Effects of size and nuptial gifts on butterfly reproduction: can females compensate for a smaller size through male-derived nutrients?

In butterflies and other insects, fecundity generally increases with female adult weight. Hence, most butterflies are essentially "capital breeders", because nutrients acquired during the larval stage are stored and subsequently used for egg production during the adult stage. However, in some species, males transfer a large nutritious ejaculate to the female at mating. These females can partly be characterized as "income breeders", and female mass can potentially be decoupled from fecundity to some extent. In the gift-giving green-veined white butterfly Pieris napi, it has been shown that female fecundity and longevity increase with number of matings and also that females mature at smaller size under poor food conditions compared to males. So it has been suggested that females can compensate for their smaller size through nuptial feeding. Here we test this hypothesis in P. napi by assessing female fecundity and longevity in relation to female mass and polyandry. The results showed no support for the hypothesis. Smaller females were not capable of increasing their mating rate to compensate for a low weight at eclosion. Instead, larger females remated sooner. Also, smaller females suffered from both a reduced daily and total fecundity compared to larger females and this decrease in fecundity was independent of female mating status, i.e. females allowed to mate only once and multiply mated females suffered to the same extent from their smaller size.     

Maturation and diel reproductive periodicity of round scad (Carangidae: Decapterus punctatus)

The diel reproductive periodicity of Carangidae is poorly known but appears to be highly variable between species. Some species spawn during the day, others are believed to spawn at night, and it is demonstrated here that round scad, Decapterus punctatus, spawn at dusk. We collected D. punctatus in the eastern Gulf of Mexico during three April cruises (1995, 1996, and 1997). Based on histological criteria, size at 50% maturity was 113 mm fork length (FL) for males and 128 mm FL for females. The gonad-somatic index (GSI) of mature males was significantly different between hours and appeared to show diel periodicity. Diel periodicity was also observed in changes in female GSIs, whole oocyte diameters, and ovarian histology. The average GSI of mature females fluctuated two-fold between day and night, and the size distribution of whole oocytes in some fish was bimodal (at 0.3-0.4 and 0.7-0.8 mm diameter) at dusk rather than unimodal during most of the diel cycle. Histological preparations revealed that these rapid changes in ovarian GSIs and oocyte size distributions were the result of final oocyte maturation. Germinal vesicle migration was observed from 0900 to 1400 hours eastern standard time (EST), germinal vesicle breakdown was evident as early as 1100 hours EST, and ovulation occurred as early as 1800 hours EST. Spawning frequency (approximately every 5 days) was similar whether calculated from the proportion of females with hydrated oocytes during the afternoon or from the proportion of females with postovulatory follicles during the morning. Batch fecundity correlated with fish size and ranged from 5,500 to 34,700 hydrated eggs per individual. These findings do not support published hypotheses that young-of-the-year D. punctatus reproduce before their first winter or that D. punctatus reproductive output is bimodal within a year.     

Male limitation of female reproductive success in a pipefish: effects of body-size differences

Summary In the pipefish Syngnathus typhle, a species with exclusive male parental care, males limit female reproductive success because of their limited brood pouch space and long pregnancy. Sexual size dimorphism is absent in these 1-year-old animals but increases with age so that older females are larger than similarly aged males. Because fecundity is related to size in both sexes and increases more rapidly with body size in females than in males, the difference in growth increases female fecundity more, relative to male fecundity, as the fish get older. We therefore predicted that male limitation of female reproductive success is even more severe when all age classes are considered. To measure a female's maximum reproductive rate, she was provided with three males. Small 1-year-old females produced as many eggs, or produced eggs at the same rate, as a male of similar size could care for. Small females filled on average 1.06 males within the time span of one male pregnancy and actually produced on average 10 eggs fewer than needed to fill a similarly sized male. Large 2-year-old females, in contrast, produced on average a surplus of 149 eggs and filled 2.7 similarly sized males within the course of one pregnancy. The difference between females of the two size classes was highly significant. Males prefer to mate with larger females if given a choice. In nature sex ratios are equal, and males limit female reproductive success in the whole population. Therefore, small females are more severely constrained by mate availability than are larger females because males choose to mate with larger females. Offprint requests to: A Berglund     

Geographical patterns in reproductive biology of the Pan-American sandy beach isopod<Emphasis Type="Italic"> Excirolana braziliensis</Emphasis>

Eleven populations of the Pan-American sandy beach isopod Excirolana braziliensis, distributed from tropical (9°N) to temperate (39°S) sandy beaches in Atlantic and Pacific Oceans, were analyzed to determine latitudinal variations in: breeding and recruitment patterns, sex ratios, size of ovigerous females and of juveniles and length–fecundity relationships. E. braziliensis exhibited strong latitudinal patterns in all reproductive traits throughout its distribution range. Breeding and recruitment shifted from continuous to seasonal from tropical to temperate beaches, having a predominance of females at higher latitudes. In agreement with the latitudinal gradient hypothesis, ovigerous females inhabiting tropical sandy beaches (low latitudes) were smaller, became sexually mature at smaller sizes and had lower individual fecundity than on temperate beaches. Juveniles were also smallest at low latitudes. Between-ocean comparisons showed very similar reproductive characteristics for roughly the same latitude. These linked reproductive parameters suggest that the intensity of breeding effort is associated with the duration of the breeding season and geographically size-related characteristics of the species. Geographic variations in the breeding and recruitment seasons, as well as in individual fecundity, size structure of mature females and sex ratios, are proposed to have major consequences in explaining local variations in population demography. Our paper also reinforces the notion that sandy beach animals are highly plastic.Communicated by O. Kinne, Oldendorf/Luhe     

Size at maturity,fecundity and reproductive potential of a protected population of the spiny lobster<Emphasis Type="Italic"> Palinurus elephas</Emphasis> (Fabricius, 1787) from the western Mediterranean

Protected lobster populations are expected to contribute to the replenishment of fished populations through increased egg production. We studied the reproductive biology and egg production potential of a population of the spiny lobster Palinurus elephas protected from fishing since 1990 in the Columbretes Islands Marine Reserve (western Mediterranean). An index of spawning potential was derived to compare egg production potential in the Reserve and in western Mediterranean exploited populations. Females' physiological maturity (ability to reproduce) and functional maturity (ability to mate and bear eggs) occurred at a carapace length (CL) of 76–77 mm. Males' physiological maturity occurred at a slightly larger size, 82.5 mm CL. In the Reserve, P. elephas' individual fecundity increases linearly with body size up to the females' maximum size, although maximum reproductive yield (eggs per body gram) was reached at intermediate sizes. Size-specific fecundity in the protected population was similar to that of lightly fished populations off Ireland and greater than that of western Mediterranean exploited populations. The female size class of 105–110 mm CL contributed most to egg production in the protected population and is well above the minimum landing size (MLS) for western Mediterranean fisheries. Newly mature females (below MLS) generate a very small fraction (1%) of the egg production from the Reserve. Given the pattern of exploitation in western Mediterranean fisheries, egg production potential depends more on the quantity than on the mean size and fecundity of the available females. The role of the greater availability of large males for mating in unfished populations is discussed in terms of the females' individual fecundity and mating success.     

Geographic variation in maternal investment: acidity affects egg size and fecundity in Rana arvalis

Environmental-stress-mediated geographic variation in reproductive parameters has been little studied in natural vertebrate populations outside the context of climatic variation. Based on life-history theory, an increase in the degree of environmental stress experienced by a population should lead to (1) a shift in reproductive allocation from fecundity to offspring quality, (2) stronger trade-offs between reproductive parameters, and (3) changes in the relationship between female phenotype and maternal investment. To test these predictions, we investigated geographic variation in maternal investment of moor frogs (Rana arvalis) in relation to breeding site acidity (pH 4-8). We found that mean egg size increased and clutch size and total reproductive output (TRO) decreased with increasing acidity among 19 Swedish moor frog populations. Tests for variation and co-variation in maternal investment and female size and age in 233 females from a subset of four acid origin (AO) and four neutral origin (NO) populations revealed that clutch size and TRO increased with female size in both acid and neutral environments. However, in AO populations, egg size also increased with female size, and clutch size and TRO with female age, whereas in NO populations, egg size increased with female age. The strength of the egg-size-clutch-size tradeoff tended to be stronger in AO than in NO females as expected if the former experience stronger environmental constraints. All in all, these results suggest that environmental acidification selects for investment in larger eggs at a cost to fecundity, imposes negative effects on reproductive output, and alters the relationship between female phenotype and maternal investment.     

Reproductive cycle of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama

The reproductive status of the holothuroid species Isostichopus badionotus (Selenka, 1867) and Holothuria mexicana (Ludwig, 1875) was studied over 16 months in Bocas del Toro (Panama), from November 1999 to February 2001. Sexual reproduction was evaluated by the gonad index method, and by histology of gonad development. In addition, population structure was assessed based on sex ratio, minimum reproductive size, and length and weight distributions of males and females. The sex ratio in both species was 1:1, with a unimodal population distribution composed mainly of mature individuals. The minimum reproductive length and weight were 13-20 cm and 150 g, respectively, for both species, although reproductive individuals 10 cm in length were also found. A consistently higher gonad index was observed in H. mexicana, due to a high proportion of mature females and males and high gonad indices in most monthly samples. Gametogenesis and spawning patterns seemed to occur throughout the year, with periods of enhanced activity. Two periods of maximum reproductive activity were tentatively identified: July-November for I. badionotus and February-July for H. mexicana, but neither species had a single, sharply defined annual spawning event. Further work on these exploited holothuroids should examine the relationships between reproduction and environmental factors and between reproductive status and recruitment.     

Egg production in the arrow squid Nototodarus gouldi (Cephalopoda: Ommastrephidae), fast and furious or slow and steady?

The process of reproductive maturation and egg release was examined in the temperate shelf squid Nototodarus gouldi. The energy allocation between somatic and reproductive growth from juvenile to mature adult was investigated throughout the life span to determine the underlying energetic strategy adopted by individuals. The relative weight of the mantle, fin and digestive gland remained unchanged during ovarian development, with no significant correlations found between the mantle length (ML)-gonad residuals and the ML-mantle (r=0.01, P>0.05), ML-fin (r=0.07, P>0.05) and ML-digestive gland (r=0.07, P>0.05) residuals. This suggested that energy was not being diverted away from somatic growth during sexual development, and consequently neither muscle nor digestive gland was being utilised as an energy store. Since squid in all maturity stages contained some food in their stomachs (e.g. 66.7% of mature animals), it is likely that the cost of maturation in this species is largely being met by food intake. The energy investment in reproductive tissues was relatively low (mean gonado-somatic index for mature individuals was 9.29% - 0.40%), indicating that only small amounts of energy were being allocated to reproduction at anyone point in time, which is characteristic of a multiple-spawning strategy. Furthermore, oviduct weight was not correlated with body size (r=0.256, P>0.05), suggesting that eggs are not stored for a single release. In all except one individual, ovary weight was consistently heavier than oviduct weight, suggesting that the ovary is not being depleted of oocytes as mature ova move into the oviducts. Additionally, the ovaries of mature females contained a range of oocyte sizes with discrete peaks, indicating a continued production and development of oocyte cohorts. The presence of some individuals with stretched empty oviducts is further evidence that the reproductive strategy of N. gouldi is slow and steady, with eggs possibly being released in discrete batches over a period of time.     

Reproduction in a simultaneous hermaphroditic shrimp, Lysmata wurdemanni: any two will do?

The caridean shrimp Lysmata wurdemanni (Gibbes) displays protandric simultaneous hermaphroditism with out-crossing, but not all males become simultaneous hermaphrodites (euhermaphrodites). In this laboratory study, we attempted to determine why some shrimp remain males. In our experiment, we grew L. wurdemanni from postlarvae to adults in several group sizes and observed their reproductive function. We found that all shrimp reared in isolation become euhermaphrodites. When cultured in a group, the proportion of shrimp remaining male decreased with increasing group size. Except for those that mated within a day, inter-molt euhermaphrodite-phase shrimp (with or without embryos) and inter-molt male-phase shrimp fertilized eggs successfully. On the other hand, euhermaphrodite shrimp can only mate as females and have their eggs fertilized during a narrow post-molt window (less than 12 h) in each molt cycle (10 days). The fertilization rate of male-euhermaphrodite pairs was similar to that of euhermaphrodite-euhermaphrodite pairs. There are at least two non-exclusive explanations for the persistence of male shrimp in a group. In certain group compositions, an individual may gain more reproductive fitness as a large male with multiple mating partners than as a small female with low clutch size. Alternatively, the presence of male-phase individuals, with variable molt-cycle duration (5-8 days), may be necessary to ensure mating. This study is the first direct experimental demonstration of social control of sex change in the decapod crustaceans.     

Reproduction in a simultaneous hermaphroditic shrimp, Lysmata wurdemanni: any two will do?

The caridean shrimp Lysmata wurdemanni (Gibbes) displays protandric simultaneous hermaphroditism with out-crossing, but not all males become simultaneous hermaphrodites (euhermaphrodites). In this laboratory study, we attempted to determine why some shrimp remain males. In our experiment, we grew L. wurdemanni from post-larvae to adults in several group sizes and observed their reproductive function. We found that all shrimp reared in isolation become euhermaphrodites. When cultured in a group, the proportion of shrimp remaining male decreased with increasing group size. Except for those that mated within a day, inter-molt euhermaphrodite-phase shrimp (with or without embryos) and inter-molt male-phase shrimp fertilized eggs successfully. On the other hand, euhermaphrodite shrimp can only mate as females and have their eggs fertilized during a narrow post-molt window (less than 12 h.) in each molt cycle (10 days). The fertilization rate of male-euhermaphrodite pairs was similar to that of euhermaphrodite-euhermaphrodite pairs. There are at least two non-exclusive explanations for the persistence of male shrimp in a group. In certain group compositions, an individual may gain more reproductive fitness as a large male with multiple mate partners than as a small female with low clutch size. Alternatively, the presence of male-phase individuals, with variable molt-cycle duration (5-8 days), may be necessary to ensure mating. This study is the first direct experimental demonstration of social control of sex change in the decapod crustaceans.     

A histological study of reproduction in the serpulids<Emphasis Type="Italic"> Pomatoceros triqueter</Emphasis> and<Emphasis Type="Italic"> Pomatoceros lamarckii</Emphasis> (Annelida: Polychaeta)

The reproductive cycles of the serpulid polychaetes Pomatoceros lamarckii (Quatrefages, 1865, as Vermilia) and P. triqueter (Linnaeus, 1767, as Serpula triquetra) were studied by histological examination during 1999 and 2000 at Bantry Bay, south-west Ireland. Gametogenesis, sex ratios and hermaphroditism were also investigated. The male/female ratio in P. lamarckii was approximately equal, but in P. triqueter it deviated significantly in favour of females. There was no significant variation in the sex ratios throughout the year in either species. Results confirm that the species are protandric hermaphrodites. In P. lamarckii, but not in P. triqueter, sexual dimorphism was exhibited in body size, the females being larger than the males of the same species. In both species, the proportion of females increased with increasing body size. Simultaneous hermaphroditism was recorded in P. lamarckii, but not in P. triqueter. There was no clearly defined annual gametogenic cycle in either species. Both species appear to have an extended reproductive season, with numerous small-scale peaks in reproductive maturity that can vary annually. Spawning was broadly synchronous between sexes. Visual observation alone (i.e. without histology) was insufficient to accurately assess reproductive condition in individuals not in, or close to, a ripe state.     

Life-history traits of Plesionika martia (Decapoda: Caridea) from the eastern-central Mediterranean Sea

Life-history traits of Plesionika martia (Milne Edwards, 1883) were studied through data collected during six seasonal trawl surveys carried out in the Ionian Sea (eastern-central Mediterranean) between July 1997 and September 1998. P. martia was found at between 304 and 676 m depth, with the highest density in the 400-600 m range. Intraspecific, size-related depth segregation was shown. Recruitment occurred in summer at the shallowest depths. Juveniles moved to the deepest grounds as they grew. The largest female and male were 26 and 25 mm carapace length, respectively. The sex ratio was slightly in favour of females at depths >400 m. Although a seasonal spawning peak was shown, the reproduction appears to be rather prolonged throughout the year. Females with ripe gonads were found from spring to autumn. Ovigerous females with eggs in late maturity stage were found year round. Large females could spawn more than one time within their annual reproductive cycle. The size at first maturity (50% of the ovigerous females) was 15.5 mm CL. Average brood size of eggs with a well-developed embryo was 2,966ǃ,521. Iteroparity, low fecundity and large egg size patterns were observed. Brood size increased according to the carapace length. Two main annual groups were found in the field population of the Ionian Sea. Estimates of the Von Bertalanffy growth parameters are: L_X=30.5 mm, k=0.44 year^-1 in females; L_X=28.0 mm, k=0.50 year^-1 in males. A negative allometry was detected mostly in the ovigerous females. The life cycle of P. martia is discussed in the light of life-history adaptations shown in other deep-water shrimp species.     

Female egg investment in relation to male sexual traits and the potential for transgenerational effects in sexual selection

Life-history theory predicts that individuals should increase their reproductive effort when the fitness return from reproduction is high. Females mated with high-quality males are therefore expected to have higher investment than females mated with low-quality males, which could bias estimates of paternal effects. Investigating the traits females use in their allocation decisions and the aspects of reproduction that are altered is essential for understanding how sexual selection is affected. We studied the potential for differential female allocation in a captive population of a precocial bird, the Chinese quail, Coturnix chinensis. Females paired with males with large sexual ornaments laid larger, but not more, eggs than females paired with males with small sexual ornaments. Furthermore, female egg mass was also significantly positively affected by male testis size, probably via some unknown effect of testis size on male phenotype. Testis size and ornament size were not correlated. Thus, both primary and secondary male sexual traits could be important components of female allocation decisions. Experimental manipulation of hormone levels during embryonic development showed that both male and female traits influencing female egg size were sensitive to early hormone exposure. Differences in prenatal hormone exposure as a result of maternal steroid allocation to eggs may explain some of the variation in reproductive success among individuals, with important implications for non-genetic transgenerational effects in sexual selection.Communicated by C. Brown     

Relationship between body size, provisioning rate, longevity and reproductive success in females of the solitary bee Osmia cornuta

Body size has often been related to reproductive success in bees and wasps. The objective of this 3-year study was to analyze the relationship between nesting female body size, provisioning rate and longevity and their effect on several traits related to parental investment and reproductive success in the solitary bee Osmia cornuta. Body size was not correlated to longevity, and it was only correlated to provisioning rate in the third year (with poor weather conditions during nesting). Variation in fecundity, offspring size and offspring mortality was not well explained by nesting female body size in any of the 3 years. However, in the third year, small females biased their investment toward males, the sex requiring smaller pollen–nectar provisions. Large females were more successful usurpers of other females' nests, but fecundity of usurpers was no higher than fecundity of nonusurpers. Large females were more likely to establish at the release site, probably in relation to size-dependent vigor at emergence. A review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees. In O. cornuta, fecundity was strongly related to longevity and provisioning rate in all 3 years. Offspring size was associated with provisioning rate in 1 year, when females with higher provisioning rates tended to produce larger sons and daughters. Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events.     

Reproduction and condition of female Octopus mimus (Mollusca: Cephalopoda)

The relationship between reproduction and condition was studied in a 15 mo sample of 919 maturing, mature and post-spawning female Octopus mimus (388 to 3714 g) caught in Iquique (North Chile). O. mimus is a semelparous species, with reproduction taking place all year round. Investment in reproductive tissues was, on average, 9.9% of mature female body weight, independent of season. However, somatic growth during maturation was dependent on season and varied between 26 and 63% of the whole-body growth in weight. The condition of females did not vary markedly until spawning, although seasonal variations were apparent, winter being the most unfavourable. Condition deteriorated dramatically after spawning, during parental care of the eggs. During this period, somatic-tissue depletion, mainly from the muscles, was >25% of the total body weight of mature females. The fecundity of O. mimus was probably limited by the costs associated with parental care of the eggs.     

Fecundity and spawning in a deep-water cirromorph octopus

 Reproductive characteristics of cirromorph octopuses, assigned to the species Opisthoteuthis grimaldii, were sampled as a commercial fishing by-catch on the Hebrides Slope, west of Scotland. A total of 254 specimens (99 female, 155 males), retrieved from bottom trawls fished at 750 to 1500 m depth, were examined. A maximum of 2097 eggs was counted in a single female ovary (mean female body weight 1242.8 g), most of them <1 mm in length. At egg lengths over 1 mm, diminishing numbers of eggs were present in 1 mm size categories up to a maximum of ≈10 mm. At body sizes >500 g (wet wt), and in every female >750 g (max. female weight recorded in the sample was 2959 g), a succession of unattached eggs was present in the proximal oviduct and a single, unattached mature egg occupied the tip of the distal oviduct. These females were assumed to be in spawning condition and the characteristics of egg distribution in the reproductive tract to be consistent with sequential release of individual eggs and continuous spawning throughout the growth period and lifespan of the mature octopus. In pre-spawning females there was a positive relationship between estimated egg numbers and maximum egg size. After the onset of spawning there was no significant further increase in estimated potential fecundity over the body-size range 500 to 3000 g. Follicular sheaths remaining in the ovary after release of eggs into the proximal oviduct were counted and used to estimate the total number of eggs released up to the time of capture. Follicular sheaths first appeared at 500 to 650 g body weight and increased steeply in number to >1000 in females >1500 g. Two individuals were found with ovarian follicular sheaths but with no terminal egg in the distal oviduct; these were assumed to have released their egg just before capture. Summation of the number of follicular sheaths counted plus the number of eggs estimated as remaining attached in the ovisac, provided a revised estimate of total potential fecundity and raised the estimate for any individual to a maximum of 3202 eggs (mean = 1396 eggs). Received: 3 February 2000 / Accepted: 17 May 2000     

Male lagoon gobies, <Emphasis Type="Italic">Knipowitschia panizzae</Emphasis>, prefer more ornamented to larger females

Female ornamentation may be directly sexually selected, by male choice or female competition, or occurs as the result of a genetic correlation, arising from sexual selection on males. However, increasing evidence supports the former hypothesis, suggesting that males actively choose their partner preferring traits indicative of female quality. In the lagoon goby, Knipowitschia panizzae, a polygynous species whose males perform parental care to eggs, body length and the size of a sex-specific yellow patch on the belly are known to be reliable indicators of female fecundity. In this paper, we tested, using dummies, the male’s mating preferences for female body and yellow belly patch sizes. The two experimental trials in which a single female trait was variable showed that males prefer a larger belly patch and a larger body size, indicating that both these characters are selected by male mate choice. However, when faced with dummies exhibiting an inverse combination of body and belly patch sizes (experiment 3), males significantly preferred the smaller ones with larger yellow belly patches. A calculation of dummy theoretical fecundity reveals that in the first two experiments, males would have received an immediate benefit from their choice in terms of egg number, whereas in the third one, males chose partners that would have provided them with fewer eggs. The male lagoon goby preference for females with larger belly patches, regardless of their size, suggests that this trait, in addition to indicating fecundity, conveys information about other aspects of female and/or egg quality.     

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.     

Reproductive Investment of a Lacertid Lizard in Fragmented Habitat

Abstract:  We studied the effect of habitat fragmentation on female reproductive investment in a widespread lacertid lizard (  Psammodromus algirus ) in a mixed-forest archipelago of deciduous and evergreen oak woods in northern Spain. We captured gravid females in fragments (≤10 ha) and forests (≥ 200 ha) and brought them to the laboratory, where they laid their eggs. We incubated the eggs and released the first cohort of juveniles into the wild to monitor their survival. Females from fragments produced a smaller clutch mass and laid fewer eggs (relative to mean egg mass) than females of similar body size from forests. Lizards did not trade larger clutches for larger offspring, however, because females from fragments did not lay larger eggs (relative to their number) than females from forests. Among the first cohort of juveniles, larger egg mass and body size increased the probability of recapture the next year. Thus, fragmentation decreased the relative fecundity of lizards without increasing the quality of their offspring. Reduced energy availability, increased predation risk, and demographic stochasticity could decrease the fitness of lizards in fragmented habitats, which could contribute to the regional scarcity of this species in agricultural areas sprinkled with small patches of otherwise suitable forest. Our results show that predictable reduction of reproductive output with decreasing size of habitat patches can be added to the already known processes that cause inverse density dependence at low population numbers.