Interference of asexual and sexual reproduction in the green hydra

The green hydra, Hydra viridissima, has three sexes: hermaphrodite, male, and female. I investigated the reproductive strategies of the green hydra and the relationship between asexual budding and sexual reproduction. The proportion of mature individuals in the asexually reproducing population increased with increasing temperature. Sexual reproduction did not interrupt asexual budding in hermaphrodites or males; sexual reproduction did interrupt asexual budding in females. Sexual reproduction also resulted in exponential population growth during the reproductive season. The number of asexual buds on each parental individual was positively correlated with the parental individual size in asexual individuals and in males. The same positive correlation was found between the number of testicles and the size of males. These correlations reflect a common tendency in asexual and sexual reproduction: larger parental individuals have a greater number of propagules or gametes. No correlation was found between size and buds or size and gonads in hermaphrodites; hermaphrodites had at most one asexual bud and were significantly larger than males, females, and asexual individuals. The larger size of hermaphrodites supports the hypothesis that producing both female and male gonads is more energetically costly than producing only one type of gamete (gonochorism).     

REPRODUCTIVE ATTRITION IN THE RARE CHAPARRAL SHRUB FREMONTODENDRON DECUMBENS LLOYD (STERCULIACEAE)

We examined reproductive attrition in Fremontodendron decumbens to characterize sexual reproduction in this rare California shrub. Reproductive individuals produced an average of 2,900 flower buds in a season, with no significant difference in bud production between two seasons. Because of intense insect predation, <;2% of initiated flower buds became mature fruits. A threefold decrease in predation of flower buds between seasons resulted in an increase in seed output the second season, indicating that seed production was partially predator-limited. Most seeds (97.8%) were dormant due to an impermeable seed coat. Breaking of the coat, mechanically or by heat, allowed high levels of germination. Chamise charate and ash added to the potting medium resulted in the highest level of germination and emergence. Rodents were more important than birds as seed predators, destroying 90% of seeds under parent shrub canopies within 8–10 months. Seeds already integrated into the seed bank were comparatively safe from predation, relative to newly added seeds. If predation was prevented, seeds were long-lived under field conditions (>;80% survived after 5.75 years). Most seedlings produced in unburned chaparral by planting heat-treated seeds in openings between shrubs were destroyed by predators (rodents and insects). All seedlings that escaped predation died during the summer drought. We concluded that sexual reproduction was limited by (in order of importance): 1) lack of fire, 2) predehiscence predation by insects, and 3) postdehiscence predation by rodents. Size distributions from two populations revealed that, despite the apparent absence of sexual reproduction in unbumed chaparral, two unbumed sites contained a large proportion of individuals in small size classes. Excavation of several small individuals demonstrated they were sprouts from the roots of nearby larger shrubs. Because asexual reproduction by rootsprouting circumvents the high attrition of sexual reproductive effort on unbumed sites, rootsprouting may be a significant reproductive strategy of some ‘sprouter’ species in chaparral.     

Fitness implications of late bud break and time of burning in Lannea edulis (Sond.) Engl. (Anacardiaceae)

Many woody plants of savannas have massive underground parts (“lignotubers”) and diminutive aboveground parts with phenologies that are triggered by fire. Lannea edulis (Anacardiaceae) represents this life form and is widely distributed in south central Africa. The phenology of L. edulis was monitored on 42 permanently marked shoots over a 3-year period, from 2002 to 2004, and under three fire treatments (early and middle dry season burning, and fire protection) to determine whether (i) fire affected time of bud break and (ii) time of fruit ripening affected seed germination. Statistical analyses revealed that shoot reproductive status and fire treatments significantly affected time of bud break and the lag between bud break and leaf production. Shoot reproductive status explained 27% (P<0.001) while the interaction between reproductive status and fire treatments explained 34% (P<0.0001) of the variation in time at bud break. The appearance of the first leaf was delayed by 4 weeks in reproductive shoots compared with vegetative shoots. Shoot reproductive status explained 28% (P<0.0001) of the variation in the lag between bud break and appearance of the first leaf suggesting the existence of resource competition between reproduction and vegetative growth. However, reproductive status and fire treatments did not affect shoot size and leaves produced because replacement shoots after dieback due to fire were as long as shoots surviving from previous years due to insignificant annual shoot extension among the latter. There was high autocorrelation in the germination rate of seed-lots with seeds from fruits ripening early having a significantly higher germination rate (47% in 2002 and 89% in 2004) than those from fruits ripening later (3% in 2002 and 53% in 2004). Since time at bud break and fruit ripening were linked, it is proposed that late bud break results in reduced fitness in L. edulis because of reduced seed germination rate. Early and late dry season fires that delay bud break and destroy reproductive structures, respectively, also reduce this measure of fitness.     

Do Blue Tits time their breeding based on cues obtained by consuming buds?

ABSTRACT.   Many temperate-zone birds must track seasonal variation in food availability to time breeding and enhance reproductive success. However, the external cues birds use to fine-tune sexual development and timing of reproduction remain unclear. One possibility is that birds may use chemical substances in plants and bud tissues as a reliable predictor of leaf flush and the appearance of caterpillars, the preferred prey of many insectivorous birds during the breeding season. However, few data on seasonal shifts in key plant material consumption are available. To test this hypothesis, we examined the gizzard contents of Corsican Blue Tits ( Cyanistes caeruleus ), a small passerine bird with a breeding schedule tightly linked to the phenology of oak trees. We quantified and compared the consumption of tree buds by birds captured during winter (nonbreeding) and early spring (prebreeding) and found that bud use by Blue Tits was limited and constant through time. Fewer than 30% of birds had buds in their gizzards during the critical prebreeding period, and these items represented less than 1% of gizzard contents. It is unlikely that this limited use of buds allowed birds to track a potential chemical signal at the time of reproductive decision making. Unless more data on the consumption of plant material become available, we suggest that other environmental factors be examined to help identify the cues used by temperate-zone birds to time their reproduction.     

Reproductive cycle of the coral-excavating sponge Thoosa mismalolli (Clionaidae) from Mexican Pacific coral reefs

Abstract. Individuals of the recently described demosponge Thoosa mismalolli are common on Mexican Pacific coral reefs, excavating burrows in living corals and in other calcareous substrata. To better understand the propagative abilities of this sponge, we conducted a histological study over an 18-month period (May 2007–November 2008) to identify sexual and asexual reproductive structures. Members of the species are viviparous and hermaphroditic, with various developmental stages of oocytes, spermatic cysts, and embryos co-occurring in the mesohyl for most of the year. This nearly continuous reproductive activity intensified during the warm season. Fertilization was internal, and embryos developed inside the parental sponge to produce an unciliated hoplitomella larva, characterized by a peculiar siliceous skeleton. In addition to the sexually generated larvae, adults of T. mismalolli formed gemmules for asexual reproduction. Gemmules occurred within the mesohyl during all months of the year, but were most abundant in the coldest months. This combination of sexual and asexual processes enables individuals of T. mismalolli to reproduce almost continuously. This strategy may facilitate both long-term persistence within reefs and effective dispersal between distant reefs.     

Microscopic anatomy and pigment characterization of coral-encrusting black sponge with cyanobacterial symbiont, Terpios hoshinota

Terpios hoshinota is a black-colored sponge encrusting both live and dead corals. On examination, we determined that in the mesohyl, unicellular cyanobacteria were distributed extracellularly in dense populations; roughly half of the area in the histological sections was occupied by the cyanobacterial cells in the mesohyl of the inner zone. Pigment extraction tests of whole sponges and microspectrophotometrical analysis of cyanobacterial symbionts showed that the black coloration of the sponge is attributable to extremely densely packed cyanobacteria expressing R-phycoerythrin. Spermatic follicles were found in the mesohyl of specimens collected in summer, indicating seasonal reproduction, which is likely to play a crucial role in long-distance dispersal. We also found a possible oocyte that did not contain cyanobacteria in the ooplasm.     

Impact of an invasive alga (Womersleyella setacea) on sponge assemblages: compromising the viability of future populations

The effects of invasive species on native fauna are understudied, even though their consequences should be taken into consideration for the proper conservation and management of marine systems. Furthermore, bioinvasions may have greater consequences if they affect key structural species with slow dynamics such as marine sponges. We propose that reproductive output could be used as a potential early warning signal to detect possible future changes in population trends of long-lived species (i.e. sponges) as a result of biological invasions. The aim of this study was to investigate the effects of invasive algal (Womersleyella setacea) overgrowth on sponge reproduction by comparing the presence of reproductive elements (spermatic cysts, oocytes, embryos, and larvae) in sponges covered by a thick carpet of the invasive algae and in sponges dwelling in the same habitat but without the invasive algae. Three variables were calculated to assess the impact of the invasive alga on sponge reproduction: the reproductive effort, the proportion of individuals in reproduction, and the size of the reproductive structures. We studied eight sponge species representing the main components of the deep rocky reefs of the area. Our results showed that W. setacea had a strong negative effect on sponge reproduction in six out of eight sponge species studied, with lower and even nil reproductive structures on the sponges subjected to the algal overgrowth. Thus, considering that sexual reproduction is necessary for the persistence of most sponge populations, a significant and constant reduction of the reproductive effort may compromise their viability and affect future trends in these benthic systems.     

Reproductive effort and future parental competitive ability: A nest box removal experiment

The life history trade‐off between current and future reproduction is a theoretically well‐established concept. However, empirical evidence for the occurrence of a fitness cost of reproduction is mixed. Evidence indicates that parents only pay a cost of reproduction when local competition is high. In line with this, recent experimental work on a small passerine bird, the Great tit (Parus major) showed that reproductive effort negatively affected the competitive ability of parents, estimated through competition for high quality breeding sites in spring. In the current study, we further investigate the negative causal relationship between reproductive effort and future parental competitive ability, with the aim to quantify the consequences for parental fitness, when breeding sites are scarce. To this end, we (a) manipulated the family size of Great tit parents and (b) induced severe competition for nest boxes among the parents just before the following breeding season by means of a large‐scale nest box removal experiment. Parents increased their feeding effort in response to our family size manipulation and we successfully induced competition among the parents the following spring. Against our expectation, we found no effect of last season's family size on the ability of parents to secure a scarce nest box for breeding. In previous years, if detected, the survival cost of reproduction was always paid after midwinter. In this year, parents did pay a survival cost of reproduction before midwinter and thus before the onset of the experiment in early spring. Winter food availability during our study year was exceptionally low, and thus, competition in early winter may have been extraordinarily high. We hypothesize that differences in parental competitive ability due to their previous reproductive effort might have played a role, but before the onset of our experiment and resulted in the payment of the survival cost of reproduction.     

Male eastern bluebirds trade future ornamentation for current reproductive investment

Life-history theory proposes that organisms must trade-off investment in current and future reproduction. Production of ornamental display is an important component of reproductive effort that has rarely been considered in tests of allocation trade-offs. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet-blue plumage that is correlated with mate acquisition and male competitive ability. To investigate trade-offs between current reproductive effort and the future expression of a sexually selected ornament, we manipulated the parental effort of males by changing their brood sizes. We found that parents provisioned experimentally enlarged broods more often than reduced broods. As predicted by life-history theory, the change in parental effort had a significant effect on the relative plumage ornamentation of males in the subsequent year: males with reduced broods significantly increased in plumage brightness. Moreover, this change in plumage coloration had a direct effect on the timing of breeding in the following season: males that displayed brighter plumage in the year following the manipulation mated with females that initiated egg laying earlier in the season. These data indicate that male bluebirds must trade-off conserving energy for production of future ornamentation versus expending energy for current reproduction.     

黄花杓兰的花芽发育

对黄花杓兰（Cypripedium flavum P.F.Hunt et Summerh.）成年植株做了一个生长季的研究,提出了一年芽、二年芽和多年休眠芽的概念。指出由芽形成到植株开花需两年时间,其具体发育路线是：第一年6－7月份,根状茎顶端二年芽基部外侧有两个新的小芽产生,即“一年芽”,至9－10月份发育出7－9片幼叶,然后随气温下降停止生长;第2年4月份复苏,即为“二年芽”,二年芽在本生长季内发育成混合芽,但一般情况下只有一个充分发育,另一个未能充分发育并且一般将来也不再有发育的机会,被称为“多年休眠芽”;第3年5月份充分发育的二年芽长出地面,形成植株,迅速开花、结果,至9月底植株枯萎。本文还讨论了黄花杓兰发育过程与环境的关系。     

Experimental increase of flying costs in a pelagic seabird: effects on foraging strategies,nutritional state and chick condition

A central point in life history theory is that parental investment in current reproduction should be balanced by the costs in terms of residual reproductive value. Long-lived seabirds are considered fixed investors, that is, parents fix a specific level of investment in their current reproduction independent to the breeding requirements. We tested this hypothesis analysing the consequences of an experimental increase in flying costs on the foraging ecology, body condition and chick condition in Cory’s shearwaters Calonectris diomedea. We treated 28 pairs by reducing the wing surface in one partner and compared them with 14 control pairs. We monitored mass changes and incubation shifts and tracked 19 foraging trips per group using geolocators. Furthermore, we took blood samples at laying, hatching and chick-rearing to analyse the nutritional condition, haematology, muscle damage and stable isotopes. Eighty-day-old chicks were measured, blood sampled and challenged with PHA immune assay. In addition, we analysed the effects of handicap on the adults at the subsequent breeding season. During incubation, handicapped birds showed a greater foraging effort than control birds, as indicated by greater foraging distances and longer periods of foraging, covering larger areas. Eighty-day-old chicks reared by treated pairs were smaller and lighter and showed a lower immunity than those reared by control pairs. However, oxygen demands, nutritional condition and stable isotopes did not differ between control and handicapped birds. Although handicapped birds had to increase their foraging effort, they maintained physical condition by reducing parental investment and transferred the experimentally increased costs to their partners and the chick. This result supports the fixed investment hypothesis and is consistent with life history theory.     

Seasonal variation in reproductive traits of the oriental shrimp Palaemon macrodactylus (Crustacea: Caridea: Palaemonidae) in a non-native population

The magnitude of variations in reproductive traits of Palaemon macrodactylus females throughout a breeding season was studied in a non-native population at Mar del Plata harbor, Argentina. Fecundity, egg size, reproductive output, weight and elemental composition of eggs, and larvae were analyzed in females collected at the beginning, in the mid point, and near the end of a reproductive season and designated as early, middle season, and late females. The highest reproductive output was observed in early females, while the highest fecundity and egg volume occurred in middle season females. Eggs and larvae showed larger body mass in early than in late females. Embryos from early females contained and consumed more carbon during development than embryos from late females, and they also used part of the available nitrogen. Differences in reproduction were observed among the three groups of females. On the one hand, late females matured early but had a poor first reproduction, with few embryos and high egg loss; however, they had longer reproductive life and an enhanced reproductive output in the following season when they became early females. On the other hand, females collected at the midpoint in the reproductive season matured later and had the highest fecundity and egg volume. In addition, larvae with different characteristics resulted from each type of female and were presumably well adapted to the conditions prevailing at the moment they hatched. The extended reproductive period and the diversity of embryos and larvae produced may favor the invading ability of the species.     

Individual variation in baseline and stress-induced corticosterone and prolactin levels predicts parental effort by nesting mourning doves

Endocrine systems have an important mechanistic role in structuring life-history trade-offs. During breeding, individual variation in prolactin (PRL) and corticosterone (CORT) levels affects behavioral and physiological processes that drive trade-offs between reproduction and self-maintenance. We examined patterns in baseline (BL) and stress induced (SI; level following a standard capture-restraint protocol) levels of PRL and CORT for breeding mourning doves (Zenaida macroura). We determined whether the relationship of adult condition and parental effort to hormone levels in wild birds was consistent with life-history predictions. Both BL PRL and BL CORT level in adults were positively related to nestling weight at early nestling ages, consistent with the prediction of a positive relationship of hormone levels to current parental effort of adults and associated increased energy demand. Results are consistent with the two hormones acting together at baseline levels to limit negative effects of CORT on reproduction while maintaining beneficial effects such as increased foraging for nestling feeding. Our data did not support predictions that SI responses would vary in response to nestling or adult condition. The magnitude of CORT response in the parents to our capture-restraint protocol was negatively correlated with subsequent parental effort. Average nestling weights for adults with the highest SI CORT response were on average 10–15% lighter than expected for their age in follow-up visits after the stress event. Our results demonstrated a relationship between individual hormone levels and within population variation in parental effort and suggested that hormonal control plays an important role in structuring reproductive decisions for mourning doves.     

Modelling compensatory regrowth with bud dormancy and gradual activation of buds

Many plants show compensatory regrowth after herbivory and dormant buds often have an important role in compensatory responses. Theoretical models have shown that herbivore damage may select for a bud bank, i.e., a pool of dormant buds that are protected from herbivory and that are activated after herbivore damage. Earlier models assumed that undamaged plants cannot activate their dormant buds without damage, although they apparently have sufficient resources for successful seed production through the additional shoots dormant buds could produce. However, many plants are able to gradually activate buds over an extended period of time without any cue from damage. The aim of this study was to analyze how herbivory imposes selection for gradual mobilization of the bud bank. I assume that selection pressures that affect the fraction of buds active at each time point include damage by herbivores, time left to the end of season, and the opportunity costs of dormant buds. I modelled bud dynamics with gradual activation when there is a single damage event and (i) when the seed set of a shoot is not dependent on the time it is active, or (ii) when the seed set of a shoot diminishes with later activation. In addition, I analyzed how (iii) risk of repeated herbivory affects selection for gradual activation. Under these models, gradual activation is optimal over a wide range of herbivory pressures. Selection appears to favour activation of all buds at the beginning of the season only when herbivore pressure is weak and when early shoots have a higher seed set than late shoots. Alternatively, strong herbivore pressure and late damage may select for a large bud bank throughout the growing season, without gradual activation; the bud bank is only mobilized after damage. In this case, damaged plants can overcompensate, i.e. they have a higher seed set than undamaged plants with the same bud activation pattern. Selection for overcompensation demands a stronger herbivore pressure in this current model than in earlier bud bank models. The model never predicts selection for overcompensation when there is a risk of repeated herbivory. This revised version was published online in July 2006 with corrections to the Cover Date.     

Costs of reproduction and terminal investment by females in a semelparous marsupial

Evolutionary explanations for life history diversity are based on the idea of costs of reproduction, particularly on the concept of a trade-off between age-specific reproduction and parental survival, and between expenditure on current and future offspring. Such trade-offs are often difficult to detect in population studies of wild mammals. Terminal investment theory predicts that reproductive effort by older parents should increase, because individual offspring become more valuable to parents as the conflict between current versus potential future offspring declines with age. In order to demonstrate this phenomenon in females, there must be an increase in maternal expenditure on offspring with age, imposing a fitness cost on the mother. Clear evidence of both the expenditure and fitness cost components has rarely been found. In this study, we quantify costs of reproduction throughout the lifespan of female antechinuses. Antechinuses are nocturnal, insectivorous, forest-dwelling small (20-40 g) marsupials, which nest in tree hollows. They have a single synchronized mating season of around three weeks, which occurs on predictable dates each year in a population. Females produce only one litter per year. Unlike almost all other mammals, all males, and in the smaller species, most females are semelparous. We show that increased allocation to current reproduction reduces maternal survival, and that offspring growth and survival in the first breeding season is traded-off with performance of the second litter in iteroparous females. In iteroparous females, increased allocation to second litters is associated with severe weight loss in late lactation and post-lactation death of mothers, but increased offspring growth in late lactation and survival to weaning. These findings are consistent with terminal investment. Iteroparity did not increase lifetime reproductive success, indicating that terminal investment in the first breeding season at the expense of maternal survival (i.e. semelparity) is likely to be advantageous for females.     

Growth and abundance of desert annuals in an arid woodland in Oman

Given the role of Scaevola plumieri as a major pioneer species in the dune environment, as well as the need forindigenous sand stabilisers for South Africa’s coastal dunes, the reproductive phenology was unravelled as a firststep to establish the potential usefulness of the species in dune stabilisation programmes. In the highly variablecoastal environment the plants must cope with many problems for successful reproduction to take place. Theplants of S. plumieri overcome these difficulties through vegetative reproduction, but sexual reproduction is byno means insignificant. Numerous peduncles, each carrying multiple buds, were produced per stem, but only fewdeveloped into ripe seeds. Several of the flowers were unfertilised and of the seeds produced many were infectedand/or aborted. There was a strong effect of dune position on the reproductive phenology of S. plumieri. Thestems situated at the landward face of the foredunes showed an overall higher reproductive performance numberof buds, flowers, unripe seeds, ripe seeds. The phenogram showed that the reproduction of stems at the landwardside of the foredunes started on average 34 days earlier and showed a tendency for a longer reproductionperiod. For the duration of the different stages in the reproduction sequence, the bud stage’ took the shortesttime and the ‘ripe seed stage’ longest time. Between the different sample years, the production of the number ofpeduncles, buds, flowers and seeds was the similar; only differences in reproduction time (duration) of the differentstages were observed for the bud, flower an unripe seeds stage between the sample years.     

Female investment in offspring size and number shifts seasonally in a lizard with single-egg clutches

The timing of reproduction strongly influences reproductive success in many organisms. For species with extended reproductive seasons, the quality of the environment may change throughout the season in ways that impact offspring survival, and, accordingly, aspects of reproductive strategies may shift to maximize fitness. Life-history theory predicts that if offspring environments deteriorate through the season, females should shift from producing more, smaller offspring early in the season to fewer, higher quality offspring later in the season. We leverage multiple iterations of anole breeding colonies, which control for temperature, moisture, and food availability, to identify seasonal changes in reproduction. These breeding colonies varied only by the capture date of the adult animals from the field. We show that seasonal cohorts exhibit variation in key reproductive traits such as inter-clutch interval, egg size and hatchling size consistent with seasonal shifts in reproductive effort. Overall, reproductive effort was highest early in the season due to a relatively high rate of egg production. Later season cohorts produced fewer, but larger offspring. We infer that these results indicate a strategy for differential allocation of resources through the season. Females maximize offspring quantity when environments are favorable, and maximize offspring quality when environments are poor for those offspring. Our study also highlights that subtle differences in methodology (such as capture date of study animals) may influence the interpretation of results. Researchers interested in reproduction must be conscious of how their organism’s reproductive patterns may shift through the season when designing experiments or comparing results across studies.     

Age and terminal reproductive attempt influence laying date in the thorn‐tailed rayadito

Age‐specific variation in reproductive effort can affect population dynamics, and is a key component of the evolution of reproductive tactics. Late‐life declines are a typical feature of variation in reproduction. However, the cause of these declines, and thus their implications for the evolution of life‐history tactics, may differ. Some prior studies have shown late‐life reproductive declines to be tied to chronological age, whereas other studies have found declines associated with terminal reproduction irrespective of chronological age. We investigated the extent to which declines in late life reproduction are related to chronological age, terminal reproductive attempt or a combination of both in the thorn‐tailed rayadito Aphrastura spinicauda, a small passerine bird that inhabits the temperate forest of South America. To this end we used long‐term data (10 years) obtained on reproductive success (laying date, clutch size and nestling weight) of females in a Chilean population. Neither chronological age nor terminal reproductive attempt explained variation in clutch size or nestling weight, however we observed that during the terminal reproductive attempt older females tended to lay later in the breeding season and younger females laid early in the breeding season, but this was not the case when the reproductive attempt was not the last. These results suggests that both age‐dependent and age‐independent effects influence reproductive output and therefore that the combined effects of age and physiological condition may be more relevant than previously thought.     

松嫩平原全叶马兰种群无性系分株的建成及其年龄结构

全叶马兰（Kalimeris integrifolia Turcz.)为根蘖型多年生草本菊种植物。在生长季后期,新生肉质细根向地表生长形成根蘖芽,根蘖芽成株后形成主根。经过对大量样本观察。创建了按形态特征对根蘖型草本植物种群无性系分株和芽的年龄划分方法,以及对主根生活年限的估计方法。分株年龄是按主根芽再生的次数划分,新根蘖芽成株为1a,1a分株死亡后从主根再萌生的分株为2a,2a分株死亡后从主根再萌生的分株为3a。新根蘖芽为0a芽,其它龄级与分株的龄级相一致,即1a分株主根芽为1a芽,2a分株主根芽为2a芽,在松嫩平原割草草场,全叶马兰种群由3个龄级无性系分株组成,呈稳定型年龄结构。分株主根的生活年限最多为4年,分株的生产力种群水平以2a最高,个体水平以1a最低,并随着龄级的增加呈增长趋势,其中,在生长季初期以3a分株的生产力最高,进入中期以后则1a最低,并随着龄级的增加呈增长趋势。其中,在生长季初期以3a分株的生产力最高,进入中期以后则1a分株的相对生长速率最大,用秋末新生根蘖数量和现时种群数量可以较准确预报翌年种群数量和年龄结构。芽库中潜在种群数量是现时种群数量的2倍以上,但不意味着种群将发生疯长,而是为替补所做的充分准备。