Correlated evolution of male and female testosterone profiles in birds and its consequences

Circulating levels of testosterone in adults have mainly evolved as a consequence of selection on males for increased levels, while levels of circulating testosterone in females may be an indirect consequence of selection on males. A review of the literature revealed that intense directional selection for high levels of circulating testosterone in birds is likely to be mainly due to direct selection on males. A comparative study of testosterone levels in birds revealed a strong positive relationship between peak testosterone levels in adult females and peak levels in males. The slope of this relationship was significantly less than unity, implying that the testosterone levels in females have been reduced in species with high levels in males. An analysis of the order of evolutionary events suggested that peak concentration of testosterone in females changed after peak concentrations of testosterone in males. Females in colonial species of birds had significantly higher circulating peak testosterone levels compared to females of solitary species, and relative levels after controlling for the effects of peak levels in males were also larger, suggesting that any costs of high testosterone levels in females are particularly likely in colonial birds. Direct selection on male circulating testosterone levels may increase the costs that females incur from high testosterone titers. For example, high female levels may negatively affect ovulation and laying and may also affect the levels of testosterone that females deposit in their eggs and hence the exposure of pre- and post-hatching offspring to testosterone. This in turn may affect not only offspring behavior, but also offspring development and the trade-offs between growth, development of immune function, and behavior in offspring.     

Ecological genetics of the barnacle,<Emphasis Type="Italic"> Hexaminius foliorum</Emphasis>, within mangrove forests near Sydney,Australia

Spatial and temporal distribution of allozyme variation at three loci in a cohort of the barnacle, Hexaminius foliorum, living on leaves of Avicennia marina was studied from recruitment to adulthood at three geographical scales. Analysis of populations shortly after recruitment showed that there were significant divergences in allele frequencies at the coarsest geographical scale studied (between estuaries, 50 km apart, Wright's F statistic=0.016) and at the finest geographic scale (between sites, 50–100 m apart, Wright's F statistic=0.018). There was, however, no significant genetic divergence at an intermediate scale (between bays, 3–4 km apart, Wright's F statistic=0.002). The genetic differences between populations decreased over time due to the selection against the null homozygotes originally present at high frequency at two loci. There was sufficient mortality (ranging from 35.5 to 80%) between seasons to account for the deaths needed for the observed changes in allele frequencies. Differences in the genetic structure between estuaries may be the result of isolation and limited mixing of cyprid larvae among estuaries. Differences in the genetic structure between sites may be due to pre- and post-settlement mortality acting on H. foliorum.Communicated by G.F. Humphrey, Sydney     

Nest concealment and parental behaviour interact in affecting nest survival in the blackcap (Sylvia atricapilla): an experimental evaluation of the parental compensation hypothesis

Nest concealment varies strongly within populations of many species. Although some studies have revealed the beneficial effects of concealment in mitigating predation pressure on nests, other studies were unable to find similar effects. One potential reason for the mixed results is that parental behaviour may compensate for the effects of nest cover, and specifically designed experimental studies are needed to reveal this compensation. I studied the effects of concealment on the probability of nest survival in the blackcap (Sylvia atricapilla), by experimentally manipulating the degree of nest-foliage cover. There was a significant effect of the treatment depending on nest type and the phase of nesting. Whereas there was no effect of concealment on nest survival in natural nests, there was a positive effect in real nests baited with plasticine clutches (i.e. without parental activity). Parents probably behaviourally compensated for poor concealment in natural nests (nest guarding, defence). In line with this, there was no effect of concealment on nest survival during incubation, whereas there was probably a positive effect in the nestling phase. Parents spent more time on the nest during incubation (80%) than during the care of nestlings (40%) and, consequently, had more opportunities to compensate for poor cover. In general, we cannot use single measures of behaviours or states (nest concealment) as an indication of predation risk because of the capacity for compensation in other behaviours.Communicated by C. Brown     

Gastrointestinal pH and development of the acid digestion in larvae and early juveniles of<Emphasis Type="Italic"> Sparus aurata</Emphasis> (Pisces: Teleostei)

Changes in digestive pH and protease activity have been determined throughout the transition from larvae to the juvenile stage in Sparus aurata in rearing conditions (from 0.04 to 100 g wet weight). Measurements of pH have been taken in the stomach and different segments along the length of the intestine using a pH microelectrode. In starved fish, the gastric pH ranged between 6.0 and 8.0 approximately, except in juveniles of intermediate size (between 1.0 and 7.0 g wet weight), which exhibited a wider pH range of 2.0–8.0. Fed fish with digestive content showed, in general, lower pH values in the stomach. A progressive decrease was observed from a pH range of 5.5–8.0 in the youngest animals (0.04 g) to a pH range of 2.0–6.2 when juveniles were approaching 1.0 g wet weight. Above this weight, the gastric pH remained constant (between 2.0 and 6.0 approximately). The pH values in the intestine ranged between 6.7 and 8.4. They were similar in the different segments and weight classes examined, and there were no significant differences between fed and starved animals. Specific acid protease activity (units per milligram soluble protein) in fed animals increased from small (0.04–1.0 g) to intermediate juveniles (1.0 and 7.0 g), but then remained similar in larger juveniles. On the contrary, specific alkaline protease activity in fed animals decreased from small to intermediate juveniles, and then remained at a similar level in larger juveniles. The results reflect a progressive transition during several months from alkaline digestion in larvae with undeveloped stomachs to the acid digestion in juveniles with fully developed stomachs. Full gastric capacity is developed in seabream juveniles of 1 g wet weight, which represents approximately 100 days post-hatching in cultured populations. Nevertheless, in the following 2.5 months, during which the intestine reaches the appropriate length, juveniles still show a transitional period in the regulatory mechanism of digestion, probably linked to the adaptation to a different feeding habit.Communicated by S.A. Poulet, RoscoffThis revised version was published in December 2003 with corrections to the legend of Fig. 3.     

Zooplankton biomass and indices of grazing and metabolism during a late winter bloom in subtropical waters

The development of the so-called late winter bloom in subtropical water was studied in an oceanic area north of the Canary Islands from January to May 2000. Zooplankton was sampled at short-term intervals (1–4 days) during the bloom (January–March), and biomass, indices of grazing (gut fluorescence) and metabolism (electron transfer system activity, ETS) were measured in four different size fractions (100–200, 200–500, 500–1000 and >1000 µm). During the bloom, ETS activity and gut fluorescence increased before the development of zooplankton biomass. At the end of February, the presence of an impressive cloud of dust formed in the Sahara desert was related to an increase in chlorophyll and small zooplankton a week later. The increments in biomass were the consequence of consumption by zooplankton as inferred from the indices of grazing and metabolism. Estimated grazing from gut fluorescence and gut evacuation rates during the period of study accounted for 55% of the assessed total ingestion from respiration and normal values of assimilation, showing the importance of the non-pigmented food in the diet of zooplankton in these waters. In contrast, the sharp decreases in zooplankton biomass observed during the bloom appeared during the dark period of the moon, the days in which the diel vertical migrants reach the shallower layers, in agreement with previous works in the area. Thus, the development of the late winter bloom in this region is suggested to be driven by the interplay between resource and consumer controls.Communicated by S.A. Poulet, Roscoff     

The costs of secondary sexual characters in the lekking great snipe (Gallinago media)

Summary Recent studies of the lekking great snipe (Gallinago media) have shown that male mating success increases with display rate and the amount of white on the tail. An obvious question then arises: what is setting the limits to male display rate and to the evolution of even whiter tails? In this study the costs of these two traits are considered. In the case of the white on the tail, almost no predation was observed during the display period. However, predation on the winter grounds and/or during migration cannot be ruled out. Males differed in the amount of white on the tail between years. It is suggested that increasing the amount of white is costly, though the reason for this is still unknown. In the case of display rate, it is shown that predation cannot be a considerable cost, again because of the near absence of predation on the leks. Instead, it is shown that male displays are energetically costly. The daily energy expenditure of displaying males was estimated to be on average 4 times the basal metabolic rate, which is close to the supposed maximum sustainable work load for birds. Furthermore, males lost on average 1.8 g per h (1.7% of the body mass) which is about 7.2 g (6.8%) during a night of display activity. Males do gain some weight during the day when no displays are performed but loose on average 9 g during an 11-day period in the early part of the breeding season. These results indicate that high energetic demands set the limits to male display performance and that males do not have to trade off the benefit of attracting females by their displays against a cost of attracting predators. Offprint requests to: J. Höglund     

Soil erosion and significance for carbon fluxes in a mountainous Mediterranean-climate watershed.

In topographically complex terrains, downslope movement of soil organic carbon (OC) can influence local carbon balance. The primary purpose of the present analysis is to compare the magnitude of OC displacement by erosion with ecosystem metabolism in such a complex terrain. Does erosion matter in this ecosystem carbon balance? We have used the Revised Universal Soil Loss Equation (RUSLE) erosion model to estimate lateral fluxes of OC in a watershed in northwestern Mexico. The watershed (4900 km2) has an average slope of 10 degrees +/- 9 degrees (mean +/- SD); 45% is >10 degrees, and 3% is >30 degrees. Land cover is primarily shrublands (69%) and agricultural lands (22%). Estimated bulk soil erosion averages 1350 Mg x km(-2) x yr(-1). We estimate that there is insignificant erosion on slopes < 2 degrees and that 20% of the area can be considered depositional. Estimated OC erosion rates are 10 Mg x km(-2) x yr(-1) for areas steeper than 2 degrees. Over the entire area, erosion is approximately 50% higher on shrublands than on agricultural lands, but within slope classes, erosion rates are more rapid on agricultural areas. For the whole system, estimated OC erosion is approximately 2% of net primary production (NPP), increasing in high-slope areas to approximately 3% of NPP. Deposition of eroded OC in low-slope areas is approximately 10% of low-slope NPP. Soil OC movement from erosional slopes to alluvial fans alters the mosaic of OC metabolism and storage across the landscape.     

Sex allocation in the solitary bee Osmia cornuta: do females behave in agreement with Fisher's theory?

Fisher's theory of sex allocation predicts that, in a panmictic population, parental investment will be equally distributed between male and female progeny. Most studies on parental investment in nesting solitary bees and wasps use offspring or provision weight as estimators of parental investment and do not corroborate Fisher's theory. The measurement of parental investment may be confounded by several factors. First, the use of offspring or provision size does not account for seasonal variation in foraging costs associated with aging of nesting females. Second, provision or offspring size do not reflect parental investment associated with nest construction. In this two-year study we measured parental investment in a solitary bee. We calculated sex allocation using both provision weight and foraging time as parental investment estimators. Investment in pollen-nectar provisions decreased, while investment in mud structures (nest construction) increased, as the nesting period progressed. Overall investment in provisions per nest was ∼25 times higher than investment in mud. Pollen-nectar foraging trips became longer as the season progressed, but mud trip duration did not vary. Due to weather differences between years, more offspring per female were produced in the first year, but progeny sex ratio and mean offspring size of both sexes were similar between years. Mortality did not differ between sexes. As predicted by Fisher's theory, production cost ratios did not differ from 1 in either year, irrespective of the currency used to estimate parental investment (provision weight or foraging time). Our results strongly support Fisher's theory.     

Early life histories in the bryozoan<Emphasis Type="Italic"> Schizobrachiella sanguinea</Emphasis>: a case study

Mediterranean populations of Schizobrachiella sanguinea seem to span a wide range of contrasting benthic habitats. We have studied adult, larval, and recruit populations of S. sanguinea from sublittoral (approximately 10 m depth) communities at Blanes (Spain, NW Mediterranean). At the study site, the species colonises semi-obscure caves and, to a lesser extent, communities of sciaphilic algae. Our first goal was to show the periods of brooding, larval release, and recruitment. Plankton was collected about twice a week between July 1998 and June 2000 over an artificial reef. A total of 303 larvae were collected in 23 of the 102 days of sampling. At the study site the species shows a single, annual larval release period, which occurs from late March to June–July every year. Recruits were found from late May to late August 1999 and in late May 2000. A recruitment peak was observed in July 1999 and again in late May 2000. The presence of brooding adults corresponded to the periods of larval supply and recruitment. Our second goal was to investigate the causes of the variable distribution of the species along several communities in the north-west Mediterranean Sea. We describe adult distribution and the dynamics of larval supply, recruitment of early (15 days from settlement) and late (4 months from settlement) recruits. Patterns of larval presence and early recruit distribution along the communities on the reef did not reflect the adult distributions, whereas this was closely matched by the distribution of late recruits. Post-recruitment mortality strongly affected recruits of the PA (photophilic algae) and SA (sciaphilic algae) communities, whereas it was very low in the communities of SOC (semi-obscure caves). Mortality of recruits was density independent and colony survivorship in the SA communities, where recruitment was the highest, appeared strongly affected by competition with brown algae. Like the majority of cheilostomate bryozoans, S. sanguinea releases a coronate larva capable of active swimming and positive reaction to light. Such swimming abilities may allow larvae to actively select the substrate on which to settle and to avoid unsuitable substrates. It seems that larval behaviour does not explain the preference of S. sanguinea for sciaphilic habitats. High post-recruitment mortality due to various factors, especially competition with fast-growing algae, seems particularly relevant in determining the adult distributions observed.Communicated by S.A. Poulet, Roscoff     

The gametogenic cycle of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Mollusca: Cephalopoda) as observed on the Atlantic coast of Andalusia (south of Spain)

The present study reports the gametogenic cycle of the common octopus, Octopus vulgaris Cuvier, 1797 in the Atlantic coast of Andalusia between May 2001 and May 2002. A total of 673 specimens of total weight ranging from 30 to 3,732 g were analysed. The gonad development was defined by means of histological methods and gonadosomatic indexes. Four gametogenic stages were identified in males (I immature, II maturing, III mature, IV spawning) and in females (I immature, II maturing, III pre-spawning, IV spawning). The sex ratio obtained was not significantly different from 1:1 (P>0.01). Mature males were found during the whole year, September being the month when the highest rate of sexually active individuals occurred (48%). In females, the gametogenic cycle commenced in December and ended in November, and spawning occurred between April and October, with two maximum peaks in August and September (48% in each). Size at sexual maturity was 850 g of total body weight for males and 1,250 g of total body weight for females. Males mature earlier than females.Communicated by S.A. Poulet, Roscoff