Agonistic and courtship behaviour patterns in the skink Chalcides viridanus (Fam. Scincidae) from Tenerife

There have been relatively few attempts to quantitatively describe behaviours in scincid lizards. Chalcides viridanus is a small body-sized skink endemic of Tenerife (Canary Islands). We describe and quantify 18 behaviour patterns (both social and agonistic) of this species, some of which have not been described before for other scincids. Video recordings of male–male, female–female, and male–female interactions were made under laboratory conditions, with controlled light–dark cycle and temperature. We describe several agonistic and courtship behaviour patterns. Within the first context, we detected a new agonistic behaviour for a scincid, “Snout to body”, that appeared at the beginning of agonistic sequences; it consisted of each animal placing its snout in contact with the other individual’s lateral side of the body. The amplitude of head movement during “Head bobbing” was lower than that described for many other lizard species. Agonistic behaviours were shown in intrasexual staged encounters both within males and females. The comparison of behaviour patterns of both types of intrasexual encounters showed that females were more active, exhibiting significantly higher frequencies of behaviour than males. Specifically, females showed the “Snout to body” pattern more frequently than males. In male–female encounters we detected courtship and copulation patterns only in April, when males performed “Bites” and “Snout to body” directed at females.     

On the reversibility of environmentally induced alterations in abstract biological systems

The environmentally induced alterations in structure of (M, ℜ) which were described previously (R. Rosen,Bull. Math. Biophysics,23, 165–171, 1961) are examined from the standpoint of determining under what circumstances they can be reversed by further environmental interactions. For simplicity we consider only the case of (M, ℜ)-systems possessing one “metabolic” and one “genetic” component. In the case of environmentally induced alteration of the “metabolic” component alone, a necessary and sufficient condition is given for the reversibility of the alteration. In the case of alteration of the “genetic” component, the situation becomes more complex; several partial results are given, but a full analysis is not available at this time. Some possible biological implications of this analysis are discussed. This research was supported by the United States Air Force through the Air Force Office of Scientific Research of the Air Research and Development Command, under Contract no. AF-49(638)-917 and Grant no. AF-AFOSR-9-63.     

Heterogeneity of heterochromatin in six species ofCtenomys (Rodentia: Octodontoidea: Ctenomyidae) from Argentina revealed by a combined analysis of C- and RE-banding

Exceptional chromosomal variability makesCtenomys an excellent model for evolutionary cytogenetic analysis. Six species belonging to three evolutionary lineages were studied by means of restriction endonuclease and C-chromosome banding. The resulting banding patterns were used for comparative analysis of heterochromatin distribution on chromosomes. This combined analysis allowed intra- and inter-specific heterochromatin variability to be detected, groups of species belonging to different lineages to be characterized, and phylogenetic relationships hypothesized from other data to be supported. The “ancestral group”,Ctenomys pundti andC. talarum, share three types of heterochromatin, the most abundant of which was also found in C. aff.C. opimus, suggesting that the latter species also belongs to the “ancestral group”. Additionally, within the subspeciesC. t. talarum, putative chromosomal rearrangements distinguishing two of the three chromosomal races were identified. Two species belong to an “eastern lineage”,C. osvaldoreigi andC. rosendopascuali, and share only one type of heterochromatin homogeneously distributed across their karyotypes.C. latro, the only analyzed species from the “chacoan” lineage, showed three types of heterochromatin, one of them being that which characterizes the “eastern lineage”.C. aff.C. opimus, because of its low heterochromatin content, is the most primitive karyotype of the genus yet described. The heterochromatin variability showed by these species, reflecting the evolutionary divergence toward different heterochromatin types, may have diverged since the origin of the genus. Heterochromatin amplification is proposed as a trend withinCtenomys, occurring independently of chromosomal change in diploid numbers.     

Male competition and coalitions in langurs (Presbytis entellus) at Jodhpur,Rajasthan, India

During a 15 month study on free ranging langurs (Presbytis entellus) at Jodhpur, Rajasthan, India, 5 adult male replacements were observed as a result of nontroop male invasions into the home ranges of 3 neighbouring one-male troops comprising 16–28 members each. Jodhpur langurs have no breeding season. Periods of instability during resident male changes lasted 11–119 days. Linear dominance hierarchies could be detected within the 3 main rival male bands of 2, 5, and 28–35 members. The respective alphas drove their allies away after their bands succeeded cooperatively at occupying a troop. During gradual replacements interim residencies alternated with multi-male stages. A large band's alpha may have had better chances to win the competition, since adult and nonadult allies functioned as “buffers” in agonistic encounters. The role of kin selection in structuring the composition of male bands and male coalitional behaviour cannot yet be quantified. Tactical “deceit” of powerful males to cause unrealistic expectations and in this way agonistic engagement of less strong males can be ruled out. “Sneaking copulations” is a proximate advantage for subordinate supporters, since they participated in 61.9% of all sexual interactions. Female promiscuity might reflect a strategy to induce male-male competition and thus select for a strong resident.     

Behaviour of Arctic charr Salvelinus alpinus during an induced mating season in captivity: how male relative size influences male behavioural investment and female preference over time

The behaviour of sexually mature Arctic charr Salvelinus alpinus specimens (fifth farm generation) was observed in captivity for four consecutive days. Only agonistic interactions between males of different size were facilitated on the first 2 days, while both agonistic and courtship interactions were possible from the third day up to the end of the experiment. The reliability of behavioural analysis was assessed in order to reduce the possibility of observer errors within the generated datasets. The behavioural investment of big males, small males and females was analysed using general linear models (two‐way repeated measures ANOVAs with time and male size as factors). A peak in the agonistic interactions between males occurred during the first day of interactions, where the agonistic investment of big males was significantly higher than that of small males. This resulted in an increased investment in submissive behaviour by the small males, who consistently performed submissive behaviours from the second day of interactions up to the end of the trial. Big males were found to invest significantly more than small males in courtship behaviours for the duration of the trial. Even though females performed inter‐sexual behaviours towards both big and small males for the entire observation period, female interaction rate towards big males was higher than towards small males. This study suggests that both male investment in mating behaviour and female preference might be related to male characteristics such as body length and that S. alpinus behavioural patterns and mate choice cues might be strongly context‐related and characterized by high levels of behavioural plasticity (i.e. presence–absence of certain behavioural units or potential reversal of a mate choice cue) within the same species. Finally, in light of this, some conservation measures are discussed. In particular, effective management plans should take into account the high level of behavioural plasticity likely to be occurring in this species.     

Regulation of arginase I activity and expression by both PD-1 and CTLA-4 on the myeloid-derived suppressor cells

An elevated number of Gr-1⁺CD11b⁺ myeloid-derived suppression cells (MDSCs) has been described in mice and human bearing tumor and associated with immune suppression. Arginase I production by MDSCs in the tumor environment may be a central mechanism for immunosuppression and tumor evasion. In this study and before, we found that Gr-1⁺CD11b⁺ MDSCs from ascites and spleen of mice bearing ovarian 18D carcinoma express a high level of PD-1, CTLA-4, B7-H1 and CD80 while other co-stimulatory molecules, namely CD40, B7-DC and CD86 are not detected. Further studies showed that PD-1 and CTLA-4 on the Gr-1⁺CD11b⁺ MDSCs regulated the activity and expression of arginase I. The blockage and silencing of PD-1, CTLA-4 or both PD-1 and CTLA4 molecules could significantly reduce arginase I activity and expression induced with tumor-associated factor. Similar results were also observed while their ligands B7-H1 and/or CD80 were blocked or silenced. Furthermore, CD80 deficiency also decreased the arginase I expression and activity. Antibody blockade or silencing of PD-1, CTLA-4 or both reduced the suppressive potential of PD-1+CTLA-4+MDSCs. Blockade of PD-1, CTLA-4 or both also slowed tumor growth and improved the survival rate of tumor-bearing mice. Thus, there may exist a coinhibitory and costimulatory molecules-based immuno-regulating wet among MDSCs. This research was supported by Nankai University grant, NSFC grant “30771967”, “985” grant,The Ministry of Science and Technology grant “2006AA020502”“06C26211200695”, Tianjin Grant “07JCZDJC03300” and “06ZHCXSH04800”.     

The complexus adhaerens of mammalian lymphatic endothelia revisited: a junction even more complex than hitherto thought

The significance of a special kind of VE-cadherin-based, desmoplakin- and plakoglobin-containing adhering junction, originally identified in certain endothelial cells of the mammalian lymphatic system (notably the retothelial cells of the lymph node sinus and a subtype of lining endothelial cells of peripheral lymphatic vessels), has been widely confirmed and its importance in the formation of blood and lymph vessels has been demonstrated in vivo and in vitro. We have recently extended the molecular and structural characterization of the complexus adhaerens and can now report that it represents a rare and special combination of components known from three other major types of cell junction. It comprises zonula adhaerens proteins (VE-cadherin, α- and β-catenin, protein p120^ctn, and afadin), desmosomal plaque components (desmoplakin and plakoglobin), and tight-junction proteins (claudin-5 and ZO-1) and forms junctions that vary markedly in size and shape. The special character and the possible biological roles of the complexus adhaerens and its unique ensemble of molecules in angiogenesis, immunology, and oncology are discussed. The surprising finding of claudin-5 and protein ZO-1 in substructures of retothelial cell-cell bridges, i.e. structures that do not separate different tissues or cell layer compartments, suggests that such tight-junction molecules are involved in functions other than the “fence” and “barrier” roles of zonulae occludentes. This work was supported by a grant from the Deutsche Forschungsgemeinschaft (DFG grant MO 345/5-2). This study is part of a thesis presented to the Faculty of Medicine of the University of Heidelberg, Germany, to fulfil the requirements of the doctoral degree (MD) of the first author.     

Visual Communication Behaviour as a Mechanism Behind Reproductive Interference in Three Pygmy Grasshoppers (Genus Tetrix, Tetrigidae, Orthoptera)

Specific mate recognition systems should enable species to recognize conspecific mates correctly. However, heterospecific matings have been observed in a variety of taxa. One of these cases is the pygmy grasshopper genus Tetrix, in which three species show sexual interactions. T. ceperoi males mount preferably females of T. subulata, but they are rejected as mates. T. subulata males prefer T. undulata females over conspecific females and heterospecific matings occur. Here, we study the underlying behavioural mechanisms of this pattern by analysing the visual courtship behaviour of the three species videographically. We test the hypothesis that the displays of T. ceperoi are highly differentiated from the other species, while the courtship of T. subulata and T. undulata is more similar. This is supported by our results: while T. ceperoi males perform a fast movement of high amplitude (“pronotal bobbing”), the other two species show only minor movements (“lateral swinging,” “frontal swinging”). The first function of a discriminant analysis of the temporal dimensions of the displays explained 96% of the variance. 100% of the T. ceperoi displays were classified correctly, whereas only 50% of the T. subulata and 81% of the T. undulata displays were grouped accurately. A two-way ANOVA revealed no significant effects of the direction of the “swinging” movements (laterally or frontally) and no interactions between direction and species either, whereas each temporal parameter differed significantly between the three species. The highest degree of differentiation was found between T. ceperoi and T. undulata, while T. subulata and T. undulata only differed significantly in two of the six temporal parameters. Our results suggest that the mismatings between T. undulata and T. subulata are caused by an insufficient specificity of the courtship behaviour. Apparently, ecological segregation of these two species could impede sexual interactions in the field.     

Engineering aspects of biological ion channels—from biosensors to computational models for permeation

Molecular sequencing has helped resolve the phylogenetic relationships amongst the diverse groups of algal, fungal-like and protist organisms that constitute the Chromalveolate “superkingdom” clade. It is thought that the whole clade evolved from a photosynthetic ancestor and that there have been at least three independent plastid losses during their evolutionary history. The fungal-like oomycetes and hyphochytrids, together with the marine flagellates Pirsonia and Developayella, form part of the clade defined by Cavalier-Smith and Chao (2006) as the phylum “Pseudofungi”, which is a sister to the photosynthetic chromistan algae (phylum Ochrophyta). Within the oomycetes, a number of predominantly marine holocarpic genera appear to diverge before the main “saprolegnian” and “peronosporalean” lines, into which all oomycetes had been traditionally placed. It is now clear that oomycetes have their evolutionary roots in the sea. The earliest diverging oomycete genera so far documented, Eurychasma and Haptoglossa, are both obligate parasites that show a high degree of complexity and sophistication in their host parasite interactions and infection structures. Key morphological and cytological features of the oomycetes will be reviewed in the context of our revised understanding of their likely phylogeny. Recent genomic studies have revealed a number of intriguing similarities in host–pathogen interactions between the oomycetes with their distant apicocomplexan cousins. Therefore, the earlier view that oomycetes evolved from the largely saprotrophic “saprolegnian line” is not supported and current evidence shows these organisms evolved from simple holocarpic marine parasites. Both the hyphal-like pattern of growth and the acquisition of oogamous sexual reproduction probably developed largely after the migration of these organisms from the sea to land.     

Basidiocarp development inCoprinus macrorhizus

The dikaryon #5026 (A2B2)+#5132 (A7B7) of the basidiomyceteCoprinus macrorhizus was treated with ultraviolet light (UV),N-methyl-N′-nitro-N-nitrosoguanidine (NG), and 5-bromouracil (BU) in order to induced developmental variations in fruiting. Among a total of 11,383 isolates, 742 were homokaryons. Fruiting test was made with the remaining 10,641 dikaryotic isolates, of which 1,594 showed abnormality in basidiocarp development. Both UV and NG were very efficient in inducing such variations, but BU was not. The induced variations were classified into seven basic types as follows: 1) “knotless”, in which hyphal knots, the first sign of fruiting, do not differentiate; 2) “primordiumless,” in which hyphal knots are formed, but no further development occurs; 3) “maturationles”, in which primordia are formed, but the maturation of primordia into adult fruit bodies does not occur; 4) “elongationless”, in which the elongation of stipe is blocked, and a basidiocarp with a very short stipe is produced; 5) “expansionless,” in which pilei do not expand normally; 6) “sporeless,” in which the formation of basidiospores is blocked and albinic pilei bearing none or only a small amount of spores are produced; and 7) “autolysisless,” in which the autolysis of pilei does not appear to occur. It has been noticed that the four steps of maturation, i.e. stipe elongation, pileus expansion, spore formation, and perhaps pileus autolysis, can proceed independently. Compound types of variations such as “elongation-expansionless,” “elongation-sporeless,” “expansion-sporeless,” and “elongation-expansion-sporeless” were also induced. UV treatment induced maturationless at the highest rate, while NG treatment sporeless. Contributions from the Division of Plant Morphology, Department of Biology, Faculty of Science, Okayama University. No. 117.     

Physics,biology and sociology: II. Suggestion for a synthesis

In continuation of previous studies (Bull. Math. Biophysics,28, 283–308; 655–661, 1966;29, 139–152, 1967) it is shown that the difference between the “metric” aspects of physics and the “relational” aspects of biological and social sciences disappear by accepting the broader definition of “relation”, such as that given in mathematics and logic. A conceptual superstructure then becomes possible from which all three branches of knowledge may be derived, though none of them can be derived from the others.     

Attributes and validity of dominance hierarchy in the female pigtail macaque

A group of six unrelated female pigtail macaques,Macaca nemestrina, of the same age, was studied in captivity over a period of five years. The animals were observed under six different situations, following changes in the social composition of the group. The following social interactions were recorded: withdrawal, attack, threat, presentation, mount, and grooming. Although behavioural rates varied for each period, their distribution network was remarkably consistent. Each subject could be assigned a rank, which remained stable over the six periods. From an analysis of the number of dyads in which one of the two animal significantly performed both paired behaviours more than the other animal (external validity), behaviours clustered into two main groups: (1) attacks, threats, and withdrawals; and (2) mounts, presentations, and grooming. Quantitative methods were performed to standardize the degree in which linearity, stability, reciprocity, and idiosyncrasy of the interactions under study may account for social variability. When considering these properties, the behaviours clustered in the same two groups (“agonistic” and “affinitive”). A method to describe the dominance style ofMacaca nemestrina was proposed, which can easily be replicated for comparisons with females of other primate species and in different conditions.     

Measuring and quantifying dynamic visual signals in jumping spiders

Animals emit visual signals that involve simultaneous, sequential movements of appendages that unfold with varying dynamics in time and space. Algorithms have been recently reported (e.g. Peters et al. in Anim Behav 64:131–146, 2002) that enable quantitative characterization of movements as optical flow patterns. For decades, acoustical signals have been rendered by techniques that decompose sound into amplitude, time, and spectral components. Using an optic-flow algorithm we examined visual courtship behaviours of jumping spiders and depict their complex visual signals as “speed waveform”, “speed surface”, and “speed waterfall” plots analogous to acoustic waveforms, spectrograms, and waterfall plots, respectively. In addition, these “speed profiles” are compatible with analytical techniques developed for auditory analysis. Using examples from the jumping spider Habronattus pugillis we show that we can statistically differentiate displays of different “sky island” populations supporting previous work on diversification. We also examined visual displays from the jumping spider Habronattus dossenus and show that distinct seismic components of vibratory displays are produced concurrently with statistically distinct motion signals. Given that dynamic visual signals are common, from insects to birds to mammals, we propose that optical-flow algorithms and the analyses described here will be useful for many researchers.Damian O. Elias and Bruce R. Land contributed equallyAn erratum to this article can be found at     

The nature and frequency of agonism in free-ranging and semi-free-ranging brown lemurs,Eulemur fulvus

Agonistic behaviour was studied in three groups each of free-ranging and semi-free-ranging brown lemurs (Eulemur fulvus) at Berenty, Madagascar and the Duke University Primate Center (DUPC) respectively. The purpose of the study was to answer questions arising from the work of other researchers regarding the frequency and intensity of agonism in this species. Authors of field studies generally concluded that agonism was rare and mild, whereas those who had studied semi-free-ranging or captive animals at the DUPC reported intense agonism during the peaks of the mating and birth seasons, with sometimes fatal wounding occurring among captive animals. I recorded 30 agonistic behaviours or “signals” which I grouped into seven general categories — cuffs, other physical contact, threats, chases, third party intervention, unprovoked submissive signals, and reciprocal aggression. The seven categories represent the types of signals which initiated or otherwise defined agonistic interactions, regardless of whether or not there was a submissive response to aggression. The relative percentages of all agonism constituted by the seven categories were not found to be significantly different between study sites. Agonistic signals were also classified as either subtle or obvious, a classification which crosscut the seven categories. At both study sites, the majority of agonistic signals initiating or defining interactions were subtle. Rates of agonism for the Berenty groups, studied during the birth season only, were significantly lower than those for the DUPC groups during the birth season, possibly due to (1) easier observation conditions at the DUPC, and (2) the impossibility of successful emigration at the DUPC, which might have resulted in social stresses translating into higher rates of agonism. In only one DUPC group was there significant variation in rates of agonism between seasons. I found agonistic behaviour to be mild, at both study sites, in the senses of subtlety of both aggressive and submissive signals, unlikelihood of response to aggression, and virtual absence of wounding; and I noted that serious wounding during other studies at the DUPC involved animals captive in caged runs. Comparing rates of the study groups with rates reported in other research for brown lemurs, other lemuriform species, and some New and Old World anthropoid species, I concluded thatE. fulvus agonism was in fact not rare except in comparison to baboons and macaques.     

Voiceprint identification and its application to sociological studies of wild Japanese monkeys (Macaca fuscata yakui)

Japanese monkeys often exchange the particular vocal sound, “coo,” especially when they feed or move as a group. It was considered that the “coo” sound had no positive social meaning, perhaps because the “coo” sound network and its function were hidden behind other behavioral observations. For identification of the vocalizer only from hearing the “coo” sound, three phonetic values, i.e., the “fundamental,” “duration,” and “formants,” plus other characteristics were used as indices of voiceprints. The results indicated that these were effective for identifying the vocalizer in two-thirds of the adults in the study troop which was composed of 12 adults and 16 immature members. The “coo” sound exchange network among the troop members (adults) was drawn on the basis of the voiceprint identification. The network showed three characteristics as follows: (1) matriarchs of the kin-groups frequently exchanged “coo” sounds with each other; (2) the other females exchanged “coo” sounds mostly within their own kin-groups; and (3) males seldom participated in the “coo” sound exchange. This suggests that “coo” sound exchange plays a central role for the matriarch of kin-groups in binding each kin-group and, ultimately, in binding all members together into an organized troop.     

Social behavior of the emperor tamarin in captivity: Components of agonistic display and the agonistic network

Agonistic behavior was studied longitudinally for 16 months in an intact family groups of captive emperor tamarins (Saguinus imperator subgrisescens) using methods from quantitative ethology and social network analysis. A motivational analysis of the components of agonistic display revealed the relative strength of each component along a continuum from strongly dominant to strongly subordinate. Tabulations of exchanges of strongly dominant and strongly subordinate components in interactions among the tamarins revealed an agonistic network (“dominance hierarchy”) that approached, but did not quite reach, the ideal state of a transitive order (“linear dominance hierarchy”). The frequency with which individual tamarins long called and scent marked was not closely correlated with their position (“dominance rank”) in the agonistic network. Instead, individuals undergoing change in status long called and scent marked frequently, irrespective of their rank.     

An ant’s eye view of culture: propagation of new traditions through triggering dormant behavioural patterns

We consider a previously unknown way of propagation of behavioural traditions in animal communities using hunting in ants as an example. We experimentally revealed that common litter dwelling ants Myrmica rubra effectively hunt jumping prey and the way the hunting behavioural pattern is distributed within ant colonies is rather sophisticated. Comparison of our results with those obtained on vertebrates enables us to suggest that “distributed social learning” plays an important role in spreading new traditions in animal communities: initial performances by a few carriers of an “at once and entirely” available behavioural pattern propagate this pattern among specimens which have only dormant “sketches” of it. Spread of these behaviours in populations is based on relatively simple forms of social learning such as social facilitation which underlies species’ predisposition to learn certain sequences of behavioural acts. To be triggered, carriers of dormant “sketches” of a relevant behavioural pattern should encounter performances of this pattern with sufficient frequency. We call this strategy triggering of dormant behavioural patterns. Integration of behaviour thus takes place not only at the individual level but at the population level as well.     

Are Blackcaps (Sylvia atricapilla) defending their nests also calling for help from their neighbours?

One hypothesized function of conspicuous mobbing of intruders by bird nest owners is to attract neighbouring birds (“calling for help” hypothesis) or third-party predators (“attract the mightier” hypothesis). These may help the nest owners by distracting and/or attacking the mobbed intruder. To date, these hypotheses have been studied solely during the mobbing of predators. Here, for the first time, I have studied mobbing attraction in the context of brood parasitism. I experimentally tested the Blackcap (Sylvia atricapilla), a small passerine with a highly aggressive and conspicuous nest defence behaviour. I elicited the aggressive responses of Blackcaps by presenting stuffed dummies of the brood parasitic Common Cuckoo (Cuculus canorus) and controls near their nests. At 32% of the nests (n = 75), the responses of the Blackcaps to dummies attracted up to 15 birds per trial from 21 passerine species. Most of the attracted birds were heterospecifics and rarely participated in mobbing; thus the “calling for help” hypothesis was not supported. No potential predators of the Cuckoo were attracted despite them living in the study area and despite prolonged mobbing by Blackcaps; thus rejecting the “attract the mightier” hypothesis. I argue that this hypothesis is unlikely to apply to typical avian predators during nest predation acts because these only last for several seconds. The number of attracted birds was a positive function of the owner’s intensity of nest defence as measured by the rates of alarm calling but not visual cues (rates of attacks). Suitable and unsuitable Cuckoo hosts did not differ in their behaviour in the vicinity of defended nests. The observed pattern of the positive correlation between the intensity of nest defence and the number of attracted birds is most likely a proximate by-product of the conspicuous nest defence by Blackcaps (but may well be adaptive for recruiting neighbours themselves). Thus, the mobbing behaviour of the Blackcap is directed towards the brood parasite and not towards other audience predators or potential recruits to the mob.     

Relationships among three Japanese <Emphasis Type="Italic">Laetiporus</Emphasis> taxa based on phylogenetic analysis and incompatibility tests

Relationships among three Japanese Laetiporus taxa (“L. sulphureus var. sulphureus” auct. jap., L. sulphureus var. miniatus, and L. versisporus) were assessed with phylogenetic analysis and incompatibility tests. Gene phylogenies inferred from the internal transcribed spacer region of nuclear ribosomal DNA, elongation factor 1α, and β-tubulin gene regions suggested that Japanese Laetiporus was divided into four groups: the yellow pore form of L. sulphureus var. miniatus, the white pore form of L. sulphureus var. miniatus, and two “L. sulphureus var. sulphureus”/ L. versisporus groups. A morphologically distinct species, Laetiporus versisporus, sharing a clade with “L. sulphureus var. sulphureus” auct. jap., was proved to be an anamorphic form of “L. sulphureus var. sulphureus” auct. jap. The “sulphureus/versisporus” isolates showed two divergent sequence types in each region. Some isolates had intraindividual polymorphism assigned to both sequence types. This finding suggests that speciation via hybridization is ongoing in the “sulphureus/versisporus” group. Single spore isolates from the “sulphureus/versisporus” group, white pore group, and yellow pore group were incompatible with each other. Our results provided strong support for the new recognition of three Laetiporus taxa in Japan.     

The metabolic rates associated with resting, and with the performance of agonistic, submissive and digging behaviours in the cichlid fish Neolamprologus pulcher (Pisces: Cichlidae)

We measured the metabolic rates as a direct estimate of energy expenditure of individual Neolamprologus pulcher, a cooperatively breeding cichlid fish, when resting and when performing agonistic, submissive or digging behaviours in a respirometer. Standard and routine metabolic rates increased linearly with body mass (range 0.9–8.4 g) when plotted on a doubly logarithmic scale (linear regression equations: standard metabolic rate: log individual oxygen consumption rate = 0.65 + 0.86 log body mass; routine metabolic rate: log individual oxygen consumption rate = 0.75 + 0.86 log body mass). Routine metabolic rates were, on average, 30% higher than standard metabolic rates. Submissive and agonistic behaviours raised routine metabolic rates by factors of 3.3 and 3.9, respectively. Digging resulted in a 6.1-fold increase of routine metabolic rates. Differences in metabolic rates between active and resting rates were statistically significant. However, those between the three behaviours were not. Mean opercular beat frequencies correlated significantly with routine metabolic rates and with metabolic rates when performing specific behaviours, which offers methodological prospects for field measurements. In N. pulcher, the high energy expenditure for submissive behaviour may indicate that this is a reliable signal. The considerable energy expenditure involved in territory defence suggests that these costs should be considered in addition to risk in cost-benefit analyses. This is the first study in which the energy expenditures of specific social and territory maintenance behaviours of individual fish were measured directly by respirometry and within the usual social setting of the fish. Accepted: 20 February 1998