Parasite burdens differ between sympatric three-spined stickleback species

The ecological theory of adaptive radiation states that differences in ecological circumstances among local populations are the cause of divergence that leads to speciation. The role of parasites in contributing to divergence has seldom been considered, despite their ubiquity and known selective effects. The potential for parasites to contribute to divergence between closely related taxa was examined by quantifying the variation in parasite burdens between sympatric three-spined stickleback species ( Gasterosteus aculeatus complex) in two lakes in coastal British Columbia, Canada. In doing so the relative importance of geographical differences between lakes and trophic or microhabitat differences between species within lakes were evaluated. The entire metazoan parasite burdens of a total of 255 limnetic and benthic sticklebacks in Paxton and Priest lakes were assayed over five time points between spring and autumn. Despite their sympatric distributions, there were large differences in parasite burdens between benthic and limnetic sticklebacks within lakes and these were consistent across both lakes. In particular, limnetics suffered greater burdens of the parasites Schistocephalus solidus and Diplostomum scudderi and benthics had much higher burdens of parasitic glochidia (mollusc larvae). Parasite burdens also differed quantitatively between lakes, but in general such differences were less pronounced than those between the stickleback species. The documented differences in parasite burdens between stickleback species have potential to contribute to divergent selection on life history, immunological and secondary sexual characters that could contribute to reproductive isolation between the species.     

Genetic divergence in adaptive characters between sympatric species of stickleback

This study explored the genetic basis of phenotypic differences between two sympatric species of ecologically and morphologically divergent sticklebacks (Gasterosteus aculeatus complex). The aim was to understand how many loci determine the differences and to what extent the differences are due to additive or nonadditive gene action. I reared the two parental species, F1 and F2 hybrids, and both backcrosses in the laboratory and measured the following quantitative characters: gill raker number and length (both involved in feeding), lateral plate number and pelvic spine length (both involved in predator defense), and growth (a fitness component). I then applied joint-scaling regression models to estimate composite additive, dominance and epistatic effects, and their contribution to divergence of parental lines. A simple additive model was sufficient for gill raker number and growth; additive and dominance effects contributed significantly to divergence in plate number and pelvic spine length; and additive, dominance, and epistatic effects contributed significantly to divergence in gill raker length. Wright's estimator for the number of loci for the four morphological characters ranged from 1 to 50. My results suggest that adaptive divergence between limnetic and benthic sticklebacks has taken place through a variety of genetic mechanisms specific to different traits. Though interspecific hybrids are completely fertile and viable in the laboratory, they are selected against in the wild. The pattern of inheritance for the traits examined here directly influences how well hybrids can exploit the two major resource environments in the wild.     

Courtship behavior,nesting microhabitat,and assortative mating in sympatric stickleback species pairs

The maintenance of reproductive isolation in the face of gene flow is a particularly contentious topic, but differences in reproductive behavior may provide the key to explaining this phenomenon. However, we do not yet fully understand how behavior contributes to maintaining species boundaries. How important are behavioral differences during reproduction? To what extent does assortative mating maintain reproductive isolation in recently diverged populations and how important are “magic traits”? Assortative mating can arise as a by‐product of accumulated differences between divergent populations as well as an adaptive response to contact between those populations, but this is often overlooked. Here we address these questions using recently described species pairs of three‐spined stickleback (Gasterosteus aculeatus), from two separate locations and a phenotypically intermediate allopatric population on the island of North Uist, Scottish Western Isles. We identified stark differences in the preferred nesting substrate and courtship behavior of species pair males. We showed that all males selectively court females of their own ecotype and all females prefer males of the same ecotype, regardless of whether they are from species pairs or allopatric populations. We also showed that mate choice does not appear to be driven by body size differences (a potential “magic trait”). By explicitly comparing the strength of these mating preferences between species pairs and single‐ecotype locations, we were able to show that present levels of assortative mating due to direct mate choice are likely a by‐product of other adaptations between ecotypes, and not subject to obvious selection in species pairs. Our results suggest that ecological divergence in mating characteristics, particularly nesting microhabitat may be more important than direct mate choice in maintaining reproductive isolation in stickleback species pairs.     

Feeding strategy of two sympatric anostracan species (Crustacea)

The transport rate of chalk, clay particles and algal cells (Scenedesmus sp.) through the digestive tract of Streptocephalus torvicornis and Branchipus schaefferi is described under experimental conditions. Differences in transport rate as well as in the degree of digestion at a fixed particle density of algae were found. In S. torvicornis, the transport rate is higher and the digestion of algae lower than in B. schaefferi. These differences might reflect trophic differences related to niche partitioning in these sympatric species. This revised version was published online in August 2006 with corrections to the Cover Date.     

Isolation between sympatric anadromous and resident threespine stickleback species in Mud Lake,Alaska

A sympatric pair of anadromous and resident freshwater threespine stickleback species (Gasterosteus aculeatus species complex) occurs in Mud Lake in the Matanuska-Susitna Valley, Alaska. The two forms differ in an array of morphological traits, including traits associated with predator defense (e.g., spine lengths) and trophic ecology (e.g., number of gill rakers). Mud Lake is only the third lake reported to have anadromous stickleback (which have a complete row of lateral plates) coexisting with low-plated resident stickleback in the absence of intermediate partially plated fish. Microhabitat and seasonal isolation appear to contribute to reproductive isolation between the two forms.     

慈姑属4种同域分布植物的人工种间杂交

泽泻科（Alismataceae）是现存最原始的单子叶植物类群之一,慈姑属（Sagittaria）是泽泻科相对进化的分类群,地域分布广泛,其物种大多集中分布于北温带,且有种间同域分布现象。为了探究同域分布物种之间的杂交可能性,我们通过选取同域分布且在系统进化树上具有代表性的冠果草（S.guayanensis subsp. lappula）、小慈姑（S. potamogetonifolia）、利川慈姑（S. lichuanensis）和野慈姑（S. trifolia）为研究对象,调研中国数字植物标本馆（CVH）的标本信息和已有文献,分析这4种慈姑的地理分布格局,并分别以这4种慈姑为母本进行人工授粉实验以检验种间杂交的可能性。基于对1 172份4种慈姑属腊叶标本的分析,表明这4种慈姑在福建省武夷山地区存在同域分布情况。通过对4种慈姑之间的人工授粉实验结果分析表明,冠果草、小慈姑、利川慈姑和野慈姑异种授粉后均出现心皮膨大现象,所有异种授粉产生的种子均不可育,败育胚种子比例各不相同,且该比例与父本和母本在系统进化树上的位置有关。     

Overlap of temporal niches among four sympatric species of shrews

Hypotheses about the dependence of circadian activity from metabolic rate and the segregation of temporal niches among competing species were verified by the study of activity patterns in a shrew community of two semiaquatic species,Neomys anomalus Cabrera, 1907 andN. fodiens (Pennant, 1771), and two terrestrial species,Sorex araneus Linnaeus, 1758 andS. minutus Linnaeus, 1766, co-existing in wet habitats of Białowieża Forest (E Poland). In ten trapping sessions, performed in early summer between 1991 and 2000, traps were open 24 hours continuously and patrolled at 1:00, 5:00, 10:00, 15:00, and 20:00. All the shrew species were most active between 20:00 and 1:00, and least active around mid-day (10:00–15:00). However, activity of the twoSorex species was lower than that of the twoNeomys species in the period 20:00–1:00, but higher in the period 15:00–20:00. BothNeomys species displayed clearly nocturnal, unimodal patterns of activity. In contrast, activity of bothSorex species was relatively evenly distributed over 24 hours and they increased their activity earlier (ie after 15:00) than bothNeomys species (after 20:00). These results confirm the idea that small shrew species with higher metabolic rate have more frequent and more equally distributed activity bouts than large species. Overlap of temporal niches was the highest within genera (99.29% between bothNeomys species and 98.36% between bothSorex species), the lowest betweenN. fodiens andS. araneus (88.26%) andS. minutus (89.34%), and intermediate betweenN. anomalus and bothSorex species (91.78 and 93.34%, respectively). Such high interspecific overlaps in activity suggest a joint-action of other mechanisms that separate ecological niches of these species also in other dimensions (eg food, microhabitat).     

Morphometry and growth of sea pen species from dense habitats in the Gulf of St. Lawrence,eastern Canada

We examined four species of sea pen (Anthoptilum grandiflorum, Halipteris finmarchica, Pennatula aculeata and Pennatula grandis) collected from the Gulf of St. Lawrence and mouth of the Laurentian Channel, eastern Canada. An exponential length–weight relationship was found for all four species, where growth in weight was progressively greater than growth in length with increasing colony size. Halipteris finmarchica, P. grandis and P. aculeata presented the better allometric fits, explaining over 80% of the variance. In addition, a count of growth increments visible in transverse sections in 86 A. grandiflorum and 80 P. aculeata samples was made. Presumed ages ranged between 5 and 28 years for A. grandiflorum and 2 and 21 years for P. aculeata. Radiocarbon assays were inconclusive and could not be used to confirm these ages; further age validation is required. Radial growth of the rod is slow during the first years, increasing at intermediate sizes of the colony and slowing down again for large colonies. Similar results were obtained from the relationship between colony length and number of growth increments where a logistic model was the best fit to the data. On average Spearman’s rank correlations showed 11% of shared variance between sea pen length or weight and environmental variables. Bottom temperature and salinity, depth and summer primary production were significantly correlated to sea pen size for most species.     

Patterns of genomic divergence and introgression between Japanese stickleback species with overlapping breeding habitats

With only a few absolute geographic barriers in marine environments, the factors maintaining reproductive isolation among marine organisms remain elusive. However, spatial structuring in breeding habitat can contribute to reproductive isolation. This is particularly important for marine organisms that migrate to use fresh‐ or brackish water environments to breed. The Japanese Gasterosteus stickleback species, the Pacific Ocean three‐spined stickleback (G. aculeatus) and the Japan Sea stickleback (G. nipponicus) overwinter in the sea, but migrate to rivers for spawning. Although they co‐occur at several locations across the Japanese islands, they are reproductively isolated. Our previous studies in Bekanbeushi River showed that the Japan Sea stickleback spawns in the estuary, while the Pacific Ocean stickleback mainly spawns further upstream in freshwater. Overall genomic divergence was very high with many interspersed regions of introgression. Here, we investigated genomic divergence and introgression between the sympatric species in the much shorter Tokotan River, where they share spawning sites. The levels of genome‐wide divergence were reduced and introgression was increased, suggesting that habitat isolation substantially contributes to a reduction in gene flow. We also found that genomic regions of introgression were largely shared between the two systems. Furthermore, some regions of introgression were located near loci with a heterozygote advantage for juvenile survival. Taken together, introgression may be partially driven by adaptation in this system. Although, the two species remain clearly genetically differentiated. Regions with low recombination rates showed especially low introgression. Speciation reversal is therefore likely prevented by barriers other than habitat isolation.     

Relationships between bird species and tree species assemblages in forested habitats of eastern North America

Aim We examined the relative influence of geographical location, habitat structure (physiognomy), and dominant plant species composition (floristics) on avian habitat relationships over a large spatial extent. Although it has been predicted that avian distributions are more likely to covary with physiognomy than with floristics at coarse scales, we sought to determine, more specifically, whether there remained a significant association between gradients in assemblages of bird species and dominant plant species within a general biome type, after statistically controlling for structural variation and geographical location of sampling sites. Location Our sample consisted of a subset of North American Breeding Bird Census survey sites that covered most of the range of eastern forests, from Florida to Nova Scotia, and west to Minnesota and North Dakota (up to c. 2500 km between sites). Methods We restricted our analyses to the single year (1981) that provided the largest sample of sites (47) for which vegetation data were available within ± 2 years of the avian surveys. We examined the relationship between avian community composition and tree species composition over this series of forested plots. Data were divided into four sets: (1) bird species abundances, (2) tree species abundances, (3) physiognomic or structural variables and (4) geographical location (latitude and longitude). We performed separate detrended correspondence analysis ordinations of birds and trees, before and after statistically partialling out covariation associated with structural variables and geographical location. To gauge the relationship between the two sets of species we correlated site scores resulting from separate ordinations. We also compared continental‐scale patterns of variation in bird and tree assemblages to understand possible mechanisms controlling species distribution at that scale. Results Both bird and tree communities yielded strong gradients, with first‐axis eigenvalues from 0.75 to 0.97. All gradients were relatively long (> 4.0), implying complete turnover in species composition. However, geographical location accounted for < 10% of the total variation associated with any ordination. Prior to partialling out covariation resulting from location and physiognomy, bird species ordinations were strongly correlated with tree species ordinations. The strength of association was reduced after partialling, but one bird and one tree axis remained significantly correlated. There was a significant species–area effect for birds, but not for trees. Main conclusions There was a significant relationship between bird species assemblages and tree species assemblages in the eastern forests of North America. Even after partialling out covariation associated with spatial location and forest physiognomy, there remained a significant correlation between major axes from bird and tree ordinations, consistent with the hypothesis that floristic variation is likely to be important in organizing assemblages of birds within a general biome type, albeit over a much larger spatial extent than originally predicted. Forest tree species ordinations differed from bird species ordinations in several ways: trees had a higher rate of turnover along underlying environmental gradients; trees appeared more patchily distributed than birds at this scale; and tree species were more spaced out along the underlying ecological gradients, with less overlap. By understanding the relationship between bird assemblages and forest floristics, we might better understand how avian communities are likely to change if tree species distributions are altered as a result of climatic changes.     

Modelling sympatric speciation by means of biologically plausible mechanistic processes as exemplified by threespine stickleback species pairs

We investigate the plausibility of sympatric speciation through a modelling study. We built up a series of models with increasing complexity while focussing on questioning the realism of model assumptions by checking them critically against a particular biological system, namely the sympatric benthic and limnetic species of threespine stickleback in British Columbia, Canada. These are morphologically adapted to their feeding habits: each performs better in its respective habitat than do hybrids with intermediate morphology. Ecological character displacement through disruptive selection and competition, and reinforcement through mating preferences may have caused their divergence. Our model assumptions include continuous morphological trait(s) instead of a dimorphic trait, and mating preferences based on the same trait(s) as selected for in food competition. Initially, morphology is intermediate. We apply disruptive selection against intermediates, frequency-dependent resource competition, and one of two alternative mating preference mechanisms. Firstly, preference is based on similarity where mating preference may result from “imprinting” on conspecifics encountered in their preferred foraging habitat. Here, speciation occurs easily—ecological hybrid inferiority is not necessary. Hybrid inferiority reinforces the stringency of assortative mating. Secondly, individual preferences exist for different trait values. Here, speciation occurs when linkage disequilibrium between trait and preference develops, and some hybrid inferiority is required. Finally, if the morphology subject to disruptive selection, frequency-dependent competition, and mate choice, is coded for by two loci, linkage disequilibrium between the two loci is required for speciation. Speciation and reinforcement of stringency of choosiness are possible in this case too, but rarely. Results demonstrate the contingency of speciation, with the same starting point not necessarily producing the same outcome. The study resulted in flagging issues where models often lack in biological realism and issues where more empirical studies could inform on whether assumptions are likely valid.     

Reproductive characteristics and population decline of four species of skate (Rajidae) off the eastern coast of Canada

Four of the most common species of skate (Rajidae) were studied off eastern Canada to determine if their reproductive characteristics were linked to their population trajectories. The fecundity of the winter skate Leucoraja ocellata , the little skate Leucoraja erinacea , the thorny skate Amblyraja radiata and the smooth skate Malacoraja senta averaged between 41 and 56 egg cases per year for each species. For all species but L. ocellata , males matured at larger sizes and at later ages than females. Theoretical rates of population increase for non-equilibrium populations of L. ocellata ( c . 0·07), M. senta ( c . 0·14) and L. erinacea and A. radiata ( c . 0·20) were low compared to most fishes, indicating that north-west Atlantic skates are intrinsically unproductive, yet are theoretically capable of supporting low-level fisheries. Nevertheless, the results of 36 years of research surveys indicate that the abundance of mature L. ocellata , A. radiata and M. senta all decreased by >90% since 1970, indicating that past fishing mortality (both directed and undirected) has outstripped the net productivity of the skate populations on the eastern Scotian Shelf. The relationship between maximum age ( t _max) and age of maturity ( t _mat) was a better predictor of population growth rate than was body size, with the species exhibiting the highest ratios of t _mat : t _max ( L. ocellata = 0·68, M. senta = 0·66) having the lowest predicted population growth rates. L. ocellata appears to have the lowest productivity and has experienced the greatest population decline, thus raising concerns over its future status.     

Floral fungal-bacterial community structure and co-occurrence patterns in four sympatric island plant species

Flowers’ fungal and bacterial communities can exert great impacts on host plant wellness and reproductive success—both directly and indirectly through species interactions. However, information about community structure and co-occurrence patterns in floral microbiome remains scarce. Here, using culture-independent methods, we investigated fungal and bacterial communities associated with stamens and pistils of four plant species (Scaevola taccada, Ipomoea cairica, Ipomoea pes-caprae, and Mussaenda kwangtungensis) growing together under the same environment conditions in an island located in South China. Plant species identity significantly influenced community composition of floral fungi but not bacteria. Stamen and pistil microbiomes did not differ in community composition, but differed in co-occurrence network topological features. Compared with the stamen network, pistil counterpart had fewer links between bacteria and fungi and showed more modular but less concentrated and connected structure. In addition, degree distribution of microbial network in each host species and each microhabitat (stamen or pistil) followed a significant power-law pattern. These results enhance our understanding in the assembly principles and ecological interactions of floral microbial communities.     

The species concept in diatoms: Evidence for morphologically distinct,sympatric gamodemes in four epipelic species

Reproductive isolation is demonstrated between morphologically distinct, sympatric demes of four epipelic diatom species inhabiting a small pond. The six demes ofSellaphora pupula, four of them allogamous, one probably autogamous, and one apparently asexual, differ morphologically with respect to size, shape, stria density and pattern. In the pond,Caloneis silicula, Cymatopleura solea, andNeidium ampliatum each contain two allogamous demes, differing principally in size. There are intrinsic barriers to hybridization between the demes, which operate at the earliest stages of sexual reproduction. Such gamodemes are perhaps best treated taxonomically as separate species.Dedicated to Prof. DrLothar Geitler on the occasion of his 90th birthday. I gratefully acknowledge the debt I owe Prof.Geitler for the insights and inspiration I have gained from his papers. I account myself fortunate to have visited him briefly in 1983, at Lunz, and to have seen the rivers where his work on diatom species began in 1922.     

Feeding strategies and diet composition of four Antarctic cnidarian species

The diet of four species of Antarctic cnidarians, two hydroids and two anthozoans, was investigated. One hydroid, Tubularia ralphii, and one anthozoan, Anthomastus bathyproctus, seem to basically consume zooplankton whereas the other hydroid species, Oswaldella antarctica, has a diet mainly based on the fine fraction of seston. The last cnidarian investigated, the stoloniferan alcyonacean, Clavularia cf. frankliniana, feeds mainly on resuspended material. The wide range of diets of these Antarctic cnidarians indicates opportunistic behaviour by feeding on different sources and taking advantage of the available food sources. Data related to low C/N ratios in the sediment and high NO₂ concentrations in areas with dense communities of benthic suspension feeders, along with data on capture rates, lead us to hypothesize that these organisms play an important role in the recycling processes of organic matter in Antarctic benthic ecosystems.