Small-scale association measures in epibenthic communities as a clue for allelochemical interactions

The small-scale associations in a rocky subtidal community in the northwestern Mediterranean were studied by a development of the continuous line transect method. This method allowed the overall measurement of non-randomness in interspecific contacts and the assignment of an association index to each species-pair, whose, significance was tested by Monte Carlo procedures. At the same time, the continuous recording allowed the study of the weakening of the interactions with increasing distances. Our purpose was to uncover evidence for allelochemical mechanisms of space occupation and maintenance. A strong non-randomness was found in the interspecific associations. This was mostly due to the interactions of the poecilosclerid sponge Crambe crambe (Schmidt) with its neighbours, especially its negative associations with other sponge species. The strength of the relationships fell drastically over the first few centimeters from the contact borders of the different species. The results pointed strongly to an allelochemical mechanism. The extracts of this sponge featured high bioactivity in laboratory assays, and field experiments demonstrated that the sponge can inhibit the growth of species in the community studied. Standard sampling techniques would have overlooked the spatial structure present in the data. The study emphasizes the need for both contact data and distance data in order to identify the underlying processes reliably. The line transect method provides both types of information easily and allows testing of models and identification of organisms likely to use chemical defenses in space competition. Its use as a preliminary step in studies of chemical ecology might help to detect presumptive allelochemical processes prior to experimental work on the potentially active species.     

Functional morphology of genitalia of four species of julidan millipedes (Diplopoda: Nemasomatidae; Julidae)

The functional anatomy of the genitalia of Nemasoma varicorne (Nemasomatidae), Brachyiulus lusitanus, Unciger foehdus and Cylindroiulus boleti (Julidae) was investigated by shock freezing of animals in copula followed by serial semithin sectioning of the engaged genitalia. The species show conspicuous differences in the functional mechanism of their genitalia. In M varicorne and B. lusitanus the vulvae remain within the vulval sac during copulation while in U.foetidus and C. boleti parts of the gonopods (promerite and mesomerite) form clasper-like structures that pull out the vulvae from the vulval sacs. With the exception of C. boleti all investigated species have a 'central funnel' on the vulva which leads into the receptaculum seminis. The sperm-transferring part of the male gonopods (solenomerite) is introduced into this funnel during copulation. In B. lusitanus and C. boleti a projection of the posterior gonopods (end-projection, brachite) fits into a slit anterior to the openings of the receptacula. The results are discussed with regard to sexual selection theory and a hypothesis is proposed that explains the evolutionary change of millipede genitalia by a combination of female choice and sperm competition phenomena.     

Differential impact of egg predation by Zetzellia mali (Acari: Stigmaeidae) on Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

The differential impact of Zetzellia mali on the phytoseiids Metaseiulus occidentalis and Typhlodromus pyri was studied in the laboratory and by analysis of population from orchard plots that contained either phytoseiid, similar numbers of prey mites and high or low densities of Z. mali. Five hypotheses were evaluated to explain why Z. mali had more impact on M. occidentalis in the field than on T. pyri. Given equal opportunity, Z. mali adult females did not consume more M. occidentalis eggs than T. pyri eggs nor did adult females of either phytoseiid inflict greater mortality on Z. mali eggs or larvae through attack or consumption. There was no difference in the within-tree association of Z. mali adult females with eggs of either phytoseiid species nor were there differences in the way prey mites (all life stages) were spatially partitioned between adult female Z. mali as compared with adults and deutonymphs (combined) of either phytoseiid. The foraging area of adult female Z. mali and the oviposition locations of the two phytoseiids from both field and laboratory data were compared using spatial statistics. Metaseiulus occidentalis laid significantly more eggs in the primary foraging area of adult female Z. mali than T. pyri. This was the only factor identified which may explain the greater impact of Z. mali on M. occidentalis. The impact of these interspecific effects on the persistence of predatory mite guilds and biological control are discussed.     

Asymmetric competition between plant species

Despite extensive interest in the role of plant size in competition, few formal attempts have been made to quantify the magnitude of asymmetric competition, particularly for interactions between members of different species. This paper introduces the concept of asymmetric interspecific competition at the population livel (i.e. mean plant performance) in mixtures of species. It proposes an index of interspecific competitive asymmetry which allows for a progressively greater asymmetric effect as the average size differences between competing species increase, and allows for such an effect whether individuals of focal species are larger or smaller, on average, than competitors. This index of competitive asymmetry is evaluated in the study of interactions between two widely coexisting annuals of disturbed habitats, Stellaria media and Poa annua. An experiment was conducted in which the density, relative frequency and relative seedling sizes (emergence times) of Poa and Stellaria individuals were varied. The relative growth rate (RGR) for both species was measured over a 22-day period. An inverse linear model was fitted for each species, relating the RGR of the focal species to the initial biomass of each species. Each response model included an asymmetry coefficient () to assess whether the impact of a unit of initial biomass of the associate species changed with the relative sizes of seedlings of the two species. A zero value of implies symmetric competition between the two populations; i.e. the competitive effect of a unit of associate species biomass does not change with its initial seedling size. If is positive the smaller the initial relative size of seedlings of the associate species, the smaller their per unit biomass effect on the response of the focal species. The model fitted our data for Stellaria and Poa well and was validated by an alternative modelling approach. Asymmetry coefficients were estimated as 0.508 (P<0.05) for the effect of Poa in the Stellaria model, and 0.0001 (NS) for the effect of Stellaria in the Poa model; i.e. the effect of Poa on Stellaria was asymmetric while the effect of Stellaria on Poa was symmetric. Differences in interspecific species asymmetric competitive effects are discussed within the context of shoot architecture, and the relative importance of competition for light versus soil resources. Finally, we discuss the relationship of this model to earlier models of competitive asymmetry, and consider the implications of interspecific competitive asymmetry for a number of current theories of plant competition and community organisation.     

Influence of drought on competition between selected Rhizobium meliloti strains and naturalized soil rhizobia in alfalfa

Drought is an important environmental factor that can affect rhizobial competition and N₂ fixation. Three alfalfa (Medicago sativa L. and M. falcata L.) accessions were grown in pots containing soil from an irrigated (Soil 1) and a dryland (Soil 2) alfalfa field in northern Utah, USA. Mutants of three strains of Rhizobium meliloti Dang. from Pakistan (UL 136, UL 210, and UL 222) and a commercial rhizobial strain 102F51a were developed with various levels of resistance to streptomycin. Seeds inoculated with these individual streptomycin-resistant mutants were sown in the two soils containing naturalized rhizobial populations. Soils in the pots were maintained at −0.03, −0.5, and −1.0 MPa. After 10 weeks, plants were harvested and nodule isolates were cultured on agar medium with and without streptomycin to determine nodule occupancy (proportion of the nodules occupied by introduced rhizobial strains). Number of nodules, nodule occupancy, total plant dry weight, and shoot N were higher for Soil 1 than Soil 2. Number of nodules, plant dry weight, and shoot N decreased as drought increased from −0.03 to −1.0 MPa in the three alfalfa accessions. Rhizobial strains UL 136 and UL 222 were competitive with naturalized alfalfa rhizobia and were effective at symbiotic N₂ fixation under drought. These results suggest that nodulation, growth, and N₂ fixation in alfalfa can be improved by inoculation with competitive and drought-tolerant rhizobia and may be one economically feasible way to increase alfalfa production in water-limited environments. Joint contribution from USDA-ARS and the Utah Agric. Exp. Sta., Utah State Univ., Logan, UT 84322-4810, USA. Journal Paper No. 4931. Joint contribution from USDA-ARS and the Utah Agric. Exp. Sta., Utah State Univ., Logan, UT 84322-4810, USA. Journal Paper No. 4931.     

The effect of genome and sex on recombination rates in Pennisetum species

The effects of homoeology and sex on recombination frequency were studied in crosses between cultivated pearl millet, Pennisetum glaucum, and two wild subspecies, P. violaceum and P. mollissimum. For the two wild x cultivated crosses, reciprocal three-way crosses were made between the F₁ hybrid and an inbred line (Tift 23DB₁). The three-way cross populations were mapped to produce a female map of each wide cross (where the F₁ was the female) and a male map (where the F₁ was the male). Total genetic map lengths of the two inter-subspecies crosses were broadly similar and around 85 % of a comparable intervarietal map. In the P. glaucumxP. mollissimum crosses, the map was further shortened by a large (40 cM) inversion in linkage group 1. Comparison of the recovered recombinants from male and female meiocytes showed an overall trend for the genetic maps to be longer in the male (10%) in both inter-subspecific crosses; however, analysis of individual linkage intervals showed no significant differences. Gametophytic selection was prevalent, and sometimes extreme, for example 121 in favour of wild alleles in the P. glaucumxP. mollissimum male recombinant population. One of the loci which determines panicle type in cultivated pearl millet and wild relatives, H, was mapped 9 cM from Xpsm812 on linkage group 7 in the P. violaceum cross.     

Sperm competition and sex allocation in simultaneous hermaphrodites

Summary Sex allocation theory is developed for hermaphrodites having frequent copulations and long-term sperm storage. Provided the sperm displacement mechanisms are similar to those known in insects, the ESS allocation to sperm versus eggs satisfies a rather simple rule. There are no data to test this rule, as yet.     

Vertical bars on male Xiphophorus multilineatus: a signal that deters rival males and attracts females

We examined the function of the vertical bar pattern on maleswordtails (Xiphophorus multilinneatus) as a signal in bothmale-male competition and female choice. This pattern had previouslybeen described as an aggressive signal because males intensifiedthe bars during male-male encounters in the laboratory. Ourfield observations supported this observation and also showedthat bars intensified when males courted females. The intensityof bars was correlated with access to females in the field.Within the size range of males that have bars, however, neitherbar number nor male size appeared to influence access to females.We used freeze-branding to remove the bars from males in thelaboratory so that we could control for characters correlatedwith bar intensity, and tested males and females separatelyso that we could separate the influence of these two componentsof sexual selection. We compared the responses of males andfemales to males that had their bars removed and control malesfreeze-branded between the bars. Test males responded more aggressivelyto males without bars as compared to control males. In addition,females showed a preference for control males over males thathad their bars removed. These results suggest that the barsmay function as a signal that deters rival males and attractsfemales.     

Dominance, competition, and energetic reserves in the European starling, Sturnus vulgaris

We investigated the relationships between social dominance,competition for food, and strategies of body mass and fat regulationin the European starling (Sturnus vulgaris). In birds housedin groups of three, subdominant birds stored more fat than dominants.A removal experiment established a causal link between socialdominance and fat reserves; in groups that had the dominantindividual removed, the remaining birds reduced body mass andfat, relative to control groups that had the subordinate removed.In a second experiment, we investigated the influences of degreeof competition for food and dominance on body mass and fat reserves.Birds under high competition increased fat reserves and tendedto have higher body mass than birds under low competition. Theincrease in fat reserves was higher in the subdominants thanin the dominants. These results are consistent with hypothesesconcerning dominance-dependent access to food; subdominant birds,or birds under increased competition, may store more fat asan insurance against periods when food cannot be obtained. However,relations between dominance, body mass, and fat reserves mayalso arise through other proximate factors relating to dominance-dependentcosts and benefits of fat storage, such as predation risk andenergetic expenditure.     

Parental effort of kestrels (Falco tinnunculus) in nest defense: effects of laying time, brood size, and varying survival prospects of offspring

We studied the nest defense behavior of Eurasian kestrels (Falcotinnunculus) towards a stuffed pine marten (Martes martes) througha 3-year vole cycle (1990–92) in western Finland. Survivalprobability of offspring decreases with a later start of breeding,and, therefore, early breeders should protect their offspringmore than late ones. We found this true for males during theincubation period, but not for females. In addition, we expectedthe nest defense intensity to increase with offspring number.During the incubation period, this was true for females, butnot for males. During the nestling phase, parents did not adjusttheir defense effort to natural or manipulated (by one to twoyoung) brood size. Survival prospects of kestrel offspring werehighest in the increasing vole year 1991 and lowest in the decreasingvole year 1992, and, therefore, we expected the defense activityof kestrels to follow the same trend. However, the oppositeresult appeared true for females with a similar tendency formales. Most hypotheses predicting avian nest defense behaviorwere not supported by our data. Temporally heterogeneous environmentand low degree of nest-site tenacity of migratory kestrels maymake them unfamiliar with environmental variation and survivalprospects of their offspring. Therefore, fitness benefits ofparental care are not predictable, and kestrels may thus adjusttheir parental effort to their own future reproductive potential(i.e., number of future breeding attempts), rather than to somecurrent investment indicator, like offspring age and number.