scholarly journals Pollinating fig wasps’ simple solutions to complex sex ratio problems: a review

2022 ◽  
Vol 19 (1) ◽  
Author(s):  
Jaco M. Greeff ◽  
Finn Kjellberg
Keyword(s):  
Natural Selection ◽  
Sex Ratios ◽  
Fig Wasps ◽  
Conditional Response ◽  
Wasp Species ◽  
Simple Strategy ◽  
Passive Response ◽  
Single Foundress ◽  
Mating Opportunities ◽  
Female Bias

AbstractLocal mate competition (LMC) favours female biased clutch sex ratios because it reduces competition between brothers and provides extra mating opportunities for sons. Fig wasps seem to fit LMC model assumptions and lay female-biased sex ratios as predicted. These female biased sex ratios increase fitness greatly. In line with predictions, their sex ratios become less female-biased as the number of mothers laying in the same fig increases. However, this variation results in comparatively small fitness benefits compared to just biased ratios and data suggest substantial mismatches with LMC theory. The mismatches are due to several factors. (1) Multiple foundresses typically lay too many daughters. (2) Single foundress sex ratios are explained by sequential oviposition and ladies-last models. (3) Mortality that typically exceeds 10% may decouple the link between primary sex ratios, the focus of model predictions, and secondary sex ratios of adult wasps that are counted by researchers. (4) Model assumptions are frequently violated: (a) clutch sizes are unequal, (b) oviposition may not be simultaneous (c) cryptic/multiple wasp species inhabit the same host, (d) foundress numbers are systematically undercounted, (e) inbreeding coefficient calculations are inaccurate, and (f) male wasps sometimes disperse. These data and calculations suggest that alternative explanations must be considered seriously. Substantial data show that wasps typically lay most of their male eggs first followed by mostly female eggs require a new approach. These “slope” strategies result in more accurate sex ratios that are automatically adjusted to foundress number, own and relative clutch sizes and to sequential clutches. This effect will alter sex ratios in all species once the egg capacity of a fig is crossed or when interference reduces clutch sizes. In addition to this passive response, the females of about half the studied species have a conditional response that reduces female bias under higher foundress numbers by laying more sons. Therefore, wasps seem to use a very simple strategy that increases their fitness. Natural selection could have optimized parameters of the slope strategy and possibly the existence of the slope strategy itself. Variation in the slope strategy that is the result of natural selection is adaptive. Research should therefore focus on quantifying variables of this slope strategy. Currently, it is unclear how much of the variation is adaptive as opposed to being coincidental by-products. Graphical Abstract

scholarly journals Sexual Selection on Population-level Mating Opportunities Drives Morph Ratios in a Fig Wasp with Extreme Male Dimorphism

2020 ◽  
Author(s):  
James M. Cook
Keyword(s):  
Population Level ◽  
Quantitative Prediction ◽  
Fig Wasp ◽  
Wing Polymorphism ◽  
Fig Wasps ◽  
Conditional Strategy ◽  
Male Dimorphism ◽  
Wasp Species ◽  
Mating Opportunities ◽  
Morph Ratios

Abstract Background Alternative mating tactics are widespread in animals and associated with extreme morphological polymorphism in some insects. Some fig wasps have both highly modified wingless males and dispersing winged males. Wingless males mate inside figs before females disperse, while winged males mate elsewhere after dispersal. Hamilton proposed a model for this system with morphs determined by alternative alleles. This has an equilibrium where the proportion of winged males equals the proportion of females dispersing unmated; i.e. the proportion of matings they obtain. Previously, we have shown qualitative support for this prediction across nine fig wasp species. Here I test the quantitative prediction in a population of the fig wasp Pseudidarnes minerva. In addition, while Hamilton envisaged simple Mendelian strategies, some fig wasp species with two wingless male morphs (but no winged males) show a conditional strategy with morph determination influenced by the number of wasps developing in a patch - I also test for this pattern in P. minerva. Results I sampled 114 figs that contained a mean of 2.1 P. minerva wasps from 44 trees across four sites in Sydney, Australia. At the whole population level, the proportion of winged males (0.84 or 0.79 corrected for sampling bias) did not differ significantly from the proportion of unmated females (0.84), providing strong quantitative support for the prediction of Hamilton’s model. In addition, there was no evidence for other factors, such as local mate competition or fighting between wingless males, that could violate simplifying assumptions of the model. Meanwhile, the proportion of winged males was not correlated with the number of wasps per fig, providing no evidence for a conditional strategy. Conclusions Morph ratios in P. minerva are consistent with Hamilton’s simple Mendelian strategy model, where morph ratios are set by average mating opportunities at the population level. This contrasts with some fig wasps from another subfamily that show conditional morph determination, allowing finer scale adaptation to fig-level mating opportunities. However, these conditional cases do not involve wing polymorphism. Male polymorphism is common and variable in fig wasps and has evolved independently in multiple lineages with apparently different underlying mechanisms.

scholarly journals The efficacy of natural selection in producing optimal sex ratio adjustments in a fig wasp species

2020 ◽  
Vol 287 (1934) ◽  
pp. 20201377 ◽  
Author(s):  
Jaco M. Greeff ◽  
Karina Pentz ◽  
Marié Warren
Keyword(s):  
Natural Selection ◽  
Sex Ratio ◽  
A Priori ◽  
Fine Tuning ◽  
Fig Wasp ◽  
Fig Wasps ◽  
Wasp Species ◽  
Local Mate ◽  
Sex Ratio Adjustment ◽  
Dual Mechanism

Ever since Darwin's discovery of natural selection, we expect traits to evolve to increase organisms' fitness. As a result, we can use optimization models to make a priori predictions of phenotypic variation, even when selection is frequency-dependent. A notable example is the prediction of female-biased sex ratios resulting from local mate competition (LMC) and inbreeding. LMC models incorporate the effects of LMC and inbreeding. Fig wasp sex ratio adjustments fit LMC predictions well. However, the appropriateness of LMC models to fig wasps has been questioned, and the role that a coincidental by-product plays in creating the apparent fit has been clearly illustrated. Here, we show that the sex ratio adjustments of a fig wasp are the result of a dual mechanism. It consists of a standard facultative LMC response favoured by natural selection, as well as a mechanism that may be the result of selection, but that could also be a coincidental by-product. If it is a by-product, the fitness increase is coincidental and natural selection's role was limited to fine-tuning it for higher fitness returns. We further document a case of an apparent fitness-reducing sex ratio adjustment. We conclude that the use of the adaptationist approach demands that our understanding of traits must be remodelled continually to rectify spurious assumptions.

scholarly journals Effect of local mate competition on fig wasp sex ratios

2006 ◽  
Vol 66 (2b) ◽  
pp. 603-610 ◽  
Author(s):  
R. A. S. Pereira ◽  
A. P. Prado
Keyword(s):  
Sex Ratio ◽  
Sex Ratios ◽  
Local Mate Competition ◽  
Mate Competition ◽  
Fig Wasp ◽  
Fig Wasps ◽  
Brood Sex Ratio ◽  
Local Mate ◽  
Female Bias ◽  
The Relationship

In fig wasps, mating takes place among the offspring of one or a few foundress mothers inside the fig from which mated females disperse to found new broods. Under these conditions, related males will compete with each other for mating and several studies have shown female bias in brood sex ratios as a response to Local Mate Competition (LMC). Studying Pegoscapus tonduzi which pollinates Ficus citrifolia in Brazil, we analysed the effect of LMC (number of foundresses) on the sex ratio of the offspring of pollinating wasps. The relationship between the foundress number and brood sex ratio qualitatively followed the theory, however the empirical sex ratio was more female biased than expected from theoretical values. The model for an optimal sex ratio considers that each foundress wasp contributes the same number of eggs to be bred and that the violation of this assumption may explain the lack of adjustment in relation to the theory.

scholarly journals Sexual selection on population-level mating opportunities drives morph ratios in a fig wasp with extreme male dimorphism

2021 ◽  
Vol 21 (1) ◽  
Author(s):  
James M. Cook
Keyword(s):  
Population Level ◽  
Quantitative Prediction ◽  
Fig Wasp ◽  
Wing Polymorphism ◽  
Fig Wasps ◽  
Conditional Strategy ◽  
Male Dimorphism ◽  
Wasp Species ◽  
Mating Opportunities ◽  
Morph Ratios

Abstract Background Alternative mating tactics are widespread in animals and associated with extreme morphological polymorphism in some insects. Some fig wasps have both highly modified wingless males and dispersing winged males. Wingless males mate inside figs before females disperse, while winged males mate elsewhere after dispersal. Hamilton proposed a model for this system with morphs determined by alternative alleles. This has an equilibrium where the proportion of winged males equals the proportion of females dispersing unmated; i.e. the proportion of matings that they obtain. Previously, we have shown qualitative support for this prediction across nine wing-dimorphic fig wasp species. Here I test the quantitative prediction in the fig wasp Pseudidarnes minerva. In addition, some fig wasp species that lack winged males, but have two wingless morphs, show a conditional strategy with morph determination influenced by the number of wasps developing in a patch. I also test for this alternative pattern in the wing-dimorphic P. minerva. Results I sampled 114 figs that contained a mean of 2.1 P. minerva wasps from 44 trees across four sites in Sydney, Australia. At the whole population level, the proportion of winged males (0.84 or 0.79 corrected for sampling bias) did not differ significantly from the proportion of unmated females (0.84), providing strong quantitative support for the prediction of Hamilton’s model. In addition, there was no evidence for other factors, such as local mate competition or fighting between wingless males, that could violate simplifying assumptions of the model. Meanwhile, the proportion of winged males was not correlated with the number of wasps per fig, providing no evidence for a conditional strategy. Conclusion The morph ratio in P. minerva is consistent with Hamilton’s simple Mendelian strategy model, where morph ratios are set by average mating opportunities at the population level. This contrasts with some fig wasps from another subfamily that show conditional morph determination, allowing finer scale adaptation to fig-level mating opportunities. However, these conditional cases do not involve wing polymorphism. Male polymorphism is common and variable in fig wasps and has evolved independently in multiple lineages with apparently different underlying mechanisms.

Factors Influencing Realized Sex Ratios in Fig Wasps: Double Oviposition and Larval Mortalities

2011 ◽  
Vol 25 (3) ◽  
pp. 254-263 ◽  
Author(s):  
Salah Ghana ◽  
Nazia Suleman ◽  
Stephen G. Compton
Keyword(s):  
Sex Ratios ◽  
Fig Wasps ◽  
Factors Influencing

Marion Island elephant seals: the paucity-of-males hypothesis tested

1999 ◽  
Vol 77 (10) ◽  
pp. 1547-1554 ◽  
Author(s):  
I S Wilkinson ◽  
R J van Aarde
Keyword(s):  
Sexual Activity ◽  
Sex Ratios ◽  
Elephant Seal ◽  
Southern Elephant Seal ◽  
Marion Island ◽  
Mirounga Leonina ◽  
Breeding Sites ◽  
Research Activities ◽  
Mating Opportunities ◽  
The Relationship

The southern elephant seal, Mirounga leonina, population at Marion Island has declined since the start of research activities there in 1973, as have populations at other Indian Ocean breeding sites. One suggested mechanism for the decline at Marion Island is a shortage of males, resulting in low insemination rates. We tested this "paucity-of-males" hypothesis by looking at (i) adult sex ratios, (ii) male sexual activity, and (iii) the relationship between levels of sexual activity and the probability of a cow pupping in the following season. Sex ratios were similar to those at other sites where populations are either stable (South Georgia) or increasing (Peninsula Valdes). Dominant bulls monopolised mating opportunities, achieving 98.1% of 629 observed copulations over three seasons. Of the 138 cows coming ashore, only 4 (2.9%) were not seen mating and 89% copulated with only one bull; dominant bulls copulated with all but 1 of the 134 (99.3%) cows that did mate. Cows were mated more frequently than at sites with stable and increasing populations. There was no difference in the observed level of sexual activity between cows seen pupping in the following season and those who failed to pup. This indicates that bulls monopolising harems are capable of inseminating all the cows. Such findings refute the paucity-of-males hypothesis.

scholarly journals Adaptation of Fig Wasps (Agaodinae) to Their Host Revealed by Large-Scale Transcriptomic Data

Insects ◽  
2021 ◽  
Vol 12 (9) ◽  
pp. 815
Author(s):  
Lianfu Chen ◽  
Simon T. Segar ◽  
Bhanumas Chantarasuwan ◽  
Da-Mien Wong ◽  
Rong Wang ◽  
...  
Keyword(s):  
Host Specificity ◽  
Large Scale ◽  
Sequence Data ◽  
Fig Wasp ◽  
Fig Wasps ◽  
Wasp Species ◽  
Functional Changes ◽  
Evolutionary Diversification ◽  
Species Specific ◽  
Close Relatives

Figs and fig wasps are highly species-specific and comprise a model system for studying co-evolution and co-speciation. The evolutionary relationships and molecular adaptations of fig wasps to their fig hosts are poorly understood, and this is in part due to limited sequence data. Here, we present large-scale transcriptomic datasets of 25 fig wasp species with the aim of uncovering the genetic basis for host specificity. Our phylogenetic results support the monophyly of all genera associated with dioecious figs, and two genera associated with monoecious figs, Eupristina and Platyscapa, were revealed to be close relatives. We identified gene loss and gain, potentially rapidly evolving genes, and genes under positive selection. Potentially functional changes were documented and we hypothesize as to how these may determine host specificity. Overall, our study provides new insights into the evolutionary diversification of fig wasps and contributes to our understanding of adaptation in this group.

scholarly journals Extraordinarily precise nematode sex ratios: Adaptive responses to vanishingly rare mating options

2021 ◽  
Author(s):  
Justin Van Goor ◽  
Edward Allen Herre ◽  
Adalberto Gomez ◽  
John D Nason
Keyword(s):  
Sex Ratio ◽  
Sex Ratios ◽  
Nematode Species ◽  
Environmental Sex Determination ◽  
Adaptive Responses ◽  
Wasp Species ◽  
Population Structures ◽  
Subdivided Populations ◽  
Sex Ratio Theory ◽  
Evolutionary Persistence

Sex ratio theory predicts both mean sex ratio and variance under a range of population structures. Here, we compare two genera of phoretic nematodes (Parasitodiplogaster and Ficophagus spp.) associated with twelve fig-pollinating wasp species in Panama. The host wasps exhibit classic Local Mate Competition: only inseminated females disperse from natal figs, and their offspring form mating pools that consist of scores of the adult offspring contributed by one or a few foundress mothers. In contrast, in both nematode genera, only sexually undifferentiated juveniles disperse, and their mating pools routinely consist of eight or fewer adults. Across all mating pool sizes, the sex ratios observed in both nematode genera are consistently female-biased (~0.34 males), which is markedly less female-biased than is often observed in the host wasps (~0.10 males). In further contrast with their hosts, variances in nematode sex ratios are also consistently precise (significantly less than binomial). The constraints associated with predictably small mating pools within highly subdivided populations appear to select for precise sex ratios that contribute both to the reproductive success of individual nematodes, and to the evolutionary persistence of nematode species. We suggest that some form of environmental sex determination underlies these precise sex ratios.

scholarly journals Phenological observations and sex ratios in Marchantia chenopoda L. (Hepaticae: Marchantiaceae)

1992 ◽  
Vol 6 (1) ◽  
pp. 161-168 ◽  
Author(s):  
María T. Moyá
Keyword(s):  
Puerto Rico ◽  
Rainy Season ◽  
Sex Ratios ◽  
Dry Season ◽  
Reproductive Structures ◽  
Female Bias ◽  
Young Sporophytes

Sex ratios were determined for 15 populations of Marchantia chenopoda L. along Puerto Rico. Sex was determined for twenty randomly selected individuals from each population. A female bias was observed in most populations. A census of reproductive structures of M. chenopoda was performed every two weeks in order to determine phenology. The highest archegoniophore count was observed to occur in January and February; minimum and maximum fertilization distances were 0.7 cm and 65 cm, respectively. Young sporophytes were most frequently observed at the beginning of the rainy season, in April and May. The highest frequency of mature sporophytes occurred during the dry season, in January and February. Capsules opened mostly during the heavy rainy season, from August to November. Protonemata were observed in the periphery of the population. Most young sporophytes aborted at the end of the moderate to heavy rainy season, from June to August. Immature gemmae cups were most frequent at the beginning of the rainy season, in April and May. Mature gemmae cups were observed throughout the rainy season, but were most frequent in September.

scholarly journals Genome-Wide Analysis of Gene Families of Pattern Recognition Receptors in Fig Wasps (Hymenoptera, Chalcidoidea)

Genes ◽  
2021 ◽  
Vol 12 (12) ◽  
pp. 1952
Author(s):  
Hong-Xia Hou ◽  
Da-Wei Huang ◽  
Zhao-Zhe Xin ◽  
Jin-Hua Xiao
Keyword(s):  
Pattern Recognition ◽  
Gene Families ◽  
Pattern Recognition Receptors ◽  
Molecular Evidence ◽  
Immune Recognition ◽  
Fig Wasp ◽  
Fig Wasps ◽  
Specific Domain ◽  
Wasp Species ◽  
Functional Studies

Pattern recognition receptors (PRRs) play important roles in detecting pathogens and initiating the innate immune response. Different evolutionary histories of pollinators and non-pollinators may result in different immune recognition systems. A previous study had reported that there were significant differences in peptidoglycan recognition proteins (PGRPs) between pollinators and non-pollinators in gene number and lineage of specific genes. In this study, based on the genomic data of 12 fig wasp species, with seven pollinators and five non-pollinators, we investigated the evolution patterns of PRRs, such as Gram-negative bacteria-binding proteins (GNBPs), C-type lectins (CTLs), scavenger receptors class B (SCRBs), fibrinogen-related proteins (FREPs), galectins, and thioester-containing proteins (TEPs). Our results showed that pollinators had no GNBP, but non-pollinators all had two gene members, which were clustered into two different clades in the phylogenetic tree, with each clade having specific domain and motif characteristics. The analysis of CTL and SCRB gene families also showed that there were lineage-specific genes and specific expansion in non-pollinators. Our results showed that there were significant differences in immune recognition between pollinators and non-pollinators, and we concluded that they had undergone flexible adaptive evolution in different environments. Our study can provide more molecular evidence for future functional studies on the immune system of fig wasps.

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