL AMERICAN APHELANDRA (ACANTHACEAE)

Controlled pollinations, followed by germination studies of seeds resulting from self- and cross-pollination, were carried out using plants from 12 populations of nine species of Aphelandra (Acanthaceae), a genus of neotropical shrubs and herbs. These results are combined with data from field studies of flower morphology, phenology, and pollinator relationships to estimate breeding system of each species. All species have floral morphological traits that prevent autogamy. Plants from five populations of five species experience low levels of geitonogamous pollen transfer; they produce few flowers daily and are pollinated by traplining hummingbirds. Excepting A. storkii, these plants are fully self-compatible (SC), and seeds from selfing are as viable as crossed seeds. Aphelandra storkii is partially self-incompatible (SI) and produces seeds from selfing that tend to germinate less successfully than crossed seeds. Plants from the remaining populations are profusely flowering shrubs, and even those pollinated by traplining hummingbirds should experience higher levels of geitonogamy. Aphelandra deppeana is pollinated by territorial hummingbirds, which should further increase the incidence of geitonogamy. All shrub species are partially SI, and two species (A. leonardii and A. sinclairiana) show significant reduced germination of selfed vs. crossed seeds. The breeding system of these species is thus modified by postpollination factors that favor the formation and maturation of outcrossed seeds. It is suggested that Aphelandra species, like other herbs and shrubs of tropical forest understory, possess a combination of breeding system traits that promote outcrossing but do not exclude geitonogamy.     

Evaluating costs for the <Emphasis Type="Italic">in vitro</Emphasis> propagation and preservation of endangered plants

In vitro methods provide opportunities for propagating and preserving endangered plant species when seed-based methods are not adequate. Such species include those that produce few or no seeds, as well as species with recalcitrant seeds. Tissue culture propagation methods can be used to produce such plants for reintroduction, research, education, display, and commerce. They can also be the basis for tissue banking as a way to preserve genetic diversity when seeds cannot be banked. With some recalcitrant species, embryo banking, a method which also utilizes in vitro culture for recovery germination, is possible. The number of endangered species that will require in vitro methods is estimated to be at least 5,000 worldwide. Further information is needed to identify these species, and the ongoing collection of information into databases on endangered species and recalcitrant species will help provide this. The costs of these methods are higher than for traditional propagation and preservation, but they may be necessary for species under higher threat. The multiplication rate of a culture, as well as the rates of rooting and acclimatization, has a major effect on the number of transfers needed for producing plants or tissue for banking, and improvements that will increase the efficiency of these steps can help lower costs. Further research into factors affecting the growth of tissues in vitro, as well as coordination of efforts among institutions with infrastructure for in vitro work, should facilitate the application of in vitro methods to the endangered species that cannot be propagated or preserved using seeds.     

THE ROOT APEX OF LINUM

Several developmental stages of primary roots of 13 species of Linum were studied. The basic pattern of root apex organization in three species consisted of a single tier of cortical initials. Ten species had a basic pattern of two tiers of cortical initials. Variations, manifested by either an increase or a decrease in the tiers of cortical initials, were observed in some roots in those species that had a basic pattern of two tiers of cortical initials. Although these variations were interpreted as being ontogenetic, there was no total reorganization of the root apex. There was anatomical and cytological evidence that a quiescent center is established in Linum roots.     

Land use history, hurricane disturbance, and the fate of introduced species in a subtropical wet forest in Puerto Rico

Tropical forests are suffering from increasing intensities and frequency of disturbances. As a result, non-native species accidentally introduced or intentionally planted for farming, plantations, and ornamental purposes may spread and potentially invade undisturbed native forest. It is not known if these introduced species will become invasive, as a result of recurrent natural disturbances such as hurricanes. Using data from three censuses (spanning 15 years) of a 16-ha subtropical wet forest plot, we investigated the impact of two hurricanes on populations of plant species that were planted in farms and plantations that were then abandoned and from the natural spread of species introduced into Puerto Rico in the past. The populations of four species (Citrus paradis, Mangifera indica, Musa sp., and Simarouba glauca) changed little over time. Six species (Artocarpus altilis, Calophyllum calaba, Genipa americana, Hibiscus pernambucensis, Syzygium jambos, and Swietenia macrophylla) declined between the first two censuses after Hurricane Hugo, then increased again in Census 3 after Hurricane Georges. Spathodea campanulata gradually increased from census to census, while Coffea arabica declined. These introduced species represent only a small part of the forest basal area and few show signs of increasing over time. The number of stems per plant, new recruits, and the growth rates of these introduced species were within the ranges of those for native plant species. The mortality rates over both census intervals were significantly lower for introduced species (<5% year⁻¹) than for native ones (15% year⁻¹). Many new recruits established after Hurricane Hugo (prior to this study) had opened the forest canopy and these high mortality rates reflect their death as the canopy recovered. Only Swietenia macrophylla and Syzygium jambos showed an increase in stem numbers in the closed canopy area of forest that had suffered limited human disturbance in the past. A future increase in frequency of disturbance may enable greater stem numbers of introduced species to establish, while lower-mortality rates compared to native species, may allow them to persist during inter-hurricane intervals. An increase in the population of introduced species, especially for those that grow into large trees, may have an impact on this tropical forest in the future.     

Role of respiration and glutathione in cadmium-induced oxidative stress in Escherichia coli K-12

Cadmium is a widespread pollutant that has been associated with oxidative stress, but the mechanism behind this effect in prokaryotes is still unclear. In this work, we exposed two glutathione deficient mutants (ΔgshA and ΔgshB) and one respiration deficient mutant (ΔubiE) to a sublethal concentration of cadmium. The glutathione mutants show a similar increase in reactive oxygen species as the wild type. Experiments performed using the ΔubiE strain showed that this mutant is more resistant to cadmium ions and that Cd-induced reactive oxygen species levels were not altered. In the light of these facts, we conclude that the interference of cadmium with the respiratory chain is the cause of the oxidative stress induced by this metal and that, contrary to previously proposed models, the reactive oxygen species increase is not due to glutathione depletion, although this peptide is crucial for cadmium detoxification.     

Patterns of vegetation surrounding springs in Goshen Bay,Utah County,Utah U.S.A.

Saline meadows in Goshen Bay, Utah County, Utah, USA, occur interspersed among large areas of salt playa. Springs irrigate these meadows which show concentric vegetational zones surrounding each spring. Each vegetational zone demonstrates its own unique dominant vascular plant species. The central zone is dominated primarily by Scirpus americanus, the middle zone by Eleocharis palustris and the outer zone by Juncus balticus, Distichlis spicata and Muhlenbergia asperifolia. As distance increases from the springs, pH and soluble salt concentration increase, while percent organic matter, percent moisture and phosphorus decrease. With the increase in salt levels and decrease in water levels, halophytic plant species generally increase with distance from the springs.