United in adversity: Aridity and cold influence aggregation behaviour in a social lizard,Egernia stokesii

Sociality has evolved independently many times in a diverse array of animal taxa. While sociality in some invertebrates, birds and mammals is relatively well understood, complex social behaviour in Squamate reptiles is a comparatively recent discovery. The extent to which social behaviour is phylogenetically constrained, or free to respond to environmental conditions is a key question in understanding sociality. We sampled 74 aggregations involving 137 individuals of a social lizard (Egernia stokesii) from 13 sites across a 96 250 km² area showing a wide range of environmental conditions. Over all locations, 70% of the lizards were found in aggregations, supporting the previous suggestion that aggregation in this species is phylogenetically constrained. However, the size of aggregations was negatively related to spatial variation in aridity and daily variation in maximum temperature, suggesting that social behaviour also varied in response to harsh environmental conditions. Lizards aggregated more in arid areas and on cold days. Our results show that it is overly simplistic to regard social behaviour in E. stokesii as either phylogenetically constrained or environmentally determined. A more nuanced appreciation of the extent to which social behaviour is free to vary in response to environmental conditions improves our understanding of social behaviour in Squamate reptiles.     

Fine‐scale genetic structuring in a group‐living lizard,the gidgee skink (Egernia stokesii)

By constraining gene flow, group living and natal philopatry can result in fine‐scale genetic structure. Although the genetic structure of some group‐living lizards has been characterised, studies are few compared with those for group‐living bird and mammal species. The Egerniinae group of lizards exhibits a high diversity of social structures, making it a useful group for comparative studies of genetic structure across a broader range of social taxa. A well‐studied member of Egerniinae is Egernia stokesii, a lizard that forms long‐term pair bonds and stable social groups and exhibits natal philopatry and limited dispersal. Evidence exists for consistent E. stokesii social structure across seven close but disconnected rocky outcrops within a 40 × 10 km area. We used summary statistics, analysis of molecular variance, Bayesian clustering, and discriminant analysis of principal components to assess if E. stokesii exhibit a consistent pattern of fine‐scale genetic structure across the same seven outcrops. Due to E. stokesii social structure and constrained dispersal, we predicted significant genetic structuring – based on microsatellite markers – among outcrops. We found significant fine‐scale genetic structuring and evidence for two genetic clusters. We discuss features of E. stokesii biology and ecology that may explain our findings. Some rocky outcrops, and some social groups, contained lizards from both genetic clusters. An examination of the composition of mixed cluster social groups did not detect any notable patterns. Therefore, further work is necessary to identify how the observed patterns may have arisen. Future investigations in E. stokesii and other group‐living lizard species are likely to contribute greatly to our understanding of the genetic consequences of group living.     

Group Structure and Stability in Social Aggregations of White's Skink, Egernia whitii

Most lizards display relatively simple social systems, but more complex and stable social aggregations appear to be common in one lineage of Australian skinks, the Egernia Group. Previous studies on this lineage have focused on species inhabiting crevices in large and disjunct rocky outcrops. Here, we describe the social organization of White's skink, Egernia whitii, a burrowing species that inhabits rocky habitats in southeastern Australia. We examined social group size, composition and stability over two field seasons using a capture‐mark‐recapture study, behavioral observations and genetic analyses. Twenty‐four social groups, each comprising two to six individuals, were present at our study site, with 75% of lizards belonging to a social grouping. A higher proportion of adults than juveniles were part of a group, while more adult females belonged to a group than adult males. Groups generally comprised a single adult pair or an adult pair with juveniles. However, groups comprising one male with multiple females and multiple individuals of both sexes also were present. Groups were highly stable throughout the study, although individual group members were observed singly on half of all observations. Paternity analysis using four microsatellite loci revealed that juveniles within groups were closely related to adults in the group, with 38% living in groups with both parents and 71% in groups with at least one parent. Our data demonstrate the presence of complex sociality in a burrowing Egernia species and, together with previous studies, suggest that stable social organization is widespread across different habitats and phylogenetic groupings within the Egernia Group.     

Nesting in a thermally challenging environment: nest‐site selection in a rock‐dwelling gecko,Oedura lesueurii (Reptilia: Gekkonidae)

In egg‐laying species, maternal oviposition choice can influence egg survival and offspring phenotypes. According to the maternal‐preference offspring‐performance hypothesis, females should choose oviposition sites that are optimal for offspring fitness. However, in thermally challenging environments, maternal oviposition behaviour may be constrained by the limited availability of suitable oviposition sites. We investigated nest‐site selection in a nocturnal lizard [velvet gecko Oedura lesueurii (Duméril and Bibron)] that inhabits a thermally challenging environment in south‐eastern Australia. The viability of these gecko populations is critical for the persistence of an endangered snake species (Hoplocephalus bungaroides Wagler) that feeds heavily on velvet geckos. Female geckos chose nest sites nonrandomly, with 87% of nests (N = 30) being laid in deep crevices. By contrast, only 13% of clutches were laid under rocks, which were the most readily available potential nest sites. Nest success in crevices was high (100%), but no eggs hatched from nests under rocks. Temperatures in nest crevices remained relatively low and constant throughout the incubation period (mean = 22.7 °C, range 21.0–24.5 °C), whereas thermal regimes under rocks showed large diurnal fluctuations. Geckos selected crevices that were deeper, had less canopy cover, and were warmer than most available crevices; in 85% of cases, such crevices were used simultaneously by more than one female. The thermally distinctive attributes of nest sites, and their frequent communal use, suggest that nest sites are a scarce resource for female velvet geckos, and that the shading of rock outcrops through vegetation encroachment may influence nest success in this species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 250–259.     

<Emphasis Type="Italic">Drosophila</Emphasis> pupation behavior in the wild

We investigated pupa distributions of D. simulans, D. buzzatii, D. melanogaster, D. immigrans and D. hydei on a number of natural breeding sites. Pupae of all five species showed aggregated distributions, which prompted us to examine these aggregations in a more detail for two species that commonly co-occur in breeding sites, D. simulans and D. buzzatii. We found that pupae of both species tend to be aggregated in conspecific clusters. Subsequent experiments revealed that both species are attracted to the odors of other larvae, though only D. buzzatii differentiated between conspecifics and heterospecifics (they preferred conspecific). Furthermore, third instar larvae of both species preferred more alkaline substrates. Altogether, our results demonstrate that Drosophila species form conspecific pupa aggregations in natural breeding sites, and that pupation site selection depends on interactions among conspecific and heterospecific larvae and on chemical characteristics of the breeding sites.     

Habitat selection by dispersing yellow-headed blackbirds: evidence of prospecting and the use of public information

In migratory birds individuals may prospect for potential breeding sites months before they attempt to breed and should use the cues most predictive of future reproductive success when selecting a breeding site. However, what cues individuals use when prospecting and which cues are used in selecting a breeding site are unknown for most species. I investigated whether yellow-headed blackbirds (Xanthocephalus xanthocephalus) prospect for future breeding sites and whether they select breeding habitats based on food availability, male or female density, or the average number of young produced per female in the previous year. Although it is often assumed that migratory birds prospect for potential breeding sites at the end of the breeding season, I investigated this by recording all visits to sites early and late in the breeding season. I found that males and females who visited sites other than the site at which they bred were more likely to disperse than individuals only observed at the site where they bred, and that males and females were more likely to prospect late in the breeding season. Both food availability and density in year ^x were not predictive of the number of young per female in year ^x+1; however, the number of young produced per female at a site in year ^x was predictive of the number of young per female in year ^x+1. As expected, dispersers used the most informative cue, the number of young per female and moved to sites with a relatively high number of young per female. This study suggests that individuals prospect for potential breeding sites late in the breeding season when they can use information gathered from the reproductive success of other individuals (i.e., public information) to select a breeding site.     

Communal Roost Site Selection in a Neotropical Harvestman: Habitat Limitation vs. Tradition

Many species have been reported to form roosting (resting, sleeping) aggregations at ‘traditional’ sites, but the alternative hypothesis that specific sites are used repeatedly because of habitat limitation is rarely tested. We studied the roosting behavior of a species of harvestman (Opiliones, Prionostemma sp.) at a lowland rainforest site in Nicaragua. Both sexes roosted by day in spiny palm trees, dispersed at dusk to forage, and rejoined aggregations just before dawn. The distribution of harvestmen among spiny palms was significantly clumped, and harvestman density did not correlate with spiny palm density. Aggregations formed repeatedly in a small subset of the available spiny palms and the same sites were used in two different years (2001, 2003). Nevertheless, the membership of aggregations was fluid; individual harvestmen were found at multiple roosts and moved up to 0.2 km per night. Translocated animals often returned to the roost where they had been released or nearby roosts but were never found at previously unused sites. The high consistency of site use but low site fidelity of individuals suggests that roost sites differed conspicuously (to the harvestmen) from sites that were not used. We found no univariate or multivariate differences between used and unused sites, however, in the characteristics of the trees or microclimate. These results conflict with the habitat limitation hypothesis but are consistent with the traditional site use hypothesis. The tradition may be mediated by a site‐labeling chemical, a mechanism that does not require individual site fidelity. We discuss these results in relation to the proposed functions of roosting aggregations.     

Female‐biased sleeping aggregations of Amegilla florea urens (Hymenoptera: Apidae)

Although male bees and wasps aggregate in specific sites overnight, few studies have addressed the behaviors of female hymenopterans during the night. It is generally understood that they spend the night in a burrow or nest structure. We found female‐biased sleeping aggregations of Amegilla florea urens in populations on Iriomote Island. The sex ratio and number of sleeping individuals in the sex‐mixed aggregations changed daily. This species used dead branches as its main sleeping places, and the maximum number of individuals observed per place was 15. Females arrived at the aggregation place later and left it earlier in the morning than did males. Our result suggests that females used the sleeping place regularly but avoided a long stay with males. The use of the same sleeping place by the two sexes may be due to a limited number of sites suitable for sleeping in the forest.     

Host location by larvae of the reptile tick Amblyomma vikirri (Acari: Ixodidae)

The newly described tick Amblyomma vikirri has a narrow host range, being found mainly on the Australian lizard, Egernia stokesii and rarely on another lizard, Tiliqua rugosa. Both hosts are in the family Scincidae. Larvae of A. vikirri were as successful in locating E. stokesii as T. rugosa from a range of release distances between 20 and 120 mm from the host. Over this range the proportion of ticks which successfully located hosts declined and the time taken by successful ticks to locate hosts increased with increasing release distance. From 60 mm, larvae of A. vikirri located four other lizard species from the families Scincidae and Agamidae and two non-living targets as successfully as they did E. stokesii. The only evidence that there was any host specificity in the searching behaviour of larvae of A. vikirri was that A. vikirri larvae spent less time paused and questing when they were searching for E. stokesii than when they were searching for T. rugosa. Aponomma hydrosauri, a tick which commonly infests T. rugosa but not E. stokesii, spent less time paused and questing when it was searching for T. rugosa than when it was searching for E. stokesii. However, the results overall suggest that the narrow host range of A. vikirri cannot be explained by any ability of the larvae of that species to discriminate between their natural host and other reptile species.     

Predator protection versus rapid growth in a montane leaf beetle

Summary Adults and larvae of Chrysomela aenicollis (Coleoptera: Chrysomelidae) feed on foliage of Salix species (Salicaceae) between 2,400–3,400 m above sea level in the eastcentral Sierra Nevada mountains of California. We predicted that (1) cold climatic conditions would be a more frequent source of mortality at higher elevations, (2) mildweather agents of mortality such as predation should be more severe at lower elevations, and (3) populations of C. aenicollis would be adapted to the local selective regime at each elevation. We tested these predictions in 1984 and 1985 by transferring over 6,000 eggs and larvae within and between two sites at 2,810 and 3,240 m elevation above sea level. During mild summer weather at both sites, survivorship on Salix branches isolated by a barrier of sticky resin was similar to that on control branches, and we concluded that aerial predators were the primary cause of mortality. At least one major predator, a solitary wasp (Symmorphus sp., Hymenoptera: Eumenidae), was specifically associated with C. aenicollis at the lower site, where beetle mortality was highest. At both sites in 1984 and 1985, larvae originating from the lower site remained in aggregations and survived more frequently than larvae from the upper site, suggesting that they are better defended against predators. During a storm with cold weather late in the 1984 season, larvae and pupae died more frequently at the upper site, and there was a marginally significant trend (P<0.1) for the lower site individuals to die more frequently than upper site larvae during the cold storm. Upper site larvae grew approximately 10% faster than lower site larvae at the lower site and under controlled conditions in the laboratory. These findings indicate that upper and lower site populations were adapted to the local selective regime, which suggest how populations of montane phytopagous insects may adapt to changing elevations.     

Diel Habitat Partitioning by Bull Charr and Cutthroat Trout During Fall and Winter in Rocky Mountain Streams

We used underwater observation to determine diel habitat partitioning between bull charr, Salvelinus confluentus, and cutthroat trout, Oncorhynchus clarki, during fall and winter (0.1–8.3°C) in two Rocky Mountain streams that differed in habitat availability. The majority (>70%) of both species emerged from concealment cover at night, though bull charr exhibited a greater tendency for nocturnal behavior than cutthroat trout. Differences in day and night counts were most pronounced at temperatures <3°C, when very few fish of either species were observed in the water column during the day, but both species were common at night. Both species used concealment cover of large woody debris and boulder substrate crevices in deep pools during the day. At night, fish emerged from cover and habitat use shifted to shallow water with low cover. Microhabitat partitioning among species and size classes occurred at night, cutthroat trout moving into shallower, faster water that was farther from cover compared to bull charr. Smaller fish of both species occupied focal positions in slower, shallower water closer to the substrate than larger fish. Large, mixed-species aggregations also were common in beaver ponds both day and night. High variation in diel and site-specific winter habitat use suggests the need for caution in developing habitat suitability criteria for salmonids based solely on daytime observations or on observations from a few sites. Our results support the need to incorporate nocturnal habitat use and partitioning in studies of salmonid ecology.     

Whether to release Oxyops vitiosa from a second Australian site onto Florida's melaleuca? A molecular approach

Quarantine colonies of the melaleuca snout beetle Oxyops vitiosa, the first melaleuca-feeding insect released in Florida, consisted of individuals collected from the Burpengary region (near Brisbane) of Queensland and their descendants. Constraints on the number of weevils available at this site motivated us to import weevils from a second site (Bundaberg). While individuals from the two sites could not be differentiated using taxonomic characters, the weevils from Bundaberg were consistently larger. This prompted concern that Bundaberg individuals might differ genetically from the quarantine population thus confounding the original host specificity studies. In a worse case scenario, the Bundaberg population might represent a separate sub-species or even a separate (cryptic) species. RAPD analysis was conducted on individuals from the two sites and on an outlier species, Gonipterus scutellatus. Neighbor Joining analysis clearly delineated the two species and also indicated some differentiation of individuals from the two O. vitiosa sites. Bootstrap analysis, however, indicated little statistical support for this intraspecific differentiation. Principal Coordinate analysis distinguished individuals from the two populations, and even distinguished two sub-populations from Bundaberg. AMOVA analysis indicated most of the variation occurred within populations but that the between population variance was significant. We conclude that further limited specificity testing and field host range observations should be conducted on the Bundaberg population before release into Florida.     

Ecological life history of Polymnia canadensis,a monocarpic species of the North American Temperate Deciduous Forest: Demography

A demographic investigation was conducted to assess variation in life history of Polymnia canadensis (Asteraceae), a geographically-widespread, herbaceous species of deciduous forests in eastern North America. During 1985-1994, 23,063 seedlings of P. canadensis were monitored at five central Kentucky study sites. Numbers at the end were: biennials, 554; triennials, 142; winter annuals, 16; monocarpic perennials, 2; tricarpic perennials (three years), 3; and dicarpic perennials (some skipped years) that matured in the first year of life, 23; in the second year, 60; and in the third year, 9. Weekly cohorts of P. canadensis generally exhibited Deevey Type III survivorship with highest seedling mortality in summer associated with low soil moisture. Wide spatial variation in life history was displayed by the fact that fall germination cohorts at dry sites generally had greater germination and survivorship than at mesic sites during seedling establishment, while the reverse was true for spring cohorts. This led to more reproductive individuals in fall germination cohorts than spring cohorts at dry sites and generally the opposite case at mesic sites. Forest shade in mesic sites caused slower growth and a greater frequency of longer-lived reproductive individuals than in open, dry sites, but it also resulted in higher survivorship than at dry sites during moderate drought. Annual population growth rate averaged across four years was not significantly different between a dry site and mesic site, indicating that despite forest shade, P. canadensis persisted in the mesic site as well as it did in the dry site. Population structure varied among years, seasons, and study sites; at two study plots, a 2-year flowering cycle of mass seeding and senescence persisted for 4 years. At a smaller scale, there was little difference in survivorship between study plots within sites or between quadrats within study plots, while in a few instances there were large differences in the number of reproductive individuals. Weekly cohorts that germinated early within seasonal cohorts had greater number of reproductive individuals than later weekly cohorts, but not consistently greater survivorship. Several droughts induced temporal variation that was as important as spatial variation. During these droughts, the population size of all cohorts that germinated prior to the droughts declined to zero at all sites, and biennials were the longest lived type of reproductive individuals. During drought, Deevey Type I survivorship was prevalent, and lack of seed rain led to dependence on persistent seed banks for recruitment in some seasonal cohorts.     

Male-biased sex ratios,female promiscuity,and copulatory mate guarding in an aggregating tropical bug,Dysdercus bimaculatus

The ecological and social bases of the mating system of the seed-feeding bug, Dysdercus bimaculatus(Hemiptera: Pyrrhocoridae), were studied in the lab and in aggregations at the host tree, Sterculia apetala(Malvales: Malvaceae), in Panama. On theoretical grounds, two factors are predicted to be of importance in determining the evolution of male mating tactics in Ms species: the operational sex ratio and the probability that undefended females will mate with other males, subjecting the gametes of deserters to sperm competition. Results of a study of a related species suggested that sperm displacement is probably substantial. Adult sex ratios at numerous sites were significantly male biased, and females whose mates were removed remated before oviposition (i. e., sperm utilization). These results predict that a mate defense tactic is likely to be superior to a nondefense tactic. The biological significance of the parameters is supported by observations that captive pairs often remained in copulafor several days, until just before oviposition. However, substantial variation in copulation duration was also observed, and possible causes of this variation are considered. Causes of male biased adult sex ratios were investigated by monitoring demographic changes within a single aggregation over 2 months. Both female juvenile and adult mortality rates were greater than male. In addition, dissections of reproductive adults showed that the flight muscles of females, but not males, had histolyzed, so that female reproduction is physiologically limited to a single site. Greater rates of immigration among both mature and young males suggests that an excess of males may also be found in the populations of bugs that subsequently colonize other host plants, so that female scarcity is typical of aggregations in all stages of development. The evolution of sex-limtied flight muscle histolysis may be explained by greater patchiness of females than males as mating resources, plus a lower energetic benefit/cost ratio of histolysis for males.     

Environmentally induced variation in uric acid concentration and xanthine dehydrogenase activity in Littorina saxatilis (Olivi) and L. arcana Hannaford Ellis

Littorina saxatilis and Littorina arcana collected from a boulder field low in the intertidal zone had a uric acid concentration significantly higher than snails collected from nearby cliff crevices that were at the upper limit of their vertical range on the shore. The absolute concentrations varied with different collections, suggesting a possible seasonal fluctuation in uric acid. L. arcana had a greater concentration of uric acid than did L. saxatilis when both were from the boulder field, but the two did not differ significantly when taken from crevices. Samples from the two sites were marked and transplanted; by four weeks the transplanted animals showed a tendency to develop a uric acid concentration similar to that of natives of the site, suggesting a physiological rather than a genetic adaptation. Xanthine dehydrogenase activity differed between animals from the two sites in the same manner as the uric acid concentration.     

Relationships between soil heavy metal concentration and mycorrhizal colonisation in<Emphasis Type="Italic"> Thymus polytrichus</Emphasis> in northern England

A study was conducted to establish whether the wild thyme [Thymus polytrichus A. Kerner ex Borbás ssp. britannicus (Ronn.) Kerguelen (Lamiaceae)] growing in the metal-contaminated soils along the River South Tyne, United Kingdom, is colonised by arbuscular mycorrhizal (AM) fungi, and whether the degree of colonisation increases (perhaps suggesting increasing mycorrhizal dependence) or decreases (indicating possible inhibition of AM growth) with increasing degree of soil contamination. Seasonal changes in AM colonisation were also assessed. The AM fungal communities colonising T. polytrichus were also investigated, using the polymerase chain reaction with restriction fragment length polymorphism and sequencing of fungal DNA to establish whether AM species richness varied between sites, and whether fungal ecotypes specific to sites with different amounts of metal contamination could be identified. All plants examined were heavily colonised by AM fungi, and mean percentage root length colonised did not increase significantly with increasing soil metal contamination. However, AM vesicle abundance (percentage of mycorrhizal root length containing vesicles) at the most contaminated site was significantly greater than at the other sites. No significant seasonal variation in degree of colonisation or vesicle abundance was found. Glomus was the predominant AM genus detected at all sites. The number of AM genotypes colonising T. polytrichus roots was similar at all sites but, although some were common to all sites, certain strains appeared to be specific to either the most- or the least-contaminated site. This variation in species may account for the difference in vesicle abundance between sites. The consistently heavy AM colonisation of T. polytrichus found suggests that these fungi are not inhibited by soil heavy metals at these sites, and that the host derives some benefit from its AM symbiont.     

Scat on the doorstep: Refuge choice in a group‐living lizard is influenced by the presence of scat piles

Group living often requires strong levels of communication between individuals. This communication is usually studied in the context of visual or auditory communication. However, chemical communication is the most widely used form of communication. We examined the role of chemical communication in mediating social decisions in a group‐living lizard, Egernia stokesii. Specifically, we examined the extent to which scat‐piling, a behaviour by which individuals deposit scat in a communal area, affected the refuge choice of individual E. stokesii. To achieve this, we examined individual refuge choice in response to scat piles or single scats and against two types of scat stimuli, one being their own scat and the other being scat belonging to an unrelated and unfamiliar conspecific. We show that lizards behave differently when presented with a scat pile compared with a single scat, and whether the scat stimulus was their own or sourced from an unfamiliar conspecific. When scats were in piles, individuals spent more time inspecting, more time in, and more often chose the treatment refuge as their final refuge choice, at a trial’s end, when the treatment was their own scat compared with when the treatment was the refuge with the unfamiliar scat. In contrast, for individual scat treatments, individuals spent more time inspecting and more often ended up in the treatment refuge with an unfamiliar scat compared with when the treatment was their own scat. These results suggest that individuals are responding to information contained within multiple components of the scats – both their volume and their source. These results have implications for understanding how social aggregations are maintained within squamates, where sociality has evolved independently from other vertebrate lineages.     

Aggregations of <Emphasis Type="Italic">Plectropomus areolatus</Emphasis> and <Emphasis Type="Italic">Epinephelus fuscoguttatus</Emphasis> (groupers,Serranidae) in the Komodo National Park,Indonesia: Monitoring and Implications for Management

Synopsis We identify fishery management implications from a long-term monitoring program focusing on spawning aggregations of high valued reef fish in Komodo National Park (KNP), Eastern Indonesia. Management objectives of KNP are not only to protect biodiversity, but also to conserve spawning stocks of high-valued commercial species for the replenishment of surrounding fishing grounds. We monitored two sites twice monthly over five years for two species of grouper, Epinephelus fuscoguttatus and Plectropomus areolatus. One site had an aggregation of both E. fuscoguttatus and P. areolatus, whereas the other site contained an aggregation of P. areolatus only. Over the five years monitoring period, aggregations typically formed during each full moon between September and February. Additionally, P. areolatus occasionally aggregated during new moons between April and July. We observed spawning only once, but because formation of aggregations were correlated to a higher incidence of behavior and signs indicative of reproduction and because most fish present were adults, it is likely that the formation of aggregations was associated with spawning. Over the five years monitoring period there was a reduction in mean fish size of up to 8 cm for P. areolatus, and a reduction in numbers of aggregating E. fuscoguttatus. Despite limited protection initiated in 2001, both sites are still heavily fished by local artisanal fishers. Because the observed reductions in size and in numbers could be caused by fishing pressure, managers should follow the precautionary principle by putting additional protective management in place. Since both species are relatively long-lived, at least five years of continued monitoring may be necessary to determine the outcome of management intervention. The variability in timing of aggregation in respect to season and moon phase in P. areolatus indicates that long-term monitoring must cover the entire year and both moon phases.     

A spawning aggregation of Nassau grouperEpinephelus striatus (Pisces: Serranidae) in the Mexican Caribbean

Synopsis An annual spawning aggregation of Nassau grouper,Epinephelus striatus, observed off the southern coast of Quintana Roo, Mexico, on full moon days in December and January, was surveyed during the winters from 1991 to 1993. A fish aggregation had been appearing at one traditional site off Mahahual for more than 80 years, but currently it forms at undetermined sites far from the original site. Fish migration was northwards in the fore reef border along the 14 m isobath and isolated groups were observed. Through visual assessment fish aggregations were found to include 200 to 500 individuals. The fishing on Nassau grouper aggregations has been active since 1910 by using hook and line; however, recently fishermen used gill nets. It is necessary to continue the research on this aggregation and establish management guidelines to protect the species.     

Climatic signals in tree rings of <Emphasis Type="Italic">Burkea africana</Emphasis> and <Emphasis Type="Italic">Pterocarpus angolensis</Emphasis> from semiarid forests in Namibia

Tree-ring studies contribute worldwide to the understanding of climate and its relation to tree growth. Long tree-ring chronologies serve as climate proxies for the reconstruction of past, pre-instrument climate and its recent change. In tropical regions, the availability of exactly dated tree-ring chronologies is limited. The dendroclimatic potential of two dominant species from dry forests in northern Namibia was examined in the study presented in this paper. Both species (Burkea africana Hook and Pterocarpus angolensis DC) were sampled at two sites (ca. 900 km apart), and the response to several climatic variables, including ENSO indices, is studied. All specimens showed distinct growth rings and cross-dating between radii was successful for all trees. Species-specific mean curves were built for both sites. The mean curves of different species of the same site synchronised significantly, allowing the construction of a site-specific chronology. Synchronisation between sites was not possible, but spectral analysis of the chronologies implied that both show similar long-term (6.7 year) oscillation patterns. B. africana is more sensitive to rainfall variation than P. angolensis at both sites. Growth response to rainfall was positive, but a time-lag in the reaction occurred between the sites, corresponding to the time-lag of the beginning of the rainy season. Air temperature showed a negative correlation with stem increment at both sites. The response at the westernmost site to two ENSO indices indicates a tree growth decrease during El Niño years, which are generally dry in southern Africa.