Hatching and viability of rotifer diapausing eggs collected from pond sediments

1. Planktonic rotifers inhabiting variable environments produce diapausing eggs that accumulate in the sediment of lakes and ponds, forming egg banks that may withstand adverse periods. A common assumption in zooplankton diapausing egg bank studies is to count as viable all eggs in the sediment that look healthy. This assumption should be challenged by asking how effectively ‘healthy‐looking’ eggs represent viable eggs. 2. In this study, viability of more than 1100 ‘healthy‐looking’ diapausing eggs belonging to the Brachionus plicatilis species complex was assessed in a laboratory hatching experiment. Eggs were collected at different depths from sediment cores obtained from 15 ponds located in coastal and inland areas of Eastern Spain. 3. Only approximately one half of the ‘healthy‐looking’ diapausing eggs hatched after incubation in experimental conditions. Almost all the hatchlings (99.4%) survived to maturity. The proportion of ‘healthy‐looking’ diapausing eggs that hatched varied among areas and among ponds within area, and substantially declined with sediment depth. Most of the hatchlings (88%) were obtained from the uppermost 2 cm of sediment. ‘Healthy‐looking’ eggs from upper sediment layers hatched after significantly shorter incubation times than eggs recovered from deeper layers. 4. Both decreased hatching success and increased incubation time for hatching with sediment depth suggest that older ‘healthy‐looking’ eggs are less responsive to hatching stimuli and could become unviable. However, the strong correlation found between the number of ‘healthy‐looking’ eggs and the number of hatchlings indicates that the abundance of ‘healthy‐looking’ eggs is a good index of egg bank viability.     

Fish as vectors in the dispersal of Bythotrephes cederstroemi: diapausing eggs survive passage through the gut

• 1 Bythotrephes cederstroemi (Crustacea: Onychopoda: Cercopagidae) is an introduced invertebrate predator currently spreading through the Laurentian Great Lakes region of North America. We examined a previously unsuspected way in which B. cederstroemi may be dispersed by fish by their consumption of diapausing eggs.
• 2 Ninety‐four percentage of the mature B. cederstroemi diapausing eggs consumed by fish were egested apparently intact. This proportion is considerably above previous estimates for the ephippial eggs of Daphnia. The hatching success of diapausing eggs was compared among four categories: (a) eggs released naturally by B. cederstroemi (control, 73% hatched) (b) eggs released during ‘stressful confinement’ (46% hatched) (c) eggs dissected from dead females (13% hatched) and (d) eggs recovered from faecal pellets following consumption by fish (viable gut passage experiment, 41% hatched).
• 3 Samples of small fish and B. cederstroemi were collected simultaneously. Examination of gut contents revealed that fish contained B. cederstroemi diapausing eggs and that B. cederstroemi bearing resting eggs were consumed selectively over those without eggs. Moreover, fish selected B. cederstroemi bearing mature rather than immature diapausing eggs.
• 4 The fact that diapausing eggs survive gut passage is important for the dispersal of B. cederstroemi. Fish often move between the pelagic and littoral zones of lakes and may thus disperse diapausing eggs widely. Fish may also move between lakes connected by river systems and can be caught and passively dispersed by anglers or piscivorous birds. Our results demonstrate the potential for fish to act as vectors in the spread of B. cederstroemi.

     

Seasonal dynamics of rotifer and crustacean zooplankton populations in a eutrophic,monomictic lake with a note on rotifer sampling techniques

The abundances, biomass, and seasonal succession of rotifer and crustacean zooplankton were examined in a man-made, eutrophic lake, Lake Oglethorpe, over a 13 month period. There was an inverse correlation between the abundance of rotifers and crustaceans. Rotifers were most abundant and dominated (>69%) the rotifer-crustacean biomass during summer months (June–September) while crustacean zooplankton dominated during the remainder of the year (>89%). Peak biomasses of crustaceans were observed in the fall (151 µg dry wt l^–1 in October) and spring (89.66 µg dry wt l^–1 in May). Mean annual biomass levels were 46.99 µg dry wt l^–1 for crustaceans and 19.26 µg dry wt l^–1 for rotifers. Trichocerca rousseleti, Polyarthra sp., Keratella cochlearis and Kellicottia bostoniensis were the most abundant rotifers in the lake. Diaptomus siciloides and Daphnia parvula were the most abundant crustaceans. Lake Oglethorpe is distinct in having an unusually high abundance of rotifers (range 217–7980 l^–1). These high densities can be attributed not only to the eutrophic conditions of the lake but also to the detailed sampling methods employed in this study.The research was supported by National Science Foundation grants DEB 7725354 and DEB 8005582 to Dr. K. G. Porter. It is lake Oglethorpe Limnological Association Contribution No. 25 and Contribution No. 371 of the Harbor Branch Foundation, Inc.     

Induction of diapausing amictic eggs in Synchaeta pectinata

Amictic females of a clone of S. pectinata from Star Lake (Norwich, Vermont) may produce diapausing as well as non-diapausing (subitaneous) eggs. The proportion of diapausing eggs produced in cultures was unaffected by temperature (12 vs 19 °C) or rotifer population density (minima of 0.33 vs 3 ind. ml^–1) at 19 °C. However, at 19 °C this proportion was higher in cultures maintained at a low food level suppressing reproduction (5 × 10³ cells ml^–1 Cryptomonas erosa) than in those maintained at a high food level (2 × 10⁴ cells ml^–1); the treatment effect was marginally significant (p=0.067). Consistent with the effect of low food availability, a period of starvation was very effective in inducing the development of diapausing eggs. None of 19 females cultured individually from hatching at 19 °C on C. erosa (2 × 10⁴ cells ml^–1) in 1-ml volumes produced any diapausing eggs in 4 days (0 out of 349 eggs), while 13 out of 16 females subjected to a 15-hour starvation period 6 hours after birth produced one or more diapausing eggs during that time (34% of the 158 eggs produced by the 16 females were diapausing). Diapausing eggs produced and left at 19 °C hatched after 4 to 13 days. Those produced in cultures with a low food level took significantly longer to hatch (9.7 days) than those produced in cultures with a high food level (8.1 days) (p=0.022). In natural communities, S. pectinata should be able to respond directly and rapidly to poor food conditions by producing eggs that undergo an obligatory dormant period before resuming development.     

Decline of Clear-Water Rotifer Populations in a Reservoir: The Role of Resource Limitation

The relative importance of and changes in resource limitation of herbivorous rotifers were assessed during the clear-water phase in the Rímov Reservoir, Czech Republic, using in situ manipulative experiments. Resource limitation was tested experimentally as the difference in population growth rate (Δr) among various experimental treatments on four occasions. The reservoir community of rotifers was exposed to three treatments: (i) control, (ii) diluted and (iii) diluted and fertilized. Significant responses to these experimental manipulations were shown by Synchaeta spp., Polyarthra spp. and Keratella cochlearis. Growth rate was usually highest during the spring rotifer maximum and decreased during the clear water phase. The highest intensity of food limitation (expressed as ‚Chlorophyll-a’ limitation) was found in Synchaeta spp. K. cochlearis had low food limitation during the spring peak, high food limitation during the second experiment and low food limitation, again, during the later experiment. In contrast, Polyarthra spp. had the same Chlorophyll-a limitation throughout the whole experimental period. Linear regression was used to estimate the relative proportion of Δr variability explained by Chlorophyll-a concentration and rotifer density in all of the experiments. Chlorophyll-a concentration explained 89, 97 and 92% of the resource limitation in Synchaeta spp., Polyarthra spp. and K. cochlearis, respectively. The proportion of variability explained by rotifer density-dependent factors was lower: 60% for Synchaeta spp. and 68 % for Polyarthra spp.     

Females from resting eggs and parthenogenetic eggs in the rotifer Brachionus calyciflorus: lipid droplets, starvation resistance and reproduction

1. In the heterogonic life cycle of monogonont rotifers, amictic (female‐producing) females develop from two types of eggs: fertilised resting (diapausing) eggs and parthenogenetic subitaneous eggs. Females hatched from resting eggs initiate clonal populations by female parthenogenesis and are called stem females. This study compares females from resting and parthenogenetic eggs that were produced under identical culture conditions and were of similar birth order. 2. Newborn stem females had many more lipid droplets in their tissues than similar‐sized, newborn females from parthenogenetic eggs. When neonates were stained with Nile Red and viewed under epifluorescent illumination, these droplets were shown to be sites of neutral‐lipid storage products. 3. Stem females had no posterolateral spines and short anterior spines, while their mothers and offspring in subsequent, parthenogenetic generations typically had long posterolateral spines and elongated anterior spines. 4. Newborn stem females survived starvation significantly longer than newborn females from parthenogenetic eggs. 5. When females from resting and parthenogenetic eggs were cultured from birth to death at a high food concentration, the reproductive potential (r day^?1) of the stem females was significantly higher (0.82–0.88 versus 0.70), primarily because of egg production at an earlier age. The mean lifetime fecundity (R_o) of stem females was significantly greater than that of females from parthenogenetic eggs. 6. Extensive lipid reserves should increase the ability of stem females to colonise new habitats. Firstly, compared with females from parthenogenetic eggs, stem females are more likely to experience starvation or food limitation. Resting eggs hatch in response to physical and chemical factors that are not directly related to food availability, and from sediments that may be far from food‐rich surface waters. Secondly, when food is abundant, stem females have a greater reproductive potential.     

Analysing threshold effects in the sexual dynamics of cyclically parthenogenetic rotifer populations

Sex in some cyclically parthenogenetic rotifers is triggered when a threshold population density is achieved. In Brachionus, accumulation of a mixis-inducing protein (MIP) that the rotifers excrete into the medium is the proximal signal. Models explicitly accounting for MIP dynamics suggest that either (1) a positive feedback of MIP concentration on MIP production or (2) a switching relationship between the proportion of sexual females and MIP concentration is needed to produce a threshold response of mixis to population density. In order to examine both possibilities, experiments were conducted to estimate the sexual response to a dilution series of medium containing the MIP. The resulting sexual response patterns are consistent with the existence of a threshold response of mixis, but are insufficient to reject one hypothesized mechanism for the threshold response in favor of the other. However, the observed smooth increase of sexual offspring with MIP concentration is consistent with the hypothesis of positive feedback on MIP production, and deserves further investigation.     

Differences in predation among morphotypes of the rotifer Asplanchna silvestrii

1. Interactions were observed between three morphotypes of the predatory rotifer Asplanchna silvestrii and six different prey (Brachionus plicatilis, B. rotundiformis, B. pterodinoides, B. satanicus, Hexarthra jenkinae and copepod nauplii) to understand the differences in feeding abilities among morphotypes that may have led to the evolution of this predator polymorphism. The outcome of predation events was affected significantly, both by predator morphotype and prey type. Predator morphotypes also interacted differently with different prey types. 2. The two smaller morphotypes, the saccate and the cruciform, responded similarly to prey overall, except that the smallest morphotype (saccate) was unable to ingest the most mobile prey (nauplii) and less able to ingest relatively large prey (B. plicatilis). The largest morphotype, the campanulate, had the highest encounter rate with prey, but the lowest probability of attack after encounter, so that it consumed far fewer prey per feeding bout than did the smaller morphotypes. This may have been because campanulates prefer larger prey than used in this study. 3. Highly mobile prey (H. jenkinae and copepod nauplii) were much less susceptible to predation than the less mobile Brachionus species. While evasiveness reduced attacks by saccates and cruciforms, campanulates did not have a significantly lower attack rate on H. jenkinae and copepod nauplii than on less evasive prey. Large body size moderately defended B. plicatilis against ingestion by saccates only. The short-spined B. satanicus was the only prey that was rejected after capture, resulting in lower ingestion probabilities for B. satanicus than other brachionid prey.     

Two hundred years of a diverse Daphnia community in Lake Naivasha (Kenya): effects of natural and human-induced environmental changes

1. We used fossil diapausing eggs extracted from ²¹⁰Pb‐dated sediment cores to reconstruct historical changes in the Daphnia community of Lake Naivasha, a climate‐sensitive lake in Kenya which over the past 200 years has experienced a series of well‐documented natural and anthropogenic environmental changes. 2. Contiguous sampling and analysis of four cores yielded ephippial capsules of eight Daphnia species. Only two of these had been recorded previously in live collections from Lake Naivasha, and one species is new to science. The four more common species (Daphnia barbata, D. laevis, D. magna, and D. pulex) show striking differences in abundance patterns and population dynamics through time. Four other species (D. lumholtzi, D. curvirostris, D. longispina s.l., and Daphnia sp. nov. type Limuru.) appear to have been present only occasionally. Nevertheless, between 1895 and 1915 seven species of Daphnia inhabited Lake Naivasha simultaneously. 3. Despite considerable natural environmental change associated with climate‐driven lake‐level fluctuations, the Daphnia community of Lake Naivasha has been severely affected by human activities over the past century, especially the introduction of exotic fishes and water‐quality changes because of agricultural soil erosion. The recent reappearance of large‐bodied Daphnia species (D. magna, D. barbata, D. lumholtzi, Daphnia sp. nov. type Limuru) after 20–110 years of absence can be explained by their release from fish predation, following a dramatic increase in turbidity caused by excess clastic sediment input from eroded catchment soils. The small‐bodied species D. laevis has fared less well recently, presumably because the benefit of lowered predation pressure is counteracted by more pronounced negative effects of increased turbidity on this species and loss of submerged macrophyte beds which formerly served as predation refuge. 4. Our results suggest that, despite considerable environmental instability and the absence of specialised zooplanktivores, top‐down control of fish on large zooplankton is important in Lake Naivasha. Predation pressure from fish has led to clear‐cut shifts in local Daphnia species composition, but failed to drive the larger taxa to extinction.     

Population dynamics of hypolimnetic rotifers in the Pluss-see (North Germany)

The hypolimnetic rotifer populations of the Pluss-see (Keratella hiemalis, Filinia terminalis, Filinia hofmanni) show similar patterns of changing population parameters indicating that they respond to particular environmental signals in the same way.     

Salinity and temperature influence in rotifer life history characteristics

A review of temperature and salinity effects on rotifer population dynamics is presented together with original data of these effects for three clones of Brachionus plicatilis. There is a clear relationship between temperature and the intrinsic rate of increase, r: an increase of temperature — within the natural environmental range — produces an exponential increase of r, and the slope of the response depends on the genotype. The effect of salinity is also genetically dependent; the highest r for each clone is observed at the salinity close to that of its environmental origin. The response of r to temperature is mainly a consequence of the response of the individual rates of development and reproductive timing. The effect of temperature on fecundity (number of descendents per individual life time) is negligible when temperature values are within the normal habitat ranges. On the other hand, salinity seems to affect primarily fecundity. The interaction salinity-temperature may be important in clones or species living in fluctuating environments with positive response to the more frequent combinations found in the corresponding habitats.     

Succession of planktonic rotifer populations in some lakes of the Eastern Rift Valley,Kenya

The quantitative succession of the spring and summer rotifer plankton and its biomass in six lakes of the Eastern Rift Valley of Kenya is discussed. The lakes can be divided into two groups: the low conductivity slightly alkaline lakes Naivasha, Oloidien and the Winam Gulf of Lake Victoria; and the high conductivity, highly alkaline-saline lakes Nakuru, Elmenteita and Bogoria. The former three show a qualitatively rich, typical warmwater rotifer association dominated by Brachionids and Filinia. The saline soda lakes are dominated by several populations of Brachionus dimidiatus, which can reach enormous numbers. The taxonomy and biometry of this species was subjected to statistical analysis. Some interesting and rare species are described: Anuraeopsis coelata, Lepadella triptera f. deconincki, Trichocerca gracilis and T. mus.The chemical limnology of the lakes is discussed and compared with other soda lakes in Central Africa, Europe and North America.     

Patterns and dynamics of rapid local adaptation and sex in varying habitat types in rotifers

Local adaptation is an important principle in a world of environmental change and might be critical for species persistence. We tested the hypothesis that replicated populations can attain rapid local adaptation under two varying laboratory environments. Clonal subpopulations of the cyclically parthenogenetic rotifer Brachionus calyciflorus were allowed to adapt to two varying harsh and a benign environment: a high‐salt, a food‐limited environment and untreated culture medium (no salt addition, high food). In contrast to most previous studies, we re‐adjusted rotifer density to a fixed value (two individuals per ml) every 3–4 days of unrestricted population growth, instead of exchanging a fixed proportion of the culture medium. Thus our dilution regime specifically selected for high population growth during the entire experiment and it allowed us to continuously track changes in fitness (i.e., maximum population growth under the prevailing conditions) in each population. After 56 days (43 asexual and eight sexual generations) of selection, the populations in the harsh environments showed a significant increase in fitness over time relative to the beginning compared to the population in untreated culture medium. Furthermore, the high‐salt population exhibited a significantly elevated ratio of sexual offspring from the start of the experiment, which suggested that this environment either triggered higher rates of sex or that the untreated medium and the food‐limited environment suppressed sex. In a following assay of local adaptation we measured population fitness under “local” versus “foreign” conditions (populations adapted to this environment compared to those of the other environment) for both harsh habitats. We found significantly higher fitness values for the local populations (on average, a 38% higher fitness) compared to the foreign populations. Overall, local adaptation was formed rapidly and it seemed to be more pronounced in the high‐salt treatment.     

Genetic variability and transgenerational regulation of investment in sex in the monogonont rotifer Brachionus plicatilis

In cyclical parthenogens such as aphids, cladocerans and rotifers, the coupling between sexual reproduction and the production of resting stages (diapausing eggs) imposes strong constraints on the timing of sex. Whereas induction of sex is generally triggered by environmental cues, the response to such cues may vary across individuals according to genetic and nongenetic factors. In this study, we explored genetic and epigenetic causes of variation for the propensity for sex using a collection of strains from a Spanish population of monogonont rotifers (Brachionus plicatilis) in which variation for the threshold population density at which sex is induced (mixis threshold) had been documented previously. Our results show significant variation for the mixis threshold among 20 clones maintained under controlled conditions for 15 asexual generations. The effect of the number of clonal generations since hatching of the diapausing egg on the mixis ratio (proportion of sexual offspring produced) was tested on 4 clones with contrasted mixis thresholds. The results show a negative correlation between the mixis threshold and mixis ratio, as well as a significant effect of the number of clonal generations since fertilization, sex being repressed during the first few generations after hatching of the diapausing egg.     

Effect of incubation temperature on the hatching of rotifer resting eggs collected from sediments

Sediment samples from Loch Leven, Scotland, were incubated at 5°C, 10°C and 15°C to induce hatching of rotifer resting eggs. The emergent animals were identified and counted. The temperature which induced hatching varied among the nine species studied. These results are discussed in relation to seasonality and temperature preferences previously recorded for the most abundant species. Resting egg densities of 2.2–13.9 eggs cm^–3 were recorded in the upper 5 cm of sediment.     

Egg diapause and clonal structure in parthenogenetic populations of Heterocypris incongruens (Ostracoda)

A study of the clonal structure of parthenogenetic populations of Heterocypris incongruens from rice-fields in Northern Italy carried out over two-year period is summarized. Significantly different levels of genetic polymorphism were found among populations. The coexistence of different electrophoretic clones and similar patterns of clonal seasonal succession have been observed in at least two different rice fields. The clone or group of clones present in fall, winter and early spring is substituted by other multilocus genotypes in late spring and summer. Different egg diapause induction mechanisms drive the clonal substitution in different clones. Here we report a new laboratory experiment designed to test the effect of temperature and photoperiod found in winter (12 °C 8:16 L:D), spring (24°C 12:12 L:D) and summer (28°C 16:8 L:D) conditions on the deposition and hatching of diapausal eggs in different multilocus genotypes. Clones respond in a way compatible with their sustained presence in the field in different seasons.     

The impact of salinity pulses on the emergence of plant and zooplankton from wetland seed and egg banks

1. In this study we compared the emergence of aquatic biota from sediments under 14‐day pulses of high (5000 mg L⁻¹) and low (1000 mg L⁻¹) salinity with emergence under freshwater and equivalent constant salinity levels. We tested the hypothesis that pulses of high salinity and short duration have no impact on the emergence of aquatic plants and zooplankton from wetland sediment. 2. The way salt is moved through the landscape may alter the response of biota to increases in salinity. Under natural hydrological regimes in rivers and floodplains salinity pulses occur often at concentrations that exceed predicted tolerance levels for aquatic biota. The impacts of natural pulses of high salinity followed by rapid return to fresh conditions may be used to inform management guidelines for the potential release of non‐natural saline water into river systems with minimal impact. 3. For both aquatic plants and zooplankton the abundance and richness of the emerging taxa decreased at higher salinities kept at constant levels. In contrast, pulses of salinity followed by return to freshwater conditions did not have a negative impact on the emergence of aquatic plants or zooplankton. For many taxa of zooplankton a positive impact was demonstrated with higher emergence following the salinity pulse. 4. The responses of aquatic plant and zooplankton taxa are grouped into five response types. Type 1: negatively impacted by all salt regimes. Type 2: preference for constant salinities. Type 3: no difference between fresh and either pulse regime. Type 4: preference for high concentration pulses. Type 5: emergence higher under a low concentration pulse. 5. Although previous studies indicate that constant high‐level salinity in rivers and wetlands can decrease the species richness of aquatic communities, this current study shows pulses may not have the same impact. Our results support the hypothesis that pulses of high salinity and short duration do not impact on the emergence of aquatic plants and zooplankton from wetland sediments. For zooplankton, pulses of salt may trigger emergence. 6. These trends may be used to explore the potential to use managed water releases to move salt through the landscape with minimal impact of salinity on aquatic biota. However, before such preliminary results are applied in management of saline water releases we need to determine the implications for interacting processes in natural ecosystems.     

An individual-based model for predicting the emergence period of sea trout fry in a Lake District stream

The objective of this study was to predict interannual fluctuations in the emergence period of sea trout fry, using models developed from field data for 70 excavated redds, and laboratory data on egg and alevin development at 30 constant temperatures (range 1·5–10·5° C with 100 naturally fertilized eggs at each temperature). Egg weight and numbers per redd both increased with female length; a power function described the relationship. Early spawners were the largest females laying the largest and most numerous eggs, whilst late spawners were the smallest females laying the smallest and least numerous eggs; middle spawners being intermediate between these two extremes. Mean values for egg weight and numbers of eggs per redd were obtained for these three groups. Hatching and emergence times in the laboratory decreased with increasing temperature. Of five models tested for hatching time, the best fit was provided by a three-parameter hyperbolic model which formed the asis of the individual-based model used to predict egg hatching and fry emergence. Model development was described in detail and the final equations predicted the times taken for 5, 50 and 95% of the fry to emerge, and hence the period over which 90% of the fry emerged. Analogous models were obtained for egg hatching. All models were excellent fits to the laboratory data. Hatching times for eggs kept in perforated boxes in the stream were almost identical to those kept at similar mean temperatures in the laboratory. Model predictions of fry emergence times were validated by field data for 8 years (1967–1971, 1974, 1975, 1980). The chief objective was therefore fulfilled, and predictions for the 30-year study (1967–1996) revealed a large variation in the timing of emergence (extremes: 11 March–4 April 1989, 15–20 May 1979). Most of the variation in median emergence date was due to variations in water temperature, with spawning dates as a secondary factor; the latter, however, had a greater effect on the length of the emergence period.     

The significance of mating processes for the genetics and for the formation of resting eggs in monogonont rotifers

Crossbreeding experiments with three geographically distinct strains (E, S, and L) of the rotifer Brachionus plicatilis have been carried out in an attempt to elucidate the apparent male sterility of strain L, which is unable to produce resting eggs.The 9 crossing possibilities of the three strains have been investigated in 27 experiments. The results refute the concept of male sterility. L-males copulate successfully with mictic as well as with amictic females of strains E and S. Fertilized amictic E and S females produce defective resting eggs, which have only one thin shell and, which disintegrate after deposition. L-females cannot, as a rule, be fertilized. Moreover, crosses between strain E and S are succesful only in one direction; the reciprocal crosses failed.To explain the present results a hypothesis is suggested that the thickness of the body wall of newborn females differs in the three strains, and between mictic and amictic individuals. A sequence of gradually increasing body wall thickness of all types of females involved, together with a comparably increasing penetration ability of the males of strain E, S, and L explains the success or failure of all crosses, including the unilateral cross E × S.     

Age‐specific recruitment and natality of California sea lions at San Miguel Island,California

We conducted a 15 yr mark‐resight study of branded California sea lions (Zalophus californianus) at San Miguel Island, California, to estimate age‐specific recruitment and natality of the population. We used the Schwarz and Stobo model to estimate sighting, survival, recruitment, timing of births, abundance, and age‐specific natality from sighting histories of 1,276 parous females. The advantage of this approach was that the reproductive status of females did not have to be known for all females of reproductive age. Probability of recruitment into the reproductive population began at age 3 or 4, peaked between ages 5 and 7, and slowly declined. Age‐specific natality was similar for ages 4–16 but declined after age 17, suggesting that reproductive senescence occurs in older females. The average annual natality for parous females 4–16 yr of age was 0.77 (SE = 0.03); natality declined to 0.56 (SE = 0.10) for parous females 17–21 yr of age. Natality for both age classes was reduced during El Niño conditions by 24% and 34%, respectively. In addition to reducing natality, El Niño events may result in a delay of recruitment if females experience El Niño conditions before they turn 4 yr of age.