Reproductive ecology of red-chinned lizards (Sceloporus undulatus erythrocheilus) in Southcentral Colorado: comparisons with other populations of a wide-ranging species

Summary We studied the reproductive ecology of a population of Sceloporus undulatus erythrocheilus near Walsenburg, CO during the summers of 1987–1990. Reproductive activity commences soon after emergence in early May and continues until the middle of July. Females mature in their second year following hatching at an age of 20–21 months and a size of 60–63 mm snout-vent length (SVL). Two clutches, averaging10.9 eggs, are produced per year. Larger females produce larger clutches with an increase of one egg per 3 mm SVL. Average SVL of females was 71 mm. Eggs are reasonably large (0.32 g) and relative clutch mass was the largest reported for the species (34% of body mass). A principal component analysis was used to examine relationships among reproductive characters in 12 populations of Sceloporus undulatus. The analysis suggested that patterns of reproduction in this wide-ranging species result from several factors reflecting both adaptive and phylogenetic sources of variation.     

Maternal body size as a morphological constraint on egg size and fecundity in butterflies

It is a widespread notion that in arthropods female reproductive output is strongly affected by female size. In butterflies egg size scales positively with female size across species, suggesting a constraint imposed by maternal size. However, in intraspecific comparisons body size often explains only a minor part of the variation in progeny size. We here include representatives of various butterfly families to test the generality of this phenomenon across butterflies. Phenotypic correlations between egg and maternal body size were inconsistent across species: correlations were non-significant for Pararge aegeria and Lycaena tityrus, significantly positive for Papilio machaon, significantly negative for Araschnia levana, and contradictory for Pieris napi. Thus, there was no general pattern linking egg size to maternal size, e.g., caused by an allometric relationship. Consequently, there was at best limited evidence for maternal size acting as a morphological constraint on egg size within butterfly species. Realized fecundity depended on maternal size in P. napi and A. levana, but not in P. aegeria, suggesting that maternal size may affect egg number more strongly than egg size. Yet, variation in fecundity was primarily explained by variation in longevity as is expected for income breeders. Heritability estimates across species were rather similar for pupal mass (ranging between 0.14 and 0.19), but more variable for egg size (0.17–0.31).     

Reproductive variation and the egg size-clutch size trade-off within and among populations of painted turtles (Chrysemys picta bellii)

Interpopulation variation in egg size, clutch size and clutch mass was studied 3 years in four populations of painted turtles (Chrysemys picta bellii) from western Nebraska. Body size varied among all populations and was larger in two large (56–110 ha), sandhills lake populations than in two populations in smaller habitats (1.5–3.6 ha) of the Platte River floodplain. Reproductive parameters (egg mass, clutch mass, and clutch size) generally increased with maternal body size within populations. Clutch wet and dry mass varied among populations but largely as a function of maternal body size. Clutch size was largest in the sandhills lake populations, both absolutely and relative to maternal body size. Egg mass was smallest in the sandhills lakes and varied annually in one population. Over all populations, an egg sizeclutch size trade-off was detected (a negative correlation between egg mass and clutch size) after statistically removing maternal body size effects. Egg wet mass and clutch size were negatively correlated over all years within the sandhills populations and in some years in three populations. Although egg size varied within populations, egg size and clutch size covaried as expected by optimal offspring size models. Thus, patterns of egg size variation should be interpreted in the context of proximate or adaptive maternal body size and temporal effects. Comparisons among populations suggest that large egg size relative to maternal body size may occur when juvenile growth potential is poor and mean maternal body size is small.     

Effects of crowding and different food levels on growth and reproductive investment of Daphnia

The effects of daphniid crowding on juvenile growth rate, length at first reproduction, clutch size and egg size of four species of Daphnia were compared with the effects of food level. Juvenile Daphnia were grown to primipary in a flow-through system in water conditioned by different densities of the same, or another, species. At high ambient food levels, water from Daphnia that had been crowded at densities 150 l^–1 depressed growth rate and lowered body size and clutch size of D. hyalina and D. galeata; effects on the same traits of D. magna and D. pulicaria were variable (stimulation, depression, or no effect). D. hyalina and D. galeata responded to gradients of increasing daphniid density (0–300 l^–1) by altering egg mass, somatic mass and clutch size to maintain a relatively constant reproductive investment; egg mass increased with crowding and then decreased in a pattern consistent with Glazier's (1992) hypothetical model of changes in offspring size in relation to food quantity and maternal demand. Effects of crowding by conspecifics were indistinguishable from those of other species. This study, which uncouples the effect of crowding per se from ambient resource depletion, shows that chemical substances released by high densities of Daphnia can cause changes in life-history traits comparable to those that occur in response to low food levels.     

Dynamics of heat transfer to cold eggs in incubating bantam hens and a black grouse

Incubating birds transfer large amount of heat from the brood patch to the eggs during rewarming of cold eggs. If a vasoconstriction is present in the brood patch as in other parts of the body, it could possibly limit heat transfer to the eggs. To investigate this, heat transfer to water-circulated eggs was measured in incubating bantam hens (Gallus domesticus) and a black grouse hen (Lyrurus tetrix) during exposure to cold eggs. Egg temperature, egg surface temperature, heat production and cloacal temperature were also measured. At all levels of egg cooling, egg surface temperature and heat transfer to the eggs was stable throughout an exposure, except during resettling movements, which often changed egg surface temperature and the level of heart transfer. Egg surface temperature decreased linearly with egg temperature in both species, but was lower and more variable at low egg temperature in black grouse than in bantam hens. A higher proportion of the heat production was transferred to the eggs in the black grouse (corresponding to 109–118% of the increase above resting level) than previously reported in bantam hens. Clutch size did not affect this efficiency of heat transfer in black grouse. It is concluded that a vasoconstruction of the brood patch does not occur even under strong cold stress from the eggs. Heat transfer to the eggs is probably controlled more by behavioural adjustments than circulatory changes. An increase in brood patch blood flow probably occurs at relatively high egg temperature at the onset of egg rewarming. The efficiency of heat transfer, and thus the energetic cost of rewarming eggs, depends on the insulation of the bird and nest structure. The boreal/subarctic black grouse was able to reduce heat loss to the environment and transfer a higher proportion of its heat production to the eggs than the tropical bantam hen.Abbreviations AVAs arteriovenous anastomoses - HP heat production - HT heat transfer - T _a ambient temperature - T _b cloacal temperature - T _brp brood patch temperature - T _e egg temperature - T _es egg surface temperature     

Nonlinear continuum of egg size-number trade-offs in a snake: is egg-size variation fitness related?

The relationship between offspring size and offspring number is crucial to life history evolution. To examine how these two life history variables are coupled and whether an altered balance between them will result in changes in maternal fitness, we manipulated clutch size of the Chinese cobra (Naja atra) by using the techniques of hormonal manipulation and follicle ablation. Females receiving exogenous follicle-stimulating hormone produced more but smaller eggs, and females undergoing follicle ablation produced fewer but larger eggs. Neither body size (body mass and snout-vent length) at hatching nor egg mass at oviposition had a role in determining hatchling survival and growth. Female hatchlings were more likely to die in early post-hatching days and grew more slowly than male hatchlings. Our data show that: (1) there is a nonlinear continuum of egg size-number trade-offs in N. atra within which there is a single inflexion where the rate at which egg size decreases with increasing clutch size, or clutch size increases with decreasing egg size, is maximized; (2) there is a fixed upper limit to egg size for a given-sized female, and the limit is not determined by her body volume; (3) egg size has no role in determining hatchling survival and growth; and (4) the extent to which females may enjoy reproductive benefits in a given reproductive episode depends on how well egg size and egg number are balanced.     

Abundance and life history ofNeomysis intermedia Czerniawsky in Lake Kasumigaura

Weekly observations ofNeomysis intermedia in Lake Kasumigaura showed two major peaks in abundance during spring and autumn (more than 10⁴ individuals m^–2) and minimum levels in summer and winter (less than 10³ individuals m^–2). Their increase in abundance followed a high egg ratio, suggesting that the increase in abundance was caused by a high reproductive rate. Major contributors to mysid population decreases include fish predation and commercial fisheries, and possible horizontal migration of the mysids. N. intermedia showed two types of life history in the lake. One type (overwintering generation) has a life span of about 6–7 months and produces about 27 eggs per brood. Another, appearing from spring to autumn, matures in 3–6 weeks at a smaller size, and produces 12 eggs per brood. The reproductive season ofN. intermedia was continuous from March through November.     

Use of swimming speed and egg ratio as predictors of the status of rotifer cultures in aquaculture

This study evaluated the use of egg ratio (eggs rotifer^–1) and swimming speed (mm min^–1) as prediction criteria for production and culture quality in mass cultures of the rotifer Brachionus plicatilis. Egg ratio was determined to be a suitable predictor of rotifer growth and production in the cultures. Low egg ratios (i.e., 0–0.17 eggs rotifer^–1) indicate reduced rotifer population over time (i.e., negative net population growth rates). However, at this time egg ratio dynamics are not suitably understood to predict in advance a sudden population collapse.Swimming speed of reproductive, egg-carrying females in the exponential growth phase was 40–45 mm min^–1. During exponential growth swimming speed was independent of the food used. Lower swimming speeds were obtained in late stationary phase (10–25 mm min^–1) when yeast was used as a food source. Both environmental factors (e.g., accumulating metabolites) and changes in nutritional state of the rotifers may have affected the swimming speed, but environmental factors appear to be the most important. We believe that swimming speed has the potential of becoming an accurate predictor of culture quality in mass cultures of rotifers.     

It takes two: Seasonal variation in sexually dimorphic weaponry results from divergent changes in males and females

Sexually dimorphic weaponry often results from intrasexual selection, and weapon size can vary seasonally when costs of bearing the weapon exceed the benefits outside of the reproductive season. Weapons can also be favored in competition over nonreproductive resources such as food or shelter, and if such nonreproductive competition occurs year‐round, weapons may be less likely to vary seasonally. In snapping shrimp (Alpheus angulosus), both sexes have an enlarged snapping claw (a potentially deadly weapon), and males of many species have larger claws than females, although females are more aggressive. This contrasting sexual dimorphism (larger weaponry in males, higher aggression in females) raises the question of whether weaponry and aggression are favored by the same mechanisms in males and females. We used field data to determine whether either sex shows seasonal variation in claw size such as described above. We found sexual dimorphism increased during the reproductive season due to opposing changes in both male and female claw size. Males had larger claws during the reproductive season than during the nonreproductive season, a pattern consistent with sexual selection. Females, however, had larger claws during the nonreproductive season than during the reproductive season—a previously unknown pattern of variation in weapon size. The observed changes in female weapon size suggest a trade‐off between claw growth and reproduction in the reproductive season, with investment in claw growth primarily in the nonreproductive season. Sexually dimorphic weaponry in snapping shrimp, then, varies seasonally due to sex differences in seasonal patterns of investment in claw growth, suggesting claws may be advantageous for both sexes but in different contexts. Thus, understanding sexual dimorphisms through the lens of one sex yields an incomplete understanding of the factors favoring their evolution.     

Annual cycle of energy and time expenditure in a golden-mantled ground squirrel population

Summary We have analyzed seasonal shifts of energy and time allocation in a population of golden-mantled ground squirrels (Spermophilus saturatus) by directly measuring total daily energy expenditure (DEE) with an isotopic technique (doubly labeled water=dlw), and by estimating components of total DEE through an integration of field behavioral observations with laboratory-measured rates of energy expenditure (oxygen consumption) associated with major behavioral and physiological states. Hibernation laster about 7 1/2 months, and the 4 1/2-month activity season consisted of mating, a 28-d gestation of 3–5 young, 5 1/2 weeks of postnatal growth building to a peak in lactation just before the young emerged above ground, an additional 2–3-week period of maternal care before dispersal, and finally restoration of body mass preceding hibernation. Although the hibernation season comprised nearly two-thirds of the year, it involved only 13–17% of annual energy expenditure, leaving about 85% of energy expenditure for the active season. Ground squirrels were actually present on the surface for only about 11% of the year's time, and the foraging time required to obtain the total annual energy supply amounted to only about 2% of the year's time. The squirrels fed mainly on herbs in the early season and hypogeous fungi later; both were used extensively during peak lactation when female energy expenditure and demand were maximal. Average daily foraging time increased steadily throughout the season to a maximum of 28% of aboveground time as availability of greens diminished and fungus predominated in the diet; time availability did not limit foraging since the animals sat on average for 65% of the daily surface time of about 7 h. Timing of reproduction is apparently optimized such that peak reproductive energy demands are matched with maximal food availability and moderate thermal conditions that minimize energy demand. Despite the greater body mass of males, the greatest total DEE (measured by dlw) of any squirrels at any time of year was that of females during peak lactation. For production of young and lactation through above-ground emergence of an average litter of 2.7, females required a total energy increase of 24% above annual nonreproductive metabolism. Yearling females all bred and performed similarly to older females, yet some costs were greater because the yearlings began and ended hibernation at smaller mass, compensated by giving birth later, and finally showed a greater absolute increase in body mass over the active season than older females. Annual metabolic energy expenditure of breeding males was about 18% greater than that of females, due to greater male body mass. Yet the annual energy intake requirement for both sexes was essentially identical (about 42MJ) due to the greater reproductive export by females in the form of newborn and milk. During the mating season males showed wide-ranging exploratory behavior and social interactions, including aggression, that involved considerable locomotory energy expenditures. Although we did not directly account for the energetics of these specific reproductive behaviors, they are critical to male reproductive success and on a daily basis they probably involved much greater energy expenditure than sperm production. Some yearling males avoided these costs by foregoing testicular development, yet they allocated four times as much energy to growth as older males, thereby increasing somatic condition for the future.     

Mating system and protandrous sex change in the lizard flathead, <Emphasis Type="Italic">Inegocia japonica</Emphasis> (Platycephalidae)

Mating system and protandrous sex change of the lizard flathead Inegocia japonica were investigated off Nagashima, Kagoshima, Japan. The reproductive season of this species was estimated to be from June to September by a fluctuation of gonadsomatic index from monthly sampling data. The hermaphrodite fish, having testicular and ovarian parts, were collected from February to December, indicating that the sex change occurred in both reproductive and nonreproductive seasons. Females were larger than males and hermaphrodites. Moreover, five cases of the sex change from male to female were directly confirmed during the 1995–1997 field census in the behavioral observation area. Home ranges, where no aggressive interaction took place among any individuals, overlapped within and between sexes. Seven pair spawnings observed in the area were not of the same individual combination, and the size of pair fish was likely to be random. This species may thus have a promiscuous mating system, i.e., random pairing, which favors protandry as predicted in the size-advantage model.     

Reproduction of the calanoid copepod Calanus propinquus in the southern Weddell Sea,Antarctica: observations in laboratory

Reproduction of the dominant Antarctic copepod Calanus propinquus was studied in February–April, 1989 aboard the R.V Dmitry Mendeleev during cruise N° 43 to the Weddell Sea. Single females were kept at 0 °C in the laboratory for 56 days with abundant food concentration (above 300 µg C l^–1 of Platymonas viridis). Females released clutches at night at 2–3 day intervals. Most clutches contained from 10 to 40 eggs, mean 37.3 eggs female^–1. Average carbon content of an egg was 0.37 ± 0.05 µg C. The maximum daily egg production rate of 30–50 eggs female^–1 d^–1 was observed for the first 3 days of the laboratory incubation, corresponding to 3.7–6.2% body C. The state of gonadal development of females showed the decline of the reproductive season in late February. The data suggest that egg laying in the region under study starts in December and lasts until March. The state of ovarian maturation, changes in vertical distribution and biochemical body composition of females suggest the possibility of two-year life cycle in C. propinquus in the southern Weddell Sea.     

International study on Artemia LXIII. Field study of the Artemia urmiana (Günther, 1890) population in Lake Urmiah, Iran

Lake Urmiah is a large (total surface 4750–6100 km² in recent times) thalassohaline hypersaline lake (150–180 g l^–1 in the period 1994–1996), located in northwestern Iran. It is the habitat of the endemic Artemia urmiana. Over the period July 1994–January 1996 a sampling campaign was organized: 36 fixed sampling stations, distributed over the entire lake's area, were sampled weekly to determine water temperature, salinity and transparency. At each occasion a filter net was dragged over a distance of 400 m in the superficial water layer to assess the density and composition of the Artemia population. A more limited sampling campaign focused on the annual fluctuations in chlorophyll concentration and on the reproductive behaviour of the brine shrimp population. Several stages of brine shrimp survived during winter months (water temperature 3°C) at low densities. Compared to available data for the Great Salt Lake, USA, Lake Urmiah shows a low algal biomass and overall low Artemia density. The increasing grazing pressure of the developing brine shrimp population in spring seems to prevent the phytoplankton from reaching high blooming concentrations, and oviparity is the dominant reproductive mode throughout the reproductive season.     

Social groupings in 18 species of butterflyfish and pair bond weakening during the nonreproductive season

I surveyed social groupings in 18 butterflyfish species of genus Chaetodon, focusing on how many individuals usually swim together in the reproductive season. Social groupings in 4 of the 18 species were first reported. Six of the 18 species were also observed in the nonreproductive seasons. Three species, of which 2 species were found to be monogamous, spent less time swimming with their partner during the nonreproductive season than the reproductive season. This observation is consistent with the hypothesis that the function of pair swimming is to increase the egg production of the female.     

Reproductive success and nestling diet in the Lesser Spotted Woodpecker ( Picoides minor ): the early bird gets the caterpillar

Populations of Lesser Spotted Woodpecker (Picoides minor) are decreasing in size, necessitating management strategies. However, data on the reproductive biology of this species are scarce. The aim of this study was to investigate the influence of timing of breeding and nestling diet on the reproductive success of the Lesser Spotted Woodpecker and thereby contribute to an understanding of the determinants influencing its reproductive success. During 6 study years between 1996–2003, we investigated various variables of reproductive success in a Lesser Spotted Woodpecker population in the Taunus low mountain range, Germany. We observed nestling feeding at breeding holes to assess the composition of nestling diet and how it changes during the season. Clutch size, number of fledglings as well as body mass of the nestlings declined with the start of egg laying. Pairs composed of individuals that had breed together in previous years started egg laying earlier than newly established pairs. Nestling diet consisted mainly of aphids, caterpillars, craneflies and wood-living larvae. The composition changed considerably within the breeding season in both early and late broods. However, early broods were provided with caterpillars more frequently, whereas late broods received wood-dwelling larvae more often. Our results suggest that Lesser Spotted Woodpeckers time their breeding so as to coincide with the early nestling stage with the highest availability of caterpillars, since this is the time of highest energy demand. Moreover, food availability declines during the breeding season, and the decline in reproductive performance seems to be an effect of this development.     

Reproductive strategies of sub-Antarctic lithodid crabs vary with habitat depth

We present the first fecundity and egg size data for three species of lithodid crab from Antarctic waters south of the Polar Front, caught in the fisheries operating around the island of South Georgia. In all species, fecundity was observed to increase with body size, but reproductive allocation was found to differ significantly between species. The highest relative fecundity (eggs^−g body weight) and smallest egg size was found in Paralomis spinosissima, which is found in shallower waters, whereas the lowest relative fecundity values and largest eggs were recorded in the deeper living species, Neolithodes diomedeae. Evidence is presented that closely related sympatric species may employ quite different reproductive strategies that suit their depth distribution whilst indicating the overriding evolutionary adaptation of reproductive traits to temperature and food availability.     

Morphometric and allometric studies on eggshells and embryos of free-living Canada geese (<Emphasis Type="Italic">Branta c. canadensis</Emphasis> Linnaeus, 1758)

A total of 289 eggs laid by free-living Canada geese (Branta canadensis canadensis Linnaeus, 1758) in the northwestern part of Germany were collected in spring 2002 and 2003 and examined for shell and embryo parameters. Mean egg mass was 160.88±19.68 g. Approximately 90% of all eggs had a mass between 140 and 185 g. About 90% of all clutches consisted of four to seven eggs. The form index of 90% of all eggs was in the range 62–74. The mean eggshell thickness on the blunt pole was 0.52±0.08 and 0.53±0.07 mm on the long axis. Embryo development follows a square function in the first half of development. Mean embryo length was correlated (P<0.01) with the mean body mass and shell thickness on the blunt pole and on the long axis of the eggs. No significant correlations were found between egg mass and form index and egg mass and shell thickness. Eggs with no visible fertilisation had a mean form index of 68.69±4.53 and fertile eggs had a mean form index of 68.50±3.73. Only 3 of 160 fertile eggs contained embryos with malformations of the beak, vertebral column, or leg. The question of regulating prevailing Canada goose populations is discussed.     

Endocrine Correlates of Reproductive Status in Breeding and Nonbreeding Wild Female Moustached Tamarins

In callitrichid primates, reproduction is usually restricted to a single female per group. Reproductive rate is high and the occurrence of a postpartum estrus can lead to simultaneous lactation and pregnancy. In contrast, nonreproductive females often show ovarian inactivity. However, most studies on callitrichid reproductive physiology have been conducted in captivity, where conditions differ considerably from those in the wild, so that reproductive conditions may be strongly modified. Using fecal estrogen and progestogen measurements to monitor female reproductive status in 2 groups of wild moustached tamarins (Saguinus mystax), we examined 1) whether reproductive females in free-ranging groups also show postpartum estrus and 2) whether nonreproductive females demonstrate signs of ovarian activity. In both reproductive females, clear changes in the excretion pattern of progestogen and estrogen metabolites over time in combination with information on parturition dates allowed us to differentiate between pregnancy, a period of postpartum ovarian inactivity lasting for 54 and 64–82 days, and a period of ovarian activity before conception. Nonreproductive females demonstrated temporal fluctuations in hormone concentrations and absolute hormone levels that were similar to ones in the breeding females during the phase of ovarian activity. The results suggest that, in contrast to most captive female tamarins, reproductive females in wild groups of moustached tamarins do not have a postpartum estrus and that nonreproductive females show ovarian activity despite the presence of a breeding female. We therefore conclude that findings from captivity should be only carefully compared to the situation in the wild.     

Demographic correlates of variable egg survival for a tropical lizard

Summary Anolis limifrons is a small iguanid lizard that is found in lowland rainforest in Central America. Important life attributes include early maturity (3–4 mo), multiple clutches of a single egg per breeding season, and low survival. Population turnover is essentially annual as less than 5% adults live more than one year. My objectives were to resolve the following specific questions: 1) Where are eggs laid? 2) Does survival differ among different types of oviposition site? 3) How is the survival of eggs related to the density and age distribution of lizards at different areas? Most eggs are laid beneath litter on the ground. Eggs laid aboveground are placed in accumulations of soil and litter in stumps, crevices in buttressed trees and vines, holes in trees, and on the tops of logs. Eggs were found in aboveground sites in proportion to the abundance of those sites. The major cause of mortality was predation by Solenopsis ants. Judging by the ratio of eggs known to have hatched to those that were predated, survival did not differ among the various types of oviposition site. Study areas with high egg survival in the middle of the wet season had relatively more young (<4–6 mo old) lizards at the end of the wet season than areas with low egg survival. The strong correlation between the relative number of young individuals and egg survival is a consequence of the seasonal cycle of growth and reproduction. Most eggs are laid in the wet season (May–December). Hatchlings complete growth during the dry season (January–April) and become reproductive adults in their second wet season. However, because hatchlings can grow to reproductive size in approximately 3 months, individuals that hatch early in the wet season may themselves produce offspring that will be <4–6 mo old in the late wet season. The relative survival of juveniles (eggs) and adults is an important datum for many theories of life history evolution. For A. limifrons, survival of eggs was two to three times more variable than survival of lizards on both temporal and spatial scales. Comparative data for reptiles are not available to put this information into an evolutionary prespective.     

Factors influencing offspring traits in the oviparous multi-clutched lizard,<Emphasis Type="Italic">Calotes versicolor</Emphasis> (Agamidae)

The determinants of offspring size and number in the tropical oviparous multi-clutched lizard,Calotes versicolor, were examined using both univariate and multivariate (path) analyses. InC. versicolor maternal snout-vent length (SVL) and body condition influence clutch mass and clutch size but have no significant influence on offspring size. The positive effect of maternal SVL and body condition on offspring number is counterbalanced by a negative effect of breeding time on egg mass. In fact, breeding time directly influences the offspring body mass and condition through variation in the egg mass. There is a trade-off between offspring mass and condition with offspring number, and breeding time influences both. Offspring hatched from the eggs of early (May–June) or mid (July–August) breeding periods invariably show lower mass and condition than those hatched from the eggs of late breeding season (September–October). Yet, there is no variation in offspring SVL among early, mid and late clutches. Thus, inC. versicolor offspring SVL is optimized while body mass and condition are not optimized.