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.     

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.     

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     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Transformation of soft coral (Coelenterata: Octocorallia) terpenes by Ovula ovum (Mollusca: Prosobranchia)

The faecal pellets from specimens of the prosobranch mollusc Ovula ovum found feeding on the soft coral Sarcophyton sp. at Eclipse Island, Palm Island Group (18°46S; 146°33E) in November 1980 were analysed. The only terpene present in the faeces, 7,8-deoxysarcophytoxide, differed from the major constituent of the soft coral, sarcophytoxide, suggesting that the latter had been transformed into the former within the cowrie. This transformation is not trivial, and could not be produced simply by acid catalysis. Subsequent analysis of tissues dissected from different regions of O. ovum indicates that the transformation is probably effected by enzymes in the digestive diverticula stomach region of the prosobranch. The transformed compound is significantly less toxic to the mosquito fish Gambusia affinis Baird and Girard than the ingested compound.     

Predator inspection behaviour in minnow shoals: differences between populations and individuals

Summary When minnows (Phoxinus phoxinus) detect a stalking pike (Esox lucius) one of their first responses is to perform inspection behaviour during which individuals or small groups approach the predator. This paper compares the inspection behaviour of two contrasting groups of minnows: Dorset minnows which have been heavily predated by pike for many thousands of years and Gwynedd minnows which have spent an equivalent period of time in a pike free environment. Minnows sympatric with pike inspected a realistic model pike more frequently and in larger shoals. Although they commenced inspection earlier they were more timid and kept a greater distance between themselves and the predator. After an inspection they were less likely to recommence foraging than minnows from the Gwynedd population. Individual differences in inspection found within the two populations suggest that selfish behaviour was present.     

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     

Feeding,growth and bioluminescence of the heterotrophic dinoflagellate Protoperidinium huberi

Feeding, growth and bioluminescence of the thecate heterotrophic dinoflagellate Protoperidinium huberi were measured as a function of food concentration for laboratory cultures grown on the diatom Ditylum brightwellii. Ingestion of food increased with food concentration. Maximum ingestion rates were measured at food concentrations of 600 g C l^-1 and were 0.7 g C individual^-1 h^-1 (1.8 D. brightwelli cells individual^-1 h^-1). Clearance rates decreased asymptotically with increasing food concentration. Maximum clearance rates at low food concentration were ca. 23 l ind^-1 h^-1, which corresponds to a volume-specific clearance rate of 5.9x10⁵ h^-1. Cell size of P huberi was highly variable, with a mean diameter of 42 m, but no clear relationship between cell size and food concentration was evident. Specific growth rates increased with food concentration until maximum growth rates of 0.7 d^-1 were reached at a food concentration of 400 g C l^-1 (1000 cells ml^-1). Food concentrations as low as 10 g C l^-1 of D. brightwellii (25 cells ml^-1) were able to support growth of P. huberi. The bioluminescence of P. huberi varied with its nutritional condition and growth rate. Cells held without food lost their bioluminescence capacity in a matter of days. P. huberi raised at different food concentrations showed increased bioluminescence capacity, up to food concentration that supported maximum growth rates. The bioluminescence of P. huberi varied over a diel cycle, and these rhythmic changes persisted during 48 h of continuous darkness, indicating that the rhythm was under endogenous control.     

Risks and benefits of lethal male fighting in the colonial,polygynous thrips Hoplothrips karnyi (Insecta: Thysanoptera)

Summary Males of Hoplothrips karnyi (Hood) (Insecta: Thysanoptera), a colonial fungus-feeding thrips, fight each other in defense of communal egg mass sites, where they mate with females that come to oviposit. Fighting males stab each other with their enlarged, armed forelegs and hit each other with their abdomens. Escalated fights occur between large males of similar size. Fights are often lethal; males that died during observations fought more frequently than other males, were stabbed more often and more severely than other males, and were relatively large, but somewhat smaller than their opponents. Large males tend to win fights and guard egg masses, and they secure about 80% of last matings before ovipositions. Guarding males apparently assess female reproductive condition by putting their forelegs partially around females' abdomens; guarding males, but not nonguarding males, mate preferentially with females that have yet to oviposit. Non-guarding males mate with females away from egg masses, sneak matings at egg masses, and occasionally challenge guarding males. Challenges tend to follow matings by non-guarding males at egg masses. Each of four observed or inferred takeovers was followed by the death of the guarding male that lost. Male fighting strategies are discussed in terms of the consistency of lethal fighting with game theory models. Guardin males appear to pursue a classical hawk strategy of escalate until injured or victorious. This strategy may be advantageous because only large males become guarders, the mating success of guarders greatly exceeds that of non-guarders, and high population viscosity ensures that benefits from killing an opponent accrue directly to gaurders. The occurrence of challenges by large non-guarders implies that fighting ability and resource value asymmetries between males change over time; such changes may result from the energetic costs of guarding, injury to guarding males, or depletion of guarding males' supply of sperm.     

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.     

Preliminary report on the digestive enzymes present in the digestive gland ofPerna viridis

Quantitative data on digestive enzymes of bivalves are very limited and so this study was performed to obtain such information, and to find out whether some of these enzymes are well adapted to conditions in the digestive gland. The green musselPerna viridis was obtained from Clementi West Market in 1985. The results show that various carbohydrases are present in the digestive gland. The activities of-amylase, cellulase and-glucosidase are higher than that of other carbohydrases, whilst the activities of both lipase and protease, especially the former, are low. The pH optima of the following enzymes are also given:-amylase = 5.8; laminarinase = 5.5;-glucosidase = 5.8; trehalase = 4.8;-glucosidase = 5.5. The only enzyme which was not well adapted to conditions in the digestive gland (pH 6.2) was trehalase. The wide spectrum of enzymes present in this bivalve indicates that it has the ability to utilize a wide range of nutrients efficiently.