Tamarin polyspecific associations: Forest utilization and stability of mixed-species groups

Niche separation is likely to play a key role in the formation of mixed-species groups. Saddle-backed tamarins (Saguinus fuscicollis) were studied at three sites with different primate communities in northern Bolivia: (1) with red-bellied tamarins,S. labiatus; (2) with emperor tamarins,S. imperator; and (3) without a congeneric species. The degree of association is higher betweenS. labiatus andS. fuscicollis than betweenS. imperator andS. fuscicollis and is related to differences in forest utilization between associating pairs. Niche separation is found to be greater betweenS. labiatus andS. fuscicollis than betweenS. fuscicollis andS. imperator. The mean height and habitat utilization ofS. fuscicollis does not differ greatly across the three sites, nor does the height of tamarins in and out of association. It is concluded that combined with differences in body size and dietary overlap, vertical segregation plays an important role in tamarin polyspecific associations (increasing the potential of both foraging and anti-predatory benefits) and that this is not a consequence of vertical displacement ofS. fuscicollis by its dominant congeners.     

Space,sympatry and speciation

Sympatric speciation remains controversial. ‘Sympatry’ originally meant “in the same geographical area”. Recently, evolutionists have redefined ‘sympatric speciation’ non‐spatially to require panmixia (m = 0.5) between a pair of demes before onset of reproductive isolation. Although panmixia is a suitable starting point in models of speciation, it is not a useful definition of sympatry in natural populations, because it becomes virtually impossible to find or demonstrate sympatry in nature. The newer, non‐spatial definition fails to address the classical debate about whether natural selection within a geographic overlap regularly causes speciation in nature, or whether complete geographic isolation is usually required. We therefore propose a more precise spatial definition by incorporating the population genetics of dispersal (or ‘cruising range’). Sympatric speciation is considerably more likely under this spatial definition than under the demic definition, because distance itself has a powerful structuring effect, even over small spatial scales comparable to dispersal. Ecological adaptation in two‐dimensional space often acts as a ‘magic trait’ that causes pleiotropic reductions of gene flow. We provide examples from our own research.     

Distribution,bioclimatic niche and sympatry of two Erinaceidae in Tunisia

Numerous mammalian species of North Africa have an enigmatic distribution between the Mediterranean and Saharan area. In order to study the distribution of hedgehogs in Tunisia, field data were collected from several protected areas and various natural and peri-urban environments during 4 years 2015–2018 covering the all bioclimatic zones present in the country. The present work provides a detailed geographical and ecological distribution of hedgehogs with a total of 168 occurrence data of Atelerix algirus and Paraechinus aethiopicus. Atelerix algirus showed a wide distribution range from north to south being absent only in the extreme Saharan environment. Paraechinus aethiopicus has been recorded in the centre and the south with a specialisation in the arid and Saharan environment. A parapatry distribution is also shown with a sympatric area between arid and Saharan bioclimates. Our results improve knowledge of the bioclimatic niche and habitat characteristic of these taxa not clearly defined in previous research. The presence of the two hedgehogs on Djerba Island (A. algirus and P. aethiopicus) is also confirmed.     

Host–pathogen coevolution,secondary sympatry and species diversification

The build-up of species locally within a region by allopatric speciation depends on geographically separated (allopatric) sister populations becoming reproductively incompatible followed by secondary sympatry. Among birds, this has happened frequently in remote archipelagos, spectacular cases including the Darwin''s finches (Geospizinae) and Hawaiian honeycreepers (Drepanidinae), but similar examples are lacking in archipelagos nearer to continental landmasses. Of the required steps in the speciation cycle, achievement of secondary sympatry appears to be limiting in near archipelagos and, by extension, in continental regions. Here, I suggest that secondary sympatry might be prevented by apparent competition mediated through pathogens that are locally coevolved with one population of host and are pathogenic in sister populations. The absence of numerous pathogens in remote archipelagos might, therefore, allow sister populations to achieve secondary sympatry more readily and thereby accelerate diversification. By similar reasoning, species should accumulate relatively slowly within continental regions. In this essay, I explore the assumptions and some implications of this model for species diversification.     

Trophic associations and specialization of phytophagous beetles (Coleoptera: Chrysomeloidea,Curculionoidea) in the east of the Russian Plain

Data on the trophic associations of beetles with plants in the east of the Russian Plain are summarized and comparative analysis of host specialization of different groups of phytophagous beetles is performed. In terms of the width of the regional trophic spectrum, monophages and narrow oligophages prevail among the Curculionoidea as a whole and in the families Curculionidae and Apionidae in particular, while moderate and broad oligophages prevail in the Chrysomeloidea and in the family Chrysomelidae. Two-thirds of the regional fauna (66%) of Curculionoidea are closely associated with plants of one genus; by contrast, in Chrysomeloidea almost 40% of the species can develop on plants from different genera of one family, the fraction of the narrowly specialized forms comprising only 43%. The higher level of trophic specialization of weevils (Curculionidae, Apionidae) and seed beetles (Bruchidae), as compared to leaf beetles (Chrysomelidae), is probably due to the larval endophagy of most species of these families. Analysis of the distribution of beetles over host plants has shown that the specialized forms are associated with plants of 65 families (about 60% of the regional flora in the east of the Russian Plain). Distribution of beetles over plant families is very non-uniform. Most of the specialized forms (78%) are associated with plants of 15 families, three of which (Asteraceae, Fabaceae, and Brassicaceae) include hosts of more than onethird of the beetle species (37%). Monophages and narrow oligophages are recorded on 201 genera of plants from 59 families. Polyphagous species are recorded on plants of 58 families. The specific features of the distribution of phytophagous beetles over host plants (as compared to other insects) is a high fraction of species developing on coenophobes (particularly those of the family Brassicaceae) typical of the pioneer stages of successions with sparse herbaceous cover, and a small number of species associated with grasses and sedges. These features are most conspicuous in the fauna of Curculionidae.     

Electrical conductivity in a mixed-species biofilm

Geobacter sulfurreducens can form electrically conductive biofilms, but the potential for conductivity through mixed-species biofilms has not been examined. A current-producing biofilm grown from a wastewater sludge inoculum was highly conductive with low charge transfer resistance even though microorganisms other than Geobacteraceae accounted for nearly half the microbial community.     

Decomposition dynamics in mixed-species leaf litter

Literature on plant leaf litter decomposition is substantial, but only in recent years have potential interactions among leaves of different species during decomposition been examined. We review emerging research on patterns of mass loss, changes in nutrient concentration, and decomposer abundance and activity when leaves of different species are decaying in mixtures. Approximately 30 papers have been published that directly examine decomposition in leaf mixtures as well as in all component species decaying alone. From these litter‐mix experiments, it is clear that decomposition patterns are not always predictable from single‐species dynamics. (Characteristics of decomposition in litter‐mixes that deviate from responses predicted from decomposition of single‐species litters alone are designated "non‐additive"; "additive" responses in mixes are predictable from component species decaying alone.) Non‐additive patterns of mass loss were observed in 67% of tested mixtures; mass loss is often (though not always) increased when litters of different species are mixed. Observed mass loss in some mixtures is as much as 65% more extensive than expected from decomposition of single‐species litter, but more often mass loss in mixtures exceeds expected decay by 20% or less. Nutrient transfer among leaves of different species is striking, with 76% of the mixtures showing non‐additive dynamics of nutrient concentrations. Non‐additive patterns in the abundance and activity of decomposers were observed in 55% and 65% of leaf mixes, respectively. We discuss some methodological details that likely contribute to conflicting results among mixed‐litter studies to date. Enough information is available to begin formulating mechanistic hypotheses to explain patterns in litter‐mix experiments. Emerging patterns in the mixed‐litter decomposition literature have implications for relationships between biodiversity and ecosystem function (in this case, the function being decomposition), and for potential mechanisms through which invasive plant species could alter carbon and nutrient dynamics in ecosystems.     

Dispersal and the transition to sympatry in vertebrates

Under allopatric speciation models, a key step in the build-up of species richness is population dispersal leading to the co-occurrence of previously geographically isolated forms. Despite its central importance for community assembly, the extent to which the transition from spatial segregation (allopatry or parapatry) to coexistence (sympatry) is a predictable process, or alternatively one governed by chance and the vagaries of biogeographic history, remains poorly understood. Here, we use estimated divergence times and current patterns of geographical range overlap among sister species to explore the evolution of sympatry in vertebrates. We show that rates of transition to sympatry vary predictably according to ecology, being faster in marine or strongly dispersive terrestrial clades. This association with organism vagility is robust to the relative frequency of geographical speciation modes and consistent across taxonomic scales and metrics of dispersal ability. These findings reject neutral models of dispersal assembly based simply on evolutionary age and are not predicted by the main alternative view that range overlap is primarily constrained by biotic interactions. We conclude that species differences in dispersal limitation are fundamental in organizing the assembly of ecological communities and shaping broad-scale patterns of biodiversity over space and time.     

Evidence for facilitation among avian army‐ant attendants: specialization and species associations across elevations

Mixed‐species assemblages can involve positive and negative interactions, but uncertainty about high‐value patchy resources can increase the value of information sharing among heterospecific co‐foragers. I sampled species composition of bird‐flocks attending army‐ant raids in three adjacent elevation zones in Costa Rica, across multiple years, to test for positive and negative associations among raid‐attending bird species. My goal was to test whether the most frequent and specialized raid‐attending species showed evidence of facilitating or excluding other bird species. I quantified elevational variation in avian community composition at raids, then asked whether species composition was associated with variation in flock characteristics (flock size and species richness). I identified the most frequent raid‐attending species (those that attended raids most frequently relative to their mist‐net capture rates), and bird species that performed specialized army ant‐following behavior (bivouac‐checking, which allows birds to memorize and track mobile army‐ant colonies). There was significant turnover of bird species among zones (including the frequent and specialized attendants); patterns of species overlap suggested a gradual transition from a Pacific‐slope to an Atlantic‐slope raid‐attending bird fauna. Raid‐attendance frequency was positively correlated with bivouac‐checking behavior. With few exceptions, the most frequent raid‐attending bird species, and the bivouac‐checking species, also participated in the most species‐rich flocks. High species‐gregariousness suggests many of the frequently attending and/or bivouac‐checking species functioned as core flock members. However, some bird species pairs were significantly negatively associated at raids. Despite species turnover, per‐flock numbers of birds at raids did not differ among geographic zones, but flocks on the Pacific‐slope were heavier because larger bodied bird species attended raids. Previous studies showed that the size (biomass) of bird‐flocks corresponds to the amount of food the birds kleptoparasitize from ant raids, and the heavier Pacific‐slope bird‐flocks could have greater negative kleptoparasitic impacts.     

Ecology of endangered Prunus korshinskyi Hand.-Mazz. in Jabal Al-Lauz,Saudi Arabia: Plant associations,size structure,and nutritional screening

The wild Prunus korshinskyi has a restricted distribution in small scattered areas of Jabal Al-Lauz (2580 m above sea level), a mountain in northwestern Saudi Arabia. Major objective of current study is to provide information about P. korshinskyi by analyzing its ecology in terms of floristic diversity, plant associations, environmental features, and its size structure and nutritional value. For this purpose, 12 stands of 20 m² were selected along the Jabal Al-Lauz mountain ridge. Three plant associations and their environmental variables were identified and characterized after application of a two-way indicator species analysis (TWINSPAN), detrended correspondence analysis (DCA). And canonical correspondence analysis (CCA) as follows: VGI: P. korshinskyi-Astracantha echinus, VGII: Artemisia siberi-P. korshinskyi, and VGIII: Retama raetam-Artemisia sieberi. The edaphic factor affecting the distribution of the associations were pH, potassium and manganese content. The size class frequency distribution of P. korshinskyi shows a J-shape in spring and fall for the whole population; there was no regeneration via seed. Nutritional evaluation showed the fruits had a higher content of total carbohydrate, fats, crude protein, phosphorus, and potassium, and lower content of ash, Fe, Ca, Mn, Mg, and Zn than that in stems and leaves. This ecological knowledge gained through this study would be beneficial for managing and conserving of P. korshinskyi in a Jabal Al-Lauz area with its distinct and unique vegetation.     

Composition and structure of a chronosequence of young, mixed-species forests in southeastern Ohio, USA

Clearcutting, a commonly used silvicultural practice in southeastern Ohio, often results in a forest stand with a different species composition than the parent stand. The time frames during which shifts in species composition occur on different sites are unclear. While some studies have documented species composition at specific points in time, none have attempted to examine differences throughout the first decades of stand development. This study focused on the early successional dynamics of young, mixed-species forests of southeastern Ohio. Species compositions were examined across a chronosequence of sixteen stands that developed following clearcutting. Stand ages ranged from six to 26 years. The sample was limited to dry-mesic hardwood forests on southerly aspects and on soils derived from residuum or colluvium. Across the chronosequence, stand density ranged from 17 636 stems ha^-1 at age 6 to 2759 stems ha^-1 at age 26, and basal area ranged from 8.2 m² ha^-1 to 22.1 m² ha^-1. Clumps comprised a substantial portion of the total stand density and basal area. At age 6–8 years after clearcutting, clumped stems accounted for 35.1% of the density and 48.2% of the basal area. At age 26 years, these proportions were 25.7% and 29.4%, respectively. Clumped stems were significantly larger (p<0.05) than non-clumped stems at each age group except 26 years. Total Quercus spp. density was greatest at age 6–8 years (3386 stems ha^-1), and least at age 26 years (581 stems ha^-1). When considered as a proportion of the total stand, however, the proportion was relatively stable, averaging 21.3%. However, importance value (IV=[relative density + relative basal area]/2) of Quercus in the upper canopy (dominant and codominant crown classes) was twice as much (72%) at age 26 years compared to age 6–8 years (35%). Quercus prinus L. was the major species across the chronosequence. For all age groups except 18–20 years. Q. prinus IV was the highest of any individual species in the upper canopy, and it ranged from 27 in the youngest stands (6–8 years) to 69 in the oldest stand (26 years). Within the intermediate crown class, the IV of Q. prinus equaled or exceeded those of all other species, except for the 18–20 year age group where it was second to A. rubrum. Quercus alba L. and Quercus velutina Lam. were minor components at age 26 years, although they dominated a comparison sample of six mature stands of the same ecosystem type. Liriodendron tulipifera L. was abundant 6–8 years after clearcutting, but nearly absent at age 26 years. Acer rubrum L. was the major species in both the intermediate and overtopped crown classes throughout the chronosequence. As gaps in the canopy occur. A. rubrum may become a more common species within the dominant-codominant crown class.     

Surface charge influences enterococcal prevalence in mixed-species biofilms

AIM: To study the influence of 15 microbial isolates on the prevalence of charge-heterogeneous and charge-homogeneous Enterococcus faecalis strains, all isolated from biliary stents, in mixed-species biofilms. METHODS AND RESULTS: Six Enterococcus faecalis strains were paired with 15 other microbial isolates to form mixed-species biofilms in a microtitre plate assay. Charge-heterogeneous Enterococcus faecalis strains display two subpopulations with different surface charges, expressed as a biomodal zeta potential distribution. It was found that, in general, the prevalence of the charge-heterogeneous, biofilm forming Enterococcus faecalis was reduced in mixed-species biofilms. The prevalence of charge-homogeneous, nonbiofilm-forming Enterococcus faecalis strains was increased only in the presence of Citrobacter freundii BS5126, Stenotrophomonas malthophilia BS937, and Candida lusitaniae BS8256, all of which introduced sizeable charge heterogeneity in the mixed microbial population. CONCLUSIONS: Charge-homogeneous Enterococcus faecalis strains are stimulated to form biofilm only by the presence of another microbial species with a considerably less negative zeta potential, thereby creating a charge-heterogeneous microbial population. SIGNIFICANCE AND IMPACT OF THE STUDY: Enterococcus faecalis is one of the predominant species isolated from mixed-species biofilms in clogged biliary stents. The current study shows how charge-homogeneous Enterococcus faecalis strains form more biofilm in the presence of other microbial species.     

Reproductive interference between honeybee species in artificial sympatry

Reproductive isolation between closely related species is often incomplete. The Western honeybee, Apis mellifera, and the Eastern hive bee, Apis cerana, have been allopatric for millions of years, but are nonetheless similar in morphology and behaviour. During the last century, the two species were brought into contact anthropogenically, providing potential opportunities for interspecific matings. Hybrids between A. mellifera and A. cerana are inviable, so natural interspecific matings are of concern because they may reduce the viability of A. cerana and A. mellifera populations – two of the world's most important pollinators. We examined the mating behaviour of A. mellifera and A. cerana queens and drones from Caoba Basin, China and Cairns, Australia. Drone mating flight times overlap in both areas. Analysis of the spermathecal contents of queens with species‐specific genetic markers indicated that in Caoba Basin, 14% of A. mellifera queens mated with at least one A. cerana male, but we detected no A. cerana queens that had mated with A. mellifera males. Similarly, in Cairns, no A. cerana queens carried A. mellifera sperm, but one‐third of A. mellifera queens had mated with at least one A. cerana male. No hybrid embryos were detected in eggs laid by interspecifically mated A. mellifera queens in either location. However, A. mellifera queens artificially inseminated with A. cerana sperm produced inviable hybrid eggs or unfertilized drones. This suggests that reproductive interference will impact the viability of honeybee populations wherever A. cerana and A. mellifera are in contact.     

Arctic charr in sympatry with burbot: ecological and evolutionary consequences

The trophic niche and parasite infection of Arctic charr (Salvelinus alpinus) were explored in two lakes with sympatric burbot (Lota lota) and two lakes without burbot in subarctic Norway. The CPUE of burbot and charr were similar in one lake, but burbot had a low population density in the other. Burbot were benthivorous in both lakes. Other co-occurring species like brown trout (Salmo trutta), Atlantic salmon parr (Salmo salar), grayling (Thymallus thymallus) and minnow (Phoxinus phoxinus) were also benthivores. At high densities, benthivorous burbot forced the whole Arctic charr population to utilise mainly the limnetic trophic niche. In contrast, at low burbot density or without burbot present, Arctic charr were primarily benthivorous in the littoral zone. Thus, a clear interactive segregation in diet was observed between Arctic charr and burbot at high burbot densities. There was also a high predation pressure from burbot on young Arctic charr along the benthic zones. The extensive use of zooplankton as prey caused a high parasite infection pressure of copepod transmitted Diphyllobothrium spp. larvae, with the potential for high negative impact on the Arctic charr population. As the benthivore trophic niche was occupied by burbot, the ecological opportunities for polymorphism with benthivorous ecotypes or morphs of Arctic charr were probably prevented. Therefore, the sympatry with burbot seems to have large ecological and evolutionary consequences for this Arctic charr population compared with neighbouring lakes where burbot is absent.     

Associations among Antarctic seabirds in mixed-species feeding flocks

We studied ten mixed-species feeding flocks of seabirds and seals off South Georgia (55̀S, 35̀W) to assess the factors that influenced the species composition of the flocks. The ten flocks were distributed between two oceanographic regions; four flocks were observed off the northwestern end of South Georgia, and six off the island's southeastern end. The flocks differed dramatically in size and species composition. The northwestern flocks were ten to 20 times larger and contained more seals and penguins. We tested whether these differences were a consequence of location, because the two regions were populated by different assemblages of species, and determined that this was not the case. Differences in flock size and species composition reflected differences in the depth distribution of the birds' and seals' prey. The northwestern flocks were associated with deep swarms of Antarctic krill Euphausia superba with a significant quantity of krill also present at the surface. We also tested for differential flock participation by the species found in the surrounding waters and classified species by their proclivity to participate in the flocks. Our results indicate that Black-browed Albatross Diomedea melanophris have a strong tendency to join, and diving-petrels Pelecanoides spp. to avoid, mixed-species feeding flocks. We postulate that Black-browed Albatrosses serve as visual cues to the presence of food for one another and for other species, and that diving-petrels avoid mixed-species flocks in part because of the risk of predation.     

Biofilm development and enhanced stress resistance of a model,mixed-species community biofilm

Most studies of biofilm biology have taken a reductionist approach, where single-species biofilms have been extensively investigated. However, biofilms in nature mostly comprise multiple species, where interspecies interactions can shape the development, structure and function of these communities differently from biofilm populations. Hence, a reproducible mixed-species biofilm comprising Pseudomonas aeruginosa, Pseudomonas protegens and Klebsiella pneumoniae was adapted to study how interspecies interactions affect biofilm development, structure and stress responses. Each species was fluorescently tagged to determine its abundance and spatial localization within the biofilm. The mixed-species biofilm exhibited distinct structures that were not observed in comparable single-species biofilms. In addition, development of the mixed-species biofilm was delayed 1–2 days compared with the single-species biofilms. Composition and spatial organization of the mixed-species biofilm also changed along the flow cell channel, where nutrient conditions and growth rate of each species could have a part in community assembly. Intriguingly, the mixed-species biofilm was more resistant to the antimicrobials sodium dodecyl sulfate and tobramycin than the single-species biofilms. Crucially, such community level resilience was found to be a protection offered by the resistant species to the whole community rather than selection for the resistant species. In contrast, community-level resilience was not observed for mixed-species planktonic cultures. These findings suggest that community-level interactions, such as sharing of public goods, are unique to the structured biofilm community, where the members are closely associated with each other.