Resource partitioning by three species of newts during their aquatic phase

This paper describes the multidimensional niche of syntopic smooth newts Triturus vulgaris. alpine newts Triturus alpestris. and Italian crested newts Triturus carnifex. which constitute a complex community of eight components (the three species and their larvae, plus juvenile and neotenic alpine newts) All the potentially limiting resources (seasonal time, diel time, habitat and food) are analyzed as single niche dimensions, and then the real multidimensional niche is calculated Resource states are objectively defined, using an algorithm that groups the significant resource categories The larvae were segregated by prey, season, habitat, and water depth, in order of decreasing importance, their overlap was very low when their complete multidimensional niche was considered, and therefore they have few opportunities for present–day competition The adults were largely segregated by season and water depth, and to a lesser degree by habitat and prey, their multidimensional niches overlapped to a certain degree, thus leaving conditions for ongoing competition Considenng all the species and the life stages together, the order of importance for segregation was season, prey, habitat, water depth and diel activity, with a low overlap in the complete niche Differences m prey size were probably due to morphological constraints, differences in diel activity had very little effect on segregation except m separating larvae from the adults, and were likely promoted by need to avoid mutual predation and cannibalism It remains to be tested whether seasonal, habitat, and water depth differences are due to competitive pressures or to ecophysiological constraints     

Comparison of gemma production among three Lophozia species during the growing season

The extent and seasonal pattern of asexual reproduction and ability to germinate in the rare liverwort Lophozia ascendens and the common liverworts L. ventricosa and L. longiflora were studied in the Boubínský prales National Nature Reserve in Šumava Mts. (Bohemian Forest), South Bohemia, Czech Republic. Asexual reproduction was quantified as the number of gemmae produced per individual shoot. Numbers of gemmae per shoot among sampling months differed significantly; increase of gemma production was delayed in L. ascendens in comparison with gemma production of L. ventricosa and L. longiflora. We suggest that gemma production is influenced by environmental factors, mainly air humidity. Germinability of gemmae was low in early spring, highest in August and September and slightly depressed in October. This pattern suggests that rather mild winters in the Czech Republic cause the lower mortality of shoots during winter and the environmental pressure towards the production of dormant gemmae is not a prominent factor affecting the population dynamics of the species under study.     

Foraging behaviour of the middle spotted woodpecker Dendrocopos medius in Sweden

During winter and spring the oak Quercus robur , was the tree species most utilized by middle spoiled woodpeckers. In March, saptrees, mainly maples Acer platanoides , were preferred for a short period. In winter the birds foraged in the crowns of oaks but in spring they preferred lower stem parts. Dead branches were utilized more in winter than in spring. The birds mostly used a gleaning and probing technique, which may render the species more senstitive to severe weather conditions than other woodpecker species. Arthropods collected from trees were mostly spiders, beetles, thrips or springtails. The nestlings were mainly fed on scale insects and larvae of moths and sawflies. The oak habitat specialization is believed to be caused by competition for food with other gleaning forest birds. Food shortage might possibly occur during periods of bad weather, causing increased mortality.     

Resource partitioning by three species of hemipteran herbivores on the basis of host plant density

Summary Three hemipteran herbivores partition a common resource plant, Senecio smallii, on the basis of bud density and plant patch size. The monophagous herbivore, Neacoryphus bicrucis, is most abundant in larger patches where bud density is greatest. The oligophagous herbivore, Harmostes reflexulus, is most abundant in small patches of high bud density. The polyphagous species, Lygus pratensis, is most abundant in small patches where abundance is independent of bud density. The aggressive nature of the monophagous herbivore appears to mediate the responses of the other species.     

Bill morphology and biometrics of three sibling woodpecker species from sympatric populations

ABSTRACT

Capsule Different methods of comparing bill morphology between woodpecker species show different, but not contradictory results.

Aims Differences among similar, closely related species which co-occur are still highly debated. In such a scenario, species should diverge morphologically to reduce competition. We studied this phenomenon, in three closely related woodpecker species that co-occur in eastern Slovakia: Great Spotted Woodpecker Dendrocopos major, Syrian Woodpecker Dendrocopos syriacus and White-backed Woodpecker Dendrocopos leucotos. The Great Spotted Woodpecker has the widest niche and lives sympatrically with both Syrian and White-backed Woodpeckers, while the distributions of the latter two species do not overlap because of their use of different habitats. We predicted that Great Spotted Woodpeckers should differ morphologically from both sympatric species: Syrian and White-backed Woodpeckers.

Methods Comparisons were made between species, based on dorsal and lateral views of maxilla using both geometric morphometric techniques and traditional measurements.

Results We confirmed our hypothesis, and found significant differences in bill shape between Great Spotted Woodpecker and both congeners. This is in contrast to the observed higher similarity of Great Spotted Woodpecker and Syrian Woodpecker in traditional body measurements. However, surprisingly, Syrian Woodpecker and White-backed Woodpecker do not differ significantly in bill shape.

Conclusions This work indicates that geometric morphometry appears to be a promising tool for the investigation of how interspecific competition influences the shape of the bill between co-occurring species.     

Ecological partitioning among parapatric cryptic species

Geographic range differences among species may result from differences in their physiological tolerances. In the intertidal zone, marine and terrestrial environments intersect to create a unique habitat, across which physiological tolerance strongly influences range. Traits to cope with environmental extremes are particularly important here because many species live near their physiological limits and environmental gradients can be steep. The snail Melampus bidentatus occurs in coastal salt marshes in the western Atlantic and the Gulf of Mexico. We used sequence data from one mitochondrial (COI) and two nuclear markers (histone H3 and a mitochondrial carrier protein, MCP) to identify three cryptic species within this broad‐ranging nominal species, two of which have partially overlapping geographic ranges. High genetic diversity, low population structure, and high levels of migration within these two overlapping species suggest that historical range limitations do not entirely explain their different ranges. To identify microhabitat differences between these two species, we modelled their distributions using data from both marine and terrestrial environments. Although temperature was the largest factor setting range limits, other environmental components explained features of the ranges that temperature alone could not. In particular, the interaction of precipitation and salinity likely sets physiological limits that lead to range differences between these two cryptic species. This suggests that the response to climatic change in these snails will be mediated by changes to multiple environmental factors, and not just to temperature alone.     

An increase in parental investment during the breeding season

The intensity of nest-defence aggression by female redwinged black birds (Agelaius phoeniceus) with eggs decreased late in the breeding season, while aggression at this time increased amongst females defending broods. High predation pressure late in the breeding season decreased the probability of survival of the late nesting attempts, so that females with young had a much greater chance of success than females with eggs at this time. The observed changes in aggressive responses are predicted by the theory of parental investment.     

Ecological association of habitats and bird species during the breeding season in southeastern New South Wales

Bird communities in southeastern New South Wales, as determined by classification and ordination techniques, were seldom discrete. The most cohesive assemblage was associated with wet sclerophyll and rainforest habitats. These moist forests also supported the richest avifaunas and had high population densities. Although most birds occurred throughout the region, most species were uncommon. Of the 116 species recorded, 51 (44%) were found on fewer than 10% of the 143 plots censused. Only 19 species occurred on more than half the plots, with six of these recorded on more than 90% of the plots. Other than a small group of birds primarily restricted to moist forest, only the most common and widely distributed species regularly occurred together. Species were grouped with habitats on an elevational gradient from coastal forests to the tablelands. Within this elevational range, species were associated with major structural features of the vegetation. In terms of bird assemblages, open habitats were distinguished from those with dense vegetation. Plots which had been recently logged were separated from mature forest. The results indicate that plans for wildlife management should consider both the assemblages of species which are associated with major forest habitats and the requirements of individual species.     

Habitat saturation and communal breeding in the acorn woodpecker

The acorn woodpecker (Melanerpes formicivorus) typically lives in social groups that breed communally: all group members help to feed the young of a single nest. The behaviour of acorn woodpeckers was compared in two locations (New Mexico and California) to test the hypothesis that saturation of the habitat by established groups may lead to more than a single male-female pair living in groups during reproduction. Several lines of evidence indicated that the habitat was less saturated in New Mexico. This difference was reflected in a lower tendency for juveniles and adults to remain in groups, higher reproductive rates and, most important, smaller group sizes. These results suggest that habitat saturation plays an important role in communal breeding in this species.     

Resource dynamics and seasonal changes in competitive interactions among three cladoceran species

A new approach to measuring zooplankton feeding success on naturalseston assemblages enabled us to test laboratory studies inthe field without being confined to radiotracers and their associatedproblems. We compared changes in the nutritional status of threespecies of cladoceran - Daphnia rosea Sars, D. middendorffianaFischer and Holopedium gibberum Zaddach - exposing them to differentnatural seston compositions. A modified version of the lipid-ovary index was applied in time-series experiments at CastleLake, California, throughout the summer of 1982. We used thesame index in competition experiments which were designed todetect shifts in competitive interactions among all of the abovespecies as a consequence of changes in the resource base. Thetime-series experiments with various seston compositions indicatedthat temporal and vertical distribution patterns of grazerswere strongly affected by the availability of suitable food.Daphnia rosea and H. gibberum had no noticeable effect on eachother's growth in early and midsummer, whereas the decline ofthe D. rosea population appeared to be accelerated by H. gibberumlater in the season. Daphnia middendorffiana, spatially separatedfrom the other two species, was unable to survive on food preferredby D. rosea and H. gibberum, whereas the latter two cladoceranswere negatively affected by D. middendorffiana when placed inwater containing hypolimnetic seston. Our results suggest thatresource abundance and composition has a much larger influenceon some zooplankton communities than previously acknowledged. ¹Present address: Lockheed EMSCO, Environmental Programs Office,1050 E. Flamingo Road, Las Vegas, NV 89109, USA     

The breeding season of three species of kiwi (Apteryx) in captivity as determined from egg-laying dates

Records from kiwi ( Apteryx spp.) breeding centres were used to obtain laying dates of 403 North Island brown kiwi ( Apteryx australis mantelli ) eggs, 24 great spotted kiwi ( Apteryx haastii ) eggs and 25 little spotted kiwi ( Apteryx owenii ) eggs. North Island brown kiwi outdoors had an annual cycle of egg-laying with 88.5% of eggs laid from June to January. Eggs were laid in every month of the year, with fewest eggs in April (2.1 % of all eggs). The greatest number of eggs was laid in July (15.6% of all eggs), with a second peak of laying in October (13.5% of all eggs). Both the great spotted kiwi and little spotted kiwi had annual cycles of egg-laying, with the main egg-laying seasons being August-January (87.5% of eggs) for the former and July-December (96.0% of eggs) for the latter. The highest peaks of laying were in October and August, respectively. Kiwi eggs were considered to belong to the same clutch if they were laid no more than 40 days apart. Clutch size in the North Island brown and little spotted kiwi was most commonly one or two eggs. In the great spotted kiwi only clutches of one egg were found. The mean clutch size of North Island brown kiwi outdoors (1.51±0.05) did not differ from that of free-living North Island brown kiwi (1.33±0.09). Similarly, there was no difference in the mean clutch size of captive and free-living little spotted kiwi (1.39±0.14 and 1.10±0.07). The mean interval between eggs for kiwi outdoors did not differ between North Island brown kiwi and little spotted kiwi (27.4±0.5 and 30.7±1.8 days). The results indicate that all three species of kiwi maintain annual cycles of egg-laying in captivity, with the main egg-laying season being longer than in free-living birds.     

PERMANENT GENETIC RESOURCES: Isolation and characterization of 12 microsatellite markers in the middle-spotted woodpecker (Dendrocopos medius)

Twelve polymorphic microsatellite loci were developed for Dendrocopos medius. Polymorphism was assessed for 27 individuals from the southwesternmost population of this woodpecker species. The number of alleles per locus ranged from three to seven, with observed heterozygosity values from 0.444 to 0.852. Genotypic frequencies conformed to Hardy-Weinberg equilibrium, and no evidence for linkage disequilibrium was observed. Multilocus genotypes resulting from this set of markers will be useful to determine genetic diversity and differentiation within and among habitat patches inhabited by D. medius. Three of the loci were polymorphic for Picoides articus.     

Short term, repeated exposure to rams during the transition into the breeding season improves the synchrony of mating in the breeding season

The ram effect is widely used in Mediterranean breeds of sheep but its use in temperate genotypes is restricted by breed seasonality. However, ewes from these highly seasonal genotypes are sensitive to stimulation by rams close to the onset of the natural breeding season. In this study we developed a pre-mating protocol of repeated, short-term exposure to rams (fence-line contact or vasectomised rams) beginning during late anoestrus and continuing into the breeding season. We hypothesised that this pre-mating protocol would synchronise the distribution of mating of North of England Mule ewes during the breeding season above that observed in ewes isolated from rams prior to mating. Ram-exposed ewes were given contact with rams (Experiment 1: fence-line; FR, n = 94 and Experiment 2: vasectomised rams; VR; n = 103) for 24 h on Days 0 (10 September), 17 and 34 of the experiment. Control ewes (Experiment 1; FC, n = 98 and Experiment 2; VC; n = 106) remained isolated from rams prior to mating. In Experiment 2, a subset of VR (n = 35) and VC ewes (n = 35) were blood sampled twice weekly to monitor their pre-mating progesterone profiles. At mating, harnessed entire rams were introduced, 17 or 16 days after the last ram exposure (Experiments 1 and 2) and raddle marks were recorded daily. The median time from ram introduction to mating was reduced in ewes given both fence-line and vasectomised ram contact (P < 0.001), leading to a more compact distribution of mating and lambing (At least P < 0.01). In the blood sampled VR ewes, there was a progressive decline in the number of days from ram exposure to the onset of dioestrus (at least P < 0.05). This observation indicates that the cycles in VR ewes became increasingly synchronised over the pre-mating period, a pattern not evident in VC ewes. In conclusion, repeated, short-term exposure of ewes to rams during the transition into the breeding season is an effective method of synchronising the distribution of mating during the breeding season.     

Food and habitat partitioning among juveniles of three fish species in the pelagial of a mesotrophic lake

An analysis of the diet of 0⁺ perch, smelt and roach caught at night of the pelagial of a mesotrophic lake showed that their food was composed mainly of herbivorous and predatory cladocerans, copepods and Chaoborus larvae during summer, and of herbivorous cladocerans and copepods during October. An analysis of habitat use by juveniles revealed separation among the species: roach occurred in the upper, perch and smelt in the deeper water layers. Food and habitat were alternatively partitioned among the juveniles. In early summer the lowest food overlap between perch and smelt coincided with high habitat overlap. In October high food overlap between perch and smelt corresponded with low habitat overlap. Smelt and roach fry used common food resources throughout the season, but they were segregated in habitat.     

Yeast succession in the Amazon fruit Parahancornia amapa as resource partitioning among Drosophila spp.

The succession of yeasts colonizing the fallen ripe amapa fruit, from Parahancornia amapa, was examined. The occupation of the substrate depended on both the competitive interactions of yeast species, such as the production of killer toxins, and the selective dispersion by the drosophilid guild of the amapa fruit. The yeast community associated with this Amazon fruit differed from those isolated from other fruits in the same forest. The physiological profile of these yeasts was mostly restricted to the assimilation of a few simple carbon sources, mainly L-sorbose, D-glycerol, DL-lactate, cellobiose, and salicin. Common fruit-associated yeasts of the genera Kloeckera and Hanseniaspora, Candida guilliermondii, and Candida krusei colonized fruits during the first three days after the fruit fell. These yeasts were dispersed and served as food for the invader Drosophila malerkotliana. The resident flies of the Drosophila willistoni group fed selectively on patches of yeasts colonizing fruits 3 to 10 days after the fruit fell. The killer toxin-producing yeasts Pichia kluyveri var. kluyveri and Candida fructus were probably involved in the exclusion of some species during the intermediate stages of fruit deterioration. An increase in pH, inhibiting toxin activity and the depletion of simple sugars, may have promoted an increase in yeast diversity in the later stages of decomposition. The yeast succession provided a patchy environment for the drosophilids sharing this ephemeral substrate.     

Temporal variation in breeding and cone size selection by three species of crossbills Loxia spp. in a native Scots pinewood

The breeding biology and cone size selection of crossbills was studied mainly during 1995 to 2002 at Abernethy Forest, Scotland, an ancient native Scots pine Pinus sylvestris wood, where only a single crossbill species, the Scottish crossbill Loxia scotica, was assumed to occur and to be adapted to feed on seeds in Scots pine cones. However, three crossbill species (common Loxia curvirostra, Scottish and parrot crossbills Loxia pytyopsittacus) nested in some years, with the parrot crossbill being the most abundant. Most nests were in old large pines, with the three crossbill species not differing in their use of tree size or stand density for nesting. The mean clutch and brood sizes were 3.8 and 2.9, and their mean survivals were 86 and 74%, respectively, with no significant differences among species. The timing of breeding differed between species, with parrot crossbills breeding earliest (median date 21 March, including second attempts) and common crossbills breeding last (median date 21 April), probably in response to the differing accessibility of Scots pine seeds to these species. The difference in the time of breeding may reduce mixed mating. Crossbills foraged preferentially on trees with small cones when the cones were closed. Small cones had thinner scales than large cones, suggesting that the preference for small cones was related to higher feeding rates on these cones when cones are closed. Such a preference was also found for captive crossbills with the Scottish crossbill showing a more pronounced preference for smaller cones than the larger‐billed parrot crossbill. However, crossbills selected larger cones within trees and trees with larger cones once the cones opened in April. Such a shift occurred presumably because variation in scale thickness has little impact on seed accessibility once cones open, and larger cones have larger and more seeds. The greater ability of parrot crossbills to exploit seeds in closed Scots pine cones allowed parrot crossbills to start breeding earlier and to have young when seeds were most accessible. Only after the cones opened were the smaller‐billed common crossbills able to easily access seeds and to start breeding. The time of breeding of Scottish crossbills was intermediate between common and parrot crossbills, and they probably had an intermediate ability to exploit Scots pine cones. The reason why there were few Scottish crossbills nesting in Abernethy Forest remains a puzzle, considering that native pine wood is assumed to be the ancestral habitat to which the Scottish crossbill is adapted. The breeding season for all crossbills ended in June, when most of the seed from a given cone cohort was shed. This is when starved broods were found, not associated with bad weather.     

Habitat selection of three chrysomelid species associated with Rumex spp.

Summary Habitat selection of three chrysomelid beetles feeding on the Rumex plants was studied in relation to the ecological requirements of the three species and the habitat characteristics arising in their unstable and often unpredictable environment. The different ecological requirements and bionomic characteristics of the three chrysomelids were associated not only with their differing degrees of feeding specialization; they also influenced habitat selection. For example, the small body size and lower food demands of Mantura clavareaui, which is a feeding specialist, allowed it to exploit any habitat, irrespective of habitat size and duration. The other specialist, Gastrophysa atrocyanea, selected larger habitats because of the relatively brief lifespan of its adults after overwintering, its reduced mortality, and greater food demand. On the other hand, Galerucella vittaticollis, which is a multivoltine, general feeder, selected the habitat that lasted longer, rather than one that was larger. This species' life cycle is not synchronized with Rumex phenology, but its adults are long-lived and it has a flexible life cycle. The different patterns in habitat selection among these three chrysomelids prevented serious overlapping of their habitats; thus, the habitat overlapping of 2 or 3 species was 34.62%, that of all 3 species 3.85%. Most of the habitats utilized by Gastrophysa, however, overlapped those of the other two species, because Gastrophysa's habitat utilization range was the narrowest of the three. In Mantura, the degree of overlap with the other two species was lowest because this beetle had the broadest range of habitat utilization. On the other hand, Galerucella adjusted the temporal, rather than the spatial coordinates of its habitat usage when it occupied habitats where the other two specialists were also present. Galerucella, unlike the other two species, is multivoltine. Consequently, it was able to coexist in habitats occupied by the others by exploiting the host plants at times when the other species were numerically few, or even absent. No direct interspecific competition among adults of the three species was observed. Their different patterns of habitat selection created by their different responses to available food allowed them to coexist on Rumex without direct competition.     

Resource partitioning among the fishes of rainforest streams in Sri Lanka

The fish assemblage typical of small rainforest streams of Sri Lanka was investigated to see if the fishes were characterized by a high degree of specialization, expected of equilibrium communities, or if they were relatively unspecialized, expected of more stochastic (non-equilibrium) communities. Morphological features, habitats, microhabitats and diets of the 20 most abundant fish species were measured. The fishes included 11 Cyprinidae (including seven Barbus spp. and two Rasbara spp.), two Gobiidae, two Channidae, and one species each in the Cobitidae, Cyprinodontidae, Belontidae, Belonidae, and Mastacembelidae. Microhabitats were determined by measuring (for 6760 fish) water column depth, distance of fish from bottom, mean water column velocity, water velocity at fish, and substrate. Morphologically, the species showed a high degree of specialization, especially in structures related to feeding. Although several species were habitat generalists, most species occurred in distinct habitats. Within habitats, microhabitat overlap among co-occuring species was low, particularly in relation to position in the water column. Principal component analysis of the microhabitat measurements produced three new variables. Most species not clearly segregated by the overlap analysis showed segregation on the new variables. Fishes not segregated by habitat or microhabitat tended to show low dietary overlaps. Specialization in feeding habits was more pronounced among the Sri Lankan fishes than noted for fishes in small rainforest streams elsewhere, in part because of the greater reliance of the Sri Lanka fishes on autochthonous foods. Overall, the fish assemblage had the characteristics expected of an equilibrium (deterministic) assemblage.