Research Article
Mate guarding and social mating system in male common chameleons (Chamaeleo chamaeleon)
- Mariano Cuadrado
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- 26 November 2001, pp. 425-435
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Male common chameleons Chamaeleo chamaeleon actively followed the female closely as a typical mate guarding behaviour during the mating season. Guarding behaviour and spatial organization were studied in 34 radio-tagged males in a population in southern Spain during three summers. Timing and extent of guarding were highly variable among males. Overall, guarding started on 16 August (range 28 July–5 September, n=34) and ended on 4 September (range 21 August–15 September, n=24) lasting 13.4 days (range 0–46 days, n=30). Males showed a polygynous serial-type mating system where males sequentially guarded (and mated) several females during the breeding season but only one at a time. The number of guarded females (mean 2.8, range 0–8, n=34) and number of mates per male (mean 0.7, range 0–3, n=34) were also highly variable among males. In general, guarding males abandoned the females soon after mating in association with clear signs that females were no longer receptive. Males showed a remarkably complex spatial organization in close association with the home range of guarded females. Guarding behaviour resulted in territorial behaviour, and ‘stable’ as well as ‘mobile’ territories were recorded. In ‘stable’ territories, males defended non-overlapping areas where the home range of one female or more was included. Although monogamy was the predominant mating system recorded in this population, social polygyny occurred in all study years. Polygynous groups were only feasible because females aggregated in small-sized areas without apparent symptoms of intrasexual competition. Breeding asynchrony of females seems to influence this complex mating system as more competitive (large) males mated with large (early reproducing) females and small (late reproducing) ones (cf. Cuadrado, 1999). Most aggressive interactions were male–male chasing of a solitary intruder by a guarding male. A removal experiment of guarding males revealed that newcomers were small-sized males, suggesting that an important fraction of males were acting as floaters searching for mating opportunities. Male characteristics (especially size and body weight) were the main factors influencing the spatial organization and the reproductive success of males in this population.
Debate on the authenticity of Pseudonovibos spiralis as a new species of wild bovid from Vietnam and Cambodia
- John H. Brandt, Maurizio Diolo, Alexandre Hassanin, Richard A. Melville, Link E. Olson, Arnoult Seveau, Robert M. Timm
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- 26 November 2001, pp. 437-444
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In the February 2001 issue of the Journal of Zoology a paper was published by Robert Timm and John Brandt (2001) that provided a new identification for two bovid frontlets held in the Kansas Natural History Museum. The frontlets had been collected in 1929 from Suoi Kiet, Binh Tuy Province in Vietnam and had been previously identified as kouprey Bos sauveli. Timm and Brandt re-identified the two frontlets as adult male and female representatives of the recently described spiral-horned ox Pseudonovibos spiralis Peter & Feiler, 1994.
Prey classification by Portia fimbriata, a salticid spider that specializes at preying on other salticids: species that elicit cryptic stalking
- Duane P. Harland, Robert R. Jackson
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- 26 November 2001, pp. 445-460
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Portia fimbriata from Queensland, Australia, is an araneophagic jumping spider (Salticidae) that includes in its predatory strategy a tactic (cryptic stalking) enabling it to prey effectively on common sympatric salticids from other genera. Using standardized tests in which only optical cues were available (prey enclosed in small glass vial within large cage), the reactions of P. fimbriata to 114 salticid species were investigated. Except for Myrmarachne spp. (ant mimics), all salticids tested triggered cryptic stalking by P. fimbriata. This included not only sympatric, but also allopatric, salticids. The salticid on which P. fimbriata most commonly preys in nature is Jacksonoides queenslandicus, but cryptic stalking was triggered by salticid species with considerably different appearances, including beetle mimics, species with unusual body shapes and species with a wide variety of camouflaging markings. Portia fimbriata was also tested with lycosid, clubionid, theridiid and desid spiders and with flies and ants, but none of these arthropods triggered cryptic stalking. Optical cues used by P. fimbriata for discrimination between salticid and non-salticid prey are discussed.
Eruption rates of the mandibular incisors of naked mole-rats (Heterocephalus glaber)
- B. K. B. Berkovitz, C. G. Faulkes
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- 26 November 2001, pp. 461-466
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Naked mole-rats Heterocephalus glaber are highly social hystricomorph rodents endemic to the arid regions of East Africa. Living entirely underground, they excavate an extensive system of tunnels with their continuously growing extra-buccal incisors. The majority of the burrow is composed of foraging tunnels, constructed in search of the underground roots and tubers that form their staple diet. Subterranean foraging imposes risks associated with not finding food, and a high energetic cost, and in addition could be limited by the rate of wear of the incisor teeth. The aim of this study was to determine eruption rates for mandibular incisors in naked mole-rats to investigate whether they showed evidence of faster eruption as an adaptation to their subterranean niche and mode of digging. Impeded eruption rates (when the teeth were left in normal occlusion) were initially determined over a period of 1 week in a group of 12 naked mole-rats kept in laboratory conditions. Unimpeded eruption rates were then monitored in the same animals by cutting one mandibular incisor free of occlusion thrice weekly, and measuring growth over a 2-week period. Mean daily impeded eruption rates were c. 228.7±17.5 μm, while mean daily unimpeded rates were higher at c. 624.7±10.4 μm. While impeded eruption rates under laboratory conditions may be an underestimate of the rates occurring in the wild, unimpeded rates (usually about twice the impeded rate) indicate the maximum potential eruption rates of the teeth and are independent of functional activity. These data are not dissimilar from those determined for laboratory rodents and lagomorphs suggesting that the naked mole-rat does not display unusual patterns of eruptive behaviour. Tooth wear resulting from their subterranean lifestyle may be offset as a limiting factor by the social behaviour of the naked mole-rat, where digging activity is distributed among a large workforce during favourable burrowing conditions.
Middle ear structures in fossorial mammals: a comparison with non-fossorial species
- M. J. Mason
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- 26 November 2001, pp. 467-486
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A large data set, comprising certain measurements of middle ear structures in mammals, was compiled both from measurements made by the author and from the literature. Parameters of the middle ear apparatus believed to be important to audition were compared between fossorial and non-fossorial species, in an attempt to identify general trends among fossorial groups. Although their tympanic membranes are not of unusual size, many fossorial mammals possess enlarged stapes footplate areas, resulting in low anatomical area ratios. Low anatomical lever ratios are also common, and the reduction or loss of middle ear muscles seems to be a consistent trend. These characteristics might be associated with poor sensitivity to airborne sound. Other features of the middle ear apparatus are more variable both between and within fossorial families. The middle ear ossicles of fossorial rodents, talpid moles and some golden moles were not found to differ in mass from those of non-fossorial mammals of similar body size. The similar ossicular morphologies of these animals suggest convergent adaptation towards a subterranean environment, but the middle ear structure alone does not seem to explain the restricted hearing range observed in certain of these species. Some genera of golden moles possess extraordinarily hypertrophied auditory ossicles, which, relative to body mass, are the largest of all mammals for which data are available. These ossicles seem to be adaptations towards a form of inertial bone conduction, used for the detection of substrate vibrations. In stark contrast, the marsupial mole Notoryctes has particularly small ossicles. The unusual middle ear structures of this animal may well be degenerate.
Habitat use by wood mice (Apodemus sylvaticus) in a changeable arable landscape
- Françoise H. Tattersall, David W. Macdonald, Barbara J. Hart, Will J. Manley, Ruth E. Feber
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- 26 November 2001, pp. 487-494
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Wood mice Apodemus sylvaticus are potentially useful indicators of change in arable ecosystems. Here we focus on changes resulting from removal of land from arable production under the set-aside scheme. Wood mice were radio-tracked to compare: (a) their use of set-aside, crop and hedgerow before and after harvest; (b) set-aside configured as margins and as a 3 ha block; (c) cut and uncut 20-m wide set-aside margins. Males had larger home ranges, and were more mobile than females. Ranges were larger, and animals more mobile, before harvest than afterwards. There were no differences in range sizes of breeding and non-breeding animals after harvest, suggesting that changes in habitat use were not a function of cessation of breeding. Before harvest, wood mice used habitats within their ranges at random, and their ranges contained a high proportion of cropped area. After harvest they preferred hedgerow and avoided margin set-aside within their ranges, but did not similarly avoid the set-aside block. The proportion of cropped area within their ranges decreased after harvest, and the proportion of margin increased. Our evidence suggests wood mice avoided using the area adjacent to the hedgerow, perhaps to avoid predators. Uncut set-aside patches were favoured and cut patches avoided, possibly in response to differences in food availability and levels of protection from predators. These results confirm that wood mice are useful indicators of change in arable landscapes.
Host–parasite interactions in natural holes: marsh tits (Parus palustris) and blow flies (Protocalliphora falcozi)
- Tomasz Wesołowski
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- 26 November 2001, pp. 495-503
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Infestation of marsh tit (Parus palustris, Paridae, Passeriformes) broods by bloodsucking larvae of Protocalliphora falcozi Séguy 1928 (Calliphoridae, Diptera) was studied over an 8-year period in a population breeding in natural holes, in the primeval forest (Białowieża National Park, eastern Poland). Overall 54% of 222 marsh tit nests were parasitized. Prevalence (27–88%) significantly changed across years. This variation did not depend on the timing of the marsh tit breeding season, or winter/spring temperatures. Frequency of infestation did not depend on forest type or hole attributes. Infestation intensity was rather low (median 8, max. 75 flies/nest, 85% of nests with <3 flies/young). Intensity and prevalence were weakly, but positively, correlated. Larger broods contained significantly more blow flies, but per nestling load did not depend on brood size. No effect of infestation on nestlings was recorded – their mortality did not increase, nor was fledging delayed. In response to the presence of blow flies, parents apparently attempted to increase their feeding rate. There was a clear reproductive cost: 60–63% of female and 68–69% of male marsh tits that had none up to eight blow flies, survived to the next spring, but only 34% of females and 44% of males with more than eight flies/brood did so. It is suggested that the small clutch size of marsh tits could have evolved, inter alia, to reduce the fitness costs of ectoparasites.
A spatial analysis of a population of red fox (Vulpes vulpes) in the Dutch coastal dune area
- Jasja J. A. Dekker, Alfred Stein, Ignas M. A. Heitkönig
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- 26 November 2001, pp. 505-510
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The red fox Vulpes vulpes is usually classified as being territorial, dispersing or transient. Past studies have focused almost exclusively on territorial or dispersing foxes, leaving transient foxes out of the analysis. In this paper, we present spatial-statistical methods for the classification of free-ranging foxes, using 95% fixed kernels and 100% minimum convex polygons. By means of these procedures we classify individual foxes on the basis of their spatial behaviour, using home-range size and home range shift. Also, we make a methodological comparison between these classification procedures and interpret the composition of these classes ethologically. The procedures apply to a sample of 24 foxes, radio-tracked in the dune area of the Netherlands from January 1997 to June 1999. We analysed size of home range and successive 3-month overlap using a geographical information system (GIS). Classifying the sample using 95% fixed kernel home ranges resulted in two classes of foxes: a class of 20 territorial foxes with relatively small home ranges (<250 ha), and a class of four dispersing and transient foxes with relatively large home ranges (400–600 ha). This study shows that a fox population can be divided into different classes of individuals in a quantitative statistical way, honouring measured characteristics. This is a clear extension of more informal ways relying on expert judgement applied so far.
Non-random pairing with respect to past breeding experience in the common toad (Bufo bufo)
- C. J. Reading
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- 26 November 2001, pp. 511-518
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A 20-year study of a common toad Bufo bufo population in south Dorset, U.K. was carried out between 1980 and 1999. On average, c. 33% of all experienced males and females arriving each year did so during the first quarter of each breeding season with only 18% arriving during the last quarter. Conversely, a slightly higher proportion (27%) of all inexperienced (first-time breeders) males and females arrived during the last quarter season than during the first quarter (22%). In 18 of the 20 years, the proportion of inexperienced females found paired with inexperienced males was significantly higher than could be expected from the proportion of inexperienced males in the pond, suggesting that selective pairing was occurring. One potential effect of pairings between inexperienced male and female toads, coupled with the sex difference in the age of attaining sexual maturity, might be a reduction in the incidence of sibling matings (inbreeding) in the common toad. This is particularly relevant in a species that is both highly philopatric and which congregates in large numbers every spring to breed in very localized areas.
Intraspecific interactions among oophagous tadpoles (Chirixalus eiffingeri: Rhacophoridae) living in bamboo stumps in Taiwan
- Yeong-Choy Kam, Yahn-Jauh Su, Jin-Liang Liu, Yao-Sung Lin
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- 26 November 2001, pp. 519-524
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We studied the intraspecific interactions among oophagous Chirixalus eiffingeri tadpoles that occupied the same water-filled bamboo stumps at Chitou, Taiwan. We monitored the growth of newly-hatched tadpoles in unoccupied and occupied bamboo stumps in the field where the latter contained large tadpoles that hatched from earlier clutches. The growth of the small, late-hatching tadpoles in occupied nests was suppressed by the presence of large, early-hatching tadpoles. However, small tadpoles that were physically separated from large tadpoles in a perforated container grew at about the same rate as small tadpoles living in pools without large tadpoles. Thus, the slower growth of the late-hatching tadpoles was probably caused by behavioural interference competition with the early-hatching tadpoles. In the laboratory, we kept large and small tadpoles together in containers and did not feed them for 6 days. The large tadpoles did not cannibalize the small tadpoles. Although large tadpoles may scavenge dead tadpoles, the effects of scavenging on growth were negligible. Interactions among cohabiting C. eiffingeri tadpoles are similar to those among the oophagous tadpoles of several hylid species that use phytotelmata. Results suggest that behavioural interference competition is the principal type of intraspecific interactions among the oophagous, non-predatory tadpoles of hylid and rhacophorid frogs.
Feeding ecology of the loggerhead turtle Caretta caretta in the western Mediterranean
- J. Tomas, F. J. Aznar, J. A. Raga
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- 26 November 2001, pp. 525-532
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We studied the feeding ecology of juvenile loggerhead turtles Caretta caretta in the western Mediterranean based on the contents of the digestive tract of 54 turtles (range of CCL: 34–69 cm) seized in Barcelona (Spain) in 1991. Turtles had been captured in fishing trawls, but specific information about dates and localities is not available. Despite this limitation, we obtained interesting evidences about the foraging strategies of loggerheads, with potentially important conservation implications. We report 33 new taxa in the diet. Results indicated that western Mediterranean loggerheads feed in an opportunistic way. Numerically, fish made up the most important prey group, followed by pelagic tunicates, crustaceans, molluscs and other invertebrates. The importance of fish as a food resource has been rarely reported, and several lines of evidence indicated that fish were possibly consumed as discarded by-catch. This raises the question over whether or not western Mediterranean fisheries are an important food source for juvenile loggerheads. The number and diversity of prey increased with turtle size, this may reflect the lack of prey selectivity of juvenile loggerheads coupled with a higher retention of food remains in larger turtles. Discounting prey that could be consumed as discarded by-catch, dietary data suggest that most, if not all, loggerheads of our sample were captured in neritic habitats. However, many turtles contained remains of both pelagic and benthic-demersal prey. These observations support the existence of an intermediate neritic phase in loggerheads' developmental shift from pelagic–oceanic to benthic–neritic foraging habitats, as previously suggested. During this phase, loggerheads would feed upon both pelagic and benthic prey.
Seasonal and interspecific evaluation of predation by mammals and birds on the introduced red swamp crayfish Procambarus clarkii (Crustacea, Cambaridae) in a freshwater marsh (Portugal)
- Alexandra Marçal Correia
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- 26 November 2001, pp. 533-541
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The crayfish Procambarus clarkii is an introduced species in Portugal, becoming a new available resource that has possibly changed local trophic relationships. The purpose of the present study was to understand better how a naïve assembly of predators reacted to a recent invader. Predation on P. clarkii by nine species of mammals (Order: Carnivora) and six species of birds (Order: Ciconiiformes) in a marsh area in Portugal, was surveyed through the collection of faeces and pellets over a 24-month period. Seasonal variation in the consumption of this prey by different predators was evaluated, and estimation made of size and weight of P. clarkii consumed by them. Procambarus clarkii was readily preyed upon by four species of mammalian carnivores and five species of ciconiiform birds. The consumption of P. clarkii varied seasonally in both years with a trend between seasonal exploitation and its population structure and availability. Generally, the use of P. clarkii by all predators was more intense in spring, summer and autumn than in winter, and was directed towards larger and heavier individuals. These findings result from the population dynamics presented by P. clarkii in the marsh area studied here, since larger and heavier individuals were more available in spring, summer and autumn. Procambarus clarkii is an important food resource for mammals and birds and it seems to play a key role in the trophic interactions of the riparian and terrestrial communities of the marsh.
The reproductive biology of Ptomascopus morio, a brood parasite of Nicrophorus
- Stephen T. Trumbo, Masahiro Kon, Derek Sikes
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- 23 July 2002, pp. 543-560
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Nicrophorine beetles (Nicrophorus and Ptomascopus spp.) use small carcasses as a food source for young, a breeding ecology distinct from other silphid beetles. While adaptations to the use of small carcasses are well known for Nicrophorus (emitting sex pheromone, burying, rounding and removing hair from carcasses, regulating brood size, regurgitating to young, and preventing predation), there is little information regarding its sister group, Ptomascopus. Like Nicrophorus, Ptomascopus morio males emit pheromone to attract females. In the absence of carrion competitors Ptomascopus morio parents were found to stay with a carcass and their brood for up to 10 days. We tested six hypotheses to examine whether young benefit from this long period of parent–offspring contact. (1) There was no evidence that parents buried or otherwise pre-empted carcasses to reduce competitive pressure. (2) We found no evidence that parents influenced the decomposition of the carcass. This was supported by experimental manipulations in which brood production (number of larvae and total brood mass) was no greater on carcasses on which parents were present than on carcasses not ‘prepared’ by parents. In addition, the carcass was not rounded and little hair was removed by the adults. (3) The presence of parents benefited the brood by reducing the negative effects of competition with carrion fly larvae. This likely resulted from predatory feeding by adult beetles. (4) Females adjusted clutch size to the size of the carcass. Parents, however, did not make a second adjustment in brood size after young reached the larval stage (filial cannibalism), as occurs in Nicrophorus. (5) Although parents were observed to open feeding holes in the carcass, this was not necessary for normal larval growth and survival. Parents were not observed to feed young directly by regurgitation. (6) Lastly, parents did not reduce predation on their brood when a conspecific intruder was present. These findings suggest that after the female parent adjusts clutch size to the size of the resource, the only parental benefit is clearing the carcass of fly larvae. Other differences with Nicrophorus include an extended period of oviposition (5 days) and less pronounced changes in ovarian mass and juvenile hormone titers in response to discovery of a carcass. In a field experiment in Kyoto, Japan, 17 of 21 broods of N. concolor during August contained larvae of P. morio. Mixed Nicrophorus–Ptomascopus broods were less common at other times of the year and when N. quadripunctatus occupied carcasses. In the laboratory, P. morio was able to parasitize 19 of 20 broods of N. concolor. The pattern of oviposition, the absence of explicit parental behaviours, and the interactions with N. concolor in the field suggest that Ptomascopus morio is a brood parasite of Nicrophorus.