Scent-rubbing and status signalling by male mammals

Summary There is no consensus about the function of scent-rubbing, a widespread behaviour in which mammals rub their bodies vigorously in substances, many strong-smelling and some artificial, such as rotting meat, intestinal contents and engine oil. Here we suggest that scent-rubbing is involved in status advertisement and that, as in assessment using scent marks, the mechanism used by competitors to assess potential opponents may be scent-matching. In scent-matching a resource holder is assessed (identified) by comparing its odour with odours on or near the defended resource. In scent marking the odour originates from the resource holder (glandular secretion, urine and faeces); in scent-rubbing the odour originates in the environment.A prerequisite of unambiguous scent-matching is that the odour of scent-marks should be uniquely characteristic of one individual. This may be why marking substances are very complex chemically. Scent-rubbing often occurs with scent-marking and, rather than acting independently of scent-marking, the odours acquired may either (i) add to the complexity of the signal, thus reducing signal ambiguity, or (ii) increase the range of the signal by adding a strong smelling component.Subordinates could potentially cheat by rubbing in the same odours as the resource holder. Resource holders could prevent cheating (i) by checking other status cues and by testing competitors whose scent matches, then escalating contests when the competitor's fighting ability (more formally, Resource Holding Power) proves to be lower than that of a resource holder and (ii) by mixing the substances used for scent-rubbing with the unique substances used in scent-marking.     

Studies on the nutritional requirements of the larval stages of Penaeus japonicus using microencapsulated diets

Fatty acid biosynthesis in the larval stages of Penaeus japonicus Bate was examined by feeding microencapsulated diets containing (1-¹⁴C) palmitic acid, and fat-free diets supplemented with defined fatty acids. Highest larval growth rates were achieved on diets containing Tapes philippinarum lipid and, when defined fatty acids were substituted, on diets containing 20:63 fatty acid. The radioactive tracer experiments indicate that 16:17, 18:0 and 18:19 fatty acids may be synthesised from palmitic acid and that P. japonicus larvae may possess the ability to elongate 18:33 to 20:53 and 22:63, and 18:26 to 20:46. However, the rates of these reactions appear to be too slow to meet the larval requirements for essential fatty acids and the 3 series of polyunsaturated fatty acids must be provided in the diet.     

Alloimmune memory for glycoproteid recognition molecules in sea anemones competing for space

A highly specific recognition system, capable of distinguishing between allogeneic (non-clonemates) and syngeneic (clonemates) conspecifics, exists in Anemonia sulcata (Coelenterata: Anthozoa). During a competitive interaction between allogeneic individuals, specialized structures (acrorhagi) are exposed and used to sting opponents. This complex behavior is called the acrorhagial response. In order to quantify this behavior we measured the acrorhagial response time (period from first contact with opponent to full expansion of acrorhagi). More than 320 reciprocal histoincompatibility responses were recorded. These histoincompatibility (H) markers are free components of the sea anemone mucus: they were characterized as glyco-proteins of a molecular weight <18 KD (Kilo-Dalton) containing terminal -d-glucose and/or -d-mannose. In alloimmune memory testing experiments, the memory trace in A. sulcata was found to persist for 5 d.     

Dwarf mongoose and hornbill mutualism in the Taru desert,Kenya

Summary Dwarf mongooses in the Taru desert region of Kenya form foraging communities with a variety of endemic bird species, especially hornbills. The prey spectra of the mongooses and hornbills overlap almost completely. For the other bird species forming the foraging community only partial overlap exists. The association between the birds and mongooses is actively sought by both parties. The birds wait in tress around the termite mound where the monogooses are sleeping for them to emerge and the mongooses delay their foraging departure if no birds are present. There is a positive relationship between the number of mongooses in the group and the number of birds accompanying them. A true mutualism only exists between the mongooses and the two hornbill species Tockus deckeni and T. flavirostris since their presence or arrival affects the subsequent start of foraging. These two hornbill species have also been observed to influence the start of foraging actively by means of two behaviour patterns termed chivvying and waking. Both the mongooses and birds are exposed to a high predator pressure from raptors with an overlap in the birds of prey predating the various species. This predator pressure is counteracted behaviourally by the mongooses by means of an altruistic behaviour pattern, guarding. Both mongooses and birds warn vocally and flee when a raptor is sighted. The mongooses modify their guarding behaviour to compensate for the warning behaviour of the birds in two ways: (a) fewer mongooses guard when large numbers of birds are present and vice versa, (b) the frequency of the mongooses' intraspecific warning calls is significantly reduced in cases where birds are present in comparison with those where they are absent. The birds also sight and respond to the raptor first on significantly more occasions than the mongooses. In addition, the birds also warn for raptor species which do not predate them but which are mongoose predators, not, however, for raptors which are not mongoose predators. This mutualistic association with its high degree of compensatory behaviour by both parties appears to be unique for free-living vertebrates and has its closest parallel in the trophobiosis described for ants and aphids.     

Are communal pukeko caught in the prisoner's dilemma?

Summary The prisoner's dilemma of game theory provides a possible explanation for communal breeding in the pukeko (Porphyrio porphyrio, Aves) where group breeding birds are at a reproductive disavantag compared to pairs. Territorial defence is largely the realm of males. A male who defect on neighbours by taking in an extra male defender and so becoming communal stands to gain territorial area and possibly higher reproductive success. Neighbours are forced to follow suit if additional males are available or lose their territory and therefore their breeding opportunities. If neighbours also admit an extra defender, the initial defector loses his advantages and all males end up with the lower breeding success of communal groups. Such defection to a communal habit appears to be the only stable solution.     

Binary individual recognition in hermit crabs

One mechanism that permits the maintenance of dominance hierarchies is individual recognition, defined as the ability of an animal to recognize a conspecific on the basis of one or more identifying cues, and to associate it with experiences of victories or defeats that the animal has gained from preceding encounters with that particular individual. We examined whether the long-clawed hermit crab, Pagurus longicarpus, could differentiate between unfamiliar and familiar opponents. The experimental protocol was designed to control in pairs of interacting individuals several factors together, such as status and relative size of the opponent, as well as species, quality, and fit of the inhabited shell. The hermit crabs were more reactive and their agonistic level was higher in unfamiliar than in familiar pairs; in addition, betas were more prone to initiate an interaction with unfamiliar than with familiar alphas. The alternative explanation—that the ability to discriminate between familiar and novel shells can explain our results per se— was tested following, in part, Jackson and Elwoods (1989) protocol for Pagurus bernhardus and was, at least for this species, rejected. This study did not determine whether a true individual recognition occurs, but demonstrated that P. longicarpus categorizes the individuals into two heterogeneous subgroups, thus being capable of a binary discrimination among opponents.Communicated by T. Czeschlik     

Individual variation and prior experience affect the discrimination of a brood-parasite by its subsocial beetle host

In birds, host experience can modify response to parasites but nothing similar is known for insects. We studied two desert tenebrionid beetles, the subsocial host Parastizopus armaticeps and its obligate and ubiquitous brood-(clepto)parasite Eremostibes opacus, which mimics the hosts odour spectrum. Nearly 10% of host burrows in the field (n=214) remain unparasitised even after introducing E. opacus experimentally (n=22). In discrimination tests, 7% of naive host beetles eject the parasite, showing individual variation in olfactory discrimination ability. To test for effects of prior breeding experience, naive (first time) or experienced (second time) breeders with or without prior parasite contact were confined with E. opacus at breeding onset, their behaviour recorded and parasite number counted on breeding completion. More first- than second-time breeding pairs completely excluded the parasite and second-time breeders with no prior contact with E. opacus had fewer parasites than any others. These pairs increased burrow-guarding dramatically, mostly during early reproduction, this being the best predictor of parasite number. Prolonged guarding, however, must be traded-off against other brood-care behaviours. Hence, it might pay breeding pairs to take the risk of accepting some parasites breeding within their burrow, and we expect this effect will be more pronounced if poor feeding and digging conditions are taken into account. This might explain why apparently 90% of the P. armaticeps breeding burrows in the field contain at least one parasitic E. opacus.Communicated by D. Gwynne     

Analysis of the effects of zinc pollution on the macro-invertebrate populations of the Afon Crafnant,North Wales

The study of the input draining from mine-waste into the Afon Crafnant, N. Wales, revealed that the most significant pollutant was zinc. The concentrations of zinc were monitored during 1978 and 1979, and spot samples were concentrated and analysed for lead, copper, cadmium, iron, cobalt, manganese and nickel. Analysis was by flame atomic absorption specto-scopy. The total zinc concentration of the input was about 4 mg Zn/l which was diluted by the Afon Crafnant to give mean annual concentrations of 213 g Zn/l and 146 g Zn/l for 1978 and 1979 respectively at a site 60 metres downstream of the input. The numbers of individuals and number of taxa in the macro-invertebrate community were reduced at these concentrations. The taxa affected werePhagocata vitta,Erpobdella octoculata, Oligochaeta,Ithytrichia lamellaris, Simuliidae,Ancylus fluviatilis andRithrogena semicolorata. Several commonly used diversity indices failed to show differences between the macro- invertebrate communities at the affected and control sites whereas a modified version of Chandler's Biotic Score (detailed in this paper) revealed both the acute effects at higher concentrations and the chronic effects at lower concentrations.     

Spatial mesoscale patterns of West Pacific picophytoplankton as analyzed by flow cytometry: their contribution to subsurface chlorophyll maxima

Vertical distributions of picophytoplankton (ppp) (<2 m) were studied by ship-board flow cytometry during two cruises in Western Pacific waters to Palau and to Australia in 1990. Weak red-fluorescing small ppp, supposed to be free-living prochlorophytes (Chisholm et al. 1988), were abundant in the area surveyed. These ppp, designated the prochlorophytes, were abundant in the surface waters (>10⁴ cells ml^-1) at the northern region (27°03N; 7°11N) in November, whereas in December at the southern tropical stations (0°23.54S; 9°20.30S; 13°50.6S), they formed subsurface maximum layers (>10⁵ cells ml^-1) on a nitracline at a depth of 3.5 to 5.4% surface irradiation. Their fluorescence intensity increased with depth below 10% surface irradiation. The prochlorophytes at a depth of 1% surface irradiation had ten times higher fluorescence than those at the surface layer. The total fluorescence intensity of the prochlorophytes accounted for 32 to 63% of the sum of the total fluorescence intensity of all fluorescing phytoplankton detected at subsurface chlorophyll maxima in the tropical area. These results suggest that distribution of the prochlorophytes is greatly affected by nitracline and by light intensity and that their chlorophyll is a major contributor to the subsurface chlorophyll maximum in the pelagic West Pacific Ocean.     

Migrational orientation in two species of littoral gastropods (Littorina angulifera and Nerita senegalensis)

Littorina angulifera (Lamark) and Nerita senegalensis Gmelin, both intertidal gastropods, appear to have a preference for specific levels of the shore. In some experiments conducted both in the laboratory and in the field, displaced snails exhibited a general homing behaviour when displaced upshore or downshore. Cues involved in this migrational behaviour have not clearly been identified.     

Influence of biological and environmental factors on nitrogenous extractives of the spurdog Squalus acanthias

The influence of sex, sexual maturity, feeding habits and season on the most important nitrogenous extractives in the muscle tissue of the spurdog Squalus acanthias L. of the southern North Sea has been studied. Feeding habits and sex have no influence, whereas the contents of -amino nitrogen, -alanine, glutamic acid and glycine decrease after sexual maturity. Only peptides are influenced by season; higher amounts occur in the spring/summer period.     

Pathway and products of CO2-fixation by green prokaryotic algae in the cloacal cavity of Diplosoma virens

Light-dependent ¹⁴CO₂ fixation by the algae of Diplosoma virens (Hartmeyer) ranged between about 3 and 27 moles mg^-1 chlorophyll h^-1. The principal first products of ¹⁴C fixation were 3-phosphoglyceric acid and phosphorylated sugars, indicating that ribulose bisphosphate carboxylase was the primary carboxylation enzyme. The activity of this enzyme in crude extracts of the algae was 4 to 6 moles CO₂ mg^-1 chlorophyll h^-1. The principal end product of ¹⁴C fixation by these algae in the ascidian host was a water-soluble oligosaccharide which was an -1,4-glucan. A maximum of 7% of the ¹⁴C fixed was found in insoluble materials of the algae or its host after 60 min ¹⁴CO₂ fixation. Whether the -1,4-glucan is a product of algal or animal metabolism remains to be determined.     

Capillary supply and mitochondrial volume density in the axial muscles of the mesopelagic teleost Argyropelecus hemigymnus

Quantitative measurements have been made on the ultra-structure and capillary supply to the axial muscles of the mesopelagic hatchet fish Argyropelecus hemigymnus (Cocco, 1829). Fish were collected at Eastern North Atlantic Ocean Station 10244, 32°48N; 31°15W during November 1980, from a depth of 480 to 550 m. Mitochondria with densely packed cristae occupy 44.3% of slow-fibre volume. Each myofibril is in direct contact with a mitochondrion. Compared with other fishes studied, the capillary supply to A. hemigymnus slow fibres is poorly developed. The average number of capillaries per fibre is 0.9, such that each m of capillary contact supplies 0.011 m² of fibre cross-sectional area. The capillary surface area (m²) supplying 1 m³ of slow-fibre mitochondria is 0.17 in anchovy (Engraulis encrasicolus), 0.14 in rat-fish (Chimaera monstrosa), 0.14 in tench (Tinca tinca), 0.16 in catfish (Clarias mossambica), and only 0.025 for A. hemigymnus. It is suggested that, relative to the former species, some modifications in factors determining tissue oxygenation (e.g. myoglobin concentrations, blood flow, perfusion distribution or haemoglobin) and/or mitochondrial respiration rate are required in order to match oxygen supply and demand to the slow muscle in A. hemigymnus.     

Microgeographic differentiation in the lipid composition of the bivalve Macoma balthica from the Gulf of Gdańsk (Southern Baltic)

Differentiation in lipid content, composition of lipid classes, fatty acids and sterols of Macoma balthica (L.) collected from various parts of the Gulf of Gdask was observed. The clams taken from the Inner Puck Bay contained significantly lower amounts of lipids, particularly of triglycerides, oleopalmitic acid and 24-methylenecholesterol. It is suggested that this differentiation is related to variability of trophic conditions in the Gulf of Gdask.     

The risks and benefits of signalling aggressive motivation: a study of cave-dwelling little blue penguins

Summary Cave-dwelling little blue penguins, Eudyptula minor, use a large repertoire of agonistic displays. Both qualitative and quantitative measures suggested that some displays were more dangerous to perform than others (estimated by the risk of being injured by one's opponent). A lag sequential analysis of interactions indicated a positive relationship between the risks of performing a display and the display's effectiveness (i.e., in deterring opponents). The cost or risk asymmetries that exist between displays may allow opponents to assess how motivated the actor is to secure or defend resources. Thus, high cost displays are more effective because they reveal a willingness to take risks and also place the opponent in a higher risk situation. Communicating in this way can be an evolutionarily stable strategy (ESS) because signalling a strong motivation entails using potentially costly behavior (i.e., there are costs associated with bluffing). Also, displays that were accompanied by calls were more effective in deterring opponents than the same displays performed silently. Calls may highlight the risks being taken by actors; this may be particularly important in nocturnal species like little blue penguins. Aggressive displays could also be used to predict what the actor was and was not likely to do following the performance of the display. Generally, aggressive displays predicted that the actor would escalate to a higher risk behavior if the opponent did not retreat, but that it would use a lower risk behavior if the opponent backed down. However, risk-based displays should be seen as revealing a certain level of motivation at the time of the display and not as a way of describing how great a risk the actor is willing to take following the display. The predictive value of displays appears to be the result of an escalation process which ensures that opponents are deterred by the lowest risk behavior possible.     

Effect of ionizing radiation on haemoglobin of marine lamellibranchs

The effect of ionizing radiation on the iron-linked protein (haemoglobin) of the marine lamellibranchs Anadara granosa (Linn.) and Cardita antiquata (Lam.) from Bombay waters, India is discussed. Purified haemoglobin solutions were exposed to a ⁶⁰Cobalt source delivering a dose of about 4,600 rad/min, at the sample irradiation point. Radiation damage or degradation was measured spectrophometrically by studying changes in the absorption spectra following irradiation in the presence and absence of oxygen. Exposure to ionizing radiation in general caused a decrease in absorption of both haemoglobins, irrespective of location, viz extra-versus intracellular, at Soret (412 m), (540 to 42 m) and (574 to 76 m) peaks, and an increase in absorption at 510 and 630 m. Upon exposure to a higher dose, O₂Hb of C. antiquata showed an increase in absorption at the protein peak (280 m); exposure to a lower dose, however, resulted in decreased absorption. Furthermore, the changes following irradiation were dependent upon the initial state of the pigment. Oxyhaemoglobin, when exposed to radiation, oxidized to hemiglobin, and hemiglobin reduced to oxyhaemoglobin. The extracellular haemoglobin of high molecular weight (3x10⁶) of the false cockle C. antiquata was found to be extremely radio-resistant, whereas intracellular haemoglobin of low molecular weight (74,000) of the arcid clam A. granosa was highly radiosensitive, since it could not be exposed to doses exceeding 18,000 r.     

Modelling skewed data with many zeros: A simple approach combining ordinary and logistic regression

We discuss a method for analyzing data that are positively skewed and contain a substantial proportion of zeros. Such data commonly arise in ecological applications, when the focus is on the abundance of a species. The form of the distribution is then due to the patchy nature of the environment and/or the inherent heterogeneity of the species. The method can be used whenever we wish to model the data as a response variable in terms of one or more explanatory variables. The analysis consists of three stages. The first involves creating two sets of data from the original: one shows whether or not the species is present; the other indicates the logarithm of the abundance when it is present. These are referred to as the presence data and the log-abundance data, respectively. The second stage involves modelling the presence data using logistic regression, and separately modelling the log-abundance data using ordinary regression. Finally, the third stage involves combining the two models in order to estimate the expected abundance for a specific set of values of the explanatory variables. A common approach to analyzing this sort of data is to use a ln (y+c) transformation, where c is some constant (usually one). The method we use here avoids the need for an arbitrary choice of the value of c, and allows the modelling to be carried out in a natural and straightforward manner, using well-known regression techniques. The approach we put forward is not original, having been used in both conservation biology and fisheries. Our objectives in this paper are to (a) promote the application of this approach in a wide range of settings and (b) suggest that parametric bootstrapping be used to provide confidence limits for the estimate of expected abundance.     

Ecological studies on a tropical limpet Cellana radiata

Some aspects of the habitat zones, distribution pattern, and population density with reference to tidal levels of the tropical limpet Cellana radiata (Born) are presented. Observations have also been made on the homing behaviour of the limpet. Unlike the temperate limpets, no migration to different tidal levels during different seasons was observed in C. radiata.     

An instance of bacterial inhibition in oceanic surface water

Water from the 150 surface film 50 miles north of Puerto Rico at 19°30 N, 66°30W supported only a negligible microflora compared to that of adjacent stations. Bacteriostasis was indicated when a ten-fold dilution of the sample resulted in a ninety-fold increase in cultivable microorganisms. The surviving organisms, atypical pseudomonads, were biochemically inactive in comparison to isolates from the other stations. Net phytoplankters at this station were similar in relative composition and abundance to those at the adjacent station.     

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.