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.     

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.     

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.     

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.     

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.     

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     

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.     

Dynamics of the TIT FOR TAT strategy during predator inspection in the guppy (Poecilia reticulata)

Summary One well-known solution to the iterated Prisoner's Dilemma is the TIT FOR TAT strategy. This strategy has three characteristics associated with it. TIT FOR TAT is nice (cooperates on the first move of a game), retaliatory (plays defect against an individual that defected on the prior move), and forgiving (cooperates with an individual which has defected in the past but cooperates in the present). Predator inspection behavior in guppies (Poecilia reticulata) was examined in order to determine whether guppies displayed these three characteristics. Results indicate that while it can be quite difficult to translate the abstract concepts of niceness, retaliation, and forgiveness into measurable behaviors, the data support the hypothesis that guppies display the three characteristics associated with the TIT FOR TAT strategy.The selfish herd and dilution effect are discussed as possible alternative explanations for the observed behavior. These hypotheses alone were insufficient to explain the results.     

Compositional topography of melon and spermaceti organ lipids in the pygmy sperm whale Kogia breviceps: Implications for echolocation

The forehead of the pygmy sperm whale Kogia breviceps contains a large melon of fatty tissue in front of a small, fat-filled, cornucopia-shaped spermaceti organ. This unique anatomical structure may possibly play an acoustical role in the whale's echolocation system, similar to the fatty melon sound lens postulated for dolphins. To better understand its function, we have studied the compositional topography of the K. breviceps melon and spermaceti organ lipids. The fatty head tissues of an adult K. breviceps were serially sectioned into 9 transverse slices. Appropriate tissue samples were cut from every other slice, and analyzed for % lipid and lipid class composition. Wax esters and triglycerides were the only major lipids present; their average carbon number in each sample was determined by gas-liquid chromatography (GLC). Our topographical analyses of K. breviceps melon indicate 3 regions of distinctive lipid composition: a fat-poor melon exterior, an outer melon of medium fat content having % triglyceride>% wax ester, and a fat-rich inner melon having % wax ester>% triclyceride. The spermaceti organ contains a fat-rich core of very high wax-ester content (84 to 99%), surrounded by a fat-poor case. Average carbon numbers of both wax esters and triglycerides were lowest in the inner melon and the spermaceti organ. At the rear of the spermaceti organ lies the museau de singe, an apparent sound generator. The lipid topography data plus anatomical and acoustical considerations suggest that the K. breviceps melon/spermaceti organ system may function as an acoustical transducer, directing and refracting sound waves from this source for the purpose of echolocation.     

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     

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     

Male quality,reproduction, and survival in the great tit (Parus major)

Summary Three possible measures of male quality (social dominance, song, and size), reproduction, and survival were studied in a single population of great tits. Winter dominance position on a feeder was related to strophe length (number of phrases per strophe), inversely related to positive drift (decrease of the singing rate of the phrases in a strophe), but not related to song repertoire size. Neither winter dominance position nor song were related to size (wing length, tarso-metatarsus length, weight).Singing capacity was not correlated with individual reproductive success in a single breeding season, using a rather limited data set. However, better singers (males which sing longer strophes, show less positive drift, and have larger song repertoires) survived better and had a higher individual lifetime reproductive success (on the basis of a male's recruited offspring of all breeding seasons). Our results show that there exist measurable differences whereby birds that are dominant in winter sing better, survive longer, and produce more surviving offspring during their life time. We suggest that differences in male quality are the common cause (direct and indirect) of all these effects.     

Vigilance and foraging substrate: anti-predatory considerations in a non-standard environment

Summary The commonly studied standard anti-predatory environment presents animals with spatially-distinct feeding sites and refuges from attack, neither of which necessarily obstructs predator detection. In contrast, tree-trunks provide animals with a markedly non-standard environment in which the foraging substrate itself may be a refuge from attack that unavoidably obstructs predator detection. Thus anti-predatory behavior in this environment should be influenced not only by a perceived risk of attack, but also by the nature of the refuge/foraging substrate itself. Downy woodpeckers (Picoides pubescens) are a common tree-trunk foraging animal, and an experimental analysis of their behavior suggests that they respond appropriately to their non-standard anti-predatory environment. In particular, anti-predatory vigilance varies strongly with changes in tree trunk diameter. Two modes of vigilance were apparent. In stationary vigilance, woodpeckers maintained the position of their feet while rotating their bodies side-to-side to peer around the trunk; mobile vigilance involved movement around the trunk itself. Both the frequency and angle of rotation of stationary vigilance increased with trunk diameter, as did the frequency of mobile vigilance. The woodpeckers also held their heads farther away from the trunk surface as diameter increased. All of these measures of vigilance increased under a greater perceived risk of predation. As might be expected given these results, downy woodpeckers avoided thick trunks; they did not, however, prefer the thinnest (least obstructive) available trunks. These preferences may reflect the influence of trunk diameter on thermo-ecological and/or anti-predator considerations not related to vigilance. Overall, this arboreal environment provides an unusual perspective on anti-predator decision-making with several implications for tree-trunk foraging animals in general.     

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.     

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.     

“Free space-photosynthesis” in the algae Fucus virsoides and Laminaria saccharina

This paper deals with the role of bicarbonates in the apparent free space (AFS) and their importance for the photosynthetic process. Both the littoral alga Fucus virsoides (Mediterranean Sea) and the sublittoral alga Laminaria saccharina (Baltic Sea) show the same response. Unloading of Clions and subsequent reloading of HCO₃-ions result in a positive photosynthetic rate in distilled water (free of CO₂). Time of exposure of an algal thallus to bicarbonate solutions, as well as the concentration of these solutions, influences AFS-assimilation of the ions.     

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.     

“Copper” granules in the barnacle Balanus balanoides

Balanus balanoides (L.) collected from an area with high heavy-metal run-off contained two different types of granule within the parenchyma cells of the prosoma. X-ray microprobe analysis shows one to be the familiar zinc granule made up of concentric layers and giving major peaks for phosphorus and zinc, and the other to be homogeneous and giving peaks for sulphur and copper. This latter granule is designated the copper granule. Whilst zinc granules are known to be composed of inorganic phosphate, various tests on copper granules in sections and in a granule-rich pellet have shown that the copper is probably complexed with organic matter. The tests also demonstrated the relatively, insoluble (inert) nature of these granules. Although zinc and copper granules were present together in the prosoma, atomic absorption analyses of whole bodies (prosoma+thorax) have shown the level of zinc (50.28 g/mg dry weight) to be much higher than that of copper (3.75 g/mg dry weight).     

Upwelling-generated plankton strandlines: important predictable food sources for seabirds at Husvik,South Georgia

Sustained strong offshore winds generate upwelling in the fjords of northern South Georgia. Deepwater plankton is pumped by upwelling towards sandy beaches at the heads of the fjords; calanoid copepods, chaetognaths and pteropods are stranded by the sand filter in large quantities on falling tides. Local benthonic species (harpacticoid copepods and amphipods) are present in the plankton but do not strand. Strandlines are very rich (<4 kg wet mass m^-2) and provide large fractions of the food of kelp gulls, sheathbills and terns in the austral summer. Birds can only exploit fresh strandlines, since air-drying of the plankton soon makes it too salty to eat.     

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.