Potential for parthenogenesis of virgin females in a bisexual population of the geographically parthenogenetic mayfly Ephoron shigae (Insecta: Ephemeroptera,Polymitarcyidae)

The burrowing polymitarcyid mayfly Ephoron shigae is a geographically parthenogenetic species. Interestingly, the distributions of the bisexual and unisexual populations overlap broadly in their respective geographic ranges. In this mayfly, obligatory diploid thelytoky appears within unisexual populations. In the present study, we examined the potential for parthenogenesis or the parthenogenetic ability of females in a bisexual population aiming to understand the emergence of unisexual populations. The results obtained revealed that females in the examined bisexual populations showed a potential for diploid thelytoky as also seen in the unisexual populations, although, in females from bisexual populations, the development success rates of their unfertilized eggs were considerably lower than those of virgin females from unisexual populations. In the three bisexual reproducing species (Ephemera japonica, Ephemera strigata, and Ephemera orientalis) in the closely‐related family Ephemeridae, diploid thelytoky (i.e. tychoparthenogenesis; < 3%) was also observed. However, in this case, the parthenogenetic development success rates of unfertilized eggs were significantly lower than those of virgin females in the bisexual (Hino‐yosui Irrigation Canal) population of E. shigae. Accordingly, we suggest that parthenogenetic ability (i.e. tychoparthenogenesis or facultative parthenogenesis) in bisexual populations of E. shigae may facilitate the evolutionary transition to unisexual populations with fully obligatory parthenogenesis. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 326–334.     

Geographic parthenogenesis in the stream mayfly Eurylophella funeralis in eastern North America

Unisexual populations were found largely at the periphery of the geographic range of the mayfly Eurylophella funeralis in eastern North America. Bisexual populations generally had a normal sex ratio but at least two populations were observed with only about 2–11% males. Hatch success for unfertilized eggs depended on temperature but in general averaged >61% in unisexual populations and < 14% in bisexual populations. Eggs took about 65 d to hatch at 10°C and 19 d at 25°C; no hatching occurred at 5 or 30°C. Successful oviposition was observed for about 97% and 20% of unmated female adults from unisexual and bisexual populations respectively. The hatch success of unfertilized eggs for three geographically distinct bisexual populations was in versely correlated with the proportion of males in the populations. The genetic structure of bisexual and unisexual populations was very similar. Unisexual populations consists of clones, with as few as four recognizable phenotypes being present at a given location. All offspring had the same phenotype as the mother. Tychoparthenogenesis is suggested as the most reasonable working hypothesis concerning the origin of unisexual populations in E. funeralis .     

Automictic parthenogenesis of a geographically parthenogenetic mayfly,Ephoron shigae (Insecta: Ephemeroptera,Polymitarcyidae)

In the geographically parthenogenetic mayfly, Ephoron shigae, egg maturation and counts of chromosome number of unfertilized, parthenogenetic eggs were studied, in comparison with fertilized eggs from a bisexual population. The primary oocyte becomes mature through two successive maturation divisions. The first maturation division (meiotic division) takes place in the primary oocyte to produce a secondary oocyte and a first polar body. The second maturation division soon occurs in the secondary oocyte, in which the nucleus is divided into a mature egg nucleus (female pronucleus) and second polar body nuclei. The first polar body, in some cases, was successively divided into two polar bodies; in other instances, it was not divided. After the successive maturation division, the egg nucleus and the sister second polar body nucleus drew near to fuse into the zygote nucleus. The chromosome number was doubled in the zygote, and this conjugation initiates subsequent embryonic development. This suggests that, in E. shigae, the process of parthenogenetic recovery of diploidy is the automictic type categorized as the ‘terminal fusion’ pattern. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 335–343.     

ANDROGENETICS AND TRIPLOIDS FROM AN INTERACTING PARTHENOGENETIC HYBRID AND ITS ANCESTORS IN STICK INSECTS

Populations of unisexual organisms are often assumed to be genetically invariant (clones) and destined to a short existence on an evolutionary timescale. Unisexual organisms are most often obligate parthenogens and, by definition, ought to be completely isolated reproductively from related bisexual organisms. The assumption of complete reproductive isolation between amphimictic ancestors and thelytokous hybrids is common to most hypotheses on the evolution of sex and its adaptive significance. Stick insects of the genus Bacillus however provide evidence for reproductive interactions between allodiploid parthenogens and their ancestors, because pure species progeny (androgenetics) and triploid descendants are produced. These findings demonstrate that, through androgenesis, offspring of parthenogenetic hybrid females can contribute specimens of both sexes to the fathering species when fertilized by syntopic ancestral males and the parthenogenetic egg of strictly clonal females, when fertilized, allows a third genome to be added to the allodiploid chromosome set. These triploid genomes promote further genetic diversification and evolution of the unisexual populations through the formation of new clones by recombination during the changed maturation mode of allotriploid eggs. All this argues for much more complex breeding systems and evolutionary pathways than are usually assumed for hybrid unisexual organisms.     

Ovostatin,an endogenous trypsin inhibitor of sea urchin eggs: Purification and characterization of ovostatin from eggs of the sea urchin,Strongylocentrotus intermedius

A trypsin inhibitor, termed ovostatin, has been purified approximately 265-fold with 82% yield, from unfertilized eggs of the sea urchin Strongylocentrotus intermedius, using trypsin coupled Sepharose 4B as an affinity column for chromatography. The isolated ovostatin is homogeneous in sodium dodecyl sulfate/polyacrylamide gel electrophoresis, the estimated molecular weight being 20K–21.5K. Ovostatin inhibits preferentially trypsin-like endogenous protease purified from the eggs of the same species and bovine pancreatic trypsin and also bovine pancreatic chymotrypsin. Values of IC₅₀ (amount causing 50% inhibition of enzymes) for trypsin-like protease purified from eggs of the same species, bovine pancreatic trypsin, and bovine pancreatic chymotrypsin, are 0.91 ± 0.13 μg/ml (4.55 ± 0.65 × 10^?8 M), 3.0 ± 0.28 μg/ml (1.5 ± 0.14 × 10^?7 M), and 4.8 ± 0.2 μg/ml (2.4 ± 0.1 × 10^?7 M), respectively, in the experimental condition used. Kinetic studies indicate that ovostatin is a noncompetitive inhibitor of trypsin. The inhibitor is relatively heat labile. NaCl (0.025–0.01 M) enhances the inhibitor activity, whereas KCl is inhibitory. Ovostatin requires a low concentration of Ca²⁺ for activity. The activity is higher in unfertilized eggs than in fertilized eggs; total activity and specific activity in unfertilized eggs is about 1.67-fold and 1.85-fold higher than those in fertilized eggs, respectively. We believe that ovostatin may regulate the function of the cortical granule protease and other trypsin-like proteases that are activated in sea urchin eggs during fertilization.     

Sex allocation in a field population of an autoparasitoid

Autoparasitoid wasps lay fertilized eggs in homopteran nymphs, and these eggs develop into female primary parasitoids. Unfertilized, male-producing eggs are laid in immatures of the wasps' own or another primary parasitoid species; males then develop as secondary or hyperparasitoids. In the population of Encarsia pergandiella studied in Ithaca, NY, fertilized eggs were laid in the nymphs of the whitefly Trialeurodes packardi (primary hosts) and unfertilized eggs were laid almost exclusively in pupal females of their own species (secondary hosts). In the two years the population was studied, secondary hosts were always much less abundant than primary hosts at both sites. However, secondary hosts were parasitized at a significantly greater rate than primary hosts. In a laboratory experiment, the encounter rate of females with primary and secondary hosts was not significantly different. Moreover, there was no evidence from the field that wasps found leaves bearing secondary hosts more frequently than leaves without secondary hosts. Dissections of field-collected females showed them to be mated, and thus capable of laying both unfertilized and fertilized eggs. These results suggest that wasps did not encounter secondary hosts at a greater rate, nor were they constrained to lay unfertilized eggs, but rather secondary hosts were preferred. The oviposition sex ratios were influenced by the proportion of secondary hosts, but were less female-biased than would be predicted from the proportion of secondary hosts alone. The results do not support the predictions of Godray and Waage (1990) for either strictly host-limited autoparasitoids (sex ratio should reflect the proportion of secondary hosts) or for egg-limited autoparasitoids (sex ratio should be equal, and independent of the proportion of secondary hosts).     

Life‐cycle variation,including female production by virgin females in Frankliniella occidentalis (Thysanoptera: Thripidae)

The influence of three different temperatures on developmental time and sex ratio was investigated in the bisexual Thysanoptera species Frankliniella occidentalis. Increasing temperatures decreased developmental time and induced a more female biased sex ratio. Remarkably, there are second instars with a prolonged developmental time requiring the same number of hours as the shortest developmental time from egg to adult. Arrhenotokous reproduction in this species is based on haplodiploidy, with virgin females producing male offspring exclusively. However, at all three temperatures tested, about 0.5% of offspring from unfertilized eggs were females. The presence of Wolbachia could not be detected in Western flower thrips and can be excluded as influencing reproduction in this species.     

Geographical variation in Japan in egg development of the mayfly, Ephoron shigae (Ephemeroptera: Polymitarcyidae)

1. The effect of temperature on embryonic development was compared in four populations, two bisexual and two unisexual, of Ephoron shigae , including one each near the northern and southern periphery of the species range in Japan.
2. Eggs from every population were chilled at 4, 8 or 12 °C for diapause development after 50 days at 20 °C for pre-diapause development (experiment I). Some eggs hatched during chilling at 8 °C or 12 °C, whereas no eggs hatched at 4 °C. The rate of hatching in a given condition of chilling was higher for the eggs from warmer winter environments.
3. Chilling at 4 or 8 °C effectively facilitated diapause development. Chilling at 12 °C was, in general, not so effective, but relatively effective for the eggs from warmer winter environments.
4. Eggs were incubated at 8, 12, 15 or 20 °C after chilling at 4 °C to examine the effect of temperature on post-diapause development (experiment II). The eggs incubated at higher temperature after chilling hatched quicker and more synchronously and had higher hatching success.
5. The relationship between temperature and the days required for hatching after chilling was well described by the power function. There was no significant difference in the slope of the regression lines (i.e. temperature dependency) among local populations. However, a longer time was required for hatching at a given temperature for the population from the colder winter environment.
6. There was no detectable difference in the observed intraspecific variations between unisexual and bisexual populations.     

Population variation and ecological correlates of tychoparthenogenesis in the mayfly, Stenonema femoratum

Species in which both sex and parthenogenesis co‐occur are extremely valuable for investigating ecological conditions favouring sex. Tychoparthenogenesis is a breeding system characterized by hatching of a small proportion of unfertilized eggs (typically < 10%) from females of sexually reproducing species. With tychoparthenogenesis, both sexual and parthenogenetic reproduction co‐occur within the same population. To identify ecological conditions that may favour this breeding system, I quantified population variation in females’ capacity for tychoparthenogenesis and investigated biotic and abiotic correlates of tychoparthenogenesis. I estimated tychoparthenognetic capacity (proportion of unfertilized eggs hatching) for females from 12 Missouri populations of the mayfly, Stenonema femoratum (Ephemeroptera: Heptageniidae), across three different habitat types – temporary streams, permanent streams and lakes. Tychoparthenogenetic capacity, measured as the population mean hatch success of unfertilized eggs, ranged from 3.8 to 10.7%. Tychoparthenogenetic capacity varied among habitats in 1996, but not in 1997. In 1996, temporary streams showed hatch success of unfertilized eggs twice that of permanent streams and lakes. Tychoparthenogenetic capacity also varied among sampling dates within years. Temporary streams also showed extremely low nymph densities compared to the other two habitats. However, habitats did not differ in adult density. Furthermore, in all populations nymphs showed significantly female‐biased sex ratios. In contrast, adult sex ratios were equal or slightly male biased. Tychoparthenogenetic capacity was negatively correlated with nymph density in 1996, but not in 1997, suggesting possible reproductive assurance in some years. Adult densities also suggested that there may be certain times of year when tychoparthenogenesis may provide benefits of reproductive assurance. Although habitats differed significantly in their abiotic characteristics, tychoparthenogenetic capacity was correlated significantly with water temperature only. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75 , 101–123.     

双斑恩蚜小蜂的生殖方式及其在烟粉虱体内的发育

首次对双斑恩蚜小蜂Encarsia bimaculata Heraty et Polaszek雌蜂的胚胎发育全过程进行连续观测和数码摄像,明确了双斑恩蚜小蜂的生殖方式是两性产雌,雌蜂为初寄生蜂,孤雌产雄且雄蜂为自复寄生。雌蜂在温度(25±1)℃,相对湿度70%±10%条件下,从卵到成虫的发育历期为(13.1±2.5)d,卵期(1.98±0.38)d,幼虫1龄(0.95±0.33)d,2龄(2.48±0.40)d,3龄(3.46±0.43)d,预蛹(0.95±0.20)d,蛹(4.88±0.80)d。     

Genetic diversity,sexual condition,and microhabitat preference determine mating patterns in Sphagnum (Sphagnaceae) peat‐mosses

In bryophytes, the possibility of intragametophytic selfing creates complex mating patterns that are not possible in seed plants, although relatively little is known about patterns of inbreeding in natural populations. In the peat‐moss genus Sphagnum, taxa are generally bisexual (gametophytes produce both sperm and egg) or unisexual (gametes produced by separate male and female plants). We sampled populations of 14 species, aiming to assess inbreeding variation and inbreeding depression in sporophytes, and to evaluate correlations between sexual expression, mating systems, and microhabitat preferences. We sampled maternal gametophytes and their attached sporophytes at 12–19 microsatellite loci. Bisexual species exhibited higher levels of inbreeding than unisexual species but did generally engage in some outcrossing. Inbreeding depression did not appear to be common in either unisexual or bisexual species. Genetic diversity was higher in populations of unisexual species compared to populations of bisexual species. We found a significant association between species microhabitat preference and population genetic diversity: species preferring hummocks (high above water table) had populations with lower diversity than species inhabiting hollows (at the water table). We also found a significant interaction between sexual condition, microhabitat preference, and inbreeding coefficients, suggesting a vital role for species ecology in determining mating patterns in Sphagnum populations. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 96–113.     

Morphological features,development and reproduction of Melittobia acasta on Bombus terrestris

Morphological features, development and reproduction behavior of the parasite Melittobia acasta (Walker) were studied when reared on the pupae of the bumblebee Bombus terrestris L. in the laboratory under 23°C, 50% relative humidity and 12 h light : 12 h dark conditions. The parasites laid transparent white and elongated eggs. Newly hatched larval size and shape were very similar to eggs but they were identified by their body segments. Larvae increased their body size through moulting and transformed into a vermiform shape. Male pupae were shiny brown with dots. The female pupae were distinguished by their black shiny color, shorter size and the presence of compound eyes. Adult male pupae were dark brown and dwarf‐winged, whereas female pupae were macropterous and brachypterous. Reproduction took place by fertilization and also parthenogenetically. Mean fecundity within 5 days by mated (47.9 ± 30.5 female^?1) and virgin (7.4 ± 6.8 female^?1) females were statistically different. Mated females laid fertilized eggs that produced adult males or females, whereas virgin females laid unfertilized eggs that produced males. Development durations of the virgin female originated eggs, larvae, pupae and adults were statistically identical with those of mated females. The parasites were female‐biased and foundress number did not affect offspring sex ratio. This study shows that both mated and virgin females of M. acasta can produce many offspring on B. terrestris pupae within a short period, indicating that they are dangerous parasites of the bumblebee in a mass rearing system.     

Aspects of the mating behaviour of male mollies (Poecilia spp.)

The unisexual fish Poecilia formosa (the Amazon molly) reproduces by gynogenesis, a process in which sperm from the males of the host bisexual species activates development of its eggs. Unisexuals live with one of the host species in nature and compete with bisexual females for the males. It was long thought that male discrimination and mate selection established a balance between the unisexual and bisexual populations. Thus, hierarchies of males were set up in which dominant males mated with their conspecific females and subordinates mated with the Amazon molly. Recent evidence suggested, however, that male fish do not discriminate between their own females and the Amazon molly, and that there always are more males available than sexually receptive females of both species.
Our findings indicate that male behaviour may be more complex than suggested by either hypothesis. Mate discrimination and courtship behaviour appear to increase with age, so that large males show almost complete preference for their conspecific females, but smaller males will mate with the Amazon molly. In complex groups, small males often dart in and mate with their own females whilst the large males are engaged in courtship activities and defence of territory.     

No patrigenes required for femaleness in the haplodiploid wasp Nasonia vitripennis

The parasitoid wasp Nasonia vitripennis is an emerging model organism for developmental and behavioral genetics. It reproduces by haplodiploidy; males typically develop parthenogenetically from haploid eggs and females from fertilized diploid eggs. A polyploid mutant strain is available in which females are triploid and lay haploid and diploid eggs that normally develop into males when unfertilized. In contrast to previous reports, approximately 2% of triploid females were found to occasionally produce daughters as well as gynandromorphs from diploid unfertilized eggs. Daughter production increased with age and differed among familial lineages. This is the first report of parthenogenetic female development in Nasonia. The results show that a paternally provided genome is not required for femaleness and call for modifications of existing models of sex determination in Nasonia.     

Are individuals from thelytokous and arrhenotokous populations equally adept as biocontrol agents? Orientation and host searching behavior of a fruit fly parasitoid

Hymenopteran parasitoids generally reproduce by arrhenotoky, in which males develop from unfertilized eggs and females from fertilized eggs. A minority reproduce by thelytoky, in which all-female broods are derived from unfertilized eggs. Thelytokous populations are potentially of interest for augmentative biological control programs since the exclusive production of females could significantly lower the costs of mass rearing. Behavioral traits are a major component of parasitoid efficacy. Here, we examined orientation and host searching behavior in thelytokous and arrhenotokous populations of the fruit fly parasitoid Odontosema anastrephae Borgmeier (Hymenoptera: Figitidae). Orientation behavior to various odorant sources was studied in a two-choice olfactometer. No major differences were found between thelytokous and arrhenotokous wasps for this behavior. However, when host-searching behaviors were analyzed, some differences were found. Thelytokous females arrived sooner, foraged longer, and remained longer on non-infested guavas than arrhenotokous females. Individuals of both forms exhibited similar stereotyped behavioral sequences vis-à-vis guava treatments, with only slight deviations detected. Our results suggest that individuals from selected thelytokous and arrhenotokous O. anastrephae populations have similar abilities to search for tephritid larvae, supporting the use of thelytokous strains for augmentative releases.     

Egg Deposition Behavior in the Haplodiploid Sawfly Athalia rosae ruficornis Jakovlev (Hymenoptera: Symphyta: Tenthredinidae)

The egg deposition behavior of the turnip sawfly, Athalia rosae (Hymenoptera: Symphyta), is described. Both unmated and mated females lay eggs individually inside of fresh young leaves of cruciferous plants. During an oviposition event, females exhibit a distinct pause in abdominal contractions just before the actual egg deposition act. Unmated females show a longer pause (11.31 s on average) than mated females (4.38 s on overall average). By employing an eye color mutation, the sex of the eggs laid by females was ascertained. Females mated once lay mostly fertilized (diploid female) eggs initially but begin to lay a considerable number of unfertilized (haploid male) eggs later in life. The laying of an unfertilized egg is associated with a longer pause (6.98 s on average) than the laying of a fertilized egg (3.76 s on average). These results are in contrast to previous reports on apocritan Hymenoptera, where the presence of a pause or a longer pause during oviposition was associated with the deposition of fertilized eggs rather than unfertilized eggs. The possibility that mated Athalia rosae females control fertilization and its implications for sex allocation strategies are discussed.     

Coexistence of bisexual and unisexual populations of Nysius groenlandicus in the Zackenberg Valley,Northeast Greenland

The seed‐feeding true bug Nysius groenlandicus (Zetterstedt) (Heteroptera: Lygaeidae) has temperature and humidity preferences similar to many desert‐inhabiting insects. Nevertheless, it is widely distributed in Greenland and occurs even in high arctic areas. This seeming paradox implies that the species has developed physiological, ecological, and behavioral mechanisms that enable it to cope with the extreme arctic environment. One of these adaptations is its ability to reproduce both sexually and asexually. Sexual reproduction is most common in coastal populations, whereas asexual reproduction (parthenogenesis) occurs mainly in inland populations. Preliminary data from the Zackenberg Valley in Northeast Greenland indicated that both reproductive strategies occurred within the same area. We therefore decided to focus on this area to find evidence of (1) coexistence of unisexual and bisexual populations, (2) factors that correlate with the distribution of the two reproductive types, and (3) morphological differences between females from sexual and asexual populations. We sampled N. groenlandicus at 51 sites within the Zackenberg area. Males and females were found in almost equal numbers in only four of the samples, and only in samples taken close to the coast line, whereas samples from the inner part of the valley consisted almost entirely of females. Thus, the distribution of uni‐ and bisexual populations at the local scale agreed with the pattern previously found for Greenland as a whole. Using data obtained from two climate stations operated in the area, we showed that average summer temperatures tend to be more favorable for N. groenlandicus in the inner part of the valley than at the coast. The inland populations inhabited small isolated patches, whereas the sexual populations were found in the more homogeneous areas along the coast. Data support the hypothesis that coexistence of the two reproductive types is maintained by a combination of environmental variability, which gives sexual reproduction an advantage, and metapopulation dynamics characterized by frequent extinctions and re‐colonizations, which favors asexual reproduction. Head width of females from unisexual populations was on average 7% larger than that of females from bisexual populations. The mechanism behind this morphological difference needs further investigation.     

A direct effect of some rifamycin derivatives on the morphology of mammalian mitochondria

Protein synthesis has been investigated in cell-free preparations from mature ovarian oocytes, unfertilized and fertilized eggs, and early embryos of Drosophila melanogaster. Preparations from unfertilized eggs have a specific activity that is 5- to 6-fold higher than the activity of fractions from ovarian oocytes. There is an additional small increase in activity of preparations from fertilized eggs. The specific activity that is rapidly attained in the fertilized egg remains essentially constant for 2 to 2.5 h after fertilization, decreases sharply during blastoderm formation, and again increases during gastrulation. The activities of unfertilized eggs decline slightly during the first 2 h after oviposition, and then decrease more sharply. About 35 % of the ribosomes in preparations from both unfertilized and fertilized eggs sediment in the polyribosome region of sucrose density gradients, whereas no polyribosomes could be detected in preparations from ovarian oocytes. In both ovarian oocytes and fertilized eggs, less than 1 % of the ribosome populations were present as subunits. Additional ribonucleoprotein material of buoyant densities different from those of ribosomal subunits or ribosomes was found throughout the sucrose gradients. About 3.5 % of the ribosomes were found to be membrane-bound in preparations from both unfertilized and fertilized eggs.     

Gonad development of the sea anemone Anthopleura asiatica in clonal populations

Seasonal fluctuations in relative gonad volume and oocyte size of the sea anemone Anthopleura asiatica were examined in 3 unisexual (male) populations and one bisexual population in the Seto Inland Sea of Japan from December 1982 to December 1985. A distinct annual cycle of gonadal maturation with a peak in the summer was found in all of the populations, although they appeared to be sustained only by asexual reproduction. Spawning occured synchronously between the 2 sexes early in the fall in the bisexual population while it was one to one and a half months later in the unisexual populations.     

Elevated water temperature impairs fertilization and embryonic development of whitefish Coregonus lavaretus

The adverse effects of high temperatures on the early life stages of anadromous whitefish Coregonus lavaretus were experimentally examined by assessing fertilization success, the percentage of developmental abnormalities, cumulative mortality and the rate of embryogenesis across a range of temperatures. Temperatures ≥ 7° C increased the proportion of unfertilized and abnormally dividing eggs, deformed embryos and consequent mortality. The higher the temperature, the more severe were the effects. When eggs were fertilized and constantly incubated at various temperatures, the effective level for 50% of the eggs and embryos (EL50) of temperature was 7·6° C at the developmental stage when eye pigmentation was visible. Fewer developmental abnormalities and a lower cumulative mortality rate were observed when embryos were exposed to high temperatures from the later, gastrula stage, than from fertilization or the four‐cell stage. Irrespective of retarded development in terms of day‐degrees (i.e. the sum of daily mean temperatures), a high incubation temperature reduced the development time of C. lavaretus, leading to earlier hatching, and hatched fry were shorter than at the reference temperature of 4–5° C. Global warming will particularly pose risks for stenothermic species such as C. lavaretus, with early life stages being especially susceptible. Thus, relatively small increases and fluctuations in river water temperatures during the spawning season of this anadromous species may have substantial negative impacts on its recruitment and population persistence.