All clear? Meerkats attend to contextual information in close calls to coordinate vigilance

Socio-demographic factors, such as group size and their effect on predation vulnerability, have, in addition to intrinsic factors, dominated as explanations when attempting to understand animal vigilance behaviour. It is generally assumed that animals evaluate these external factors visually; however, many socially foraging species adopt a foraging technique that directly compromises the visual system. In these instances, such species may instead rely more on the acoustical medium to assess their relative risk and guide their subsequent anti-predator behaviour. We addressed this question in the socially foraging meerkat (Suricata suricatta). Meerkats forage with their head down, but at the same time frequently produce close calls (‘Foraging’ close calls). Close calls are also produced just after an individual has briefly scanned the surrounding environment for predators (‘Guarding’ close calls). Here, we firstly show that these Guarding and Foraging close call variants are in fact acoustically distinct and secondly subjects are less vigilant (in terms of frequency and time) when exposed to Guarding close call playbacks than when they hear Foraging close calls. We argue that this is the first evidence for socially foraging animals using the information encoded within calls, the main adaptive function of which is unrelated to immediate predator encounters, to coordinate their vigilance behaviour. In addition, these results provide new insights into the potential cognitive mechanisms underlying anti-predator behaviour and suggest meerkats may be capable of signalling to group members the ‘absence’ of predatory threat. If we are to fully understand the complexities underlying the coordination of animal anti-predator behaviour, we encourage future studies to take these additional auditory and cognitive dimensions into account.     

Should vigilance always decrease with group size?

Levels of anti-predator vigilance often decrease with group size in birds and mammals. Vigilance may also serve other purposes such as scrounging. While scrounging, individual adopt the vigilant posture to locate and eventually exploit the food discoveries of their companions. Models that combine anti-predator alertness and scrounging were developed to examine how changes in scrounger use with group size could alter the shape of the function relating vigilance to group size and whether the additional investment in vigilance while searching could be used to reduce the allocation of time to vigilance while in the food patch. Results indicate that increased allocation of time to scrounging during search can add to the overall level of vigilance and even counteract the expected decrease in vigilance with group size. However, the addition of scrounging was found to have little impact on vigilance while in the food patch. Behavioral effects that are dependent on group size, such as scrounging, may lead to changes in the shape of the vigilance function and may mitigate against any gain garnered from a reduction in individual levels of vigilance in larger groups.     

Female foraging and male vigilance in white-tailed ptarmigan (Lagopus leucurus): opportunism or behavioural coordination?

For several species of birds, high rates of male vigilance are correlated with high rates of female foraging. This relationship is thought to ultimately result in higher reproductive success for females paired with highly vigilant males. However, previous research has not examined the behavioural mechanism that produces the correlation between male vigilance and rates of female foraging. Foraging females may take advantage of vigilance that males are using for other purposes. Alternatively, the purpose of male vigilance may be to increase females' ability to forage. We examined these alternatives by testing whether vigilance preceded or followed bouts of female foraging more often than would occur by chance alone, using simultaneous behaviour observations of pre-incubation pairs of white-tailed ptarmigan (Lagopus leucurus). Our results indicate that each member of a pair may influence the behaviour of the other. Females were more likely to initiate foraging bouts after males became vigilant than if their mate remained non-vigilant. Moreover, non-vigilant males were more likely to become vigilant if their mate was foraging than if she was engaged in some other activity. Despite the possibility that a sexual conflict exists as each member of a pair attempts to maximize its fitness, both sexes behave as though a major role of male vigilance is to enhance female foraging opportunities. Received: 3 May 1999 / Received in revised form: 14 June 1999 / Accepted: 16 June 1999     

Ontogenetic shifts in fusion–rejection thresholds in a colonial marine hydrozoan,<Emphasis Type="Italic"> Hydractinia symbiolongicarpus</Emphasis>

Like many modular organisms, genetically distinct colonies of the hydrozoan Hydractinia symbiolongicarpus naturally fuse to produce chimeras. One of the principal cooperative benefits of fusion arises from the increased size of the resulting chimeric individual, which may enhance survivorship. However, fusion also promotes conflict through competition between cell lineages for representation in reproductive tissues. Previous studies on H. symbiologicarpus show that, consistent with kin selection theory, a highly polymorphic self/non-self recognition system limits fusion to close kin. However, these recognition systems are intrinsically subject to error. Conspecific acceptance threshold theory predicts that as the costs and benefits of making recognition errors change, or the frequencies of encounters between acceptable and unacceptable kin vary, the recognition system should respond. Specifically, as the benefits of acceptance decline or the frequency of encounters with unacceptable individuals increases, the acceptance threshold should become more restrictive. We tested this hypothesis by monitoring changes in the expression of fusion/rejection behaviors of H. symbiolongicarpus during colony establishment, a period of high mortality when the size-dependent benefits of fusion may be changing most rapidly, and the frequency of encounters with close kin declines. Across seven full-sib families, fusion frequencies between pairs of sibling colonies declined from 73% for 3-day-old colonies to 58% by day 12. This decline is consistent with optimal acceptance threshold theory. However, the period of maximum decline also corresponds to an interval during which the recognition effector mechanism becomes fully functional, suggesting that the shift to a more restrictive conspecific acceptance threshold may reflect an intrinsic constraint on recognition system maturation.Communicated by T. Czeschlik     

Sexual selection in darkness? Female mating preferences in surface- and cave-dwelling Atlantic mollies,<Emphasis Type="Italic"> Poecilia mexicana</Emphasis> (Poeciliidae,Teleostei)

In many species, females prefer large males as mating partners. Male size can easily be determined visually. We examined how such mate choice evolves when individuals are deprived of sensory information previously used to determine male quality. Using a cave-dwelling population of Atlantic mollies, Poecilia mexicana, as a model, we asked whether the female mating preference for large male body size still occurs in this cave fish, or whether it is lacking due to the absence of visual communication in the cave environment. In simultaneous choice tests we compared the time females spent with a large or a small male. In the river-dwelling and cave-dwelling fish, as well as the population originating from the cave entrance, we found a significant preference for large males in light. Hence, the ancestral mating preference of surface-dwelling mollies has been maintained during the evolution of the cave form. However, only the cave population exhibited a mating preference in darkness. Determination of male body size appears to have been taken over by a non-visual sensory system, probably the lateral line.Communicated by K. Lindström     

Trace element contents in fine particulate matter (PM<Subscript>2.5</Subscript>) in urban school microenvironments near a contaminated beach with mine tailings,Chañaral,Chile

Air quality in schools is an important public health issue because children spend a considerable part of their daily life in classrooms. Particulate size and chemical composition has been associated with negative health effects. We studied levels of trace element concentrations in fine particulate matter (PM_2.5) in indoor versus outdoor school settings from six schools in Chañaral, a coastal city with a beach severely polluted with mine tailings. Concentrations of trace elements were measured on two consecutive days during the summer and winter of 2012 and 2013 and determined using X-ray fluorescence. Source apportionment and element enrichment were measured using principal components analysis and enrichment factors. Trace elements were higher in indoor school spaces, especially in classrooms compared with outdoor environments. The most abundant elements were Na, Cl, S, Ca, Fe, K, Mn, Ti, and Si, associated with earth’s crust. Conversely, an extremely high enrichment factor was determined for Cu, Zn, Ni and Cr; heavy metals associated with systemic and carcinogenic risk effects, whose probably origin sources are industrial and mining activities. These results suggest that the main source of trace elements in PM_2.5 from these school microenvironments is a mixture of dust contaminated with mine tailings and marine aerosols. Policymakers should prioritize environmental management changes to minimize further environmental damage and its direct impact on the health of children exposed.     

Variation in colony geometry modulates internal light levels in branching corals, <Emphasis Type="Italic">Acropora humilis</Emphasis> and <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Colonial photosynthetic marine organisms often exhibit morphological phenotypic plasticity. Where such plasticity leads to an improved balance between rates of photosynthesis and maintenance costs, it is likely to have adaptive significance. To explore whether such phenotypic plasticity leads to more favourable within-colony irradiance for reef-building branching corals, this relationship was investigated for two coral species Acropora humilis and Stylophora pistillata, along a depth gradient representing light habitats ranging from 500 to 25 μmol photons m⁻² s⁻¹, during 2006 at Heron Island, Great Barrier Reef (23.44°S, 151.91°E). In the present study changes in flow-modulated mass transfer co-varied with light as a function of depth. In low-light (deep) habitats, branch spacing (colony openness) in A. humilis and S. pistillata was 40–50% greater than for conspecifics in high-light environments. Also, branches of A. humilis in deep water were 40–60% shorter than in shallow water. Phenotypic changes in these two variables lead to steeper within-colony light attenuation resulting in 38% higher mean internal irradiance (at the tissue surface) in deep colonies compared to shallow colonies. The pattern of branch spacing was similar for S. pistillata, but this species displayed an alternate strategy with respect to branch length: shade adapted deep and cave colonies developed longer and thinner branches, allowing access to higher mass transfer and irradiance. Corals in cave habitats allowed 20% more irradiance compared to colonies found in the deep, and had a 47% greater proportion of irradiance compared to colonies in the shallow high-light environment. Such phenotypic regulation of internal light levels on branch surfaces partly explains the broad light niches of many branching coral species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Reproduction in the deepwater squid<Emphasis Type="Italic"> Moroteuthis ingens</Emphasis>, what does it cost?

The cost of reproduction for the terminal spawning onychoteuthid squid, Moroteuthis ingens, was analysed using measures of condition and tissue biochemistry. Both males and females showed a dramatic drop in the weight of the gonad in stage 6 (spent) individuals. The mantle weight and nidamental gland weight of females also decreased during the maturation process. Males, however, had a marked increase in both the penis and spermatophoric complex weight in spent individuals, while female oviducal gland weight and nidamental gland length also increased in stage 6 individuals. Residual analysis indicated that testis growth was not developing at the expense of mantle growth, although there was a suggestion of cost to the fins. Females showed that the development of the ovary occurred at a cost to both the mantle and fins. Overall body condition also declined with maturity stage for both males and females, with stage 6 individuals of both sexes in poor condition. Very few females had eggs in the oviducts, suggesting that the oviducts are used as ducts instead of storage organs. Proximal analysis revealed a loss of constituents within the mantle during maturation, with an associated increase in water, indicating the remobilization of energy from the mantle to fuel reproduction. This study suggests that the digestive gland is not used as an energy store in this species.Communicated by M.S. Johnson, Crawley     

Does begging affect growth in nestling tree swallows,<Emphasis Type="Italic"> Tachycineta bicolor</Emphasis>?

Much of the theoretical work on the evolution of begging assumes this elaborate display is costly. The evidence for an energetic cost to begging has, however, been equivocal. Metabolic studies on nestling birds suggest that begging requires minimal energy, but some growth studies have shown that excess begging reduces growth rates. One difficulty in interpreting these results is that metabolic and growth studies have each been performed on different species. Here, we test whether high begging frequencies depress growth in nestling tree swallows, Tachycineta bicolor, a species in which the metabolic cost of begging has been measured. When we compared the growth of nestlings stimulated to beg at either high or low frequencies, we found no significant differences in their mass gained, wing growth or portion of ingested energy devoted to begging either during the experimental period or in the 24 h following the end of the experiment. We also found no significant relationship between begging intensity and growth measurements. The results of our study are consistent with previous metabolic studies on this species suggesting that the energetic cost of begging is relatively low. More generally, evidence for a fitness cost of begging via decreased growth is equivocal.Communicated by J. Dickinson     

Does sex affect both individual and collective vigilance in social mammalian herbivores: the case of the eastern grey kangaroo?

In several vertebrate taxa, males and females differ in the proportions of time they individually devote to vigilance, commonly attributed to sex differences in intra-specific competition or in absolute energy requirements. However, an effect of sex on collective vigilance is less often studied (and therefore rarely predicted), despite being relevant to any consideration of the adaptiveness of mixed- vs single-sex grouping. Controlling for group size, we studied the effect of sex on vigilance in the sexually dimorphic eastern grey kangaroo Macropus giganteus, analysing vigilance at two structural levels: individual vigilance and the group’s collective vigilance. Knowing that group members in this species tend to synchronise their bouts of vigilance, we tested (for the first time) whether sex affects the degree of synchrony between group members. We found that females were individually more vigilant than males and that their vigilance rate was unaffected by the presence of males. Collective vigilance did not differ between female-only and mixed-sex groups of the same size. Vigilance in mixed-sex groups was neither more nor less synchronous than in single-sex groups of females, and the presence of males seemed not to affect the degree of synchrony between females. Sixty-six percent of vigilant acts were unique (performed when no other kangaroo was alert), and only about one unique vigilant act in every three induced a collective wave of vigilance. The proportions of vigilant acts that were unique were 60% for females but only 46% for males. However, the sexes differed little in the rates at which their unique vigilant acts were copied. This limited study shows that the differences in vigilance between male and female kangaroos had no discernible effect upon collective vigilance.     

Source apportionment of PM<Subscript>2.5</Subscript> in Tangshan,China—Hybrid approaches for primary and secondary species apportionment

The objective of this paper is to propose a hybrid approach for the source apportionment of primary and secondary species of PM_2.5 in the city of Tangshan. The receptor-based PMF (Positive Matrix Factorization) is integrated with the emission inventory (EI) to form the first hybrid method for the source apportionment of the primary species. The hybrid CAMx-PSAT-CP (Comprehensive Air Quality Model with Extensions–Particulate Source Apportionment Technology–Chemical Profile) approach is then proposed and used for the source apportionment of the secondary species. The PM_2.5 sources identified for Tangshan included the soil dust, the metallurgical industry, power plants, coalfired boilers, vehicles, cement production, and other sources. It is indicated that the PM_2.5 pollution is a regional issue. Among all the identified sources, the metallurgy industry was the biggest contribution source to PM_2.5, followed by coal-fired boilers, vehicles and soil dust. The other-source category plays a crucial role for PM_2.5, particularly for the formation of secondary species and aerosols, and these other sources include non-specified sources such as agricultural activities, biomass combustion, residential emissions, etc. The source apportionment results could help the local authorities make sound policies and regulations to better protect the citizens from the local and regional PM_2.5 pollution. The study also highlights the strength of utilizing the proposed hybrid approaches in the identification of PM_2.5 sources. The techniques used in this study show considerable promise for further application to other regions as well as to identify other source categories of PM_2.5.

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Within-group spatial position and vigilance: a role also for competition? The case of impalas (<Emphasis Type="Italic">Aepyceros melampus</Emphasis>) with a controlled food supply

Theory predicts that individuals at the periphery of a group should be at higher risk than their more central conspecifics since they would be the first to be encountered by an approaching terrestrial predator. As a result, it is expected that peripheral individuals display higher vigilance levels. However, the role of conspecifics in this “edge effect” may have been previously overlooked, and taking into account the possible role of within-group competition is needed. Vigilance behavior in relation to within-group spatial position was studied in impalas (Aepyceros melampus) feeding on standardized patches. We also controlled for food distribution in order to accurately define a “central” as opposed to a “peripheral” position. Our data clearly supported an edge effect, with peripheral individuals spending more time vigilant than their central conspecifics. Data on social interactions suggest that it was easier for a foraging individual to defend its feeding patch with its head lowered, and that more interactions occurred at the center of the group. Together, these results indicate that central foragers may reduce their vigilance rates in response to increased competition. Disentangling how the effects of competition and predation risk contribute to the edge effect requires further investigations.     

Seasonal variations of transport pathways and potential sources of PM<Subscript>2.5</Subscript> in Chengdu,China (2012–2013)

Seasonal pattern of transport pathways and potential sources of PM_2.5 in Chengdu during 2012–2013 were investigated based on hourly PM_2.5 data, backward trajectories, clustering analysis, potential source contribution function (PSCF), and concentration-weighted trajectory (CWT) method. The annual hourly mean PM_2.5 concentration in Chengdu was 97.4 mg·m^–3. 5, 5, 5 and 3 mean clusters were generated in four seasons, respectively. Short-distance air masses, which travelled within the Sichuan Basin with no specific source direction and relatively high PM_2.5 loadings (>80 mg·m^–3) appeared as important pathways in all seasons. These short pathways indicated that emissions from both local and surrounding regions of Chengdu contributed significantly to PM_2.5 pollution. The cities in southern Chengdu were major potential sources with PSCF>0.6 and CWT>90 mg·m^–3. The northeastern pathway prevailed throughout the year with higher frequency in autumn and winter and lower frequency in spring and summer. In spring, long-range transport from southern Xinjiang was a representative dust invasion path to Chengdu, and the CWT values along the path were 30-60 mg·m^–3. Long-range transport was also observed in autumn from southeastern Xinjiang along a northwesterly pathway, and in winter from the Tibetan Plateau along a westerly pathway. In summer, the potential source regions of Chengdu were smaller than those in other seasons, and no long-range transport pathway was observed. Results of PSCF and CWT indicated that regions in Qinghai and Tibet contributed to PM_2.5 pollution in Chengdu as well, and their CWT values increased to above 30 mg·m^–3 in winter.

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Na<Superscript>+</Superscript>/K<Superscript>+</Superscript>–ATPase activity and gill ultrastructure in the hyper-hypo-regulating crab<Emphasis Type="Italic"> Chasmagnathus granulatus</Emphasis> acclimated to dilute,normal, and concentrated seawater

We studied Na⁺/K⁺ –ATPase activity and ultrastructure in gills of the hyper-hypo-regulating crab Chasmagnathus granulatus Dana, 1851 acclimated to different salinities: 10, 30 and 45, known to be hypo-, iso-, and hyper-osmotic to the hemolymph, respectively. After centrifugation of homogenates at 11,000 g, Na⁺/K⁺–ATPase activity was almost entirely found in the pellets from the posterior (6–8) and anterior (3–5) gills, whereas very little was detected in the supernatant liquid. Specific activity of gill 6 was 41.3, 30.2, and 28.2 µmol Pi h^–1 mg prot^–1 for crabs acclimated to 10, 30, and 45, respectively, the result for 10 being significantly higher than those at 30 and 45. Although the concentration of sodium at which the reaction rate is half-maximal (K _M) was similar in the three acclimation salinities, only the enzyme from crabs acclimated to 10 was inhibited by high sodium concentration. Specific activity of gill 5 increased with the increment in external salinity (10.1, 15, and 18.1 µmol Pi h^–1 mg prot^–1 for 10, 30, and 45, respectively), the only significant difference being that between the extreme salinities. The epithelium thickness of the dorsal portion of gill 6 showed a variation among salinities: 21.7, 15.8 and 17.2 µm for 10, 30 and 45, respectively. There were significant differences in epithelium thickness between the 10 and the other salinities. In all three salinities, the ultrastructure of gill 6 epithelium showed a high density of mitochondria, estimated by their volume fraction (Vv _m=0.307–0.355). These mitochondria were packed between extensive basolateral membrane interdigitations in ionocytes and pillar cells. Gill 5 showed three cell types: pillars which possess mitochondria packed between membrane folds only in their interdigitations with neighbouring cells; type-I cells 8.0 µm thick with low density of mitochondria (Vv _m=0.088), and type-II cells, 9.9 µm thick and rich in mitochondria (Vv _m=0.423), but lacking basolateral interdigitations. Vv _m of type-I cells of gill 5 was significantly lower than those of type-II cells of the same gill and the ionocytes of gill 6. No significant difference in Vv _m was detected between the latter cell types.Communicated by P.W. Sammarco, Chauvin     

Extrapair paternity in the swamp sparrow, <Emphasis Type="Italic">Melospiza georgiana</Emphasis>: male access or female preference?

Over the past two decades, the combination of molecular and field methods has revealed considerable variation in the level of extrapair fertilizations among socially monogamous birds. Models predicting extrapair young range in scale from a single population to multiple Orders, and there is no single, unifying theory for these reproductive tactics. We investigated proximate explanations of extrapair fertilizations in two subspecies of the swamp sparrow, Melospiza georgiana georgiana and Melospiza georgiana nigrescens, across a range of social and environmental conditions. The presence of extrapair young was best predicted by the size of two male plumage badges (one correlated with parental care and one with territorial aggression) relative to the badge size of their immediate neighbors, the interaction of these two measures, mean territory size, and the maximum size of the aggression badge among neighbors. The size of the male’s parental care badge (relative to neighbors) was negatively correlated with the probability of lost paternity. The relative size of the aggression badge was positively correlated with the presence of extrapair young when the parental care badge was small and negatively correlated when the badge was large. Controlling for these crown measures, males with larger territories were less likely to suffer losses in paternity. There was no effect of breeding density, breeding synchrony, their interaction, subspecies, or weather during the fertile period on the presence of extrapair young. These results suggest that female preference for males that provide more parental care (or preference for genes that convey this trait) plays a dominant role in extrapair interactions among swamp sparrows. Models based on female assessments of relative mate quality offer a promising explanation of patterns in extrapair fertilizations among bird species.     

Differences in life-cycle traits of<Emphasis Type="Italic"> Calanus finmarchicus</Emphasis> originating from 60°N and 69°N,when reared in mesocosms at 69°N

Two different Calanus finmarchicus (Gunnerus) cohorts originating from 60°N (Bergen) and 69°N (Tromsø) were investigated in equal environmental conditions to study their different physiological responses to the same environment. A two-plus-two-bag mesocosm study was carried out between March and July 1998, in Håkøybotn, Tromsø, in order to determine development and mortality rates of the two parallel cohorts of C. finmarchicus. For practical reasons, the cohort from Bergen was incubated 10 days earlier than the Tromsø cohort. Consequently, they were exposed to elevated food conditions as compared to the Tromsø cohort. A high initial mortality among the Bergen cohort could clearly be ascribed, by genetic discrimination, to "contamination" with C. helgolandicus. After this initial mass mortality, the mortality was constantly 0.03-0.04 day^у. In cohorts starting from naupliar stage I, there was no significant difference in development or growth, the median development time (NI-CIV) differing by only 7 days (~6%). The difference in development time can be explained to a large extent (~4 days) by temperature differences. This is substantiated with model simulations using a physiological model developed for C. finmarchicus. There was a time lag in incubation between the two cohorts, resulting in elevated temperature during incubation of the Tromsø cohort. A fraction of both cohorts differentiated sexually at stage CV, with males differentiating before females. Females from both cohorts produced eggs, but specific egg production differed significantly (P>0.001, t-test). This was supported by elevated RNA:DNA ratio in females from the Bergen cohort. Both cohorts demonstrated quite similar development and physiological growth rates and, consequently, are considered to belong to the same genetic population inhabiting the Norwegian Shelf. The study demonstrates that C. finmarchicus is capable of adaptation to changes in environment and, thereby, demonstrates a significant physiological plasticity.     

Cannibalize or care? The role of perceived paternity in the sand goby, <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>

Parental care is a costly part of reproduction. Hence, natural selection should favor males which avoid caring for unrelated young. However, the decision to abandon or reduce care requires cues which are evaluated to give information on potential reproductive value of the offspring. The prediction that male sand gobies, Pomatoschistus minutus, care for foreign eggs as long as they were spawned in their own nest and at least some of such cues are fulfilled was tested. Egg-guarding males that had recently taken part in a spawning event were given a clutch of eggs that was sired either by themselves or another male, in either their own or another male’s aquarium. Males that had not taken part in a spawning event were used as controls and were given eggs sired by another male. We measured the amount of filial cannibalism and nest building. Control group males did not care for eggs and ate them all before rebuilding the nest. In the other treatments, there were no significant effects of paternity, though males moved to another male’s aquarium increased their clutch area threshold and completely consumed larger clutches than males that were not moved. There was no intermediate response in any treatment in the form of increased partial filial cannibalism or less well-constructed nests. Our results suggest that egg-guarding males cannot distinguish between eggs sired by themselves and those sired by other males but are able to react to cues indicating paternity state. Males do not adopt eggs to attract females in P. minutus.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and the Galápagos Islands (Ecuador). VI. Agariciidae, <Emphasis Type="Italic">Pavona clavus</Emphasis>

The reproductive ecology of the zooxanthellate reef coral Pavona clavus was investigated at several sites in Costa Rica, Panamá, and the Galápagos Islands (Ecuador) over the period 1985–2009. Pavona clavus displayed stable gonochorism as only five hermaphrodites were found in 590 samples. At four of five locations, sex ratios were skewed toward female dominance; however, at Saboga Island (Panamá) male colonies predominated. In Panamá, sexual maturity was observed in an estimated eight-year-old female colony, and several colonies of 10–20 years of age demonstrated gametogenesis. Sexual activity was observed at all study sites, but gamete development occurred in only 14–31% of colonies sampled sporadically. Seasonality of gametogenic activity occurred predominantly during the warm/wet season, June to August, at mainland sites (Caño Island, Costa Rica, and Gulfs of Chiriquí and Panamá, Panamá). This pattern was repeated in the Galápagos Islands, but mainly from March to May when seasonally high sea temperatures and rainfall prevailed there. Histological sampling and field observations indicated that spawning was centered around the full moon, most frequently on lunar day 17, and near sunset (1,800 h). Mean fecundity (mature ova cm^?2 live tissue) estimates were significantly different for two sites and ranged from ~1,780 (Saboga Island, Gulf of Panamá, seasonally upwelling) to ~4,280 (Uva Is, Gulf of Chiriquí, non-upwelling). Assuming three annual spawning events colony^?1 (August, September, October), extrapolation of minimum and maximum fecundities yield 5,340 and 12,840 ova cm^?2 year^?1. Seasonal, lunar, and diel spawning patterns in nine zooxanthellate species at Uva Island indicate asynchronous coral community spawning.     

The life cycles of two coexisting copepods, <Emphasis Type="Italic">Calanus chilensis</Emphasis> and <Emphasis Type="Italic">Centropages brachiatu</Emphasis>s,in the upwelling zone off northern Chile (23°S)

A time series study of the copepods Calanus chilensis and Centropages brachiatus was carried out at the coastal upwelling zone of Mejillones (23°S, northern Chile), to analyze their annual life cycles in association with upwelling variation. These species co-exist in the upwelling zone. Weekly sampling of zooplankton and oceanographic variables including Chlorophyll-a and phytoplankton composition were obtained during January–December 2002 at a fixed station (ca. 90 m depth). Stages of abundances, their proportions, changes in body length of adult females, sex ratio and egg production rate (EPR), were used as proxies to examine copepods’ demography. Upwelling, assessed by weekly Ekman transport and oceanographic conditions, was intermittent throughout the year with lack of periodicity components. Populations of both copepod species did not correlate with these non-predictable upwelling events. C. chilensis reproduced year-round and the population showed ca. 15 peaks of adults with an average time interval between peaks of 20 days. C. brachiatus showed a similar life cycle, also having 15 peaks of adults at about 22 days of time intervals. Cross-correlation functions and spectral analysis showed that both populations correlated positively through time, but not in phase, evidencing a time lag for their reproductive cycles. The lag was also evident in their population abundances. Both species differ in their development rates and this may result in non-in phase life cycles. Our findings suggest that species-dependent attributes, such as development rates, modulated by adaptations to temperature, might impose constraints in the species life cycles determining the population cycles. Such attributes must be considered when modeling and understanding population dynamics and secondary production of copepods.