Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems
RESEARCH ARTICLE (Open Access)

Sex and flowers: testing the resource-dependent selection hypothesis for flower sex allocation

Jonathan T. D. Finch https://orcid.org/0000-0002-6862-0308 A * , Alexander Watson-Lazowski B and James M. Cook C
+ Author Affiliations
- Author Affiliations

A Tasmanian Institute of Agriculture, University of Tasmania, Sandy Bay Campus, Private Bag 98, Hobart, Tas. 7001, Australia.

B John Innes Centre, Norwich Research Park, Norwich, NR4 7UH, UK.

C Hawkesbury Institute for the Environment, Western Sydney University, Locked Bag 1797, Penrith, NSW 2751, Australia.

* Correspondence to: jonathan.finch@utas.edu.au

Handling Editor: Lynda Prior

Australian Journal of Botany 70(4) 323-333 https://doi.org/10.1071/BT22015
Submitted: 8 February 2022  Accepted: 30 May 2022   Published: 18 July 2022

© 2022 The Author(s) (or their employer(s)). Published by CSIRO Publishing. This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License (CC BY-NC-ND)

Abstract

Context: Monoecious plants can adjust their proportional investment in male and female flowers to maximise reproductive fitness. The female reproductive function (seeds) often has greater resource costs than the male (pollen). Larger plants are generally thought to have greater resource availability and should have a female biased sex ratio, referred to as the size-dependent selection hypothesis. However, empirical tests of this hypothesis have found mixed support. This may be because size alone is not always a reliable proximate value for resource availability, which can be influenced by other abiotic factors.

Aims: Breynia oblongifolia (Phyllanthaceae) is a perennial monoecious plant with unisexual moth-pollinated flowers from eastern Australia. Fruit production in Breynia is heavily influenced by rainfall, which is highly variable. We hypothesised that where soil moisture limits female function, Breynia would produce more male flowers (i.e. resource-dependent selection).

Methods: We used a multi-year observational dataset to look for evidence of resource-dependent flower sex ratios in a wild population and conducted a manipulative glasshouse experiment to test alternative hypotheses for flower sex selection.

Key results: In both our manipulative glasshouse experiment and observed wild population, decreasing soil water content resulted in higher proportions of male flowers, supporting the resource-dependent sex selection hypothesis.

Conclusions: Soil moisture influences flower sex ratios but plant size does not.

Implications: Future studies should not assume that height equates to resource wealth, as this is often overly simplistic and ignores the potential for key resources, like soil moisture or light, to fluctuate.

Keywords: Breynia oblongifolia, flower sex selection, monoecious, Phyllanthaceae, plant height, resource-dependent sex selection, size-dependent sex selection, soil moisture.


References

Abe T (2002) Flower bud abortion influences clonal growth and sexual dimorphism in the understorey dioecious shrub Aucuba japonica (Cornaceae). Annals of Botany 89, 675–681.
Flower bud abortion influences clonal growth and sexual dimorphism in the understorey dioecious shrub Aucuba japonica (Cornaceae).Crossref | GoogleScholarGoogle Scholar | 12102522PubMed |

Ackerly DD, Jasieński M (1990) Size-dependent variation of gender in high density stands of the monoecious annual, Ambrosia artemisiifolia (Asteraceae). Oecologia 82, 474–477.
Size-dependent variation of gender in high density stands of the monoecious annual, Ambrosia artemisiifolia (Asteraceae).Crossref | GoogleScholarGoogle Scholar | 28311470PubMed |

Andrieu E, Debussche M, Thompson JD (2007) Size-dependent reproduction and gender modification in the hermaphroditic perennial plant Paeonia officinalis. International Journal of Plant Sciences 168, 435–441.
Size-dependent reproduction and gender modification in the hermaphroditic perennial plant Paeonia officinalis.Crossref | GoogleScholarGoogle Scholar |

Ashman T-L, Knight TM, Steets JA, Amarasekare P, Burd M, Campbell DR, Dudash MR, Johnston MO, Mazer SJ, Mitchell RJ, Morgan MT, Wilson WG (2004) Pollen limitation of plant reproduction: ecologocial and evolutionary causes and consequences. Ecology 85, 2408–2421.
Pollen limitation of plant reproduction: ecologocial and evolutionary causes and consequences.Crossref | GoogleScholarGoogle Scholar |

Ashok K, Guan Z, Yamagata T (2003) Influence of the Indian Ocean Dipole on the Australian winter rainfall. Geophysical Research Letters 30, 1821
Influence of the Indian Ocean Dipole on the Australian winter rainfall.Crossref | GoogleScholarGoogle Scholar |

Asikainen E, Mutikainen P (2005) Pollen and resource limitation in a gynodioecious species. American Journal of Botany 92, 487–494.
Pollen and resource limitation in a gynodioecious species.Crossref | GoogleScholarGoogle Scholar | 21652426PubMed |

Barker PA, Carl D, Kimball F, Harper T (1982) Variation in the breeding system of Acer grandidentatum. Available at https://academic.oup.com/forestscience/article/28/3/563/4656635

Bernacchi CJ, VanLoocke A (2015) Terrestrial ecosystems in a changing environment: a dominant role for water. Annual Review of Plant Biology 66, 599–622.
Terrestrial ecosystems in a changing environment: a dominant role for water.Crossref | GoogleScholarGoogle Scholar | 25621516PubMed |

Bickel AM, Freeman DC (1993) Effects of pollen vector and plant geometry on floral sex ratio in monoecious plants. American Midland Naturalist 130, 239–247.
Effects of pollen vector and plant geometry on floral sex ratio in monoecious plants.Crossref | GoogleScholarGoogle Scholar |

Blake-Mahmud J, Struwe L (2020) When the going gets tough, the tough turn female: injury and sex expression in a sex-changing tree. American Journal of Botany 107, 339–349.
When the going gets tough, the tough turn female: injury and sex expression in a sex-changing tree.Crossref | GoogleScholarGoogle Scholar | 32086802PubMed |

Cao G-X, Kudo G (2008) Size-dependent sex allocation in a monocarpic perennial herb, Cardiocrinum cordatum (Liliaceae). Plant Ecology 194, 99–107.
Size-dependent sex allocation in a monocarpic perennial herb, Cardiocrinum cordatum (Liliaceae).Crossref | GoogleScholarGoogle Scholar |

Chailakhyan MKh (1979) Genetic and hormonal regulation of growth, flowering, and sex expression in plants. American Journal of Botany 66, 717–736.
Genetic and hormonal regulation of growth, flowering, and sex expression in plants.Crossref | GoogleScholarGoogle Scholar |

Charnov EL (1982) ‘The theory of sex allocation.’ (Princeton University Press) Available at https://press.princeton.edu/books/paperback/9780691083124/the-theory-of-sex-allocation-mpb-18-volume-18

Charnov EL, Bull J (1977) When is sex environmentally determined? Nature 266, 828–830.
When is sex environmentally determined?Crossref | GoogleScholarGoogle Scholar | 865602PubMed |

Clay K (1993) Size-dependent gender change in green dragon (Arisaema dracontium; Araceae). American Journal of Botany 80, 769–777.
Size-dependent gender change in green dragon (Arisaema dracontium; Araceae).Crossref | GoogleScholarGoogle Scholar |

Costich DE (1995) Gender specialization across a climatic gradient: experimental comparison of monoecious and dioecious ecballium. Ecology 76, 1036–1050.
Gender specialization across a climatic gradient: experimental comparison of monoecious and dioecious ecballium.Crossref | GoogleScholarGoogle Scholar |

Crowley PH, Harris W, Korn E (2017) Optimal sex allocation under pollen limitation. Theoretical Ecology 10, 417–431.
Optimal sex allocation under pollen limitation.Crossref | GoogleScholarGoogle Scholar |

Cruden RW, Lyon DL (1985) Patterns of biomass allocation to male and female functions in plants with different mating systems. Oecologia 66, 299–306.
Patterns of biomass allocation to male and female functions in plants with different mating systems.Crossref | GoogleScholarGoogle Scholar | 28311603PubMed |

Delesalle VA (1992) Architecture and gender allocation: floral sex expression along branches of the Monoecious cucurbit, Apodanthera undulata. International Journal of Plant Sciences 153, 108–116.
Architecture and gender allocation: floral sex expression along branches of the Monoecious cucurbit, Apodanthera undulata.Crossref | GoogleScholarGoogle Scholar |

Delesalle VA, Mazer SJ (2009) Size-dependent pollen:ovule ratios and the allometry of floral sex allocation in Clarkia (Onagraceae) taxa with contrasting mating systems. American Journal of Botany 96, 968–978.
Size-dependent pollen:ovule ratios and the allometry of floral sex allocation in Clarkia (Onagraceae) taxa with contrasting mating systems.Crossref | GoogleScholarGoogle Scholar | 21628249PubMed |

Dey R, Lewis SC, Arblaster JM, Abram NJ (2019) A review of past and projected changes in Australia’s rainfall. WIREs: Climate Change 10, e577
A review of past and projected changes in Australia’s rainfall.Crossref | GoogleScholarGoogle Scholar |

Dodson CH (1962) Pollination and variation in the subtribe Catasetinae (Orchidaceae). Annals of the Missouri Botanical Garden 49, 35–56.
Pollination and variation in the subtribe Catasetinae (Orchidaceae).Crossref | GoogleScholarGoogle Scholar |

Dorken ME, Barrett SCH (2003) Gender plasticity in Sagittaria sagittifolia (Alismataceae), a monoecious aquatic species. Plant Systematics and Evolution 237, 99–106.
Gender plasticity in Sagittaria sagittifolia (Alismataceae), a monoecious aquatic species.Crossref | GoogleScholarGoogle Scholar |

Dorken ME, Barrett SCH (2004) Phenotypic plasticity of vegetative and reproductive traits in monoecious and dioecious populations of Sagittaria latifolia (Alismataceae): a clonal aquatic plant. Journal of Ecology 92, 32–44.
Phenotypic plasticity of vegetative and reproductive traits in monoecious and dioecious populations of Sagittaria latifolia (Alismataceae): a clonal aquatic plant.Crossref | GoogleScholarGoogle Scholar |

Dorken ME, Pannell JR (2008) Density-dependent regulation of the sex ratio in an annual plant. The American Naturalist 171, 824–830.
Density-dependent regulation of the sex ratio in an annual plant.Crossref | GoogleScholarGoogle Scholar | 18429672PubMed |

Ellsworth DS, Anderson IC, Crous KY, Cooke J, Drake JE, Gherlenda AN, Gimeno TE, Macdonald CA, Medlyn BE, Powell JR, Tjoelker MG, Reich PB (2017) Elevated CO2 does not increase eucalypt forest productivity on a low-phosphorus soil. Nature Climate Change 7, 279–282.
Elevated CO2 does not increase eucalypt forest productivity on a low-phosphorus soil.Crossref | GoogleScholarGoogle Scholar |

Finch JTD, Power SA, Welbergen JA, Cook JM (2018) Two’s company, three’s a crowd: co-occurring pollinators and parasite species in Breynia oblongifolia (Phyllanthaceae). BMC Evolutionary Biology 18, 193
Two’s company, three’s a crowd: co-occurring pollinators and parasite species in Breynia oblongifolia (Phyllanthaceae).Crossref | GoogleScholarGoogle Scholar | 30547744PubMed |

Finch JTD, Power SA, Welbergen JA, Cook JM (2019) A non-pollinating moth inflicts higher seed predation than two co-pollinators in an obligate pollination mutualism. Ecological Entomology 44, 780–791.
A non-pollinating moth inflicts higher seed predation than two co-pollinators in an obligate pollination mutualism.Crossref | GoogleScholarGoogle Scholar |

Finch JTD, Power SA, Welbergen JA, Cook JM (2021a) Staying in touch: how highly specialised moth pollinators track host plant phenology in unpredictable climates. BMC Ecology and Evolution 21, 1–13.
Staying in touch: how highly specialised moth pollinators track host plant phenology in unpredictable climates.Crossref | GoogleScholarGoogle Scholar |

Finch JTD, Power SA, Welbergen JA, Cook JM (2021b) Testing for apomixis in an obligate pollination mutualism. Journal of Pollination Ecology 29, 167–178.
Testing for apomixis in an obligate pollination mutualism.Crossref | GoogleScholarGoogle Scholar |

Fox JF (1993) Size and sex allocation in monoecious woody plants. Oecologia 94, 110–113.
Size and sex allocation in monoecious woody plants.Crossref | GoogleScholarGoogle Scholar | 28313867PubMed |

Freeman DC, Klikoff LG, Harper KT (1976) Differential resource utilization by the sexes of dioecious plants. Science 193, 597–599.
Differential resource utilization by the sexes of dioecious plants.Crossref | GoogleScholarGoogle Scholar | 17759589PubMed |

Freeman DC, Harper KT, Charnov EL (1980) Sex change in plants: old and new observations and new hypotheses. Oecologia 47, 222–232.
Sex change in plants: old and new observations and new hypotheses.Crossref | GoogleScholarGoogle Scholar | 28309476PubMed |

Freeman DC, McArthur ED, Harper KT, Blauer AC (1981) Influence of environment on the floral sex ratio of monoecious plants. Evolution 35, 194–197.
Influence of environment on the floral sex ratio of monoecious plants.Crossref | GoogleScholarGoogle Scholar | 28563448PubMed |

Geber MA, Dawson TE, Delph LF (1999) ‘Gender and sexual dimorphism in flowering plants.’ (Springer)

Ghiselin MT (1969) The evolution of hermaphroditism among animals. The Quarterly Review of Biology 44, 189–208.
The evolution of hermaphroditism among animals.Crossref | GoogleScholarGoogle Scholar | 4901396PubMed |

Gregg KB (1973) Studies on the control of sex expression in the genera Cycnoches and Catasetum, Subtribe Catasetinae, Orchidaceae. PhD thesis, University of Miami, Miami, FL, USA. Available at https://scholarship.miami.edu/esploro/outputs/doctoral/Studies-On-The-Control-Of-Sex-Expression-In-The-Genera-Cycnoches-And-Catasetum-Subtribe-Catasetinae-Orchidaceae/991031447277702976S

Guo H, Mazer SJ, Du G (2010) Geographic variation in primary sex allocation per flower within and among 12 species of Pedicularis (Orobanchaceae): proportional male investment increases with elevation. American Journal of Botany 97, 1334–1341.
Geographic variation in primary sex allocation per flower within and among 12 species of Pedicularis (Orobanchaceae): proportional male investment increases with elevation.Crossref | GoogleScholarGoogle Scholar | 21616886PubMed |

Han B, Wang X-F, Huang S-Q (2011) Production of male flowers does not decrease with plant size in insect-pollinated Sagittaria trifolia, contrary to predictions of size-dependent sex allocation. Journal of Systematics and Evolution 49, 379–385.
Production of male flowers does not decrease with plant size in insect-pollinated Sagittaria trifolia, contrary to predictions of size-dependent sex allocation.Crossref | GoogleScholarGoogle Scholar |

Hibbs DE, Fischer BC (1979) Sexual and vegetative reproduction of striped maple (Acer pensylvanicum L.). Bulletin of the Torrey Botanical Club 106, 222–227.
Sexual and vegetative reproduction of striped maple (Acer pensylvanicum L.).Crossref | GoogleScholarGoogle Scholar |

Hidalgo BF, Bazan SF, Iturralde RB, Borsch T (2020) Phylogenetic relationships and character evolution in neotropical Phyllanthus (Phyllanthaceae), with a focus on the cuban and caribbean taxa. International Journal of Plant Sciences 181, 284–305.
Phylogenetic relationships and character evolution in neotropical Phyllanthus (Phyllanthaceae), with a focus on the cuban and caribbean taxa.Crossref | GoogleScholarGoogle Scholar |

Hoffmann P, Kathriarachchi H, Wurdac KJ (2006) A phylogenetic classification of Phyllanthaceae (Malpighiales; Euphorbiaceae sensu lato). Kew Bulletin 61, 37–53.

Iqbal N, Khan NA, Ferrante A, Trivellini A, Francini A, Khan MIR (2017) Ethylene role in plant growth, development and senescence: interaction with other phytohormones. Frontiers in Plant Science 8, 475
Ethylene role in plant growth, development and senescence: interaction with other phytohormones.Crossref | GoogleScholarGoogle Scholar | 28421102PubMed |

Irish EE, Nelson T (1989) Sex determination in monoecious and dioecious plants. The Plant Cell 1, 737
Sex determination in monoecious and dioecious plants.Crossref | GoogleScholarGoogle Scholar | 12359907PubMed |

Ishii HS (2004) Increase of male reproductive components with size in an animal-pollinated hermaphrodite, Narthecium asiaticum (Liliaceae). Functional Ecology 18, 130–137.
Increase of male reproductive components with size in an animal-pollinated hermaphrodite, Narthecium asiaticum (Liliaceae).Crossref | GoogleScholarGoogle Scholar |

Iwasa Y (1991) Sex change evolution and cost of reproduction. Behavioral Ecology 2, 56–68.
Sex change evolution and cost of reproduction.Crossref | GoogleScholarGoogle Scholar |

Kawakita A, Kato M (2004) Obligate pollination mutualism in Breynia (Phyllanthaceae): further documentation of pollination mutualism involving Epicephala moths (Gracillariidae). American Journal of Botany 91, 1319–1325.
Obligate pollination mutualism in Breynia (Phyllanthaceae): further documentation of pollination mutualism involving Epicephala moths (Gracillariidae).Crossref | GoogleScholarGoogle Scholar | 21652364PubMed |

Khryanin VN (2002) Role of phytohormones in sex differentiation in plants 1. Russian Journal of Plant Physiology 49, 545–551.
Role of phytohormones in sex differentiation in plants 1.Crossref | GoogleScholarGoogle Scholar |

Klinkhamer PGL, de Jong TJ, Metz H (1997) Sex and size in cosexual plants. Trends in Ecology & Evolution 12, 260–265.
Sex and size in cosexual plants.Crossref | GoogleScholarGoogle Scholar |

Knight TM, Steets JA, Ashman T-L (2006) A quantitative synthesis of pollen supplementation experiments highlights the contribution of resource reallocation to estimates of pollen limitation. American Journal of Botany 93, 271–277.
A quantitative synthesis of pollen supplementation experiments highlights the contribution of resource reallocation to estimates of pollen limitation.Crossref | GoogleScholarGoogle Scholar | 21646188PubMed |

Korpelainen H (1994) Sex ratios and resource allocation among sexually reproducing plants of Rubus chamaemorus. Annals of Botany 74, 627–632.
Sex ratios and resource allocation among sexually reproducing plants of Rubus chamaemorus.Crossref | GoogleScholarGoogle Scholar |

Korpelainen H (1998) Labile sex expression in plants. Biological Reviews 73, 157–180.
Labile sex expression in plants.Crossref | GoogleScholarGoogle Scholar |

Liao W-J, Zhang D-Y (2008) Increased maleness at flowering stage and femaleness at fruiting stage with size in an andromonoecious perennial, Veratrum nigrum. Journal of Integrative Plant Biology 50, 1024–1030.
Increased maleness at flowering stage and femaleness at fruiting stage with size in an andromonoecious perennial, Veratrum nigrum.Crossref | GoogleScholarGoogle Scholar | 18713353PubMed |

Liu F, Chen J-M, Wang Q-F (2009) Size-dependent sex allocation in a monoecious species Sagittaria pygmaea (Alismataceae). Annales Botanici Fennici 46, 95–100.
Size-dependent sex allocation in a monoecious species Sagittaria pygmaea (Alismataceae).Crossref | GoogleScholarGoogle Scholar |

Lloyd DG (1972) Breeding systems in Cotula L. (Compositae, Anthemidae). New Phytologist 71, 1181–1194.
Breeding systems in Cotula L. (Compositae, Anthemidae).Crossref | GoogleScholarGoogle Scholar |

Lloyd DG, Bawa KS (1984) Modification of the gender of seed plants in varying conditions. Evolutionary Biology 17, 255–338.

Lloyd DG, Webb CJ (1977) Secondary sex characters in plants. The Botanical Review 43, 177–216.
Secondary sex characters in plants.Crossref | GoogleScholarGoogle Scholar |

Matsui K (1995) Sex expression, sex change and fruiting habit in an Acer rufinerve population. Ecological Research 10, 65–74.
Sex expression, sex change and fruiting habit in an Acer rufinerve population.Crossref | GoogleScholarGoogle Scholar |

McArthur ED (1977) Environmentally induced changes of sex expression in Atriplex canescens. Heredity 38, 97–103.
Environmentally induced changes of sex expression in Atriplex canescens.Crossref | GoogleScholarGoogle Scholar |

Méndez M, Traveset A (2003) Sexual allocation in single-flowered hermaphroditic individuals in relation to plant and flower size. Oecologia 137, 69–75.
Sexual allocation in single-flowered hermaphroditic individuals in relation to plant and flower size.Crossref | GoogleScholarGoogle Scholar | 12844252PubMed |

Moore JC, Pannell JR (2011) Sexual selection in plants. Current Biology 21, R176–R182.
Sexual selection in plants.Crossref | GoogleScholarGoogle Scholar | 21377091PubMed |

Nanami S, Kawaguchi H, Yamakura T (2004) Sex change towards female in dying Acer rufinerve trees. Annals of Botany 93, 733–740.
Sex change towards female in dying Acer rufinerve trees.Crossref | GoogleScholarGoogle Scholar | 15102611PubMed |

Nicotra AB (1999) Reproductive allocation and the long-term costs of reproduction in Siparuna grandiflora, a dioecious neo-tropical shrub. Journal of Ecology 87, 138–149.
Reproductive allocation and the long-term costs of reproduction in Siparuna grandiflora, a dioecious neo-tropical shrub.Crossref | GoogleScholarGoogle Scholar |

Ortiz PL, Arista M, Talavera S (2002) Sex ratio and reproductive effort in the dioecious Juniperus communis subsp. alpina (Suter) Čelak. (Cupressaceae) along an altitudinal gradient. Annals of Botany 89, 205–211.
Sex ratio and reproductive effort in the dioecious Juniperus communis subsp. alpina (Suter) Čelak. (Cupressaceae) along an altitudinal gradient.Crossref | GoogleScholarGoogle Scholar | 12099351PubMed |

Pannell J (1997) Mixed genetic and environmental sex determination in an androdioecious population of Mercurialis annua. Heredity 78, 50–56.
Mixed genetic and environmental sex determination in an androdioecious population of Mercurialis annua.Crossref | GoogleScholarGoogle Scholar | 16397637PubMed |

Pinheiro JC, Bates DM (2000) ‘Mixed-Effects Models in S and S-PLUS.’ (Springer: New York, NY, USA)
| Crossref |

Ramírez F, Davenport TL (2016) Mango (Mangifera indica L.) pollination: a review. Scientia Horticulturae 203, 158–168.
Mango (Mangifera indica L.) pollination: a review.Crossref | GoogleScholarGoogle Scholar |

Reekie E, Bazzaz F (2011) ‘Reproductive allocation in plants.’ (Eds E Reekie, F Bazzaz). (Elsevier) Available at https://books.google.com.au/books?hl=en&lr=&id=_KJdaiLKAogC&oi=fnd&pg=PP1&dq=Reekie+and+Bazzaz+(2005.+Reproductive+allocation+in+plants&ots=9e0HZrhZV7&sig=dViSU85TzeImdf1Ai71E8tIAaFI

Risbey JS, Pook MJ, McIntosh PC, Wheeler MC, Hendon HH (2009) On the remote drivers of rainfall variability in Australia. Monthly Weather Review 137, 3233–3253.
On the remote drivers of rainfall variability in Australia.Crossref | GoogleScholarGoogle Scholar |

Roach DA (1993) ‘Evolutionary senescence in plants.’ (Kluwer Academic Publishers)

Rosenheim JA, Schreiber SJ, Williams NM (2016) Does an ‘oversupply’ of ovules cause pollen limitation? New Phytologist 210, 324–332.
Does an ‘oversupply’ of ovules cause pollen limitation?Crossref | GoogleScholarGoogle Scholar | 26574903PubMed |

Sarkissian TS, Barrett SCH, Harder LD (2001) Gender variation in Sagittaria latifolia (Alismataceae): is size all that matters? Ecology 82, 360–373.
Gender variation in Sagittaria latifolia (Alismataceae): is size all that matters?Crossref | GoogleScholarGoogle Scholar |

Schlessman MA (1991) Size, gender, and sex change in dwarf ginseng, Panax trifolium (Araliaceae). Oecologia 87, 588–595.
Size, gender, and sex change in dwarf ginseng, Panax trifolium (Araliaceae).Crossref | GoogleScholarGoogle Scholar | 28313704PubMed |

Sherman DA, Dahlgren JP, Ehrlén J, García MB (2019) Sex and the cost of reproduction through the life course of an extremely long-lived herb. Oecologia 191, 369–375.
Sex and the cost of reproduction through the life course of an extremely long-lived herb.Crossref | GoogleScholarGoogle Scholar | 31428868PubMed |

Shwe EE, Wu B, Huang S-Q (2020) Both small and large plants are likely to produce staminate (male) flowers in a hermaphrodite lily. Plant Diversity 42, 142–147.
Both small and large plants are likely to produce staminate (male) flowers in a hermaphrodite lily.Crossref | GoogleScholarGoogle Scholar |

Teixido AL, Valladares F (2019) Heat and drought determine flower female allocation in a hermaphroditic Mediterranean plant family. Plant Biology 21, 1024–1030.
Heat and drought determine flower female allocation in a hermaphroditic Mediterranean plant family.Crossref | GoogleScholarGoogle Scholar | 31282088PubMed |

Teixido AL, Guzmán B, Staggemeier VG, Valladares F (2017) Phylogeny determines flower size-dependent sex allocation at flowering in a hermaphroditic family. Plant Biology 19, 963–972.
Phylogeny determines flower size-dependent sex allocation at flowering in a hermaphroditic family.Crossref | GoogleScholarGoogle Scholar | 28727278PubMed |

Tomiuk J, Hales DF, Wöhrmann K, Morris D (1991) Genotypic variation and structure in Australian populations of the aphid Schoutedenia lutea. Hereditas 115, 17–23.
Genotypic variation and structure in Australian populations of the aphid Schoutedenia lutea.Crossref | GoogleScholarGoogle Scholar |

Torices R, Méndez M (2011) Influence of inflorescence size on sexual expression and female reproductive success in a monoecious species. Plant Biology 13, 78–85.
Influence of inflorescence size on sexual expression and female reproductive success in a monoecious species.Crossref | GoogleScholarGoogle Scholar | 21134090PubMed |

Vallejo-Marín M, Rausher MD (2007) The role of male flowers in andromonoecious species: energetic costs and siring success in Solanum carolinense L. Evolution 61, 404–412.
The role of male flowers in andromonoecious species: energetic costs and siring success in Solanum carolinense L.Crossref | GoogleScholarGoogle Scholar | 17348949PubMed |

Vélez-Mora D, Ramón P, Vallejo C, Romero A, Duncan D, Quintana-Ascencio PF (2021) Environmental drivers of femaleness of an inter-Andean monoecious shrub. Biotropica 53, 17–27.
Environmental drivers of femaleness of an inter-Andean monoecious shrub.Crossref | GoogleScholarGoogle Scholar |

Wang X, Huang L, Gichira AW, Wang X (2019) The effects of density on size-dependent gender plasticity in the monoecious species Sagittaria potamogetifolia (Alismataceae). Saudi Journal of Biological Sciences 26, 413–420.
The effects of density on size-dependent gender plasticity in the monoecious species Sagittaria potamogetifolia (Alismataceae).Crossref | GoogleScholarGoogle Scholar | 31485186PubMed |

Wolfe LM, Shmida A (1995) Regulation of gender and flowering behavior in a sexually dimorphic desert shrub (Ochradentus baccatus Delile [Resedaceae]). Israel Journal of Plant Sciences 43, 325–337.
Regulation of gender and flowering behavior in a sexually dimorphic desert shrub (Ochradentus baccatus Delile [Resedaceae]).Crossref | GoogleScholarGoogle Scholar |

Wolfe LM, Shmida A (1997) The ecology of sex expression in a gynodioecious Israeli desert shrub (Ochradenus baccatus). Ecology 78, 101–110.
The ecology of sex expression in a gynodioecious Israeli desert shrub (Ochradenus baccatus).Crossref | GoogleScholarGoogle Scholar |

Wright SI, Barrett SCH (1999) Size-dependent gender modification in a hermaphroditic perennial herb. Proceedings of the Royal Society of London Series B: Biological Sciences 266, 225–232.
Size-dependent gender modification in a hermaphroditic perennial herb.Crossref | GoogleScholarGoogle Scholar |

Zhang D-Y, Jiang X-H (2002) Size-dependent resource allocation and sex allocation in herbaceous perennial plants. Journal of Evolutionary Biology 15, 74–83.
Size-dependent resource allocation and sex allocation in herbaceous perennial plants.Crossref | GoogleScholarGoogle Scholar |

Zhang L, Wang X, Du G (2011) Primary floral allocation per flower in 12 Pedicularis (Orobanchaceae) species: significant effect of two distinct rewarding types for pollinators. Journal of Plant Research 124, 655–661.
Primary floral allocation per flower in 12 Pedicularis (Orobanchaceae) species: significant effect of two distinct rewarding types for pollinators.Crossref | GoogleScholarGoogle Scholar | 21286775PubMed |

Zhang L-H, Zhang Y-W, Zhao X-N, Huang S-J, Zhao J-M, Yang Y-F (2014) Effects of different nutrient sources on plasticity of reproductive strategies in a monoecious species, Sagittaria graminea (Alismataceae). Journal of Systematics and Evolution 52, 84–91.
Effects of different nutrient sources on plasticity of reproductive strategies in a monoecious species, Sagittaria graminea (Alismataceae).Crossref | GoogleScholarGoogle Scholar |

Zhang Z-Q, Zhu X-F, Sun H, Yang Y-P, Barrett SCH (2014) Size-dependent gender modification in Lilium apertum (Liliaceae): does this species exhibit gender diphasy? Annals of Botany 114, 441–453.
Size-dependent gender modification in Lilium apertum (Liliaceae): does this species exhibit gender diphasy?Crossref | GoogleScholarGoogle Scholar | 25062885PubMed |

Zimmerman JK (1991) Ecological correlates of labile sex expression in the orchid Catasetum viridiflavum. Ecology 72, 597–608.
Ecological correlates of labile sex expression in the orchid Catasetum viridiflavum.Crossref | GoogleScholarGoogle Scholar |

Zimmerman JK, Aide TM (1989) Patterns of fruit prodcution in a neotropical orchid: pollinator vs. resource limitation. American Journal of Botany 76, 67–73.
Patterns of fruit prodcution in a neotropical orchid: pollinator vs. resource limitation.Crossref | GoogleScholarGoogle Scholar |