Nest defence in the American goldfinch

In the American goldfinch (Carduelis tristis), vocalizations during nest defence consist largely of two different types of calls: either ‘sweeet’ (SWT) or ‘bearbee’ (BB). By recording call rates and behaviour during controlled experiments, it was demonstrated that: (1) SWT calls have a rallying function and attract more goldfinches than BB calls; (2) goldfinches attempt to distract potential predators away from their nests; (3) BB calls cause a decrease in nestling conspicuousness, both audibly and visually; (4) call rates show a positive relationship with increasing conspicuousness of nestlings; (5) call rates of BB and SWT calls vary inversely with danger to the nest; (6) high call rates are given by parents with successful nests whereas low call rates are associated with failures caused by nest predation; (7) call rates show a positive relationship with clutch and brood size; (8) call rates and types did not vary with different potential predators; and (9) call rates and types used by males and females are related to their respective roles in the nesting cycle. Several hypotheses are discussed that might explain changes in call rates through the nesting cycle.     

Calling as an ownership convention on pied wagtail territories

Winter feeding territories were more profitable to owners than intruders. Even when intruders landed on a territory undetected they failed to achieve a profitable feeding rate because they fed in areas recently depleted by the owner. For intruders therefore, ownership was a sign of unprofitability. Intruder scalled ‘chis-ick’, apparently to find out if a territory was occupied. Owners replied ‘chee-wee’ and this was usually sufficient to cause the intruder to retreat. Only owners called ‘chee-wee’. Owners reacted more aggressively to tape-recorded broadcasts of the ‘chee-wee’ call played on their territory than to broadcasts of ‘chis-ick’.     

Ready to Fight: Reliable Predictors of Attack in a Cooperatively Breeding,Non‐Passerine Bird

Signal reliability is a major focus of animal communication research. Aggressive signals are ideal for measuring signal reliability because the signal referent – attack or no attack – can be measured unambiguously. Signals of aggressive intent occur at elevated rates in aggressive contexts, predict subsequent aggression by the signaler, and elicit appropriate responses from receivers. We tested the ‘predictive criterion’ in smooth‐billed anis, Crotophaga ani, by broadcasting one of two playback types (‘ahnee’ calls only or ‘ahnee + hoot’ calls), presenting a taxidermic mount, and observing the animals’ behavior. Based on the hypotheses that ‘hoot’ calls and ‘throat‐inflation’ displays signal aggressive intent, we predicted that they would be associated with attack, and that signaling rate would increase over the time period leading up to an attack. Indeed, both hoots and throat‐inflation displays reliably predicted attack. The second prediction, that signaling rate increases in the time leading up to attack, was strongly supported for throat‐inflation displays, which increased over the pre‐attack period in both treatments. Hoots increased over the pre‐attack period in ahnee playbacks but not in ahnee + hoot playbacks. Hierarchical signaling systems are characterized by early, less‐reliable predictors of attack, and later, more reliable predictors of attack. During both natural and simulated interactions, the more‐reliable throat‐inflation display tended to precede the less‐reliable hoot call, suggesting that this signaling system is not hierarchical. In a comparison of 11 putative signals of aggressive intent in birds, the throat‐inflation display had the second highest mutual information (reduction in uncertainty) among visual signals and non‐passerine signals while hoots had below‐average mutual information. Natural observations indicate that both hoots and throat‐inflation displays occur in the context of aggressive between‐group encounters, and hoots also occur during within‐group interactions. Throat‐inflation displays appear to be reliable indicators of aggressive intent, but the function of hoot calls is less clear.     

Agonistic Encounters in a Cricket Frog (Acris crepitans) Chorus: Behavioral Outcomes Vary with Local Competition and within the Breeding Season

Male cricket frogs ( Acris crepitans ) gather in breeding choruses and call to attract females. The call also serves to maintain an individual's calling space. When an intruder invades a resident male's space, the resident will display one of three behavior patterns. He will either attack the intruder, abandon calling, or tolerate the intruder's presence and continue calling. We simulated an intruder by broadcasting a stimulus from a speaker and categorized the response of the resident male. We recorded social variables relevant to local competition among males, specifically, local caller density (within 2 m) and the amplitude of the nearest neighbor's calls. In addition we recorded the size of the subject, the amplitude of the stimulus, and the time of night and the season. We used a multivariate approach to assess the relative importance of these variables. Local caller density and nearest neighbor call amplitude were the best predictors of a resident's response to the intruder; higher levels of local competition resulted in more males tolerating the intruder. In addition, behavior changed over the season: males were more likely to abandon or ignore the intruder early in the season and more likely to attack later in the season. No other variables were related to the outcome of the agonistic encounter. This study suggests that the most important variables impinging on a male's decision to fight, flee, or ignore an intruder are influences external to the individuals involved in the conflict.     

Spatial aspects of nest defence by pumpkinseed sunfish (Lepomis gibbosus): Stimulus features and an application of catastrophe theory

Using fixed stimulus dummies as territory intruders, we studied the spatial distributions of the aggressive responses made by nesting male pumpkinseed sunfish. Results replicated over three field seasons indicate that males defend two territory boundaries concurrently, depending on whether an intruder is approaching (“the defence perimeter’) or withdrawing (‘the attack perimeter’). This finding supports Zeeman's (1976) ‘cusp catastrophe model’ of nest defence. In addition, the defence perimeter remains constant over the breeding cycle while the attack perimeter varies. We discuss the ecological costs and benefits of this variation and present a motivational interpretation of the cusp catastrophe model. By varying the speed at which dummies intruded into nests, we determined that males react with a fixed latency to intruders at a fixed distance from the nest. Finally, the spatial distributions of male defence responses were partially determined by the location of the nest rim, but were unaffected by dummy size (contrary to Zeeman's model) or posture.     

Additional Information on the Relationship between Intra-specific Aggression and Anti-predator Behaviour in the Three-spined Stickleback,Gasterosteus aculeatus

18 non-breeding three-spined sticklebacks from a heavily predated site a further 24 fish from a predator-free site were screened for their responses to a realistic model pike. The behaviour of the same fish when confronted with a territorial intruder during the breeding season was also recorded, the two tests being carried out in random sequence and with an interval of at least 6 weeks. Multivariate analyses identified major behavioural dimensions in the pike test and in the intruder test, summarized by the labels ‘boldness’ and ‘aggression’ respectively. Sticklebacks from the heavily predated site were both less bold and less aggressive than those from the unpredated site. Levels of aggression for individuals within each population were positively correlated with their boldness scores, but only for the period when the subjects were recovering from a simulated predatory attack.     

Male territoriality,fighting and assessment of fighting ability in the Australian frog Uperoleia rugosa

Males of Uperoleia rugosa form aggregations around the breeding pond. Within the aggregations, each calling male maintains an exclusive zone in which it does not tolerate another calling male. However, attacks on silent intruders, satellite males or females were never observed. Territorial males warned calling intruders by changing from advertisement calls to encounter calls (a display); if the intruder persisted in calling the resident would either attack or retreat and call elsewhere. Fights were usually won by the heavier males, and males of similar weight were more likely to fight, whereas males retreated from opponents that were substantially heavier. Calling was energetically costly and territorial males lost weight and thus their fighting abilities declined, whereas silent satellite males gained weight and were able to oust lighter territorial males. There was thus a dynamic interchange between territorial and satellite males depending on their relative weights. The playback of recorded advertisement calls showed that males can assess each others' fighting abilities on the basis of their vocalizations. Males retreated from a loudspeaker broadcasting the calls of a larger male but they attacked the loudspeaker if the calls of smaller or similar sized males were played (N=39, P<0·001, one-tailed binomial test). Dominant frequency was probably the call parameter used in assessment as it was correlated with the weight (r=−0·71, P<0·001, N=100) and hence the fighting ability of the caller, but this hypothesis needs further testing.     

A conflicting-tendency model of spider agonistic behaviour: Hybrid-pure population line comparisons

Contests over web-sites in the spider, Agelenopsis aperta, have been simulated by a model in which the next act of a spider is determined by the value of two ‘drives’ or ‘tendencies’, to attack and to flee. Such a model can explain the fact that F₁ hybrids between two geographical races of A. aperta are not intermediate between their parents, but are more aggressive than either. The model also accounts for the following features of the observed contests: (1) contests consist of a series of bouts, during which the spiders perform increasingly aggressive acts; (2) contests are usually won by the larger spider; (3) if sizes are equal, contests are usually won by the owner; (4) contests over excellent-quality sites are more costly than over poor-quality sites; and (5) transition probabilities for de-escalation, matching and escalation with respect to prior acts show site quality and weight bias trends. It is argued that a two-tendency model is the simplest that would account for these observations. The model also accurately simulates spider behaviour in agonistic contexts not used in its development (i.e. round-robin contests involving opponents from different populations, and a group of contests involving unequally-matched hybrid opponents).The main discrepancy between the model and observed encounters is that the model predicts that, if the owner is larger, the cost of a contest will not vary with the value of the disputed site. This is because the smaller intruder, having no knowledge of site quality (Riechert 1984) is expected to withdraw early in the contest, regardless of site quality. In the actual contests over excellent sites, estimated costs are higher even when the owner is larger and the intruder loses. The change in intruder behaviour associated with site value occurs following the first series of actions, but within the first bout of the interactions: our analyses indicate that any information conveyed by the owner concerning site value is not through overt behaviour (i.e. through the frequencies with which specific acts are performed).     

Characterization of self-compatibility in sweet cherry varieties by crossing experiments and molecular genetic analysis

Self-compatibility is a major breeding objective in sweet cherry. The identification and characterization of new sources of self-compatibility will be useful for breeding and research purposes. In this work, self-compatibility of four local Spanish sweet cherry varieties was investigated by crossing experiments and molecular genetic analysis of two self-incompatibility loci. Crossing experiments included self- and cross-pollinations in the laboratory followed by microscopic observation of pollen tube growth and fruit set assay in the field. After crossing experiments, two accessions, ‘Son Miró’ and ‘Talegal Ahín’, were self-compatible while the other two were self-incompatible. Inheritance of S-locus and microsatellite EMPaS02 (linked to self-compatibility, Sc) were investigated in self-pollination progeny of both self-compatible genotypes. Results indicate that self-compatibility in ‘Talegal Ahín’ is similar to self-compatibility described in sweet cherry ‘Cristobalina’ and may be caused by the same mutation. That is a pollen part mutation not linked to the S-locus but linked to microsatellite EMPaS02 in cherry LG3. In ‘Son Miró’ self-compatibility seems more complex, affecting pollen and style function, and probably involving more than one mutation not described previously in sweet cherry. Together with ‘Cristobalina’, the newly described self-compatible varieties ‘Son Miró’ and ‘Talegal Ahín’ confirm the existence of unique self-compatible plant material in local germplasm from Spain that should be conserved and characterized for its use in breeding and research.     

Information transfer during cricket frog contests

We investigated information transfer during vocal interactions between cricket frogs, Acris crepitans, with a specific focus on information about size and intention. In response to opponents, cricket frogs alter both temporal and spectral (frequency) aspects of their calls. Previous work suggests that males use dominant frequency, which is correlated with size, to provide information about fighting ability, and use temporal call characters, which are independent of size, to provide information about aggressive intent. We tested this hypothesis by investigating the relationship between call characters and contest behaviour. We presented a focal male with a simulated opponent, and categorized his behavioural response as attack, abandon or tolerate. We found that information about opponent size does not appear to influence a male's decision to fight, flee or tolerate an intruder. Whether or not males use the size information that is encoded in call frequency remains unclear. In contrast, changes in call frequency, which are not correlated with size, predicted the outcome of simulated contests, suggesting that male cricket frogs signal information about agonistic intent. Temporal call characters indicated whether or not a resident tolerated an opponent, but they did not predict contest outcome (attack versus abandon). Furthermore, the difference between the temporal call characters of a focal male and the simulated opponent predicted whether the resident tolerated the opponent. We suggest, therefore, that temporal call changes may be a cooperative signal designed to facilitate assessment of size through physical contact in wrestling bouts. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Effects of residence,aggressive experience and intruder familiarity on attack shown by male mice

Two groups of 12 mature male Swiss Morini strain mice were matched for aggressiveness on the basis of their response to anosmic docile male intruders after 24 h individual housing in a large defensible cage. One group was subsequently individually housed without disturbance for a further 12 days whereas the second group was exposed to a new anosmic intruder every 3 days over this same period. All mice were subsequently retested against anosmic intruders for a 10 min period. Animals which had had the opportunity to fight repeatedly showed more attack than individuals lacking such experience. A second experiment contrasted the responses of dominant isolates to the same or a different anosmic intruder, 10 min after a successful attack. Familiar intruders were attacked less vigorously than unfamiliar mice. It seems likely that the defeated mouse becomes less potent as a stimulus eliciting attack as the resident becomes habituated to it. Conversely, unfamiliar intruders evoke aggressive reactions by residents despite the prolonged fights they had had before final testing. These experiments provide little support for the appetence view of aggression in mice (i.e. the view that aggressiveness is augmented by depriving the animal of the opportunity to fight) or the view that this phenomenon is simply a consequence of “social deprivation”.     

Confusion over residency and the escalation of damselfly territorial disputes

Territorial contests among male Calopteryx maculata (Odonata) damselflies normally involve one or a few short (4–10s) pursuits of an intruder by the resident. However, in a 3-year sample of 2005 bouts, 18·5% involved escalated, spiralling aerial chases that were significantly longer, some lasting several hours. These escalations usually occurred when two males shared the same territory but were rare in contests with intruders or with neighbouring territorial males. Escalations were most probably due to situations in which two males accidentally became residents in the same territory. This removed the ‘residency’ asymmetry that appears to settle most territorial disputes in these damselflies. Using moving territories (oviposition sites), the ‘confusion over residency’ hypothesis for escalation was tested experimentally. Bouts between males on adjacent, equally sized sites were escalated significantly in duration and intensity when their territories were merged. These experiments also showed that two alternative hypotheses, the ‘male aggressiveness’ and ‘valuable resource’ hypothesis, were not important general explanations of the escalation of contests.     

Parental investment in nest defence by stonechats (Saxicola torquata)

Breeding stonechats (Saxicola torquata) made mixed sequences of two calls when a human intruder entered territories. ‘Whits’ are modulated notes with a small frequency range, and in laboratory tests caused nestlings to stop begging. ‘Chacks’ cover a wide range of frequencies, and in the field were combined with flights made so as to distract an intruder from the nest. On average male and female call-rates were similar, but varied greatly according to the intruder's distance from the nest, and at different stages of the nesting cycle. Rates increased rapidly after hatching, and this correlated most closely with the cumulative total of parents' visits to feed nestlings. This suggests that the level of defence may be adjusted to the value of the offspring to their parents. Call-rates declined about one week after fledging. A smaller peak by some pairs at the start of incubation was apparently related to probable poor condition after a previous breeding attempt, and after laying large clutches. Rates of Whits were higher at nests with larger broods, up to an asymptote, but rates of Chacks were independent of brood size. Birds suffering nest-predation showed lower call-rates before the event than equivalent successful birds, suggesting that the calls do reduce the risk of predation.     

Temporal and structural features of infant calls in relation to caregiving behaviour in common marmosets,Callithrix j. jacchus

The role of infants in the communal rearing system of marmosets and tamarins has received little attention although their demands may not be complied by inadequate helpers. As acoustic signalling plays an important role in their social communication, it is expected to be an effective way for infants to signal their demand for care. As a first approach, this study focussed on the use and quality of ‘tsik’ calls given by infants of common marmosets (Callithrix j. jacchus) in the context of caregiving. The acoustic and non-acoustic behaviour of a group of common marmosets were recorded during the first two weeks of the life of two infants (twins). Vocalizations were classified, and the structure of the infants' ‘tsik’ calls was quantified by measuring specific signal parameters. Infants gave ‘tsik’ calls in close temporal relation to caregiving behaviour. Specific parameters covaried with the subsequent behaviour of potential caretakers. Values of call duration, start frequency, peak frequency, and range were lower when infants' calls were followed by no reaction or an approach of group members, whereas these parameters showed higher values when followed by a transfer of infants. Our results suggest that the actual demand of infants is encoded in the variations of ‘tsik’ calls.     

Wrestling in the red-spotted newt (Notophthalmus viridescens): resource value and contestant asymmetry determine contest duration and outcome

When an unpaired male red-spotted newt encounters a male and female engaged in amplexus courtship, he attempts to displace the amplectant male, or ‘owner’, by wrestling with him. In this study, it was found that the amount of time an intruding male invests in wrestling is determined by both the size of the female, which seems to be a reliable indicator of her fecundity, and the length of the intruder relative to the owner. In 90% of all wrestling encounters observed, the owner retained the female. Successful displacement was rare and occurred only in encounters in which the intruder was at least as long as the owner. Similar contest behaviour in other amphibians is briefly discussed.     

Morphological and molecular characterization of somaclonal variations in tissue culture-derived banana plants

In this study, 40 000 tissue culture-derived banana plants (vitroplants) at different growth stages, i.e. acclimatization, nursery and open field of banana (Musa spp.) cultivar ‘Grand Naine’ were screened for somaclonal variations using morphological investigations and molecular characterization. The total detected variants were grouped into 25 off-types (two of them died) in addition to the normal plant. Random Amplified Polymorphic DNA (RAPD) was carried out to study the differences among the normal cultivar ‘Grand Naine’ and its 23 variants using 17 arbitrary primers. Cluster analysis results revealed that ‘winged petiole’ and ‘deformed lamina’ were more related to the normal plant. However, ‘Giant plant’ and ‘weak plant’ related to each other and clustered with normal plant. According to principal coordinate analysis, most of the variants were aggregated nearly, whereas ‘variegated plant’ was separated apart from the other variants. This may reflect the genetic difference between ‘variegated plant’ and the other variants. The results obtained from both molecular and morphological analyses were in contiguous with better resolution when using the PCOORDA analysis than cluster analysis. Thus, it can be said that molecular markers can be used to eliminate the undesirable somaclonal variants from the lab without additional culture of the vitroplants in the field in order to save time and efforts.     

EQUALITY AND THE TREATMENT‐ENHANCEMENT DISTINCTION

In From Chance to Choice, Allen Buchanan, Dan Brock, Norman Daniels and Daniel Wikler propose a new way of defending the moral significance of the distinction between genetic treatments and enhancements. They develop what they call a ‘normal function model’ of equality of opportunity and argue that it offers a ‘limited’ defence of this distinction. In this article, I critically assess their model and the support it (allegedly) provides for the treatment‐enhancement distinction. First, I argue that there is a troubling tension in the normal function model. Secondly, I argue that neither of the rationales invoked by Buchanan et al. really serves to justify this model or the results they seek to derive from it with respect to the significance of the distinction between treatments and enhancements.     

The first valid publication of the Linnaean Mnium triquetrum (Musci) and other new names for mosses proposed by N. Jolyclerc

A brief biographical sketch of Nicolas-Marie-Thérèse Jolyclerc (1746–1817), the French naturalist and adherent of C. Linnaeus, is presented. The publication history of his Système sexuel des végétaux, the first French version of the Linnaean sexual system of the plants is outlined. Apart from the well-known editions of this work of 1797, 1798 and 1810, a neglected edition of 1803 is described, and it is shown that this is not a pirated edition as has been suggested. A list of fourteen new combinations in mosses proposed by N. Jolyclerc in the 1803 edition of Système sexuel des végétaux is presented and some names are briefly annotated. Additionally, it is shown that Mnium triquetrum, basionym of the well-known moss species Meesia triquetra, was validly published for the first time by N. Jolyclerc in 1803, not by H. E. Richter in 1839 as is commonly accepted. Accordingly, the correct author citation for Meesia triquetra should be ‘(L. ex Jolycl.) Ångstr.’, not ‘(L. ex Richt.) Ångstr.’ The lectotype of Mnium triquetrum L. ex Jolycl. is selected from the Linnaean herbarium (LINN).     

How does a parent respond when its mate emphasizes the wrong role? A test using a monogamous fish

The convict cichlid fish, Archocentrus nigrofasciatus, is biparental: the male spends the majority of his time defending the territory and the female spends much of her time close to the offspring. Under natural conditions, this separation into sex-typical roles is somewhat blurred as males do spend some time with the offspring and females do attack intruders. Here we tested whether an individual selects a parental role based on the location (i.e. parental role) of its mate. For example, do females emphasize offspring care because the male is away from the offspring? Will males be more likely to care for the offspring when the female is away from the offspring? We manipulated the location of one parent by placing it in a transparent plastic box, either near the offspring or at the far end of the tank near a clear plastic compartment that held a conspecific male intruder. We tested both male and female parent under the following four conditions: boxed mate near offspring with no intruder present, boxed mate near offspring with intruder present, boxed mate near intruder compartment with no intruder present, and boxed mate near intruder compartment with intruder present. We found that both parents spent more time with the offspring and less time attacking the intruder when the mate was positioned near the offspring and more time away from the offspring and more time attacking the intruder when the mate was near the intruder. Males were more affected by the location of their mates than were females and we concluded that males were attracted both to their mates' location and their offspring while females were mostly attracted to their offspring. Overall, the location of the mate had little effect on the types of aggressive activities used against the intruder. We did find that males increased their aggression towards boxed females when they were positioned far from the offspring, whereas the aggressive behaviour of females towards boxed males when they were positioned near the offspring was ambiguous. We suggest that males in particular enforce the separation of sex-specific parental roles via this aggression.     

The agonistic and sexual functions of vocalizations of male brown-headed cowbirds,Molothrus ater

Functional analyses of three vocalizations of male brown-headed cowbirds, Molothrus ater, are presented and it is shown that two of these function as songs. Perched song (PS) is given by stationary birds and is used over short distances in male-male aggression and female courtship. The ‘flight whistle’ (FW) is a multisyllabic, usually pure tone, vocalization and the ‘single syllable flight call’ (SS) a monosyllabic pure tone. FWs and SSs are given occasionally by perched males and repetitively by flying males. Field playbacks of FWs showed that male cowbirds responded in the agonistic way most passerines respond to playback of conspecific song. Playbacks of the female's chatter call demonstrated that males of all three subspecies use FWs and SSs, but almost never PSs, to communicate with a female as they approach her. Thus FWs and SSs are used to communicate over long distances with both males and females. FWs are also used in one critical short-distance context as they are significantly more likely than PSs to be given just before copulations in nature, at least in our western study area. By contrast, studies of captive eastern cowbirds showed that PSs regularly precede copulations whereas FWs rarely do so. These different results for western and eastern birds may be an artefact of captivity and/or a result of geographical variation in behaviour.