Populational distribution I: The digital simulation

This paper presents a method of investigating the distributional pattern of a biological population, using a technique of simulation. The method consists of the comparison of the empirical frequency curve of a population, obtained using the Method of Quadrats, with a simulated one, since the pattern of the simulated curve depends on the simulated distribution. An example is given.     

Suspension-feeding by herbivorous calanoid copepods: A cinematographic study

Many planktonic calanoid copepods are commonly described as filter-feeders. Direct observations using high-speed micro-cinematography indicate that these animals are suspension-feeders. They capture and handle the food particles not passively according to size and shape but, in most cases, actively using sensory inputs for detection, motivation to capture, and ingestion.     

Male capacity as related to sperm production,pregnancy initiation,and sperm competition in deer mice (Peromyscus maniculatus)

Summary Limitations on male capacity may be important to the evolution of mating strategies and behavior. Sperm counts in successive ejaculates of deer mice (Peromyscus maniculatus) decreased progressively through six ejaculations and remained depressed the following day. Males were as effective in inseminating females in cycling estrus with their second two ejaculates as with their first two and as effective in inseminating postpartum females with their fifth as with their first. In sperm competition situations males competed with rested males as effectively with their fourth, fifth, and sixth ejaculates as with their first. Thus, although sperm counts decrease in successive ejaculates, males suffer no apparent functional deficiency as long as they continue ejaculating. It is not clear whether or not deer mice are honest salesmen but they appear to be honest ejaculators and usually are honest intromitters.     

Exosmosis and “free space” in marine benthic algae

In distilled water, marine benthic algae immediately lose ions. This can be demonstrated by means of chloride titration or conductivity measurements. The rapidity and the complete reversibility of this process show that ion movements out of and into the free space are involved. When Laminaria saccharina thalli, exposed to a series of increasing NaCl-concentrations, are subsequently transferred into distilled water, the external concentration increases proportionally to rate of ion loss. Through its free space, the alga establishes an ionic equilibrium with its external medium. If the alga thallus is killed (20 sec boiling in isosmotic sea water), the extent of chloride loss is much higher. Because of thermal destruction of the osmotic space, the chloride can then escape from the entirc thallus. Since sublittoral algae die upon drying, chloride loss from dry thalli is much higher than that from wet thalli: this difference is small in littoral algae, which tolerate short periods of dryness. A close relationship exists between extent of chloride loss and degree of resistance to drying.     

Siblicide,parent-offspring conflict,and unequal parental investment by egrets and herons

Summary O'Connor (1978) generated behavioral predictions from Trivers's (1974) theory of parent-offspring conflict (POC) in relation to avian brood reduction, especially for species using escalated sibling aggression. In general, when daily mortality rates fall between certain values (the mortality difference thresholds for parents and surviving offspring: O'Connor 1978), POC is expected. At such times parents and the early-hatching senior offspring are expected to disagree on the necessity, or at least timing, of the victim chick's death. Because that death is promoted directly by the senior chicks and because parents have realistic means of nullifying those efforts, behavioral conflict is expected. I examined the activities of all family members in one siblicidal (great egrets, Casmerodius albus) and one nonsiblicidal brood-reducing species (great blue herons, Ardea herodias) of ardeidae in Texas nesting colonies. Daily mortality rates over two breeding seasons approximated the predicted POC zone for four-chick (B/4) heron broods and three-chick (B/3) egret broods. As predicted, egret sibling aggression was significantly lower in two-chick broods. Herons seldom fought, regardless of brood size. Special attention was paid to the egret B/3 data for behavioral manifestations of POC. Parents did not interfere overtly with sibling fights (99.2% of 2,829), but parents seemed to have a subtle mollifying effect on the aggression (fewer sever fights occurred when parents were present). Parents showed no favoritism in food distribution to dying victim chicks. The mixed results are consistent with two rather different POC interpretations: (1) that significant conflict exists and the offspring are winning or (2) that there is no significant evolutionary conflict. In the latter case, parents are regarded as creating asymmetries among offspring (e.g., via hatch asynchrony), then leaving competition among siblings to run its course without further intervention (laissezfaire policy). This view explicitly recognizes that siblicide and other investment-skewing selfish chick behaviors may serve parental interests well-i.e., that equal allocation of resources is not likely to be the parental optimum. Using the relative reproductive values of each nestling as an index of the parent's unequal parental investment optimum produces a closer fit to the observed food distribution skew, but does not resolve the question of whether conflict remains.     

Primary production in Dumbell Bay in the Arctic Ocean

Photosynthesis, chlorophyll, inorganic nutrients, and related factors were measured throughout a productive season in a small coastal inlet of the Arctic Ocean. Significant production was confined to 5 m for a very limited time period beacause of both light and nutrient limitations. Maximum chlorophyll was 8.2 mg Chl a m^-3 and maximum gross and net photosynthesis rates were 830 and 550 mg C m^-2 d^-1. Annual gross and net photosynthesis is estimated at about 12 and 9 g C m^-2 respectively. The effect of light stimulation on assimilation is described, sources of coastal zone nutrient enrichment are considered, and discrepancies between gross and net photosynthesis are discussed with reference to the growth characteristics of the phytoplankton population.Bigelow Laboratory for Ocean Sciences. Contribution No. 76008     

Alternative mating tactics and the law of Diminishing Returns: The Satellite Threshold Model

Summary The mating systems of many species comprise at least 2 alternative male mating tactics — broadly termed Dominant and Satellite tactics here. But, there exists no satisfactory conceptual framework in which to explain their co-occurrence and persistence.A Dominant male's ability to copulate is a positive, but decelerating, function of either his territory's attractiveness or his own attractiveness to females. This Diminishing Returns Effect affords mating opportunities for Satellite males associated with the most attractive sites (or males). A graphical Satellite Threshold Model, analogous to the Polygyny Threshold Model (Orians 1969), is developed to predict quantitatively the presence or absence of Satellite male tactics.Five predictions of the Model are supported quantitatively by data from dragonfly and anuran social systems and, more generally, by evidence from a wide variety of species.The Model should apply both to circumstances in which males are able to alternate between tactics and the instances in which males make an irrevocable tactical decision.     

Sorted samples and quantitative counts in appendicularian catches

Sorting centers are important for processing material collected during extensive oceanic expeditions. Based on information obtained from samples of appendicularians, this paper demonstrates that usual procedures adopted in sorting centers produce samples which contain only a fraction of the organisms present in the original catch (or subsample). In addition, the proportion of individuals sorted varies considerably. In four Indian Ocean Standard Net catches collected in the Arabian Sea, subsamples (called sorted samples in this paper) are compared with careful microscopical counts (quantitative counts) performed by the author. The numbers of appendicularians found in quantitative counts were 2.4 to 19.2 times higher than those in the sorted samples. Sorting efficiency varies in different forms of appendicularians; it is particularly low in Fritillaria sp. Bad preservation and small body size further reduce the portion sorted out. Under certain conditions (e.g. net clogging), numerous small appendicularians are retained which normally would pass easily through the meshes of the plankton net.     

Temporal variation and distribution of trace metals in freshwater and fish from Calabar River, S.E. Nigeria

An investigation on the abundance and distribution of trace metals (Fe, Cu, Zn, Mn, Cr, Cd and Pb) in water, and nine species of fish samples from Calabar river was carried out in 1992. The concentrations of iron (6000–7240gl^–1), zinc (4910–7230gl^–1), and cadmium (3–7gl^–1) showed moderate pollution while those of copper (420–630gl^–1), manganese (23–48gl^–1), chromium (<10–20gl^–1) and lead (<1–10gl^–1) in water were well below WHO permissible levels. Significant seasonal changes (0.001p0.25) were obtained for iron, copper, zinc, manganese and cadmium in water. Furthermore, iron, zinc and cadmium showed statistically significant spatial changes (0.005p0.10). Of the nine fish species studied, no statistically significant relationship between body weight and the concentrations of the metals was observed. The concentrations of the metals per mean total body weight apparently decreases in the order Fe>Zn>Cu>Mn>Pb>Cd=Cr and were within the limits that were safe for consumption.     

Groups confuse predators by exploiting perceptual bottlenecks: a connectionist model of the confusion effect

Aggregation is a well documented behaviour in a number of animal groups. The confusion effect is one mechanism thought to mitigate the success of predators feeding on gregarious prey and hence favour aggregation. An artificial neural network model of prey targeting is developed to explore the advantages prey species might derive through a tendency to group. The network illustrates how an abstract model of the computational mechanisms mediating the perception of prey position is able to show a degradation in performance as group size increases. The relationship between group size and predator confusion has a characteristic decreasing decelerating shape. Prey oddity is shown to reduce the impact of the confusion effect, thereby allowing predators to target prey more accurately. Hence shoaling behaviour is most profitable to the prey when prey phenotypes are visually indistinguishable to a predator. Futhermore it is shown that prey oddity is relatively more costly in large groups than in small groups and the implications for assortative schooling are discussed. Both the model and the results are intended to make the general point that cognitive constraints will limit the information that a nervous system can process at a number of different levels of neural organization.     

The zooplankton of the inshore waters of Dar es Salaam (Tanzania,S. E. Africa) with observations on reactions to artificial light

From 18th December, 1968 to 5th January, 1970, zooplankton samples were taken in darkness and using artificial light in selected areas of the Dar es Salaam coast (Tanzania, S. E. Africa). Surface sea temperature was measured on most occasions, salinity for the first 7 months only. The neritic waters of Dar es Salaam experience a warm period during January to March and a cool period during July to September. The annual salinity cycle is not known. Thirtysix dark and 17 light zooplankton samples were analysed; where possible, organisms were identified to species, others to generic or higher taxonomic level. The principal taxa and their mean percentage proportions (figures in parentheses) in the dark samples were: Calanoida (49. 1), Larvacea (11. 9), Corycaeus spp. (6.4), Cypridina sinuosa (5.6), Oithona spp. (4.8), caridean larvae (4.0), Sagitta spp. (3.8), Euterpina (2.1), Lucifer (1.2), Oncea (1.2), calyptopis larvae (1.0), Hydromedusae (1.0), Euconchoecia chierchiae (1.0), Creseis acicula (1.0), brachyuran zoeae (0.8), Ctenophora (0.5), Mysidacea (0.5), fish eggs (0.5), postlarval bivalves (0.5), postlarval gastropods (0.5), Cumacea (0.1), Gammaridea (0.1) and Hyperiidea (0.1). Evadne tergestina and Thalia democratica abound in waters at certain times of the year only and then virtually disappear. The remaining groups were numerically unimportant most of the time. Almost every major group showed an annual cycle of abundance; greater numbers were recorded either for the entire or part of the period February to August, compared to the period September to November. From the end of January to early August, 1969, the average numbers per haul of total zooplankton were about three times greater than for the period mid-August to mid-November. During February-March and July-August, several oceanic indicators were observed together in the neritic waters. Artificial light induced the following changes in night zooplankton: Mysidacea, Leptochela sp., Hyperiidea, Cypridina sinuosa, brachyuran megalopae and fish larvae were attracted towards artificial light and aggregated densely under the lamp; Calanoida, Corycaeus spp., Macrosetella, Microsetella and Euterpina avoided regions of strong illumination and aggregated in dim-lit areas; Lucifer, Creseis acicula and postlarval gastropods were more abundant in dim-lit samples compared to dark hauls; caridean larvae, brachyuran zoeae and larvaceans were less abundant in high-intensity light samples compared to dark hauls on the same nights.This work was performed at the Department of Zoology, University of Dar es Salaam, Tanzania, S.E. Africa.     

Age-structure dependence of the “specific production” in zooplankton populations

This account represents the results of investigations on the production of some zooplankton populations. For comparing different populations the index specific production (the production rate of the unit of the biomass) is used. The correlation between specific production and age-structure for some populations is described. On the basis of the growth theory of Von Bertalanffy (1938) a theoretical model which helps to explain the relation between specific production and age-structure of the population is proposed.     

Aqua regia extractable trace elements in surface soils of venezuela

Surface soils (0–15 cm) were sampled at 10–20 km intervals along two transects in Venezuela. One (1162 km, 70 samples) ran west to east parallel with the Caribbean coastline, the other (920 km, 92 samples) ran south to north from the frontier with Brazil to the Caribbean shore. Sampling took place in both a wet and a dry season. Trace metals were extracted from dried, sieved (<2 mm) soil with boiling aqua regia followed by analysis by ICP or flame AAS. Metal values did not differ significantly between the two seasons and dates were averaged. Geometric mean values for the west–east transect were: Cr=41.5, Cu 17.9, Cs 3.6, Li=13.9, Mn=294, Ni=21.3, Pb=17.4, Sr=39.4, V=60.4 and Zn = 83.7g g^–1, respectively. Similarly, for the south–north transect Cr=21.3, Cu=4.3, Cs=1.1, Li=2.0, Mn=55.7, Ni=4.4, Pb=6.1, Sr=13.3, V=28.2 and Zn=16.7g g^–1, respectively. A classification of samples by lithology showed surface soil composition to be related to rock composition. Metal values were low in the soils in the south of the country, in the Guyana highlands (Gran Sabana). Low Zn contents were prevalent. Lead contents were affected by roadside fallout from vehicles using leaded petrol except that high Pb contents of soils in the Gran Sabana were of more complex origin.     

Hypo-osmotic regulation coupled with reduced metabolic urea in the dogfish Poroderma africanum: An analysis of serum osmolarity,chloride, and urea

Pyjama sharks (Poroderma africanum) were exposed to a wide range of salinities, over which blood serum was analysed for osmolarity, chloride and urea concentrations. Fish were divided into two groups, those fed twice weekly (high intake), and those fed once a month (low intake). Both groups were exposed to the same salinity range. High intake fish showed the characteristic elasmobranch osmolarity picture, with serum values slightly hyper-osmotic at all times. Low intake fish, however, showed a degree of hypo-osmotic regulation. Serum values for both groups overlapped at very low salinities. Serum urea was also affected by diet, so that again two distinct sets of values were produced, again with overlap at the lower salinities. When previously well-fed fish were starved over a period of one month, serum urea and osmolarity decreased simultaneously. Consequently, it is felt that serum osmolarity is directly related to serum urea levels. Serum chloride was not found to be affected by diet, both groups showing the same change in blood values when exposed to the same change in salinity. It is shown, however, that a reduction in food intake, over a period of more than a fortnight, can reduce metabolic urea to the extent of depressing serum osmolarity and, hence, shift the ionic and osmotic equilibrium between the fish and the sea water. This may result in varying degrees of hypoosmotic regulation.     

Allozyme and colour differences between two sibling species of the heteronemertean Lineus torquatus from the Sea of Japan

Brown and cherry colour morphs of the heteronemertean Lineus torquatus Coe, 1901 from the Vostok Bay of the Sea of Japan were compared using allele frequencies at 18 isozyme loci. These morphs shared no common alleles at 9 of the 18 loci. The observed genetic differences are strong evidence that the brown and cherry colour morphs, which occur sympatrically, are different species. Nei's genetic distance between these species (D=1.135) is considerably higher than typical D values for congeneric animal species. As with much previous work on nemerteans, intraspecific allozymic variability in both species was high, with mean expected heterozygosity values 0.286±0.049 and 0.159±0.051 for L. torquatus (brown) and L. torquatus (cherry) species, respectively.     

The effects of operational sex ratio and food deprivation on copulation duration in the water strider (Gerris remigis Say)

Summary A North American water strider, Gerris remigis (Hemiptera) copulates for prolonged periods of time, apparently beyond the time necessary for insemination (Rubenstein 1984; Wilcox 1984; Clark 1987), I tested predictions from two different hypotheses of the adaptive significance of this behavior. Manipulations of the operational sex ratio in artificial streams revealed that water striders copulate significantly longer when the sex ratio is male-biased. This result is consistent with the hypothesis that prolonged copulation functions as a type of contact mate guarding, which reduces sperm competition by preventing the female from remating prior to oviposition. The finding is also consistent with the hypothesis that copulation is prolonged to enhance female foraging efficiency. In a second experiment, sex ratio and total population density were held constant, and the period of food deprivation of females was manipulated. Contrary to the predictions of the female foraging hypothesis, starved females did not copulate longer than fed females. Of the two hypotheses considered, only the copulatory guarding hypothesis explains both results. This does not mean, however, that there is no selective benefit from the enhanced foraging efficiency resulting from prolonged copulation. Nor does it exclude the possibility that this benefit has been necessary for the evolution of prolonged copulation in water striders.     

Metabolic diversity in epibiotic microflora associated with the Pompeii wormsAlvinella pompejana andA. caudata (Polychaetae: Annelida) from deep-sea hydrothermal vents

Specimens of alvinellid polychaetes (Alvinella pompejana Desbruyères and Laubier, 1980 andA. caudata Desbruyères and Laubier, 1986) and their tubes were sampled from deep-sea hydrothermal vents at 13°N from the manned submersible Nautile during the Hydronaut cruise (October to November 1987) on the East Pacific Rise. Samples were subjected to bacterial analysis aboard the mother ship Nadir to detect bacteria involved in the nitrogen and sulphur cycles, in non-specific heterotrophic processes, and displaying resistance to selected heavy metals. Cultures were incubated at different temperatures under atmospheric and in situ (250 atm) pressures. Bacterial growth was observed in enrichment cultures for most metabolic types screened. Heavy-metal-resistant bacteria were also detected in many samples. No filamentous bacterial form was observed in the cultures. The results demonstrate a high metabolic diversity in episymbiotic flora, indicating that the worm (A. pompejana orA. caudata), its tube and its epiflora represent a complex micro-ecosystem.     

Growth of the Atlantic surfclam,Spisula solidissima,from Georges Bank to the Delmarva Peninsula,USA

Age/shell length data for offshore surfclam, Spisula solidissima (Dillwyn, 1817), populations were used to estimate the parameters of the von Bertalanffy growth model by time period and region, from Georges Bank in the north to the Delmarva Peninsula in the south. Randomization tests were used to compare curves. We tested the a priori hypothesis that growth curves would change over time in the south (i.e., New Jersey and Delmarva) but remain constant in the north (i.e., Long Island and South New England). This hypothesis was proposed because surfclam population structure in the south had been altered by the hypoxic event of 1976, and possibly by intense, long-term commercial harvesting. Northern regions, unaffected by these factors, served as natural controls. Based on a comparison of data collected in 1980 with pooled data from 1989 and 1992, the hypothesis was supported. Both the growth coefficient (k) and maximum shell length (L ) declined between two time periods in the two southern regions, while during the same time interval, no change occurred in the two northern regions. Differences in growth between regions were often statistically significant. For example, compared with the southern regions, the growth coefficient on Georges Bank was larger, and those clams attained a smaller maximum length. In a comparison of adjacent regions from Delmarva to S. New England, k increased from south to north. This could imply faster growth in cooler water, as well as no relationship between growth and primary productivity. Alternatively, size-selective mortality, imposed by the commercial fishery, was discussed as a mechanism that might account for this unexpected pattern.     

Night irradiance and synchronization of lunar release of planula larvae in the reef coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus), which is known to release planula larvae on a monthly cycle, was grown in full daytime solar irradiance, but with four treatments of night irradiance: (1) natural night irradiance, (2) shifted-phase (total darkness during nights of full moon with artificial irradiance at lunar intensity on nights of new moon), (3) constant full moon (full lunar irradiance every night), and (4) constant new moon (total darkness every night). The reproductive cycle of the corals held in the shifted-phase treatment moved out of synchrony with the cycle of corals exposed to a natural lunar cycle of night irradiance. Two previously described types of P. damicornis were tested. The Type Y normally start releasing larvae at full moon, with peak production at third quarter. In the shifted-phase treatment they began releasing planulae at new moon (artificial full moon), with peak production at first quarter. The Type B corals, that normally start releasing planulae at new moon with peak production at first quarter, began to release planulae at full moon (artificial new moon), with peak production at third quarter. Populations of corals grown either in the constant full moon or constant new moon treatment quickly lost synchronization of monthly larva production, although production of planulae continued. Thus spawning is synchronized by night irradiance.Contribution No. 702 of the Hawaii Institute of Marine Biology     

Badges of status and the cost of aggression

Summary The badges of status hypothesis suggests that frequency-dependent selection can maintain honest signalling of aggressiveness. Analysis of a simple ESS model reveals that this is only possible if aggressive individuals incur a cost independent of any particular fights they become involved in. Without such a cost, an honest population is vulnerable to invasion by a modest mutant bearing a dishonestly small badge. Analysis of a second model reveals that where individuals differ in their ability to bear the contest-independent cost of aggressiveness, badge size should reflect this ability. Under such circumstances, plumage badges which serve as honest status signals may have functional consequences beyond their role in settling contests. Thus females, when choosing a mate, could make good use of the information that plumage badges provide. Analysis of a third model shows that the existence of contest-independent benefits to badge size, such as increased mating success, is compatible with the maintenance of honest status signalling.Correspondence to: R.A. Johnstone