Dispersal and development of bagworm larvae (Lepidoptera: Psychidae) on three host plants

The present study investigated the development and dispersal of bagworm larvae, Thyridopteryx ephemeraeformis (Haworth) (Lepidoptera: Psychidae), on three hosts: arborvitae, Thuja occidentalis, blue spruce, Picea pungens, and juniper, Juniperus virginiana. The biomass of larvae was recorded at repeated intervals at 21 field sites. The relative quality of plants varied during larval development: early in the season, larvae were heavier on juniper and arborvitae than on blue spruce; late in the season, in contrast, larvae and pupae were lighter on juniper than on the two other plants. Larvae on blue spruce pupated later than those on arborvitae. In the laboratory, feeding rate and larval growth were not influenced by the density of larvae on the foliage of arborvitae up to a range which greatly exceeded that observed at any of our experimental sites. The incidence of dispersal among neonates emerging on arborvitae was high, as indicated by the high number of larvae captured on sticky traps, and the relatively low number of early instars on arborvitae foliage in relation to the reproductive output of females in the parental generation. The high rate of dispersal among early instars may be due to potential resource depletion over successive generations of bagworms.     

Transgenerational marking of marine fish larvae: stable-isotope retention, physiological effects and health issues

This study examined the toxicological and physiological responses of a commercially important coral-reef grouper, Plectropomus leopardus (Serranidae), to injection of enriched stable-isotope barium chloride (BaCl₂) solution. Thirty adult P. leopardus were subject to one of two ¹³⁸BaCl₂ injection treatment groups (corresponding to dosage rates of 2 and 4 mg ¹³⁸Ba kg⁻¹ body mass), and a control group in which fish were injected with 0·9% sodium chloride (NaCl) solution. Fish from each group were sampled at post-injection intervals of 48 h and 1, 3, 5 and 8 weeks, at which time blood and tissue samples were removed from each fish. Residual concentrations of Ba and ¹³⁸Ba:¹³⁷Ba ratios were measured in muscle, gonad, liver and bone tissues of each experimental fish. Elevated Ba concentrations were detected in all treatment fish tissue samples within 48 h post injection. Residual Ba concentrations decreased throughout the remainder of the 8 week experimental period in all tissues except bone. The BaCl₂ injection had no significant effects on measured whole blood variables or on the plasma concentrations of steroid hormones. Enriched Ba stable isotopes can therefore be used at low dosages to mark larvae of commercially important marine fishes, without adverse effects on the health of the fishes or on humans who may consume them.     

Pheromone release by the two forms of Callosobruchus maculatus females: effects of age, temperature and host plant

Abstract. The influence of age, temperature and host plant on pheromone release was investigated in females of two forms of Callosobruchus maculatus (Coleoptera: Bruchidae), referred to as the flightless form and the flight form. The emission of pheromone seemed to be correlated with reproductive status: the flightless-form females which were sexually mature at emergence began to release pheromone on the first day following emergence. The flight-form females which exhibited no reproductive activity at emergence also showed a delayed emission of pheromone. The temperature conditions and the availability of host plant seeds had no apparent effect on pheromone release by the flightless form. In flight-form females pheromone release was stimulated by the presence of host plant seeds. Temperature seemed to affect the age at which females initiated pheromone release for the first time after emergence, pheromonal emission being brought forward slightly by higher temperature. All these results are discussed relative to the specific environment of each form.     

寄主种类、距离和种群密度对烟粉虱扩散的影响

在笼罩条件下,分别以黄瓜、棉花、蕹菜为虫源寄主和诱集寄主,研究寄主植物、空间距离、虫源密度等因子对B型烟粉虱扩散的影响。结果发现,从虫源寄主上扩散的成虫量,与烟粉虱对诱集寄主和虫源寄主的嗜性差异程度有关,诱集寄主相对于虫源寄主的嗜性越强,扩散出来的成虫数量就越多,反之扩散的成虫数量就越少。成虫扩散量与诱集寄主的距离呈负相关,并且诱集寄主的嗜性越强,随着距离的增大,诱集量下降的速率越快。成虫扩散量与虫源种群密度呈正关,但非嗜好寄主上的虫口承载量相对较低,在虫口密度相对较高时,非嗜好寄主上烟粉虱向外扩散的相对量更大。烟粉虱的扩散量与寄主营养状况呈负相关,寄主营养胁迫时,非嗜好寄主上可以获得相对更高的诱集量。     

Oviposition preference of Lycaena salustiusfor,and larval performance on,a novel host plant: an example of ecological fitting

1. ‘Ecological fitting’ is the process whereby the suites of traits an organism carries from previous evolutionary relationships are used to enable colonisation of novel environments or resources. 2. The concept has much explanatory power in studies of novel host associations, particularly when data suggest a deviation from optimal foraging theory, but is often overlooked in studies of herbivore host selection behaviour in favour of evolutionary hypotheses. 3. In the present study, the concept was used to explain the unusual host selection behaviour of the New Zealand endemic oligophagous butterfly Lycaena salustius Fabricius, the larvae of which feed on endemic Polygonaceae species and the introduced and closely related Fagopyrum esculentum Moench. 4. In field cage oviposition choice assays involving only endemic plants, females preferred to oviposit on the rare Muehlenbeckia astonii Petrie. However, the novel host F. esculentum was overwhelmingly preferred in an additional greenhouse assay. In larval no‐choice performance assays, fitness indicators were variable for the novel host. 5. This imperfect relationship between oviposition preference and larval performance is discussed as a possible example of ecological fitting and highlights the potential use of the concept as an explanatory tool in novel host selection behaviour studies.     

幼虫密度对甜菜夜蛾生长发育与繁殖的影响

为了研究甜菜夜蛾Spodoptera exigua(Hübner)幼虫的密度对其发育及繁殖的影响,本实验观察了5种幼虫密度下（1,5,10,20,30头/瓶）,幼虫发育和成虫繁殖情况。结果表明：幼虫和蛹历期、存活率和蛹重均差异显著。幼虫和蛹历期均以20头/瓶的最短,1头/瓶的最长,其余随幼虫密度增加而延长;幼虫至蛹存活率以10头/瓶的最高,其余随幼虫密度增加而降低;1头/瓶的蛹最重,显著高于其他密度的,其余随幼虫密度增加而下降。尽管密度间成虫羽化率和产卵前期均无显著差异,但成虫产卵量、寿命和畸形率差异显著。1头/瓶的产卵量最多,其次为10头/瓶的,其余随幼虫密度增加而减少,30头/瓶的产卵量显著少于其他密度的; 密度在1～20头/瓶范围内,雌蛾寿命均较短,显著短于30头/瓶的,而雄蛾寿命以5头/瓶的最短,显著短于其他密度的（10头/瓶除外）,10头/瓶的次之,其余密度间差异不显著;不同幼虫密度下羽化的成虫畸形率差异显著,10头/瓶的最低,其余随幼虫密度增加而升高; 生命表结果表明甜菜夜蛾在10头/瓶下世代存活率和种群增长指数均最高,幼虫密度过低或过高均不利于种群增长;世代存活率（S）和种群增长指数（I）与幼虫密度之间的关系均呈抛物线关系：S =－0.2087x²+2.5694x+211.52 (R²=0.88),I=－0.0552x²+0.9166x+54.168 (R²=0.95)。结果提示幼虫密度影响甜菜夜蛾种群动态的重要生态因子之一。     

Behavior of newly hatched potato tuber moth larvae,Phthorimaea operculella Zell. (Lepidoptera: Gelechiidae), in relation to their host plants

A study was made of neonate larvae of Phthorimaea operculella.Host finding from soil-laid eggs and dispersal from hosts and nonhosts were first examined. Of first-instar larvae hatching from soil-laid eggs, 80% found the potato plant while roughly 50% found each of the other three plants (datura, tobacco, and tomato). Dispersal from potato, datura, and tobacco was very low, while on tomato it was higher and a high mortality was observed in the 24 h period. Dispersal from nonhost plants was high. Behavior on leaves of hosts and non-hosts is described and some host-plant specific behaviors are identified. Effects on behavior of some of the physical and chemical factors are described. Methylene dichloride extracts of potato leaf wax reduced locomotion rates and the number of turns during locomotion. Also methylene dichloride extract and, to a lesser extent, methanol extract caused biting behavior.     

Dispersal and population structure of the rufous bettong,Aepyprymnus rufescens (Marsupialia: Potoroidae)

Abstract Many species of herbivorous mammals declined to extinction following European settlement of inland Australia. The rufous bettong, Aepyprymnus rufescens (a macropodoid marsupial), is ecologically similar to many of these species. We used analysis of microsatellite markers to determine dispersal patterns and mating system characteristics in a cluster of local populations of A. rufescens, with the aim of gaining a better understanding of regional population dynamics in such species. Particularly, we asked whether the rufous bettong showed source‐sink dynamics, as Morton (1990) hypothesized that many mammals may have been made vulnerable to extinction through such processes. We compared populations separated by distances of up to 12 km, and detected significant genetic differentiation among local populations (F_ST = 0.016). Females displayed greater genetic structuring than males, suggesting that females dispersed over shorter distances or less frequently than males. Geographic distance was weakly related to genetic distance between populations suggesting some gene flow at this scale, and paternity assignment indicated that dispersal can occur over distances of up to 6.5 km. Our study populations varied widely in density, but density did not explain the pattern of genetic differentiation observed. These findings of significant structure among populations, some influence of distance on genetic divergence and that density explains little of the divergence among populations, suggested that source‐sink dynamics did not play a large role among these populations. Variance in male mating success was low (maximum assigned paternity for an individual male was 14% of offspring). While data on multiple maternity were limited, roughly half of repeat maternity was assigned to the same male, suggesting that the mating system of the rufous bettong is not purely promiscuous.     

The effects of host plant species and larval density on immune function in the polyphagous moth Spodoptera littoralis

Immune functions are costly, and immune investment is usually dependent on the individual''s condition and resource availability. For phytophagous insects, host plant quality has large effects on performance, for example growth and survival, and may also affect their immune function. Polyphagous insects often experience a large variation in quality among different host plant species, and their immune investment may thus vary depending on which host plant species they develop on. Larvae of the polyphagous moth Spodoptera littoralis have previously been found to exhibit density‐dependent prophylaxis as they invest more in certain immune responses in high population densities. In addition, the immune response of S. littoralis has been shown to depend on nutrient quality in experiments with artificial diet. Here, I studied the effects of natural host plant diet and larval density on a number of immune responses to understand how host plant species affects immune investment in generalist insects, and whether the density‐dependent prophylaxis could be mediated by host plant species. While host plant species in general did not mediate the density‐dependent immune expression, particular host plant species was found to increase larval investment in certain functions of the immune system. Interestingly, these results indicate that different host plants may provide a polyphagous species with protection against different kinds of antagonisms. This insight may contribute to our understanding of the relationship between preference and performance in generalists, as well as having applied consequences for sustainable pest management.     

Variation in dispersal mortality and dispersal propensity among individuals: the effects of age, sex and resource availability

1.  Dispersal of individuals between habitat patches depends on both the propensity to emigrate from a patch and the ability to survive inter-patch movement. Environmental factors and individual characteristics have been shown to influence dispersal rates but separating the effects of emigration and dispersal mortality on dispersal can often be difficult. In this study, we use a soil mite laboratory system to investigate factors affecting emigration and dispersal mortality.
2.  We tested the movement of different age groups in two-patch systems with different inter-patch distances. Differences in immigration among age groups were primarily driven by differences in emigration but dispersal mortality was greater for some groups. Immigration declined with increasing inter-patch distance, which was due to increasing dispersal mortality and decreasing emigration.
3.  In a second experiment, we compared the dispersal of recently matured males and females and tested the impact of food availability during the developmental period on their dispersal. Dispersal was found to be male biased but there was no significant sex bias in dispersal mortality. There was some evidence that food availability could affect emigration and dispersal mortality.
4.  These results demonstrate that both emigration and dispersal mortality can be affected by factors such as individual age and resource availability. Understanding these effects is likely to be important for predicting the fitness costs and population consequences of dispersal.     

The integrative effects of population density, photoperiod, temperature, and host plant on the induction of alate aphids in Schizaphis graminum

The wheat aphid Schizaphis graminum (Rondani) displays wing dimorphism with both winged and wingless adult morphs. The winged morph is an adaptive microevolutionary response to undesirable environmental conditions, including undesirable population density, photoperiod, temperature, and host plant. Here we studied the integrative effects of population density, photoperiod, temperature, and host plant on the induction of alate aphids in S. graminum. The present results show that these four factors all play roles in inducing alate aphids in S. graminum but population density is the most important under almost all circumstances. In importance, population density is followed by photoperiod, host plant, and temperature, in that order. These results indicate that ambient environmental factors are highly important to stimulation of alate aphids in S. graminum, especially when population density reaches 64 individuals per leaf.     

Parasite species richness in carnivores: effects of host body mass, latitude, geographical range and population density

Aim  Comparative studies have revealed strong links between ecological factors and the number of parasite species harboured by different hosts, but studies of different taxonomic host groups have produced inconsistent results. As a step towards understanding the general patterns of parasite species richness, we present results from a new comprehensive data base of over 7000 host–parasite combinations representing 146 species of carnivores (Mammalia: Carnivora) and 980 species of parasites.
Methods  We used both phylogenetic and non-phylogenetic comparative methods while controlling for unequal sampling effort within a multivariate framework to ascertain the main determinants of parasite species richness in carnivores.
Results  We found that body mass, population density, geographical range size and distance from the equator are correlated with overall parasite species richness in fissiped carnivores. When parasites are classified by transmission mode, body mass and home range area are the main determinants of the richness of parasites spread by close contact between hosts, and population density, geographical range size and distance from the equator account for the diversity of parasites that are not dependent on close contact. For generalist parasites, population density, geographical range size and latitude are the primary predictors of parasite species richness. We found no significant ecological correlates for the richness of specialist or vector-borne parasites.
Main conclusions  Although we found that parasite species richness increases instead of decreases with distance from the equator, other comparative patterns in carnivores support previous findings in primates, suggesting that similar ecological factors operate in both these independent evolutionary lineages.     

Tethered flight activity of pear psylla, Cacopsylla pyricola: Seasonal, host, and morphotypic effects

Pear psylla, Cacopsylla pyricola Förster (Homoptera: Psyllidae), was flown in the laboratory to test hypotheses suggested by field observations. Flight durations of the same insects flown on two different days were highly correlated, suggesting that there were biolgical differences among insects in flight tendencies. Flight durations were similar between summerform and winterform morphotypes. Flight frequencies and durations increased between September and November collections of winterforms, results that are consistent with field observations. Long-duration (60+ min) flights were three to four times as frequent in winterforms restricted to senescing pear foliage compared to insects provided access to pear seedlings. Winterforms collected during the fall flight period from a pear orchard and an adjacent apple orchard (an overwintering habitat) showed similar flight durations. There was no correlation between flight durations and any of five body and wing measurements; body size of winterforms increased between September and November. It has been proposed elsewhere that much of the fall movement into adjacent non-pear habitats by winterforms is due to short duration non-migratory flights rather than to a true migratory flight. The lack of differences between apple- and pear-collected winterforms may be consistent with this hypothesis.     

幼虫密度对草地螟生长发育及繁殖的影响

为了明确幼虫密度对草地螟Loxostege sticticalis种群增长的影响, 对室内条件下（温度22±1℃, RH 70%±5%, 光周期16L∶ 8D）不同幼虫密度[1, 10, 20, 30和40头/瓶（650 mL）］饲养的草地螟生长发育及繁殖进行了研究。结果表明: 幼虫密度对草地螟幼虫体色、 发育历期和存活率, 以及蛹重和成虫生殖等有显著影响。随着幼虫密度的增加, 幼虫体色黑化程度呈增强趋势, 幼虫密度大于10头/瓶时的体色黑化值均显著大于幼虫密度为1头/瓶时的体色黑化值（P<0.05）。20头/瓶的幼虫和蛹历期最短, 且随幼虫密度的增加而显著延长（P<0.05）。幼虫存活率以10头/瓶最高, 其他幼虫密度的幼虫存活率显著较低（P<0.05）。蛹重以1头/瓶的最重, 并随幼虫密度增加而显著下降（P<0.05）。成虫产卵量和交配率分别以1和20头/瓶的幼虫密度最高, 幼虫密度升高则产卵量、 交配率逐渐降低。成虫产卵历期随着幼虫密度的增加逐渐缩短。雌、 雄蛾寿命分别以10和20头/瓶幼虫密度时最长, 幼虫密度过高时雌雄蛾寿命又显著缩短（P<0.05）。生命表分析表明, 幼虫密度对草地螟种群增长指数有显著影响, 以10头/瓶幼虫密度的种群增长指数最高, 幼虫密度过高或过低时种群增长指数下降。据此认为, 幼虫密度是影响草地螟种群增长的重要因子之一。     

Differential utilization of ash phloem by emerald ash borer larvae: ash species and larval stage effects

• 1 Two experiments were performed to determine the extent to which ash species (black, green and white) and larval developmental stage (second, third and fourth instar) affect the efficiency of phloem amino acid utilization by emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae.
• 2 EAB larvae generally utilized green ash amino acids more efficiently than those of the other two species. For example, the concentrations of only six (two essential) and seven (two essential) amino acids were lower in frass from EAB that fed upon black and white ash than in the corresponding phloem, respectively. By contrast, concentrations of 16 (eight essential) amino acids were lower in the frass from EAB that fed upon green ash than in the phloem. In addition, in green ash, the frass : phloem ratios of 13 amino acids were lower than their counterparts in black and white ash.
• 3 The concentrations of non‐essential amino acids glycine and hydroxylproline were greater in frass than in phloem when EAB fed on black ash, although not when EAB fed on green or white ash.
• 4 The concentration of total phenolics (a group of putative defensive compounds to EAB, expressed as antioxidant activity of acetone extraction) was high in EAB frass but even higher in the phloem samples when the data were pooled across ash species and EAB larval stages. This suggests EAB larvae may eliminate phenolics through a combination of direct excretion and enzymatic conversion of phenolics to nonphenolics before excretion. Because the ratio of frass total phenolics to phloem total phenolics in white ash was lower than the ratios in black and green ash, the ability to destroy phenolics or convert them to nonphenolics was greater when EAB larvae fed on white ash.
• 5 Fourth‐instar EAB extracted phloem amino acids, including threonine, more efficiently than third‐instar EAB. The different larval developmental stages of EAB did not differ in their apparent ability to destroy phenolics or convert them to nonphenolics.

     

Linking hydrological,geomorphological and biological properties in hyper‐arid conditions: Distribution processes of a dominant clonal cactus by run‐off

The objective was to analyse the asexual spread of Tephrocactus aoracanthus (Cactaceae) related to run‐off triggered by torrential summer rains in a hyper‐arid desert. Through a manipulative essay, 30 joints (asexual dispersing stems) were placed in the mainstream of an alluvial fan and 30 joints out of the alluvial fan, without run‐off influence (control). All joints were labelled, and their weight and diameter were measured. After each rain event with run‐off, joints were surveyed identifying their spatial coordinates and environmental context. For this, two factors with two levels each one was selected: position respect to both microrelief (channels with concentrated run‐off, or inter‐channels as flat areas between channels, without run‐off) and vegetation (entangled or not entangled in shrubs). Through a geographic information system, changes in slope angle and rugosity of terrain were analysed. Linear regressions were performed between the partial displacement of joints and their weight and diameter, grouped according to environmental context before each rainfall. None of the joints of the control treatment were moved, but all joints located in the mainstream were relocated. Three rainfall events triggered a mean displacement of 480.4 ± 210.0 m (SD); 103.0 ± 138.9 m (SD) and 33.2 ± 52.3 m (SD), respectively. The mean accumulated displacement of all joints during the experience was 577.12 ± 220.0 m (SD). As they moved, the joints passed through several different environmental contexts, increasing the proportion of joints entangled in vegetation and in inter‐channel position. Until rooted, joints could be displaced by run‐off in an anisotropic process, with an evident effect of slope angle and rugosity of terrain. On the other hand, no direct relationship was found between the displacement and the magnitude of the event, or both the size and the weight of joints. We assess the strategies related to the dispersal mechanism of clonal plants, related to conditions and enabling its agamic dispersal over long distances.     

The effects of host plant phenology on the demography and population dynamics of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae)

Abstract.

• 1 We examined the effects of variation in the timing of spring leaf production and autumn leaf fall on the survival, mortality and abundance of Cameraria hamadryadella on Quercus alba and Q.macrocarpa.
• 2 We monitored and manipulated the timing of foliation on field and potted Q.alba trees and observed the abundance of C.hamadryadella on those trees. We also monitored and manipulated the timing of leaf fall on Q.alba and Q.macrocarpa trees in the field and observed its effects on survival, mortality and abundance of C.hamadryadella.
• 3 Variation in the timing of spring leaf production has no effect on C. hamudryadella abundance. However, a warm winter and spring in 1991 led to accelerated development and the imposition of a facultative third generation in one out of ten years of observation.
• 4 In 1989, leaves fell relatively early and leaf fall in the autumn accounted for more than 50% of the mortality of C.hamudryadella. in 1990 and 1991 leaves fell relatively late and leaf fall induced mortality was substantially reduced and overwinter survival was markedly increased.
• 5 The abundance of C.hamadryadella remained constant in the spring and summer of 1990 following the previous autumn's relatively early leaf fall, but increased by 10-fold in the spring of 1991 following the relatively late leaf fall of autumn 1990. The abundance of C.hamadryadella also increased 4-fold between the summer of 1991 and the spring of 1992 after another autumn of relatively late leaf fall. We attribute these increases in abundance in part to reduced mortality because of later leaf fall.
• 6 Variation in the timing of autumn leaf fall may be responsible for initiating outbreaks of C.hamadryadella.

     

First record of the host plant and larvae of Pachyrhynchus sonani (Coleoptera: Curculionidae) on Lanyu Island,Taiwan

Pachyrhynchus sonani Kôno, 1930, a legally protected species in Taiwan, is endemic to Lanyu Island (Orchid Island) and Ludao Island (Green Island). Because of its unclear life history, the host and immature stages of this species in the wild have not so far been recorded. On Lanyu Island, we recently identified Barringtonia asiatica as a host plant of P. sonani, with several larvae feeding on its xylem. Herein, we describe its biology and the habitat surrounding the host plant.     

Avoidance of natural enemies by adult whiteflies, Bemisia argentifolii, and effects on host plant choice

In this study we asked whether, in the context of a trap crop system, differential predation risks among plants influence host choice patterns of adult whiteflies, Bemisia argentifolii. We investigated whether adult whiteflies avoid natural enemies inhabiting poinsettia (a cash crop) and whether this behavior can be used to increase the movement of whiteflies to cucumber (a trap crop). The potential of cucumber as a trap crop was first evaluated and we found that significantly more whiteflies were attracted to cucumber when the whiteflies were released between the two plants. However, the accumulation of whiteflies on cucumber substantially diminished if the insects had first settled on poinsettia. Under such circumstances, we investigated whether movement of adult whiteflies to cucumber could be increased by creating conditions that would cause the whiteflies that had settled on poinsettia to leave the plant. A mechanical disturbance, consisting of shaking the plant, was first used to test this hypothesis. The shaking caused more whiteflies to leave poinsettia and move onto the trap crop, compared to undisturbed whiteflies. We then asked whether the presence of natural enemies on the cash crop could induce whiteflies to leave the cash crop and move onto the trap crop. Three natural enemies were tested: two predators, Amblyseius swirskii and Delphastus catalinae, and a parasitoid Encarsia formosa. The presence of D. catalinae on poinsettia induced significantly more whiteflies to disperse to cucumber compared to poinsettia with no natural enemies, whereas A. swirskii and E. formosa did not result in a significant increase. Predator avoidance behavior by adult whiteflies should be investigated further in the context of trap cropping and other crop-habitat alterations designed to help manage whitefly abundance.     

Fecundity and its variability in orange roughy: effects of population density, condition, egg size, and senescence

Fecundity of orange roughy in 1987–1989 adjusted for standard length (S.L.) varied significantly between New South Wales (42 787 eggs female⁻¹) South Australia (35 339 eggs female⁻¹) and east Tasmania (31 085 eggs female⁻¹). Only 10-17% of the variability in fecundity of eastern Tasmania orange roughy was explained by s.l . in any year from 1987–1992. However, liver condition and age of the fish, in combination with s.l ., explained 27% of the variation in fecundity. Fecundity declined in fish over 60 years old. It was also significantly correlated with lipid levels in the ovary, in particular, with triacylglycerol as a proportion of the total lipid fraction. Significant interannual changes in fecundity appeared to be related to the impact of fishing. From 1987–1992, the orange roughy stock off east Tasmania was reduced by 50% by the fishery, and mean fecundity increased 20% over that period. This compensatory increase in individual fecundity, combined with an apparent increase in the proportion of females spawning annually from 54 to 71%, limited the decline in the population's egg production over this period to approximately 15%.