Annual variation in the seasonal shift in egg size and clutch size in Sceloporus woodi

Summary Early and late season clutch parameters were examined over a three year period in the Florida scrub lizard, Sceloporus woodi. Precipitation levels were monitored throughout the study. In the early and late season of 1984 and the early season of 1986 precipitation levels approximated long-term mean levels of precipitation. In 1985 a severe winter drought occurred. Clutch size was positively related to body size in all samples in every year. In 1984 and 1986, egg size was not related to clutch size, whereas, in 1985 egg size was negatively related to clutch size. In 1985, females produced large clutches of small eggs early in the season and small clutches of large eggs late in the season. In 1984, no seasonal changes in egg or clutch size occurred. In the late season of 1986, females produced the largest clutches and the smallest eggs of all the samples, but egg and clutch size were not statistically different from the early season egg and clutch size of 1986. Total clutch dry weight, an estimate of total clutch energy, was not different in any of the six sampling periods. These data do not support current adaptationist models that attempt to explain the control of clutch and egg size in lizards. It is argued in this paper that egg and clutch size may vary in response to past environments that affect a female's physical condition, as well as, current resources that may be important for maintenance and reproduction. Egg and clutch size appear to be plastic traits selected to respond to proximal environmental variation, whereas, the investment of total dry matter/clutch has been optimized.     

A comparative study of egg mass and clutch size in the Anseriformes

The factors explaining interspecific differences in clutch investment in precocial birds are poorly understood. We investigated how variations in clutch characteristics are related to environmental factors in a comparative study of 151 extant species of ducks, geese and swans (Anseriformes). Egg mass was negatively related to clutch size in a phylogenetic regression, a relationship that was much stronger when controlling for female mass. Nest placement was related to both egg size and clutch size, with cavity-nesting species laying more but smaller eggs. Egg size was positively correlated with incubation period and with female mass, and also with sexual size dimorphism (i.e. male mass relative to that of the female). Clutch size was not related to female mass. Species with long term pair bonds laid smaller clutches and larger eggs. The size of the breeding range was strongly positively correlated with clutch size and clutch mass, and its inclusion in multivariate models made other biogeographical variables (hemisphere, breeding latitude or insularity) non-significant. The small clutches in insular species appear to be a product of small range size rather than insularity per se. Our results suggest there is an evolutionary trade-off between clutch and egg size, and lend support to Lack’s resource-limitation hypothesis for the waterfowl.     

Habitat size, flora, and fauna: Interactions in a tidal saltwater marsh

Anthropogenic habitat fragmentation is increasingly problematic in both terrestrial and aquatic systems. Fragmentation reduces the size of habitat patches, so examining the effect of patch size on community structure can provide insight into the potential effects of fragmentation. In this study, we examined the effect of habitat size on the density of Spartina alterniflora shoots in tidal saltwater marshes, as well as on the two predominant macrofaunal species, the marsh periwinkle Littoraria irrorata and fiddler crabs Uca spp. We estimated the density of shoots in three different marsh habitats, (1) large island marshes, (2) small island marshes, and (3) large fringing marshes, in Indian Field Creek, York River, Chesapeake Bay. We manipulated shoot density in each of the marsh types to distinguish between the effects of marsh grass density and marsh type on crab and Littoraria densities in the system. We found significant differences in grass density among the three marsh types as well as significant species-specific effects of grass density, marsh type, and distance from edge on faunal abundance. Decreasing the shoot density resulted in a decrease in Littoraria density in the large marshes. Littoraria density increased with distance from edge in the small marshes and in the first 5 m of the fringing marshes, then decreased with distance from edge after 5 m in the fringing marshes. Shoot density had a negative effect on crabs in both the large and small marshes. These results suggest that fragmentation would have a negative effect on the community structure by lowering the densities of both the flora and fauna.     

Experimentally reducing clutch size reveals a fixed upper limit to egg size in snakes, evidence from the king ratsnake, Elaphe carinata

Snakes are free of the pelvic girdle's constraint on maximum offspring size, and therefore present an opportunity to investigate the upper limit to offspring size without the limit imposed by the pelvic girdle dimension. We used the king ratsnake (Elaphe carinata) as a model animal to examine whether follicle ablation may result in enlargement of egg size in snakes and, if so, whether there is a fixed upper limit to egg size. Females with small sized yolking follicles were assigned to three manipulated, one sham-manipulated and one control treatments in mid-May, and two, four or six yolking follicles in the manipulated females were then ablated. Females undergoing follicle ablation produced fewer, but larger as well as more elongated, eggs than control females primarily by increasing egg length. This finding suggests that follicle ablation may result in enlargement of egg size in E. carinata. Mean values for egg width remained almost unchanged across the five treatments, suggesting that egg width is more likely to be shaped by the morphological feature of the oviduct. Clutch mass dropped dramatically in four- and six-follicle ablated females. The function describing the relationship between size and number of eggs reveals that egg size increases with decreasing clutch size at an ever-decreasing rate, with the tangent slope of the function for the six-follicle ablation treatment being -0.04. According to the function describing instantaneous variation in tangent slope, the maximum value of tangent slope should converge towards zero. This result provides evidence that there is a fixed upper limit to egg size in E. carinata.     

The influence of egg size and colour on egg consumption by the Algerian hedgehog (<Emphasis Type="Italic">Atelerix algirus</Emphasis>)

We tested the hypothesis that the Algerian hedgehog (Atelerix algirus) consumes smaller eggs in preference to larger ones and that the colour of the eggs (brown or white spotted) does not influence the egg consumption rate. Eggs of three galliform species (the quail, Coturnix japonica; the red legged partridge, Alectoris rufa; the chicken, Gallus domesticus) were offered to 14 individual hedgehogs under experimental conditions. Overall, the hedgehogs consumed 107 of the 225 eggs offered (48%) in 5 days. The colour of the eggs did not influence the rates of egg consumption. However, egg size had a significant influence on the rates of egg consumption. Generalized linear mixed models showed that hedgehogs preferred to consume smaller quail eggs (91%, n = 65) than intermediate partridge eggs (44%, n = 110), whereas the largest chicken eggs (n = 50) were not consumed at all. There was a strong inter-individual variation in egg consumption, particularly of partridge eggs, from 0 to 100%, whereas individual variation in consumption of quail eggs was lower, from 70 to 100%.     

Distribution and diversity of Gallionella-like neutrophilic iron oxidizers in a tidal freshwater marsh

Microbial iron oxidation is an integral part of the iron redox cycle in wetlands. Nonetheless, relatively little is known about the composition and ecology of iron-oxidizing communities in the soils and sediments of wetlands. In this study, sediment cores were collected across a freshwater tidal marsh in order to characterize the iron-oxidizing bacteria (FeOB) and to link their distributions to the geochemical properties of the sediments. We applied recently designed 16S rRNA primers targeting Gallionella-related FeOB by using a nested PCR-denaturing gradient gel electrophoresis (DGGE) approach combined with a novel quantitative PCR (qPCR) assay. Gallionella-related FeOB were detected in most of the samples. The diversity and abundance of the putative FeOB were generally higher in the upper 5 to 12 cm of sediment than in deeper sediment and higher in samples collected in April than in those collected in July and October. Oxygen supply by macrofauna appears to be a major force in controlling the spatial and temporal variations in FeOB communities. The higher abundance of Gallionella-related FeOB in April coincided with elevated concentrations of extractable Fe(III) in the sediments. Despite this coincidence, the distributions of FeOB did not exhibit a simple relationship to the redox zonation inferred from the geochemical depth profiles.     

Determinants of seed bank dynamics of two dominant helophytes in a tidal salt marsh

We investigated dynamics and spatial distribution of Scirpus mariqueter and Spartina alterniflora seed banks at Chongming Dongtan in the Yangtze River estuary, China. Five sites along an elevational gradient were chosen, one in each of the main zones (mudflat, Scirpus monoculture, Scirpus–Spartina mixture, Spartina monoculture and Spartina–Phragmites mixture). Three surveys were performed just after seed rain, before germination and after germination, respectively. During the period of November 2005 to May 2006, soil seed density of Scirpus mariqueter declined by 36%, and that of Spartina alterniflora by 58%. The spatial distributions of their seed banks were also different. Soil seed density of Scirpus mariqueter was not determined directly by seed production, but positively correlated with total aboveground biomass of the whole plant community. On the contrary, soil seed density of Spartina alterniflora just after seed rain (November) was significantly correlated with seed production, but had a poor relationship with the community's aboveground biomass. Our results indicated that other factors such as tidal movement might have had great influence on dispersal of Scirpus mariqueter, which would also affect its population dynamics. The understanding of this process can help us improve the conservation and restoration efforts.     

The white colour of the Ostrich ( Struthio camelus ) egg is a trade-off between predation and overheating

Most ground nesters lay pigmented eggs, and egg pigmentation generally matches the environment. Pigmentation of eggs has evolved as a protective device against predation, but dark-pigmented eggs can be susceptible to overheating when exposed to solar radiation. The Ostrich (Struthio camelus) lays white eggs that are unattended for the first few weeks before incubation, and are quite visible to predators. To evaluate the effect of colour on the surface and core temperatures, we painted some Ostrich eggs dark brown or white, and left some unpainted (control), and exposed all of them directly to the sun during the day. The surface and core temperatures of brown eggs were significantly higher than those of the white-painted and control eggs. In addition, the core temperature of brown eggs exceeded 37.5°C, which is the temperature at which embryo mortality starts to increase. In a second experiment, we placed eggs (brown-painted and control) in various types of vegetation to study their visibility to an observer walking towards them. The white eggs were discovered from a significantly longer distance than the brown eggs, indicating that the predation risk may be much higher for white eggs. The results thus suggest that white eggs minimise overheating and allow the Ostrich to leave its eggs unattended before incubation starts, but they are more susceptible to predation.     

Vegetation changes in the freshwater tidal marsh of the Delaware estuary

We examined the areal extent and changes in thefreshwater tidal wetlands along a 56.4 km and a80.6 km reach of the Delaware River between Chester,Pa. and Trenton, N.J. Most of the remainingfreshwater tidal wetlands of the Delaware River arefound along tributaries which drain the coastal plainof New Jersey. We identified polygons of marsh, mud,and open water using color infrared aerial photographyobtained at low tide in 1977 and 1978. Marsh polygonswere classified into either high marsh or low marshaccording to the dominant visual signature of thevegetation of each polygon, and placed in a geographicinformation system for subsequent analysis. The totalarea of marsh within the two reaches totaled 1416 ha,of which 71% was high marsh and 29% low marsh. Asite re-examination in 1997 and 1998 of marsh arearepresenting 32% of the total marsh area revealedthat, while the total area of wetland appears to haveremained constant, high marsh vegetation along thelower reaches of the tributaries has been replaced bylow marsh vegetation. The fraction of the sample thatwas low marsh increased from 9% in 1977–78 to 34% in1997/8.     

Encountering competitors reduces clutch size and increases offspring size in a parasitoid with female-female fighting

Understanding the size of clutches produced by only one parent may require a game-theoretic approach: clutch size may affect offspring fitness in terms of future competitive ability. If larger clutches generate smaller offspring and larger adults are more successful in acquiring and retaining resources, clutch size optima should be reduced when the probability of future competitive encounters is higher. We test this using Goniozus nephantidis, a gregarious parasitoid wasp in which the assumption of size-dependent resource acquisition is met via female-female contests for hosts. As predicted, smaller clutches are produced by mothers experiencing competition, due to fewer eggs being matured and to a reduced proportion of matured eggs being laid. As assumed, smaller clutches generate fewer but larger offspring. We believe this is the first direct evidence for pre-ovipositional and game-theoretic clutch size adjustment in response to an intergenerational fitness effect when clutches are produced by a single individual.     

On the evolution of clutch size and nest size in passerine birds

Summary I examined the hypothesis that the clutch size of some altricial birds may be limited by over-crowding of the nestlings in the nest, by comparing data on different species of European passerines. Large-sized birds build, relative to the body, larger nests than small-sized birds, both as regards the inner and the outer nest widths and as regards edge breadth; only inner nestcup depth did not change relatively to body size. Nest size also varied in relation to nesting place. Birds with open nests built off the ground had a rather narrow nestcup, whereas those with a domed nest, or which nest in a cavity, had a wide nestcup. When only open-nesters were compared, birds nesting on, or close to, the ground tended to have a wider nestcup than birds nesting above the ground. Inner nestcup width was correlated with the amount of mosses and lichens used in building the nest; the more of such materials the narrower the nestcup. The three variables: standardised body size, nesting place, and type of nesting material used accounted for 92% of the overall variation observed in inner nestcup width. When controlling for adult body size, clutch size was positively correlated with the size of the nestcup. A multiple regression analysis showed that relative nestcup depth, nest site, and type of nesting materials used, accounted for 64% of the overall variation in clutch size.     

Intra-seasonal decline of clutch size in Lesser Snow Geese

Summary Different mechanisms proposed to explain the intra-seasonal decline in clutch size of Lesser Snow Geese (Anser caerulescens caerulescens) were tested at the La Perouse Bay colony, Manitoba, Canada. Ovary examination of females collected after laying revealed that the actual number of eggs produced per female decreased over the laying period. This finding eliminates nest-parasitism, partial clutch predation and renesting attempts as sufficient explanations for seasonal clutch size decline. Follicular atresia induced by reserve depletion was also rejected since its occurrence was similar among early and late nesters. The decline in clutch size was observed within age-classes, and therefore age effects on clutch size and laying date per se do not account for the observed relationship. Clutch size and laying date were respectively positively and negatively correlated with the amount of nutrient reserves in females at the onset of laying, and also covaried within individuals observed breeding in several seasons. Laying date repeatability was estimated at 0.22. It is postulated that the seasonal decline in clutch size results mainly from a positive feed-back of female's nutrient reserves on the hormones controlling ovary development. Hatching synchrony or shortness of the breeding season may be the ultimate factors responsible for the intra-seasonal clutch size decline in Lesser Snow Geese.     

Lethal and sublethal costs of autotomy and predator presence in damselfly larvae

We studied the costs of lamellae autotomy with respect to growth and survival of Lestes sponsa damselfly larvae in field experiments. We manipulated predation risk by Aeshna cyanea dragonfly larvae and lamellae status of L. sponsa larvae in field enclosures and compared differences in numbers, size and mass of survivors among treatments. In the absence of a free-ranging A. cyanea larva, about 29% of the L. sponsa larvae died. This was probably due to cannibalism. The presence of a free-ranging A. cyanea reduced larval survival by 68% compared to treatments in which it was absent or not permitted to forage on L. sponsa damselflies. Across all predator treatments, lamellae autotomy reduced survival by about 20%. The mean head width and mass of survivors was lower in the enclosures with a free-ranging A. cyanea compared to the other two predator treatments. This suggested that larvae grew less in the presence of a free-ranging predator, indicating that increased antipredator behaviours were more important in shaping growth responses than reduced population density. Mass, but not head width, of survivors was also reduced after autotomy. The fitness consequences of these effects for the adults may be pronounced. In general, these field data strongly suggest that lamellae autotomy affects population regulation of damselflies. Received: 1 February 1999 / Accepted: 22 March 1999     

Experimental manipulation of egg quality in chickens: influence of albumen and yolk on the size and body composition of near-term embryos in a precocial bird

The importance of avian egg components in the determination of hatchling size and quality has yet to be fully evaluated. In the first experiment, 20% of the albumen and/or the yolk was removed from chicken eggs to determine the impact of each egg component on metabolism and various size measures in near-term embryos. Results show that metabolic rate, dry body mass, and internal organ mass are largely independent of egg composition. Removal of albumen resulted in a decrease in wet body mass corresponding to decreases in water content in the body and the yolk sac, and decreased tibiotarsus length. Removal of yolk resulted in no change in body mass, but decreases in both wet and dry yolk sac mass. In a second experiment, removal of 15% of either egg component led to reductions in hatchling mass similar to those observed in whole near-term embryos. Albumen, as the primary source of water in the egg, is the primary determinant of hatchling size and may influence hatchling success through size-related limiting factors. Differences in yolk content may influence neonatal quality as a nutritional supplement, but seem not to result in greater tissue formation during embryonic development. Accepted: 2 September 1997     

Fine-scale environmental heterogeneities of tidal creeks affect distribution of crab burrows in a Chinese salt marsh

Tidal creeks are an important structure of salt marshes in estuarine ecosystems, providing valuable ecosystem services to wildlife in the estuary. To determine the effects of environmental heterogeneities within tidal creeks on the features of crab burrows, we divided a typical creek section into four parts (i.e., microhabitats): bottom, slope, edge and flat, investigated the distribution of crab burrows and sediment properties on creek sections in the Yangtze River estuary, and compared the burrow distribution in tidal creeks with that in non-creek areas. Our results showed that from the creek bottom to flat soil water content declined (F_3, 60 = 93.8, p < 0.001), and the variations of other sediment physical and chemical properties associated with the change of soil water content were significant among the microhabitats on the creek sections (p < 0.001 for pH, conductivity, and grain size). No crab burrows were found at the creek bottom. The burrows on the slope were smaller in size (p < 0.001 for burrow opening diameter) while the density was higher than that at the edge and on the flat (F_2, 45 = 31.2, p < 0.001). Moreover, although the correlations between burrow distribution and sediment properties varied among the microhabitats on the creek sections, crabs generally selected relatively solid sediments to build their burrows. On the slope, there was a significantly negative relationship between burrow density and soil water content (r² = 0.53, p < 0.001). At the edge, the correlation between total burrow opening area and soil water content was significantly negative (r² = 0.44, p < 0.002). The density of small crab burrows (<10 mm) was greater, but that of large burrows (>10 mm) was lower in tidal creeks than in non-creek habitats. Therefore, sediment properties showed a gradual transition from hydrophytic to terrestrial environments on the creek section, which caused significant differences of burrow distribution among the microhabitats. The creeks of tidal salt marshes could affect ecological processes and functioning through affecting crab burrows.     

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the ‘energetic incubation constraint’ hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.