13C/12C ratios and the trophic importance of algae in Florida Syringodium filiforme seagrass meadows

Over 380 stable carbon isotope (¹³C) analyses made during 1981–82 showed that Syringodium filiforme Kutz seagrass meadows in the Indian River lagoon of eastern Florida have food webs based on algal rather than seagrass carbon. Seagrasses averaging approximately-8 were isotopically distinct from algae epiphytic on seagrass blades (X=-19.3) and particulate organic matter in the water column X=-21.6. ¹³C values of most fauna ranged between-16 and-22, as would be expected if food web carbon were derived solely from algal sources. These results counter the idea that seagrass detritus is the dominant carbon source in seagrass ecosystems. Two factors that may contribute to the low apparent importance of seagrass in the study area are high algal productivities that equal or exceed S. filiforme productivity and the high rates of seagrass leaf export from meadows.     

Optimal foraging: Some simple stochastic models

Summary Some simple stochastic models of optimal foraging are considered. Firstly, mathematical renewal theory is used to make a general model of the combined processes of search, encounter, capture and handling. In the case where patches or prey items are encountered according to a Poisson process the limiting probability distribution of energy gain is found. This distribution is found to be normal and its mean and variance are specified. This result supports the use of Holling's disc equation to specify the rate of energy intake in foraging models. Secondly, a model based on minimization of the probability of death due to an energetic shortfall is presented. The model gives a graphical solution to the problem of optimal choices when mean and variance are related. Thirdly, a worked example using these results is presented. This example suggests that there may be natural relationships between mean and variance which make solutions to the problems of energy maximization and minimization of the probability of starvation similar. Finally, current trends in stochastic modeling of foraging behavior are critically discussed.     

On the integration of individual foraging strategies with colony ergonomics in social insects: nectar-collection in honeybees

Summary We experimentally tested whether foraging strategies of nectar-collecting workers of the honeybee (Apis mellifera) vary with colony state. In particular, we tested the prediction that bees from small, fast growing colonies should adopt higher workloads than those from large, mature colonies. Queenright small colonies were set up by assembling 10 000 worker bees with approximately 4100 brood cells. Queenright large colonies contained 35 000 bees and some 14 500 brood cells. Thus, treatments differed in colony size but not in worker/brood ratios. Differences in workload were tested in the context of single foraging cycles. Individuals could forage on a patch of artificial flowers offering given quantities and qualities of nectar rewards. Workers of small colonies took significantly less nectar in an average foraging excursion (small: 40.1 ± 1.1 SE flowers; large: 44.8 ± 1.1), but spent significantly more time handling a flower (small: 7.3 ± 0.4 s ; large: 5.8 ± 0.4 s). When the energy budgets for an average foraging trip were calculated, individuals from all colonies showed a behavior close to maximization of net energetic efficiency (i.e., the ratio of net energetic gains to energetic costs). However, bees from small colonies, while incurring only marginally smaller costs, gained less net energy per foraging trip than those from large colonies, primarily as a result of prolonged handling times. The differences between treatments were largest during the initial phases of the experimental period when also colony development was maximally different. Our results are at variance with simple models that assume natural selection to have shaped behavior in a single foraging trip only so as to maximize colony growth. Offprint requests to: P. Schmid-Hempel     

Chitinolytic enzymes in the integument and midgut-gland of the shrimpPalaemon serratus during the moulting cycle

Chitinase andN-acetyl--D-glucosaminidase activity were quantified inPalaemon serratus (Pennant) integument and midgut gland during the moulting cycle. Studies were performed on specimens collected near Concarneau, France, in July 1989. The changes in specific activity are different in the two organs: in the midgut gland enzymatic activity is high throughout the whole moulting cycle with a weak peak at the early premoult Stage D1, whereas in the integument the activity of both enzymes is very low throughout post- and intermoult stages and rises only after premoult Stage D1. The highest specific activity is reached in D1 for chitinase and somewhat earlier (D1) forN-acetyl--D-glucosaminidase. The increase in specific chitinolytic activity coincides with an increase in ecdysteroids.     

Lead-based paint in dwellings: The potential for contamination of the home environment during renovation

The quantity and particle size characteristics of lead in dust released during three different paint removal techniques was determined under controlled conditions and in situ in a dwelling. Air-lead and deposited dust-lead levels were highest after sanding but burning-off and hot-air removal methods also produced significant contamination. The importance of dust particle-size and lead is discussed in relation to the potential hazard to home renovators via inhalation and to children via the hand-to-mouth route.This paper is also being published in Proceedings of the Conference on Lead in the Home Environment and is one of several selected from SEGH sponsored conferences for simultaneous publication in this journal.     

Annual cycles in worker size of the seed-harvester antVeromessor pergandei (Hymenoptera: Formicidae)

Summary The seed-harvester antVeromessor pergandei Mayr is primitively polymorphic; workers are monophasically allometric. There is a distinct annual cycle in mean worker body size that replicates across colonies and habitats (Fig. 1); this cycle occurs through alteration of the worker size distribution (Fig. 2). There is little, if any, morphspecific task specialization by workers suggesting worker size variance is a colony-level adaptation permitting maintenance of a large and constant worker force during periods of resource fluctuation. Smaller workers appear in the foraging force following the triple crunch of reduced seed availability, reduced favorable times to forage, and alate production during winter months. Adult and startingV. pergandei colonies exhibit strong intraspecific territoriality, suggesting the selective advantage for maintenance of a large and constant worker force. Such selective pressures may have provided the initial variance in worker size distributions that led subsequently to specialized castespecific task performance in more distinctly polymorphic ant species.     

Lipid biochemistry and reproductive biology in two species of Gammaridae (Crustacea: Amphipoda)

Gammarus oceanicus Segerstråle, 1947 and Echinogammarus marinus (Leach, 1815) were sampled during the breeding season from Oslofjord in 1984, and their lipid composition examined in relation to reproductive condition. In G. oceanicus, female lipid content increased as the ovary matured. Both the amount of lipid stored and the rate of accumulation were greater in spring than in winter. Spring eggs contained 12.4 g lipid, of which 63% was triacylglycerol and 27% phospholipid. Both fractions decreased steadily during embryonic development. Winter eggs contained 19.2g lipid, of which 52% was triacylglycerol and 43% phospholipid. During the early stages of embryonic development the amount of phospholipid decreased sharply, whereas that of triacylglycerol increased, suggesting that some of the fatty acid released from phospholipid was sequestered temporarily as triacylglycerol. When newly spawned, both winter and spring eggs were richer in monoenoic fatty acids than adult amphipods and these acids were the major fuel consumed during development. 6 fatty acids were utilised more slowly than 3 acids, and egg carotenoid pigment content remained constant. Female E. marinus increased in lipid content as the ovary matured. Spring eggs contained 14.7 g lipid when newly spawned and this increased to 16.6 g during the early stages of development. This increase was entirely triacylglycerol, which declined in later stages; the source of the extra lipid was unclear. Eggs contained very little phospholipid or sterol, and both of these components remained at a steady low level during development. E. marinus eggs were not significantly rich in thonoenoic acids compared with adults, and saturated, monoenoic and polyenoic acids were utilised about equally during development. Both adults and eggs were rich in 20.46, which was utilised at a slower rate than the 3 polyunsaturated acids during embryonic development; again, egg carotenoid pigment content remained constant. In both species there was a decrease in the size of the egg (and as a result, of the newly hatched juvenile), but an increase in total reproductive output (i.e., the total weight of the egg clutch) per female as the breeding season proceeded. The reproductive output of an individual female is probably related to food availability during the period of ovarian maturation, whereas the size of an individual egg is dictated largely by feeding conditions for the juveniles once they are independent of the female. The different patterns of lipid utilisation during development found in this study emphasize the flexibility of response in the reproductive biology of gammarid amphipods. It is not yet possible, however, to relate the differing patterns in a simple way either to egg size or total female reproductive output. Two outstanding problems are the source of extra triacylglycerol during the early stages of development of E. marinus and the metabolic cost of brooding eggs.     

Importance of the fatty acids 20:5ω 3 and 22:6ω 3 in the diet of plaice (Pleuronectes platessa) larvae

The requirement of plaice (Pleuronectes platessa L.) larvae for the fatty acids 20:5 3 and 22:6 3 was studied. Larvae were reared from first feeding to beyond metamorphosis on Artemia sp. nauplii (EG brand), whose nutritional content had been manipulated by enrichment. Some larvae were fed Artemia rich in 20:5 3 and 22:6 3, and others were fed nauplii with no 22:6 3 and low levels of 20:5 3 (1.9% of the total fatty acids). The differences in diet had no significant difference on growth or survival of the plaice larvae. The results indicate that the plaice larvae in this study did not require 22:6 3 in their diet, even when the levels of 20:5 3 were low.     

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.     

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.     

Vibrational communication between hitchhikers and foragers in leaf-cutting ants (Atta cephalotes)

Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Communicated by P. Pamilo     

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.     

Dominance of larval forms in euphausiid (Crustacea:Eucarida) ontogenesis

Variability and dominance of Euphausia superba Dana larval forms (Furcilia-I and Furcilia-II stages) in the plankton of the Scotia and Weddell Seas have been investigated. New forms (1, 2, 3 and 14) have been recorded. The dominant forms of Furcilia-I stage are proposed to represent the initial dominants. In accordance with the initial dominance, five types of larval form combinations have been established. During the initial dominance change — from 5 to 2, the larval form number increases. This phenomenon is due to parallel development of euphauiid larvae; it results from a developmental pathway change in the main larval mass. Such deviations are reflected in an initial dominance change. In each pathway, the larvae pass through an unequal number of moults before reaching Form 5 (in accordance with the proposed moult scheme of Furcilia-I stage larvae). In the case of initial dominance of less-developed forms, the presence of different parallel pathways increases. As a result, the total number of larval forms present in the plankton changes, causing increased variability of the total number of larval forms. Each type of larval form combination is the result of summation of all larval forms, which derive from different developmental pathways of the larvae in different localities. The most usual developmental pathway for Euphausia superba is through Form 5, which may, thus, be regarded as the main developmental pathway for this species. This pathway probably corresponds to optimal environment conditions for larval growth and morphogenesis with maximal synchronisation of both processes. Consequently, it may be suggested that changes in developmental pathways of larvae are connected with suboptimal environmental conditions.     

The structure of plankton communities of the Indian Ocean

Species diversity and trophic structure of plankton communities have been considered as most important biological features. The Fisher () and Simpson () indices and the information index H=- pi log₂ pi (Margalef) were used for evaluation of species diversity. The indices have been calculated within the group of filter feeders by numbers ( and H) and biomass ( and H) of the species; the best results were obtained in the latter case. The index H=- pi log₂ pi is proposed for the quantitative evaluation of trophic structure, where a portion of an individual trophic group is taken as pi. The plankton material was collected by the R.V. Vityaz in the northern Indian Ocean in January through February, 1960. Analysis of characteristics of zooplankton distribution, species and trophic diversity in connection with characteristics of the water regime resulted in the definition of 3 types of the plankton communities: (a) those in intensive divergence zones; (b) those in poor divergence zones; (c) those in regions with stable water stratification or with feebly marked convergence, distiguished by different values of biomass, species diversity, and trophic group ratios.     

The evolution of cheating and selfish behavior

Summary We examine the necessary conditions for the spread of genes that determine selfish and cheating behaviors and the rate of spread of these genes through structured populations, in order to address the question of the invadability of altruistic systems by anti-social mutations. We find that, although cheaters always have a higher relative fitness than altruists within groups, population structures which permit the evolution of altruism also preclude invasion by anti-social mutations. These results are related to a discussion by Hamilton (1971) concerning the limits to the evolution of altruistic and selfish behaviors.     

Response of a soft-bottom harpacticoid community to stingray (Dasyatis sabina) disturbance

The recolonization by harpacticoid copepods of patches disturbed by stingrays [Dasyatis sabina (LeSeur)] feeding in a subtidal (2–3 m depth) site off the Florida gulf coast (Lat. 29°54.55N, Long. 84°31.45W) was followed. Samples from disturbed patches could no longer be distinguished from control samples in terms of numbers of harpacticoids 29 h after disturbance. No species became significantly disproportionately abundant during the recovery suggesting that small-scale natural disturbances are not an important organizing force in this community.     

Abnormal gonadal morphology in intersex,<Emphasis Type="Italic"> Echinogammarus marinus</Emphasis> (Amphipoda): a possible cause of reduced fecundity?

Recent reports have demonstrated a cost associated with intersexuality in Amphipoda, including reduced fecundity and fertility. In this study, the gross morphology of the gonads in normal and intersex Echinogammarus marinus (Amphipoda) were compared to determine whether resource allocation to gonadal tissue accounted for this reduced fitness. Evidence for the presence of the male sex-determining hormone, androgenic gland hormone (AGH), was compared between sexual phenotypes using MALDI mass spectrometry. Two distinct intersex phenotypes (male intersex and female intersex) were found, with variation in gonadal structure corresponding with external phenotype. Examination of male intersexes revealed normal testicular development (testes, seminal vesicles and vas deferens), but also revealed the formation of an oviduct. Ovaries of intersex females showed normal ovarian development, but were reduced in length by approximately 20% due to the presence of vas deferens. The number of vas deferens in intersex females was equal to the number (one or two) of genital papillae. We hypothesise that the reduced ovarian length observed in intersex females is a likely cause of the reduced brood size previously reported in intersex females of this species. Variation in the sexual phenotype corresponded both to development of the androgenic gland and to expression of a peptide fragment corresponding to the A chain of androgenic gland hormone (AGH). Androgenic glands and a putative AGH peptide were present in males. However, in both normal and intersex females, the androgenic glands were only present in a rudiment form and the peptide was not detected. Intersex males were found to possess abnormal glands that appeared hypertrophied. However, AGH peptides were not detected, supporting the suggestion that the intersex phenotype is manifested via perturbations of AGH.An erratum to this article can be found at Communicated by J.P. Thorpe, Port Erin     

The erect ‘penis’ is a flag of submission in a female-dominated society: greetings in Serengeti spotted hyenas

In East Africa, spotted hyenas live in large clans in a highly structured society dominated by females. A clan is a fission-fusion society where members are often solitary or in small groups. Spotted hyenas have a ritualized greeting during which two individuals stand parallel and face in opposite directions. Both individuals usually lift their hind leg and sniff or lick the anogenital region of the other. The unique aspect of greetings between individuals is the prominent role of the erect penis in animals of both sex. Female spotted hyenas have fused outer labiae and a pseudo-penis formed by the clitoris which closely resembles the male penis and can be erected. During greetings subordinates signalled submission with gestures which were not necessarily reciprocated by the dominant participant. Asymmetries were most pronounced in greetings between adult females where the probability of asymmetries increased with the divergence in rank between partners. Greetings between adult females and males were uncommon and restricted to males above median rank, principally the alpha male. Models of primate affiliative behavior assume that benefits derived from social relationships with different individuals are not equal and that individuals are selected to maximize the benefits they receive from social relationships with others. The observed distribution of greetings between partners of different rank matched the predictions of these models. An examination of non-adaptive hypotheses on the evolution of the pseudo-penis demonstrated that the conventional scenario linking (initial) virilization of female genitalia with selection for female dominance does not explain either the initial virilization, nor the evolution of the pseudo-penis to its current form and use. We sketch a new scenario that links (1) initial virilization to the occurrence of neonatal siblicide amongst members of a twin litter, and (2) costs of maintenance, pseudo-penile control over copulation and male submission. Our analysis confirms previous adaptive hypotheses on the function of the pseudo-penis in greetings and suggests new hypotheses to account for hitherto unexplained features. Correspondence to: M.L. East     

Carbon-isotope ratio gradients in western arctic zooplankton

Zooplankton from 87 stations in the Bering, Chukchi and Beaufort Seas sampled in 1985, 1986 and 1987 showed a geographic gradient in stable carbon-isotope ratios (¹³C). The zooplankton most depleted in ¹³C were found in the central and eastern Beaufort Sea and those most enriched were from the Bering and Chukchi Seas. Average ¹³C values ranged from-20.9 to-26.7 for copepods and from-19.4 to-25.1 for euphausiids. Euphausiids show a minimum of 1.0 enrichment relative to copepods throughout the study area. Relative biomasses of the major zooplankton taxa varied significantly across the Alaskan Beaufort Sea in October 1986, with euphausiids dominating in the west and copepods in the east. These differences in taxonomic composition affected the weighted ¹³C values for total zooplankton and may produce an even more pronounced geographic gradient in zooplankton ¹³C than that found within a single taxon. The bowhead whale Balaena mysticetus migrates between wintering areas in the Bering Sea and summering areas in the Beaufort Sea and feeds over this geographic range. The zooplankton ¹³C gradient is the probable source of ¹³C oscillations found along the baleen plates of this planktivore.     

Trunk trails and the searching strategy of a leaf-cutter ant,Atta colombica

Summary Atta colombica uses chemical mass recruitment that allows the rapid exploitation of resources. Most foragers thus search only within patches. Accumulation of extra foragers at patches results in sampling of alternate food items and area-restricted search as patch resources are depleted.Individual workers have a higher probability of removing a leaf fragment the earlier they arrive at a bait. Workers that arrive when much of the resource is gone travel further on the bait (within the patch) but do not spend significantly more time at the patch. They give up after 50–80s.Foraging effort is centered on the extensive trail system, not on the nest a predicted by time and energy foraging models. Search effort is also trail centered. The probability that an item will be discovered decreases with distance from the trail and increasing litter depth. Trail traffic and trail quality together mave no significant effect although this may be because they act antagonistically.Economic considerations predict that trials should be built to high quality and very productive sites. If trails are built as a result of recruitment and recruitment reflects patch quality and productivity, characteristics of forage sites are physically embodied in the trail system.Leaf cutter foraging is better understood as a long term optimization that effectively exploits resources over the lifetime of the colony than as prudent predation that husbands resources.