Hostname: page-component-848d4c4894-8kt4b Total loading time: 0 Render date: 2024-07-06T15:36:34.500Z Has data issue: false hasContentIssue false
  • English
  • Français

Inbreeding depression and the cost of inbreeding on seed germination

Published online by Cambridge University Press:  09 December 2015

Jerry M. Baskin  and
Carol C. Baskin
Jerry M. Baskin
Affiliation:
Department of Biology, University of Kentucky, Lexington, KY40506-0225USA
Carol C. Baskin*
Affiliation:
Department of Biology, University of Kentucky, Lexington, KY40506-0225USA Department of Plant and Soil Sciences, University of Kentucky, Lexington, KY40546-0312USA
*
*Correspondence E-mail: ccbask0@uky.edu
Get access Rights & Permissions [Opens in a new window]

Abstract

We review the literature on effects of inbreeding depression (ID) on seed germination for 743 case studies of 233 species in 64 families. For 216 case studies, we also review the relationship between mass and germination in inbred vs. outbred seeds. Inbred seeds germinated equally well as outbred seeds in 51.1% of 743 case studies, but better than outbred seeds in only 8.1%. In c. 50.5% of 216 cases, mass of inbred seeds was equal to (38.0%) or larger than (12.5%) that of outbred seeds. The magnitude of ID spans most of the − 1 to +1 range for relative performance for germination of inbred vs. outbred seeds; in contrast to what might be expected, seed germinability often is not negatively correlated with the coefficient of inbreeding (F) or positively corrected with population genetic diversity; neither heterosis nor outbreeding depression for germination is common in crosses between populations; and ID in most endemics is low and does not differ from that of widespread congeners. Our results on the effects of ID on seed mass and germination do not agree with the limited number of comparisons Darwin (1876) made on the effects of selfing vs. outcrossing on these two life-history traits. Recommendations are made on how to improve dormancy breaking and germination procedures in order to make the results of studies on ID more relevant to the natural world.

Keywords

dormancy-breaking/germination proceduresendemic speciesheterosismagnitude of inbreeding depressionmaternal environmental effectsoutbreeding depressionoutcrossing distanceseed mass
Type
Review Article
Information
Seed Science Research , Volume 25 , Issue 4 , December 2015 , pp. 355 - 385
Copyright
Copyright © Cambridge University Press 2015 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Aegisdóttir, H.H. and Thorhallsdóttir, T.E. (2006) Breeding system evolution in the Arctic: a comparative study of Campanula uniflora in Greenland and Iceland. Arctic, Antarctic, and Alpine Research 38, 305–312.Google Scholar
Aegisdóttir, H.H., Jespersen, D., Kuss, P. and Stöcklin, J. (2007) No inbreeding depression in an outcrossing alpine species: the breeding system of Campanula thyrsoides . Flora 202, 218–225.CrossRefGoogle Scholar
Affre, L. and Thompson, J.D. (1997) Population genetic structure and levels of inbreeding depression in the Mediterranean island endemic Cyclamen creticum (Primulaceae). Biological Journal of the Linnean Society 60, 527–549.Google Scholar
Affre, L. and Thompson, J.D. (1999) Variation in self-fertility, inbreeding depression and levels of inbreeding in four Cyclamen species. Journal of Evolutionary Biology 12, 113–122.CrossRefGoogle Scholar
Ågren, J. and Schemske, D.W. (1993) Outcrossing rate and inbreeding depression in two annual monoecious herbs, Begonia hirsuta and B. semiovata . Evolution 47, 125–135.Google Scholar
Alonzo, C. and Garcia-Sevilla, M. (2013) Strong inbreeding depression and individually variable mating system in the narrow endemic Erodium cazorlanum (Geraniaceae). Anales del Jardin Botánico de Madrid 70, 72–80.Google Scholar
Anderson, N.O., Ascher, P.D. and Widmer, R.E. (1992) Lethal equivalents and genetic load. Plant Breeding Reviews 10, 93–127.Google Scholar
Angeloni, F., Ouborg, N.J. and Leimu, R. (2011) Meta-analysis on the association of population size and life history with inbreeding depression in plants. Biological Conservation 144, 35–43.Google Scholar
APG-III (The Angiosperm Phylogeny Group) (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG-III. Botanical Journal of the Linnean Society 161, 105–121.Google Scholar
Arena, G., Symes, C.T. and Witkowski, E.T.F. (2013) The birds and the seeds: opportunistic avian nectarivores enhance reproduction in an endemic montane aloe. Plant Ecology 214, 35–47.CrossRefGoogle Scholar
Armbruster, P. and Reed, D.H. (2005) Inbreeding depression in benign and stressful environments. Heredity 95, 235–242.Google Scholar
Ashman, T.-L. (1992) The relative importance of inbreeding and maternal sex in determining progeny fitness in Sidalcea oregana ssp. spicata, a gynodioecious plant. Evolution 46, 1862–1874.CrossRefGoogle ScholarPubMed
Ashworth, L. and Galetto, L. (2001) Pollinators and reproductive success of the wild cucurbit Cucurbita maxima ssp. andreana (Cucurbitaceae). Plant Biology 3, 398–404.Google Scholar
Bachmann, U. and Hensen, I. (2007) Is declining Campanula glomerata threatened by genetic factors? Plant Species Biology 22, 1–10.CrossRefGoogle Scholar
Bailey, M.F. and McCauley, D.E. (2006) The effects of inbreeding, outbreeding and long-distance gene flow on survivorship in North American populations of Silene vulgaris . Journal of Ecology 94, 98–109.CrossRefGoogle Scholar
Barnes, B.V., Bingham, R.T. and Squillace, A.E. (1962) Selective fertilization in Pinus monticola Dougl. Silvae Genetica 11, 103–111.Google Scholar
Barnes, M.E. and Rust, R.W. (1994) Seed production in Gentiana newberryi (Gentianaceae). Great Basin Natualist 54, 351–358.Google Scholar
Barringer, B.C. and Geber, M.A. (2008) Mating system and ploidy influence levels of inbreeding depression in Clarkia (Onagraceae). Evolution 62, 1040–1051.Google Scholar
Baskin, C.C. and Baskin, J.M. (2014) Seeds: Ecology, biogeography, and evolution of dormancy and germination (2nd edition). San Diego, Elsevier/Academic Press.Google Scholar
Baskin, J.M. and Baskin, C.C. (1975) Growth of Ruellia humilis plants from embryos of chilled, GA3-treated and laboratory-stored seeds. Botanical Gazette 136, 299–305.Google Scholar
Baskin, J.M. and Baskin, C.C. (2000) Evolutionary considerations of claims for physical dormancy-break by microbial action and abrasion by soil particles. Seed Science Research 10, 409–413.CrossRefGoogle Scholar
Baskin, J.M. and Baskin, C.C. (2003) Classification, biogeography, and phylogenetic relationships of seed dormancy. pp. 518–544 in Smith, R.D.; Dickie, J.B.; Linington, S.H.; Pritchard, H.W.; Probert, R.J. (Eds) Seed conservation: Turning science into practice. Kew, The Royal Botanic Gardens.Google Scholar
Baskin, J.M. and Baskin, C.C. (2004) A classification system for seed dormancy. Seed Science Research 14, 1–16.CrossRefGoogle Scholar
Baskin, J.M., Baskin, C.C and Li, X. (2000) Taxonomy, anatomy and evolution of physical dormancy in seeds. Plant Species Biology 15, 139–152.Google Scholar
Becker, U., Reinhold, T. and Matthies, D. (2006) Effects of pollination distance on reproduction and offspring performance in Hypochoeris radicata: experiments with plants from three European regions. Biological Conservation 132, 109–118.CrossRefGoogle Scholar
Belaoussoff, S. and Shore, J.S. (1995) Floral correlates and fitness consequences of mating-system variation in Turnera ulmifolia . Evolution 49, 545–556.Google Scholar
Bermingham, L.H. and Brody, A.K. (2011) Pollen source affects female reproductive success and early offspring traits in the rare endemic plant Polemonium vanbruntiae (Polemoniaceae). Plant Species Biology 26, 244–253.CrossRefGoogle Scholar
Bernhardt, P. (1999) The rose's kiss: A natural history of flowers. Chicago, University of Chicago Press.Google Scholar
Biere, A. (1991a) Parental effects in Lychnis flos-cuculi. I: Seed size, germination and seedling performance in a controlled environment. Journal of Evolutionary Biology 3, 447–465.Google Scholar
Biere, A. (1991b) Parental effects on Lychnis flos-cuculi. II: Selection on time of emergence and seedling performance in the field. Journal of Evolutionary Biology 3, 467–486.Google Scholar
Billingham, M.R., Simões, T., Reusch, T.B.H. and Serrão, E.A. (2007) Genetic sub-structure and intermediate optimal outcrossing distance in the marine angiosperm Zostera marina . Marine Biology 152, 793–801.CrossRefGoogle Scholar
Black, M. (2009) Darwin and seeds. Seed Science Research 19, 193–199.Google Scholar
Bowles, M.L., Jacobs, K.A., Zettler, L.W. and Delaney, T.W. (2002) Crossing effects on seed viability and experimental germination of the federal threatened Platanthera leucophaea (Orchidaceae). Rhodora 104, 14–30.Google Scholar
Boyd, R.S., Teem, A. and Wall, M.A. (2011) Floral biology of an Alabama population of the federally endangered plant, Xyris tennesseensis Kral (Xyridaceae). Castanea 76, 255–265.Google Scholar
Brennan, A.C., Harris, S.A. and Hiscock, S.J. (2005) Modes and rates of selfing and associated inbreeding depression in the self-incompatible plant Senecio squalidus (Asteraceae): a successful colonizing species in the British Isles. New Phytologist 168, 475–486.Google Scholar
Broadhurst, L.M., Lowe, A., Coates, D.J., Cunningham, S.A., McDonald, M., Vesk, P.A. and Yates, C. (2008) Seed supply for broadscale restoration: maximizing evolutionary potential. Evolutionary Applications 1, 587–597.Google Scholar
Brock, J.H. and Peterson, S.J. (1975) Reproductive biology of Pulsatilla patens (Ranunculaceae). The American Midland Naturalist 94, 476–478.Google Scholar
Busch, J.W. (2005) Inbreeding depression in self-incompatible and self-compatible populations of Leavenworthia alabamica . Heredity 94, 159–165.CrossRefGoogle ScholarPubMed
Busch, J.W. (2006) Heterosis in an isolated, effectively small, and self-fertilizing population of the flowering plant Leavenworthia alabamica . Evolution 60, 184–191.Google Scholar
Buza, L., Young, A. and Thrall, P. (2000) Genetic erosion, inbreeding and reduced fitness in fragmented populations of the endangered tetraploid pea Swainsonia recta . Biological Conservation 93, 177–186.Google Scholar
Byers, D.L. (1998) Effect of cross proximity on progeny fitness in a rare and a common species of Eupatorium (Asteraceae). American Journal of Botany 85, 644–653.Google Scholar
Byers, D.L. and Waller, D.M. (1999) Do plant populations purge their genetic load? Effects of population size and mating history on inbreeding depression. Annual Review of Ecology and Systematics 30, 479–513.Google Scholar
Caño, L., Escarré, J., Blanco-Moreno, J.M. and Sans, F.X. (2008) Assessing the effect of inbreeding and long-distance gene flow on the invasive potential of Senecio pterophorus (Asteraceae). Australian Journal of Botany 56, 539–549.CrossRefGoogle Scholar
Carr, D.E. and Dudash, M.R. (1995) Inbreeding depression under a competitive regime in Mimulus guttatus: consequences for potential male and female function. Heredity 75, 437–445.Google Scholar
Carr, D.E. and Dudash, M.R. (1996) Inbreeding depression in two species of Mimulus (Scrophulariaceae) with contrasting mating systems. American Journal of Botany 83, 586–593.Google Scholar
Carr, D.E. and Eubanks, M.D. (2002) Inbreeding alters resistance to insect herbivory and host plant quality in Mimulus guttatus (Scrophulariaceae). Evolution 56, 22–30.Google ScholarPubMed
Carr, D.E., Fenster, C.B. and Dudash, M.R. (1997) The relationship between mating-system characters and inbreeding depression in Mimulus guttatus . Evolution 51, 363–372.Google Scholar
Carrió, E., Herreros, R., Bacchetta, G. and Güemes, J. (2008) Evidence of delayed selfing in Fumana juniperina (Cistaceae). International Journal of Plant Sciences 169, 761–767.Google Scholar
Carvallo, G.O. and Medel, R. (2010) Effects of herkogamy and inbreeding on the mating system of Mimulus luteus in the absence of pollinators. Evolutionary Ecology 24, 509–522.Google Scholar
Chang, S.-M. and Rausher, M.D. (1999) The role of inbreeding depression in maintaining the mixed mating system of the common morning glory, Ipomoea purpurea . Evolution 53, 1366–1376.Google Scholar
Charlesworth, D. and Charlesworth, B. (1987) Inbreeding depression and its evolutionary consequences. Annual Review of Ecology and Systematics 18, 237–268.Google Scholar
Charlesworth, D. and Willis, J.H. (2009) The genetics of inbreeding depression. Nature Reviews Genetics 10, 783–796.CrossRefGoogle ScholarPubMed
Charlesworth, D., Lyons, E.E. and Litchfield, L.B. (1994) Inbreeding depression in two highly inbreeding populations of Leavenworthia . Proceedings of the Royal Society of London B 258, 209–214.Google Scholar
Cheptou, P.-O. and Donohue, K. (2011) Environment-dependent inbreeding depression: its ecological and evolutionary significance. New Phytologist 189, 395–407.CrossRefGoogle ScholarPubMed
Cheptou, P.-O. and Schoen, D.J. (2003) Frequency-dependent inbreeding depression in Amsinckia . The American Naturalist 162, 744–753.Google Scholar
Cheptou, P.-O., Berger, A., Blanchard, A., Collin, C. and Escarré, J. (2000a) The effect of drought stress on inbreeding depression in four populations of the Mediterranean outcrossing plant Crepis sancta (Asteraceae). Heredity 85, 294–302.Google Scholar
Cheptou, P.-O., Imbert, E., Lepart, J. and Escarré, J. (2000b) Effects of competition on lifetime estimates of inbreeding depression in the outcrossing plant Crepis sancta (Asteraceae). Journal of Evolutionary Biology 13, 522–531.Google Scholar
Cheptou, P.-O., Lepart, J. and Escarré, J. (2001) Inbreeding depression under intraspecific competition in a highly outcrossing population of Crepis sancta (Asteraceae): evidence for frequency-dependent variation. American Journal of Botany 88, 1424–1429.Google Scholar
Clauss, M.J. and Venable, D.L. (2000) Seed germination in desert annuals: an empirical test of adaptive bet hedging. The American Naturalist 155, 168–186.CrossRefGoogle ScholarPubMed
Clements, R.K., Baskin, J.M. and Baskin, C.C. (1999) The comparative biology of the two closely-related species Penstemon tenuiflorus Pennell and P. hirsutus (L.) Willd. (Scrophulariaceae, section Graciles): II. Reproductive biology. Castanea 64, 299–309.Google Scholar
Cohen, D. (1966) Optimizing reproduction in a randomly varying environment. Journal of Theoretical Biology 12, 119–129.Google Scholar
Coles, J.F. and Fowler, D.P. (1976) Inbreeding in neighboring trees in two white spruce populations. Silvae Genetica 25, 29–34.Google Scholar
Colling, G., Reckinger, C. and Matthies, D. (2004) Effects of pollen quantity and quality on reproduction and offspring vigor in the rare plant Scorzonera humilis (Asteraceae). American Journal of Botany 91, 1774–1782.Google Scholar
Culley, T.M., Weller, S.G., Sakai, A.K. and Rankin, A.E. (1999) Inbreeding depression and selfing rates in a self-compatible, hermaphroditic species, Schiedea membranacea (Caryophyllaceae). American Journal of Botany 86, 980–987.Google Scholar
Daehler, C.C. (1999) Inbreeding depression in smooth cordgrass (Spartina alterniflora, Poaceae) invading San Francisco Bay. American Journal of Botany 86, 131–139.Google Scholar
Dahlgaard, J. and Warncke, E. (1995) Seed set and germination in crosses within and between two geographically isolated small populations of Saxifraga hirculus in Denmark. Nordic Journal of Botany 15, 337–341.Google Scholar
Darwin, C. (1876) The effects of cross and self fertilization in the vegetable kingdom. London, John Murray.Google Scholar
Davila, Y.C. and Wardle, G.M. (2002) Reproductive ecology of the Australian herb Trachymene incisa subsp. incisa (Apiaceae). Australian Journal of Botany 50, 619–626.Google Scholar
Davis, H.G., Taylor, C.M., Civille, J.C. and Strong, D.R. (2004) An Allee effect at the front of a plant invasion: Spartina in a Pacific estuary. Journal of Ecology 92, 321–327.Google Scholar
Del Castillo, R.F. (1998) Fitness consequences of maternal and nonmaternal components of inbreeding in the gynodioecious Phacelia dubia . Evolution 52, 44–60.Google Scholar
Delesalle, V.A. and Muenchow, G.E. (1992) Opportunities for selfing and inbreeding depression in Sagittaria congeners (Alismataceae) with contrasting sexual systems. Evolutionary Trends in Plants 6, 81–91.Google Scholar
Delph, L.F. (2004) Testing for sex differences in biparental inbreeding and its consequences in a gynodioecious species. American Journal of Botany 91, 45–51.Google Scholar
de Vere, N., Jongejans, E., Plowman, A. and Williams, E. (2009) Population size and habitat quality affect genetic diversity and fitness in the clonal herb Cirsium dissectum . Oecologia 159, 59–68.Google Scholar
Donohue, K. (1998) Effects of inbreeding on traits that influence dispersal and progeny density in Cakile edentula var. lacustris (Brassicaceae). American Journal of Botany 85, 661–668.Google Scholar
Dudash, M.R. (1990) Relative fitness of selfed and outcrossed progeny in a self-compatible, protandrous species, Sabatia angularis L. (Gentianaceae): a comparison in three environments. Evolution 44, 1129–1139.Google Scholar
Dudash, M.R. and Fenster, C.B. (2001) The role of breeding system and inbreeding depression in the maintenance of an outcrossing mating strategy in Silene virginica (Caryophyllaceae). American Journal of Botany 88, 1953–1959.Google Scholar
Dudash, M.R., Carr, D.E. and Fenster, C.B. (1997) Five generations of enforced selfing and outcrossing in Mimulus guttatus: inbreeding depression variation at the population and family level. Evolution 51, 54–65.Google Scholar
Durel, C.E., Bertin, P. and Kremer, A. (1996) Relationship between inbreeding depression and inbreeding coefficient in maritime pine (Pinus pinaster). Theoretical and Applied Genetics 92, 347–356.Google Scholar
Eckert, C.G. and Barrett, S.C.H. (1994) Inbreeding depression in partially self-fertilizing Decodon verticillatus (Lythraceae): population-genetic and experimental analyses. Evolution 48, 952–964.Google Scholar
Edmands, S. (2002) Does parental divergence predict reproductive compatibility? Trends in Ecology and Evolution 17, 520–527.Google Scholar
Edmands, S. (2007) Between a rock and a hard place: evaluating the relative risks of inbreeding and outbreeding for conservation and management. Molecular Ecology 16, 463–475.Google Scholar
El-Agamy, S.Z.A., Sherman, W.B. and Lyrene, P.M. (1981) Fruit set and seed number from self- and cross-pollinated highbush (4x) and rabbiteye (6x) blueberries. Journal of the American Society for Horticultural Science 106, 443–445.Google Scholar
Ellmer, M. and Andersson, S. (2004) Inbreeding depression in Nigella degenii (Ranunculaceae): fitness components compared with morphological and phenological characters. International Journal of Plant Sciences 165, 1055–1061.Google Scholar
Emery, S.N. and McCauley, D.E. (2002) Consequences of inbreeding for offspring fitness and gender in Silene vulgaris, a gynodioecious plant. Journal of Evolutionary Biology 15, 1057–1066.Google Scholar
Eppley, S.M. and Pannell, J.R. (2009) Inbreeding depression in dioecious populations of the plant Mercurialis annua: comparisons between outcrossed progeny and the progeny of self-fertilized feminized males. Heredity 102, 600–608.Google Scholar
Evans, A.S., Mitchell, R.J. and Cabin, R.J. (1996) Morphological side effects of using gibberellic acid to induce germination: consequences for the study of seed dormancy. American Journal of Botany 83, 543–549.Google Scholar
Fenster, C.B. (1991) Gene flow in Chamaecrista fasciculata (Leguminosae). II. Gene establishment. Evolution 45, 410–422.Google Scholar
Ferdy, J.-B., Loriot, S., Sandmeier, M., Lefranc, M. and Raquin, C. (2001) Inbreeding depression in a rare deceptive orchid. Canadian Journal of Botany 79, 1181–1188.Google Scholar
Ferrer, M.M., Good-Avila, S.V., Montaña, C., Domínguez, C.A. and Eguiarte, L.E. (2009) Effect of variation in self-incompatibility on pollen limitation and inbreeding depression in Flourensia cernua (Asteraceae) scrubs of contrasting density. Annals of Botany 103, 1077–1089.Google Scholar
Finch-Savage, W.E. and Leubner-Metzger, G. (2006) Seed dormancy and the control of germination. New Phytologist 171, 501–523.Google Scholar
Fischer, M. and Matthies, D. (1997) Mating structure and inbreeding and outbreeding depression in the rare plant Gentianella germanica (Gentianaceae). American Journal of Botany 84, 1685–1692.Google Scholar
Fischer, M. and Matthies, D. (1998) RAPD variation in relation to population size and plant fitness in the rare Gentianella germanica (Gentianaceae). American Journal of Botany 85, 811–819.Google Scholar
Fischer, M., Hock, M. and Paschke, M. (2003) Low genetic variation reduces cross-compatibility and offspring fitness in populations of a narrow endemic plant with a self-incompatibility system. Conservation Genetics 4, 325–336.Google Scholar
Fishman, L. (2001) Inbreeding depression in two populations of Arenaria uniflora (Caryophyllaceae) with contrasting mating systems. Heredity 86, 184–194.Google Scholar
Footitt, S., Douterelo-Soler, I., Clay, H. and Finch-Savage, W.E. (2011) Dormancy cycling in Arabidopsis seeds is controlled by seasonally distinct hormone-signaling pathways. Proceedings of the National Academy of Sciences of the United States of America 198, 20236–20241.Google Scholar
Forrest, C.N., Ottewell, K.M., Whelan, R.J. and Ayre, D.J. (2011) Tests for inbreeding and outbreeding depression and estimation of population differentiation in the bird-pollinated shrub Grevillea mucronulata . Annals of Botany 108, 185–195.Google Scholar
Fowler, D.P. (1965a) Effects of inbreeding in red pine, Pinus resinosa Ait. III. Factors affecting natural selfing. Silvae Genetica 14, 37–46.Google Scholar
Fowler, D.P. (1965b) Effects of inbreeding in red pine, Pinus resinosa Ait. IV. Comparison with other northeastern Pinus species. Silvae Genetica 14, 76–81.Google Scholar
Fowler, D.P. and Park, Y.S. (1983) Population studies of white spruce. I. Effects of self-pollination. Canadian Journal of Forest Research 13, 1133–1138.Google Scholar
Fox, C.W., Scheibly, K.L. and Reed, D.H. (2008) Experimental evolution of the genetic load and its implications for the genetic basis of inbreeding depression. Evolution 62, 2236–2249.Google Scholar
Fox, G.A., Evans, A.S. and Keefer, C.J. (1995) Phenotypic consequences of forcing germination: a general problem of intervention in experimental design. American Journal of Botany 82, 1264–1270.Google Scholar
Franklin, E.C. (1970) Survey of mutant forms and inbreeding depression in species of the family Pinaceae. USDA Forest Service Research Paper SE-61, 1–21.Google Scholar
Galetto, L., Bernardello, G., Isele, I.C., Vesprin, J., Speroni, G. and Berduc, A. (2000) Reproductive biology of Erythrina crista-galli (Fabaceae). Annals of the Missouri Botanical Garden 87, 127–145.Google Scholar
Galeuchet, D.J., Perret, C. and Fischer, M. (2005) Performance of Lychnis flos-cuculi from fragmented populations under experimental biotic interactions. Ecology 86, 1002–1011.Google Scholar
Galloway, L.F. and Etterson, J.R. (2005) Population differentiation and hybrid success in Campanula americana: geography and genome size. Journal of Evolutionary Biology 18, 81–89.Google Scholar
Galloway, L.F. and Etterson, J.R. (2007) Inbreeding depression in an autotetraploid herb: a three cohort field study. New Phytologist 173, 383–392.Google Scholar
Galloway, L.F., Etterson, J.R. and Hamrick, J.L. (2003) Outcrossing rate and inbreeding depression in the herbaceous autotetraploid, Campanula americana . Heredity 90, 308–315.CrossRefGoogle ScholarPubMed
Garcia-Fernández, A., Iriondo, J.M. and Escudero, A. (2012) Inbreeding at the edge: does inbreeding depression increase under more stressful conditions? Oikos 121, 1435–1445.Google Scholar
Gargano, D., Gullo, T. and Bernardo, L. (2009) Do inefficient selfing and inbreeding depression challenge the persistence of the rare Dianthus guliae Janka (Caryophyllaceae)? Influence of reproductive traits on a plant's proneness to extinction. Plant Species Biology 24, 69–76.Google Scholar
Gigord, L., Lavigne, C. and Shykoff, J.A. (1998) Partial self-incompatibility and inbreeding depression in a native tree species of La Réunion (Indian Ocean). Oecologia 117, 342–352.Google Scholar
Glaettli, M. and Goudet, J. (2006) Variation in the intensity of inbreeding depression among successive life-cycle stages and generations in gynodioecious Silene vulgaris (Caryophyllaceae). Journal of Evolutionary Biology 19, 1995–2005.Google Scholar
Godoy, A.R., Oviedo, V.R.S., Castro, M.M. and Cardoso, A.I.I. (2006) Efeito da endogamia na produção de sementes de pepino caipira. Bragantia 65, 569–573.Google Scholar
González-Varo, J.P. and Traveset, A. (2010) Among-individual variation in pollen limitation and inbreeding depression in a mixed-mating shrub. Annals of Botany 106, 999–1008.Google Scholar
Goodwillie, C. (2000) Inbreeding depression and mating systems in two species of Linanthus (Polemoniaceae). Heredity 84, 283–293.Google Scholar
Goodwillie, C. and Knight, M.C. (2006) Inbreeding depression and mixed mating in Leptosiphon jepsonii: a comparison of three populations. Annals of Botany 98, 351–360.Google Scholar
Graeber, K., Nakabayashi, K., Miatton, E., Leubner-Metzger, G. and Soppe, W.J.J. (2012) Molecular mechanisms of seed dormancy. Plant, Cell & Environment 35, 1769–1786.Google Scholar
Greimler, J. and Dobeš, Ch. (2000) High genetic diversity and differentiation in relict lowland populations of Gentianella austriaca (A. and J. Kern.) Holub (Gentianaceae). Plant Biology 2, 628–637.Google Scholar
Gremer, J.R. and Venable, D.L. (2014) Bet hedging in desert winter annual plants: optimal germination strategies in a variable environment. Ecology Letters 17, 380–387.Google Scholar
Grindeland, J.M. (2008) Inbreeding depression and outbreeding depression in Digitalis purpurea: optimal outcrossing distance in a tetraploid. Journal of Evolutionary Biology 21, 716–726.Google Scholar
Groom, M.J. (1998) Allee effects limit population viability of an annual plant. The American Naturalist 151, 487–496.Google Scholar
Groom, M.J. and Preuninger, T.E. (2000) Population type can influence the magnitude of inbreeding depression in Clarkia concinna (Onagraceae). Evolutionary Ecology 14, 155–180.Google Scholar
Guillaume, P. and Jacquemart, A.-L. (1999) Early-inbreeding depression in Vaccinium myrtillus and V. vitis-idaea . Protoplasma 208, 107–114.Google Scholar
Hamilton, M.B. and Mitchell-Olds, T. (1994) The mating system and relative performance of selfed and outcrossed progeny in Arabis fecunda (Brassicaceae). American Journal of Botany 81, 1252–1256.Google Scholar
Hardner, C.M. and Potts, B.M. (1995) Inbreeding depression and changes in variation after selfing in Eucalyptus globulus ssp. globulus . Silvae Genetica 44, 46–54.Google Scholar
Hauser, T.P. and Loeschcke, V. (1994) Inbreeding depression and mating-distance dependent offspring fitness in large and small populations of Lychnis flos-cuculi (Caryophyllaceae). Journal of Evolutionary Biology 7, 609–622.Google Scholar
Hauser, T.P. and Loeschcke, V. (1995) Inbreeding depression in Lychnis flos-cuculi (Caryophyllaceae): effects of different levels of inbreeding. Journal of Evolutionary Biology 8, 589–600.Google Scholar
Hauser, T.P. and Loeschcke, V. (1996) Drought stress and inbreeding depression in Lychnis flos-cuculi (Caryophyllaceae). Evolution 50, 1119–1126.Google Scholar
Hayes, C.N., Winsor, J.A. and Stephenson, A.G. (2005a) Multigenerational effects of inbreeding in Cucurbita pepo ssp. texana (Cucurbitaceae). Evolution 59, 276–286.Google Scholar
Hayes, C.N., Winsor, J.A. and Stephenson, A.G. (2005b) Environmental variation influences the magnitude of inbreeding depression in Cucurbita pepo ssp. texana (Cucurbitaceae). Journal of Evolutionary Biology 18, 147–155.Google Scholar
Heenan, P.B., Smissen, R.D. and Dawson, M.I. (2005) Self-incompatibility in the threatened shrub Olearia adenocarpa (Asteraceae). New Zealand Journal of Botany 43, 831–841.Google Scholar
Helenurm, K. and Schaal, B.A. (1996) Genetic and maternal effects on offspring fitness in Lupinus texensis (Fabaceae). American Journal of Botany 83, 1596–1608.Google Scholar
Heliyanto, B., Veneklaas, E.J., Lambers, H. and Krauss, S.L. (2005) Preferential outcrossing in Banksia ilicifolia (Proteaceae). Australian Journal of Botany 53, 163–170.Google Scholar
Hellman, E.W. and Moore, J.N. (1983) Effect of genetic relationship to pollinizer on fruit, seed, and seedling parameters in highbush and rabbiteye blueberries. Journal of the American Society for Horticultural Science 108, 401–405.CrossRefGoogle Scholar
Hensen, I. and Oberprieler, C. (2005) Effects of population size on genetic diversity and seed production in the rare Dictamnus albus (Rutaceae) in central Germany. Conservation Genetics 6, 63–73.Google Scholar
Hensen, I. and Wesche, K. (2006) Relationships between population size, genetic diversity and fitness components in the rare plant Dictamnus albus in central Germany. Biodiversity Conservation 15, 2249–2261.Google Scholar
Hereford, J. (2009) Postmating/prezygotic isolation, heterosis, and outbreeding depression in crosses within and between populations of Diodia teres (Rubiaceae) Walt. International Journal of Plant Sciences 170, 301–310.Google Scholar
Herrera, J. (1991) The reproductive biology of a riparian Mediterranean shrub, Nerium oleander L. (Apocynaceae). Botanical Journal of the Linnean Society 106, 147–172.Google Scholar
Heschel, M.S., Hausmann, N. and Schmitt, J. (2005) Testing for stress-dependent inbreeding depression in Impatiens capensis (Balsaminaceae). American Journal of Botany 92, 1322–1329.Google Scholar
Heywood, J.S. (1993) Biparental inbreeding depression in the self-incompatible annual plant Gaillardia pulchella (Asteraceae). American Journal of Botany 80, 545–550.Google Scholar
Himes, S.L. and Wyatt, R. (2005) Costs and benefits of self-fertility in Asclepias exaltata (Apocynaceae). Journal of the Torrey Botanical Society 132, 24–32.Google Scholar
Hirao, A.S. (2010) Kinship between parents reduces offspring fitness in a natural population of Rhododendron brachycarpum . Annals of Botany 105, 637–646.Google Scholar
Hirayama, K., Ishida, K., Setsuko, S. and Tomaru, N. (2007) Reduced seed production, inbreeding, and pollen shortage in a small population of a threatened tree, Magnolia stellata . Biological Conservation 136, 315–323.CrossRefGoogle Scholar
Holtsford, T.P. (1996) Variation in inbreeding depression among families and populations of Clarkia tembloriensis (Onagraceae). Heredity 76, 83–91.Google Scholar
Holtsford, T.P. and Ellstrand, N.C. (1990) Inbreeding effects in Clarkia tembloriensis (Onagraceae) populations with different natural outcrossing rates. Evolution 44, 2031–2046.Google Scholar
Hooftman, D.A., van Kleunen, M. and Diemer, M. (2003) Effects of habitat fragmentation on the fitness of two common wetland species, Carex davalliana and Succisa pratensis . Oecologica 134, 350–359.Google Scholar
Hossaert-McKey, M. and Bronstein, J.L. (2001) Self-pollination and its costs in a monoecious fig (Ficus aurea, Moraceae) in a highly seasonal subtropical environment. American Journal of Botany 88, 685–692.Google Scholar
Hull-Sanders, H.M., Eubanks, M.D. and Carr, D.E. (2005) Inbreeding depression and selfing rate of Ipomoea hederacea var. integriuscula (Convolvulaceae). American Journal of Botany 92, 1871–1877.Google Scholar
Husband, B.C. and Gurney, J.E. (1998) Offspring fitness and parental effects as a function of inbreeding in Epilobium angustifolium (Onagraceae). Heredity 80, 173–179.Google Scholar
Husband, B.C. and Schemske, D.W. (1995) Magnitude and timing of inbreeding depression in a diploid population of Epilobium angustifolium (Onagraceae). Heredity 75, 206–215.Google Scholar
Husband, B.C. and Schemske, D.W. (1996) Evolution of the magnitude and timing of inbreeding depression in plants. Evolution 50, 54–70.Google Scholar
Husband, B.C. and Schemske, D.W. (1997) The effect of inbreeding in diploid and tetraploid populations of Epilobium angustifolium (Onagraceae): implications for the genetic basis of inbreeding depression. Evolution 51, 737–746.Google Scholar
Ishida, K. (2006) Maintenance of inbreeding depression in a highly self-fertilizing tree, Magnolia obovata Thunb. Evolutionary Ecology 20, 173–191.Google Scholar
Jacobi, C.M., do Carmo, R.M. and Oliveira, R.S. (2000) The reproductive biology of two species of Diplusodon Pohl (Lythraceae) from Serra do Cipo, southeastern Brazil. Plant Biology 2, 670–676.CrossRefGoogle Scholar
Jacobi, C.M., Ramalho, M. and Silva, M. (2005) Pollination biology of the exotic rattleweed Crotalaria retusa L. (Fabaceae) in NE Brazil. Biotropica 37, 357–363.Google Scholar
Jing, X.-M. and Zheng, G.-H. (1999) The characteristics in seed germination and dormancy of four wild species of tree peonies and their bearing on endangerment. Acta Phytophysiologica Sinica 25, 214–221.Google Scholar
Jóhannsson, M.H., Gates, M.J. and Stephenson, A.G. (1998) Inbreeding depression affects pollen performance in Cucurbita texana . Journal of Evolutionary Biology 11, 579–588.Google Scholar
Johnson, G.R., Sorensen, F.C., St. Clair, J.B. and Cronn, R.C. (2004) Pacific northwest forest tree seed zones: a template for native plants? Native Plants Journal 5, 131–140.Google Scholar
Johnson, T.R., Stewart, S.L., Kauth, P., Kane, M.E. and Philman, N. (2009) Confronting assumptions about spontaneous autogamy in populations of Eulophia alta (Orchidaceae) in south Florida: assessing the effect of pollination treatments on seed formation, seed germination and seedling development. Botanical Journal of the Linnean Society 161, 78–88.Google Scholar
Johnston, M.O. (1992) Effects of cross and self-fertilization on progeny fitness in Lobelia cardinalis and L. siphilitica . Evolution 46, 688–702.Google Scholar
Johnston, M.O. and Schoen, D.J. (1996) Correlated evolution of self-fertilization and inbreeding depression: an experimental study of nine populations of Amsinckia (Boraginaceae). Evolution 50, 1478–1491.Google Scholar
Jolivet, C. and Bernasconi, G. (2007) Within/between population crosses reveal genetic basis for siring success in Silene latifolia (Caryophyllaceae). Journal of Evolutionary Biology 20, 1361–1374.Google Scholar
Juillet, N., Dunand-Martin, S. and Gigord, L.D.B. (2007) Evidence for inbreeding depression in the food-deceptive colour-dimorphic orchid Dactylorhiza sambucina (L.) Soò. Plant Biology 9, 147–151.Google Scholar
Kahman, S. and Poschold, P. (2000) Population size, plant performance, and genetic variation in the rare plant Arnica montana L. in Rhön, Germany. Basic and Applied Ecology 1, 43–51.Google Scholar
Kalisz, S. (1989) Fitness consequences of mating system, seed weight, and emergence date in a winter annual, Collinsia verna . Evolution 43, 1263–1272.Google Scholar
Kärkkäinen, K., Koski, V. and Savolainen, O. (1996) Geographical variation in the inbreeding depression of Scots pine. Evolution 50, 111–119.Google Scholar
Kärkkäinen, K., Kuittinen, H., van Treuren, R., Vogl, C., Oikarinen, S. and Savolainen, O. (1999) Genetic basis of inbreeding depression in Arabis petraea . Evolution 53, 1354–1365.Google Scholar
Karron, J.D. (1989) Breeding systems and levels of inbreeding depression in geographically restricted and widespread species of Astragalus (Fabaceae). American Journal of Botany 76, 331–340.Google Scholar
Keller, L.F. and Waller, D.M. (2002) Inbreeding effects in wild populations. Trends in Ecology and Evolution 17, 230–241.Google Scholar
Kennedy, B.F. and Elle, E. (2008) The inbreeding depression cost of selfing: importance of flower size and population size in Collinsia parviflora (Veronicaceae). American Journal of Botany 95, 1596–1605.Google Scholar
Kephart, S.R., Brown, E. and Hall, J. (1999) Inbreeding depression and partial selfing: evolutionary implications of mixed-mating in a coastal endemic, Silene douglasii var. oraria (Caryophyllaceae). Heredity 82, 543–554.Google Scholar
Kittleson, P.M. and Maron, J.L. (2000) Outcrossing rate and inbreeding depression in the perennial yellow bush lupine, Lupinus arboreus (Fabaceae). American Journal of Botany 87, 652–660.Google Scholar
Klips, R.A. and Snow, A.A. (1997) Delayed autonomous self-pollination in Hibiscus laevis (Malvaceae). American Journal of Botany 84, 48–53.Google Scholar
Kochánková, J. and Mandák, B. (2009) How do population genetic parameters affect germination of the heterocarpic species Atriplex tatarica (Amaranthaceae)? Annals of Botany 103, 1303–1313.Google Scholar
Koelewijn, H.P. (1998) Effects of different levels of inbreeding on progeny fitness in Plantago coronopus . Evolution 52, 692–702.Google Scholar
Koelewijn, H.P. (2004) Sibling competition, size variation and frequency-dependent outcrossing advantage in Plantago coronopus . Evolutionary Ecology 18, 51–74.Google Scholar
Koelewijn, H.P. and van Damme, J.M.M. (2005) Effects of seed size, inbreeding and maternal sex on offspring fitness in gynodioecious Plantago coronopus . Journal of Ecology 93, 373–383.Google Scholar
Koelewijn, H.P., Koski, V. and Savolainen, O. (1999) Magnitude and timing of inbreeding depression in Scots pine (Pinus sylvestris L.). Evolution 53, 758–768.Google Scholar
Kuser, J. (1983) Inbreeding depression in Metasequoia . Journal of the Arnold Arboretum 64, 475–481.Google Scholar
Lammi, A., Siikamäki, P. and Mustajärvi, K. (1999) Genetic diversity, population size, and fitness in central and peripheral populations of a rare plant Lychnis viscaria . Conservation Biology 13, 1069–1078.Google Scholar
Lande, R. and Schemske, D.W. (1985) The evolution of self-fertilization and inbreeding depression in plants. I. Genetic models. Evolution 39, 24–40.Google Scholar
Latta, R. and Ritland, K. (1994) The relationship between inbreeding depression and prior inbreeding among populations of four Mimulus taxa. Evolution 48, 806–817.CrossRefGoogle ScholarPubMed
Le Cadre, S., Tully, T., Mazer, S.J., Ferdy, J.-B., Moret, J. and Machon, N. (2008) Allee effects within small populations of Aconitum napellus ssp. lusitanicum, a protected species in northern France. New Phytologist 179, 1171–1182.Google Scholar
Le Corff, J. (1996) Establishment of chasmogamous and cleistogamous seedlings of an ant-dispersed understory herb, Calathea micans (Marantaceae). American Journal of Botany 83, 155–161.Google Scholar
Leimu, R. and Mutikainen, P. (2005) Population history, mating system, and fitness variation in a perennial herb with a fragmented distribution. Conservation Biology 19, 349–356.CrossRefGoogle Scholar
Levin, D.A. and Bulinska-Radomska, Z. (1988) Effects of hybridization and inbreeding on fitness in Phlox . American Journal of Botany 75, 1632–1639.Google Scholar
Lhamo, N., Ramsey, M. and Vaughton, G. (2006) Density- and frequency-dependent inbreeding depression in the Australian annual Hibiscus trionum var vesicarius . Evolutionary Ecology Research 8, 717–730.Google Scholar
Libby, W.J., McCutchan, B.G. and Millar, C.I. (1981) Inbreeding depression in selfs of redwood. Silvae Genetica 30, 15–25.Google Scholar
Lienert, J. and Fischer, M. (2004) Experimental inbreeding reduces seed production and germination independent of fragmentation of populations of Swertia perennis . Basic and Applied Ecology 5, 43–52.Google Scholar
Liu, H. and Koptur, S. (2003) Breeding system and pollination of a narrowly endemic herb of the Lower Florida Keys: impacts on the urban-wildland interface. American Journal of Botany 90, 1180–1187.Google Scholar
Liu, H. and Spira, T.P. (2001) Influence of seed age and inbreeding on germination and seedling growth in Hibiscus moscheutos (Malvaceae). Journal of the Torrey Botanical Society 128, 16–24.Google Scholar
Lofflin, D.L. and Kephart, S.R. (2005) Outbreeding, seedling establishment, and maladaptation in natural and reintroduced populations of rare and common Silene douglasii (Caryophyllaceae). American Journal of Botany 92, 1691–1700.Google Scholar
Luijten, S.H., Oostermeijer, J.G., van Leeuwen, N.C. and den Nijs, H.C.M. (1996) Reproductive success and clonal genetic structure of the rare Arnica montana (Compositae) in The Netherlands. Plant Systematics and Evolution 201, 15–30.Google Scholar
Luijten, S.H., Dierick, S., Oostermeijer, J.G.B., Raijmann, L.E.L. and den Nijs, H.C.M. (2000) Population size, genetic variation and reproductive success in the rapidly declining self-incompatible Arnica montana in The Netherlands. Conservation Biology 14, 1776–1787.Google Scholar
Luijten, S.H., Kéry, M., Oostermeijer, J.G. and den Nijs, H.C.M. (2002) Demographic consequences of inbreeding and outbreeding in Arnica montana: a field experiment. Journal of Ecology 90, 593–603.Google Scholar
Mabberley, D.J. (2008) Mabberley's plant-book. A portable dictionary of plants, their classification and uses (3rd edition). Cambridge, Cambridge University Press.Google Scholar
Maki, M. (1993) Outcrossing and fecundity advantage of females in gynodioecious Chionographis japonica var kurohimensis (Liliaceae). American Journal of Botany 80, 629–634.Google Scholar
Mandák, B. and Pyšek, P. (1999) Effects of plant density and nutrient levels on fruit polymorphism in Atriplex sagittata . Oecologia 119, 63–72.Google Scholar
Mandujano, M.C., Montana, C. and Eguiarte, L.E. (1996) Reproductive ecology and inbreeding depression in Opuntia rastrera (Cactaceae) in the Chihuahuan Desert: why are sexually derived recruitments so rare? American Journal of Botany 83, 63–70.Google Scholar
Marshall, M. and Ganders, F.R. (2001) Sex-biased seed predation and the maintenance of females in a gynodioecious plant. American Journal of Botany 88, 1437–1443.Google Scholar
Mathiasen, P., Rovere, A.E. and Premoli, A.C. (2007) Genetic structure and early effects of inbreeding in fragmented temperate forests of a self-incompatible tree, Embothrium coccineum . Conservation Biology 21, 232–240.Google Scholar
Mayer, S.S., Charlesworth, D. and Meyers, B. (1996) Inbreeding depression in four populations of Collinsia heterophylla Nutt. (Scrophulariaceae). Evolution 50, 879–891.Google Scholar
McCall, C., Waller, D.M. and Mitchell-Old, T. (1994) Effects of serial inbreeding on fitness components in Impatiens capensis . Evolution 48, 818–827.Google Scholar
Meier, C. and Holderegger, R. (1998) Breeding system, germination, and phenotypic differences among populations of Saxifraga aizoides (Saxifragaceae) at the periphery of its alpine distribution. Nordic Journal of Botany 18, 681–688.Google Scholar
Melville, A.H., Galletta, G.J. and Draper, A.D. (1980) Seed germination and early seedling vigor in progenies of inbred strawberry selections. HortScience 15, 749–750.Google Scholar
Mena-Ali, J.I., Keser, L.H. and Stephenson, A.G. (2008) Inbreeding depression in Solanum carolinense (Solanaceae), a species with a plastic self-incompatibility response. BMC Evolutionary Biology 8, 10. doi:10.1186/1471-2148-8-10.Google Scholar
Menges, E.S. (1991) Seed germination percentage increases with population size in a fragmented prairie species. Conservation Biology 5, 158–164.Google Scholar
Mergen, F., Burley, J. and Furnival, G.M. (1965) Embryo and seedling development in Picea glauca (Moench) Voss after self-, cross- and wind-pollination. Silvae Genetica 14, 188–194.Google Scholar
Michaels, H.J., Shi, X.J. and Mitchell, R.J. (2008) Effects of population size on performance and inbreeding depression in Lupinus perennis . Oecologia 154, 651–661.Google Scholar
Molina-Freaner, F. and Jain, S.K. (1993) Inbreeding effects in a gynodioecious population of the colonizing species Trifolium hirtum All. Evolution 47, 1472–1479.Google Scholar
Molina-Freaner, F., Cervantes-Salas, M., Morales-Romero, D., Buchmann, S. and Fleming, T.H. (2003) Does the pollinator abundance hypothesis explain geographic variation in the breeding system of Pachycereus pringlei? International Journal of Plant Sciences 164, 383–393.Google Scholar
Montalvo, A.M. (1994) Inbreeding depression and maternal effects in Aquilegia caerulea, a partially selfing plant. Ecology 75, 2395–2409.Google Scholar
Mooney, E.H. and McGraw, J.B. (2007) Effects of self-pollination and outcrossing with cultivated plants in small natural populations of American ginseng, Panax quinquefolius (Araliaceae). American Journal of Botany 94, 1677–1687.Google Scholar
Moran-Palma, P. and Snow, A.A. (1997) The effect of interplant distance on mating success in federally-threatened, self-incompatible Hymenoxys herbacea = H. acaulis var glabra (Asteraceae). American Journal of Botany 84, 233–238.Google Scholar
Moreira, B. and Pausas, J.G. (2012) Tanned or burned: the role of fire in shaping physical seed dormancy. PLoS ONE 7(12), e51523.Google Scholar
Morgante, M., Vendramin, G.G., Rossi, P. and Olivieri, A.M. (1993) Selection against inbreds in early life-cycle phases in Pinus leucodermis Ant. Heredity 70, 622–627.Google Scholar
Mullarkey, A.A., Byers, D.L. and Anderson, R.C. (2013) Inbreeding depression and partitioning of genetic load in the invasive biennial Alliaria petiolata (Brassicaceae). American Journal of Botany 100, 509–518.Google Scholar
Murza, G.L. and Davis, A.R. (2005) Flowering phenology and reproductive biology of Drosera anglica (Droseraceae). Botanical Journal of the Linnean Society 147, 417–426.Google Scholar
Mustajärvi, K., Siikamäki, P. and Åkerberg, A. (2005) Inbreeding depression in perennial Lychnis viscaria (Caryophyllaceae): effects of population mating history and nutrient availability. American Journal of Botany 92, 1853–1861.Google Scholar
Mutikainen, P. and Delph, L.F. (1998) Inbreeding depression in gynodioecious Lobelia siphilitica: among-family differences override between-morph differences. Evolution 52, 1572–1582.Google Scholar
Naito, Y., Konuma, A., Iwata, H., Suyama, Y., Seiwa, K., Okuda, T., Lee, S.L., Muhammad, N. and Tsumura, Y. (2005) Selfing and inbreeding depression in seeds and seedlings of Neobalanocarpus heimii (Dipterocarpaceae). Journal of Plant Research 118, 423–430.Google Scholar
Naito, Y., Kanzaki, M., Iwata, H., Obayashi, K., Lee, S.L., Muhammad, N., Okuda, T. and Tsumura, Y. (2008) Density-dependent selfing and its effects on seed performance in a tropical canopy tree species, Shorea acuminata (Dipterocarpaceae). Forest Ecology and Management 256, 375–383.Google Scholar
Nason, J.D. and Ellstrand, N.C. (1995) Lifetime estimates of biparental inbreeding depression in the self-incompatible annual plant Raphanus sativus . Evolution 49, 307–316.Google Scholar
Navarro, L. and Guitián, J. (2002) The role of floral biology and breeding system on the reproductive success of the narrow endemic Petrocoptis viscosa Rothm. (Caryophyllaceae). Biological Conservation 103, 125–132.Google Scholar
Newman, D. and Pilson, D. (1997) Increased probability of extinction due to decreased genetic effective population size: experimental populations of Clarkia pulchella . Evolution 51, 354–362.Google Scholar
Newport, M.E.A. (1989) A test for proximity-dependent outcrossing in the alpine skypilot, Polemoniun viscosum . Evolution 43, 1110–1113.Google Scholar
Nielsen, L.R., Siegismund, H.R. and Hansen, T. (2007) Inbreeding depression in the partially self-incompatible endemic plant species Scalesia affinis (Asteraceae) from Galápagos islands. Evolutionary Ecology 21, 1–12.Google Scholar
Norman, J.K., Sakai, A.K., Weller, S.G. and Dawson, T.E. (1995) Inbreeding depression in morphological and physiological traits of Schiedea lydgatei (Caryophyllaceae) in two environments. Evolution 49, 297–306.Google Scholar
Nuñez-Farfán, J., Cabrales-Vargas, R.A. and Dirzo, R. (1996) Mating system consequences on resistance to herbivory and life history traits in Datura stramonium . American Journal of Botany 83, 1041–1049.Google Scholar
Oliviera, I., Swan, M. and Gouyon, P.-H. (1983) Reproductive system and colonizing strategy of two species of Carduus (Compositae). Oecologia 60, 114–117.Google Scholar
O'Neil, P. (1994) Genetic incompatibility and offspring quality in the tristylous plant Lythrum salicaria (Lythraceae). American Journal of Botany 81, 76–84.Google Scholar
Oostermeijer, J.G.B., van Eijek, M.W. and den Nijs, J.C.M. (1994) Offspring fitness in relation to population size and genetic variation in the rare perennial plant species Gentiana pneumonanthe (Gentianaceae). Oecologia 97, 289–296.Google Scholar
Oostermeijer, J.G.B., Altenburg, R.G.M. and den Nijs, J.C.M. (1995) Effects of outcrossing distance and selfing on fitness components in the rare Gentiana pneumonanthe (Gentianaceae). Acta Botanica Neerlandica 44, 257–268.Google Scholar
Oriani, A., Sano, P.T. and Scatena, V.L. (2009) Pollination biology of Syngonanthus elegans (Eriocaulaceae - Poales). Australian Journal of Botany 57, 94–105.Google Scholar
Ortiz-Barney, E. and Ackerman, J.D. (1999) The cost of selfing in Encyclia cochleata (Orchidaceae). Plant Systematics and Evolution 219, 55–64.Google Scholar
Ouborg, N.J. and van Treuren, R. (1994) The significance of genetic erosion in the process of extinction. IV. Inbreeding load and heterosis in relation to population size in the mint Salvia pratensis . Evolution 48, 996–1008.Google Scholar
Ouborg, N.J., Biere, A. and Mudde, C.L. (2000) Inbreeding effects on resistance and transmission-related traits in the Silene-Microbotryum pathosystem. Ecology 81, 520–531.Google Scholar
Owen, K., Vaughton, G. and Ramsey, M. (2007) Facilitated autogamy and costs of selfing in the perennial herb Bulbine bulbosa (Asphodelaceae). International Journal of Plant Sciences 168, 579–585.Google Scholar
Owens, S.J. and Miller, R. (2009) Cross- and self-fertilization of plants – Darwin's experiments and what we know now. Botanical Journal of the Linnean Society 161, 357–395.Google Scholar
Ozimec, B. and Husband, B.C. (2011) Effect of recurrent selfing on inbreeding depression and mating system evolution in an autopolyploid plant. Evolution 65, 2038–2049.Google Scholar
Parker, I.M., Nakamura, R.R. and Schemske, D.W. (1995) Reproductive allocation and the fitness consequences of selfing in two sympatric species of Epilobium (Onagraceae) with contrasting mating systems. American Journal of Botany 82, 1007–1016.Google Scholar
Parker, M.A. (1992) Outbreeding depression in a selfing annual. Evolution 46, 837–841.Google Scholar
Peakall, R. and Beattie, A.J. (1996) Ecological and genetic consequences of pollination by sexual deception in the orchid Caladenia tentactulata . Evolution 50, 2207–2220.Google Scholar
Petit, S., Jusaitis, M. and Bickerton, D. (2009) Effect of pollen load, self-pollination and plant size on seeds and germination in the endangered pink-lipped spider orchid, Caladenia behrii . Australian Journal of Botany 57, 307–314.Google Scholar
Pettersson, M.W. (1992) Advantages of being a specialist female in nodioecious Silene vulgaris s.l. (Caryophyllaceae). American Journal of Botany 79, 1389–1395.Google Scholar
Picó, F.X. and Koubek, T. (2003) Inbreeding effects on fitness traits in the heterocarpic herb Leontodon autumnalis L. (Asteraceae). Acta Oecologica 24, 289–294.Google Scholar
Picó, F.X., Ouborg, N.J. and van Groenendael, J.M. (2003) Fitness traits and dispersal ability in the herb Tragopogon pratensis (Asteraceae): decoupling the role of inbreeding depression and maternal effects. Plant Biology 5, 522–530.Google Scholar
Picó, F.X., Ouborg, N.J. and van Groenendael, J.M. (2004a) Influence of selfing and maternal effects on life-cycle traits and dispersal ability in the herb Hypochaeris radicata (Asteraceae). Botanical Journal of the Linnean Society 146, 163–170.Google Scholar
Picó, F.X., Ouborg, N.J. and van Groenendael, J.M. (2004b) Evaluation of the extent of among-family variation in inbreeding depression in the perennial herb Scabiosa columbaria (Dipsacaceae). American Journal of Botany 91, 1183–1189.Google Scholar
Piesch, R.F. and Stettler, R.F. (1971) The detection of good selfers for haploid induction in douglar-fir. Silvae Genetica 20, 144–148.Google Scholar
Platenkamp, G.A.J. and Shaw, R.G. (1993) Environmental and genetic maternal effects on seed characters in Nemophila menziesii . Evolution 47, 540–555.Google Scholar
Pluess, A.R. and Stöcklin, J. (2004) Genetic diversity and fitness in Scabiosa columbaria in the Swiss Jura in relation to population size. Conservation Genetics 5, 145–156.Google Scholar
Pounders, C., Reed, S. and Pooler, M. (2006) Comparison of self- and cross-pollination on pollen tube growth, seed development, and germination in crapemyrtle. HortScience 41, 575–578.Google Scholar
Price, M.V. and Waser, N.M. (1979) Pollen dispersal and optimal outcrossing in Delphinium nelsonii . Nature 277, 294–297.Google Scholar
Prill, N., Bullock, J.M., van Dam, N.M. and Leimu, R. (2014) Loss of heterosis and family-dependent inbreeding depression in plant performance against multiple herbivores under drought stress. Journal of Ecology 102, 1497–1505.Google Scholar
Pujol, B. and McKey, D. (2006) Size asymmetry in intraspecific competition and the density-dependence of inbreeding depression in a natural plant population: a case study in cassava (Manihot esculenta Crantz, Euphorbiaceae). Journal of Evolutionary Biology 19, 85–96.Google Scholar
Puterbaugh, M.N., Wied, A. and Galen, C. (1997) The functional ecology of gynodioecy in Eritrichium aretioides (Boraginaceae), the alpine forget-me-not. American Journal of Botany 84, 393–400.Google Scholar
Quilichini, A., Debussche, M. and Thompson, J.D. (2001) Evidence for local outbreeding depression in the Mediterranean island endemic Anchusa crispa Viv. (Boraginaceae). Heredity 87, 190–197.Google Scholar
Raabová, J., Münzbergová, Z. and Fischer, M. (2009) Consequences of near and far between-population crosses for offspring fitness in a rare herb. Plant Biology 11, 829–836.Google Scholar
Ramsey, M. and Vaughton, G. (1996) Inbreeding depression and pollinator availability in a partially self-fertile perennial herb Blandfordia grandiflora (Liliaceae). Oikos 76, 465–474.Google Scholar
Ramsey, M. and Vaughton, G. (1998) Effect of environment on the magnitude of inbreeding depression in seed germination in a partially self-fertile perennial herb (Blandfordia grandiflora, Liliaceae). International Journal of Plant Sciences 159, 98–104.Google Scholar
Ramsey, M., Seed, L. and Vaughton, G. (2003) Delayed selfing and low levels of inbreeding depression in Hibiscus trionum (Malvaceae). Australian Journal of Botany 51, 275–281.Google Scholar
Rao, G.-Y., Widen, B. and Andersson, S. (2002) Patterns of inbreeding depression in a population of Brassica cretica (Brassicaceae): evidence from family-level analyses. Biological Journal of the Linnean Society 76, 317–325.Google Scholar
Ricardo, C.T., Corrado, C.A., Mandujano, M.C. and Molina-Freaner, F. (2006) Reproductive consequences of clonal growth in Stenocereus eruca, a rare clonal cactus of the Sonoran Desert. Evolutionary Ecology 20, 131–142.Google Scholar
Richards, C.M. (2000a) Inbreeding depression and genetic rescue in a plant metapopulation. The American Naturalist 155, 383–394.Google Scholar
Richards, C.M. (2000b) Genetic and demographic influences on population persistence: gene flow and genetic rescue in Silene alba . pp. 271–291 in Young, A.G.; Clarke, G.M. (Eds) Genetics, demography and viability of fragmented populations. Cambridge, Cambridge University Press.Google Scholar
Richter, K.S. and Weis, A.E. (1998) Inbreeding and outcrossing in Yucca whipplei: consequences for the reproductive success of plant and pollinator. Ecology Letters 1, 21–24.Google Scholar
Riley, R. (1956) The influence of the breeding system on the genecology of Thlaspi alpestre L. New Phytologist 55, 319–330.Google Scholar
Robertson, A.W., Kelly, D. and Ladley, J.J. (2011) Futile selfing in the trees Fuchsia excorticata (Onagraceae) and Sophora microphylla (Fabaceae): inbreeding depression over 11 years. International Journal of Plant Sciences 172, 191–198.Google Scholar
Rosquist, G. (2001) Reproductive biology in diploid Anthericum ramosum and tetraploid A. liliago (Anthericaceae). Oikos 92, 143–152.Google Scholar
Routley, M.B., Mavraganis, K. and Eckert, C.G. (1999) Effect of population size on the mating system in a self-compatible, autogamous plant, Aquilegia canadensis (Ranunculaceae). Heredity 82, 518–528.Google Scholar
Ruan, C.-J., Li, H. and Mopper, S. (2009) Kosteletzkya virginica displays mixed mating in response to the pollinator environment despite strong inbreeding depression. Plant Ecology 203, 183–193.Google Scholar
Ruckelshaus, M.H. (1995) Estimates of outcrossing rates and of inbreeding depression in a population of the marine angiosperm Zostera marina . Marine Biology 123, 583–593.Google Scholar
Schaal, B.A. (1984) Life-history variation, natural selection, and maternal effects in plant populations. pp. 188–206 in Dirzo, R.; Sarukhan, J. (Eds) Perspectives on plant population ecology. Sunderland, MA, Sinauer Associates.Google Scholar
Schemske, D.W. (1983) Breeding system and habitat effects on fitness components in three neotropical Costus (Zingiberaceae). Evolution 37, 523–539.Google Scholar
Schemske, D.W. and Pautler, L.P. (1984) The effects of pollen composition on fitness components in a neotropical herb. Oecologia 62, 31–36.Google Scholar
Schlichting, C.D. and Devlin, B. (1992) Pollen and ovule sources affect seed production of Lobelia cardinalis (Lobeliaceae). American Journal of Botany 79, 891–898.Google Scholar
Schmidt, K. and Jensen, K. (2000) Genetic structure and AFLP variation in remnant populations in the rare plant Pedicularis palustris (Scrophulariaceae) and its relation to population size and reproductive components. American Journal of Botany 87, 678–689.Google Scholar
Schoen, D.J. (1983) Relative fitnesses of selfed and outcrossed progeny in Gilia achilleifolia (Polemoniaceae). Evolution 37, 292–301.Google Scholar
Seavey, S.R. and Carter, S.K. (1994) Self-sterility in Epilobium obcordatum (Onagraceae). American Journal of Botany 81, 331–338.Google Scholar
Seed, L., Vaughton, G. and Ramsey, M. (2006) Delayed autonomous selfing and inbreeding depression in the Australian annual Hibiscus trionum var. vesicarius (Malvaceae). Australian Journal of Botany 54, 27–34.Google Scholar
Seltmann, P., Cocucci, A., Renison, D., Cierjacks, A. and Hensen, I. (2009) Mating system, outcrossing distance effects and pollen availability in the wind-pollinated treeline species Polylepis australis BITT (Rosaceae). Basic and Applied Ecology 10, 52–60.Google Scholar
Sheridan, P.M. and Karowe, D.N. (2000) Inbreeding, outbreeding, and heterosis in the yellow pitcher plant, Sarracenia flava (Sarraceniaceae), in Virginia. American Journal of Botany 87, 1628–1633.Google Scholar
Shi, X.J., Michaels, H.J. and Mitchell, R.J. (2005) Effects of self-pollination and maternal resources on reproduction and offspring performance in the wild lupine, Lupinus perennis (Fabaceae). Sexual Plant Reproduction 18, 55–64.Google Scholar
Simons, A.M. (2011) Modes of response to environmental change and the elusive empirical evidence for bet hedging. Proceedings of the Royal Society B 279, 1601–1609.Google Scholar
Smithson, A. (2006) Pollinator limitation and inbreeding depression in orchid species with and without nectar rewards. New Phytologist 169, 419–430.Google Scholar
Snow, A.A. and Spira, T.P. (1993) Individual variation in the vigor of self pollen and selfed progeny in Hibiscus moscheutos (Malvaceae). American Journal of Botany 80, 160–164.Google Scholar
Sorensen, F. (1969) Embryonic genetic load in coastal douglas-fir, Pseudotsuga menziesii var. menziesii . The American Naturalist 103, 389–398.Google Scholar
Sorensen, F. (1971) Estimate of self-fertility in coastal douglas-fir from inbreeding studies. Silvae Genetica 20, 115–120.Google Scholar
Sorensen, F.C. and Miles, R.S. (1974) Self-pollination effects of douglas-fir and ponderosa pine seeds and seedlings. Silvae Genetica 23, 135–138.Google Scholar
Sorensen, F., Franklin, J.F. and Wollard, R. (1976) Self-pollination effects on seed and seedling traits in noble fir. Forest Science 22, 155–159.Google Scholar
Sork, V.L. and Schemske, D.W. (1992) Fitness consequences on mixed-donor pollen loads in the annual legume Chamaecrista fasciculata . American Journal of Botany 79, 508–515.Google Scholar
Squillace, A.E. and Bingham, R.T. (1958) Selective fertilization in Pinus monticola Dougl. Silvae Genetica 7, 188–196.Google Scholar
Stacy, E.A. (2001) Cross-fertility in two tropical tree species: evidence of inbreeding depression within populations and genetic divergence among populations. American Journal of Botany 88, 1041–1051.Google Scholar
Stein, K. and Hensen, I. (2013) The reproductive biology of two understory plants in the Atlantic rain forest, Brazil. Ecological Research 28, 593–602.Google Scholar
Stephenson, A.G., Leyshon, B., Travers, S.E., Haynes, C.N. and Winsor, J.A. (2004) Interrelationships among inbreeding, herbivory, and disease on reproduction in a wild gourd. Ecology 85, 3023–3034.Google Scholar
Stevens, J.P. and Bougourd, S.M. (1988) Inbreeding depression and the outcrossing rate in natural populations of Allium schoenoprasum L. (wild chives). Heredity 60, 257–261.Google Scholar
Stout, J. (2007) Reproductive biology of the invasive exotic shrub, Rhododendron ponticum L. (Ericaceae). Botanical Journal of the Linnean Society 155, 373–381.Google Scholar
Susko, D.J. and Clubb, M. (2008) Pollination effects on patterns of ovule fate in Hesperis matronalis (Brassicaceae). Botany 86, 466–474.Google Scholar
Takagawa, S., Washitani, I., Uesugi, R. and Tsumura, Y. (2006) Influence of inbreeding depression on a lake population of Nymphoides peltata after restoration from the soil seed bank. Conservation Genetics 7, 705–716.Google Scholar
Taylor, D.R., Trimble, S. and McCauley, D.E. (1999) Ecological genetics and gynodioecy in Silene vulgaris: relative fitness of females and hermaphrodites during the colonization process. Evolution 53, 745–751.Google Scholar
Teixeira, S., Foerster, K. and Bernasconi, G. (2009) Evidence for inbreeding depression and post-pollination selection against inbreeding in the dioecious plant Silene latifolia . Heredity 102, 101–112.Google Scholar
Templeton, A.R. (1986) Coadaptation and outbreeding depression. pp. 105–116 in Soulé, M.E. (Ed.) Conservation biology. The science of scarcity and diversity. Sunderland, MA, Sinauer Associates.Google Scholar
Thiele, J., Hansen, T., Siegismund, H.R. and Hauser, T.P. (2010) Genetic variation in inbreeding depression among floral and fitness traits in Silene nutans . Heredity 104, 52–60.Google Scholar
Trager, M.D., Menges, E.S, Quintana-Ascencio, P.F. and Weekley, C.W. (2005) Outcrossing effects on the reproductive performance of Hypericum cumulicola, an endangered Florida scrub endemic. Journal of the Torrey Botanical Society 132, 204–213.Google Scholar
Trame, A.-M., Coddington, A.J. and Paige, K.N. (1995) Field and genetic studies testing optimal outcrossing in Agave schottii, a long-lived clonal plant. Oecologia 104, 93–100.Google Scholar
Van Assche, J.A., Debucquoy, K.L.A. and Rommens, W.A.F. (2003) Seasonal cycles in the germination capacity of buried seeds of some Leguminosae (Fabaceae). New Phytologist 158, 315–323.Google Scholar
van Damme, J.M.M. and van Delden, W. (1984) Gynodioecy in Plantago lanceolata L. IV. Fitness components of sex types in different life cycle stages. Evolution 38, 1326–1336.Google Scholar
Vandepitte, K., Roldán-Ruiz, I. and Honnay, I. (2009) Reproductive consequences of mate quantity versus mate diversity in a wind-pollinated plant. Acta Oecologica 35, 548–553.Google Scholar
Vange, V. (2002) Breeding system and inbreeding depression in the clonal plant species Knautia arvensis (Dipsacaceae): implications for survival in abandoned grassland. Biological Conservation 108, 59–67.Google Scholar
van Treuren, R., Bijlsma, R., Ouborg, N.J. and van Delden, W. (1993) The significance of genetic erosion in the process of extinction. IV. Inbreeding depression and heterosis effects caused by selfing and outcrossing in Scabiosa columbaria . Evolution 47, 1669–1680.Google Scholar
Vegis, A. (1964) Dormancy in higher plants. Annual Review of Plant Physiology 15, 185–224.Google Scholar
Venable, D.L. (1985) The evolutionary ecology of seed heteromorphism. The American Naturalist 126, 577–595.Google Scholar
Vogler, D.W., Filmore, K. and Stephenson, A.G. (1999) Inbreeding depression in Campanula rapunculoides L. I. A comparison of inbreeding depression in plants derived from strong and weak self-incompatibility phenotypes. Journal of Evolutionary Biology 12, 483–494.Google Scholar
Wagenius, S., Hangelbroek, H.H., Ridley, C.E. and Shaw, R.G. (2009) Biparental inbreeding and inter-remnant mating in a perennial prairie plant: fitness consequences for progeny in their first eight years. Evolution 64, 761–771.Google Scholar
Walisch, T.J., Colling, G., Poncelet, M. and Matthies, D. (2012) Effects of inbreeding and interpopulation crosses on performance and plasticity of two generations of offspring of a declining grassland plant. American Journal of Botany 99, 1300–1313.Google Scholar
Wallace, L.E. (2003) The cost of inbreeding in Platanthera leucophaea (Orchidaceae). American Journal of Botany 90, 235–242.Google Scholar
Waller, D.M., Dole, J. and Bersch, A.J. (2008) Effects of stress and phenotypic variation on inbreeding depression in Brassica rapa . Evolution 62, 917–931.Google Scholar
Wang, T., Hagqvist, R. and Tigerstedt, P.M.A. (1999) Inbreeding depression in three generations of selfed families of silver birch (Betula pendula). Canadian Journal of Forest Research 29, 662–668.Google Scholar
Waser, N.M. and Price, M.V. (1989) Optimal outcrossing in Ipomopsis aggregata: seed set and offspring fitness. Evolution 43, 1097–1109.Google Scholar
Waser, NM. and Price, M.V. (1991) Outcrossing distance effects in Delphinium nelsonii: pollen load, pollen tubes, and seed set. Ecology 72, 171–179.Google Scholar
Waser, N.M. and Price, M.V. (1994) Crossing-distance effects in Delphinium nelsonii: outbreeding and inbreeding depression in progeny fitness. Evolution 48, 842–852.Google Scholar
Waser, N.M., Price, M.V. and Shaw, R.G. (2000) Outbreeding depression varies among cohorts of Ipomopsis aggregata planted in nature. Evolution 54, 485–491.Google Scholar
Watanabe, A., Goka, K. and Washitani, I. (2003) Effects of population spatial structure on the quantity and quality of seeds set by Primula sieboldii (Primulaceae). Plant Species Biology 18, 107–121.Google Scholar
Whitkus, R. (1988) Experimental hybridizations among chromosome races of Carex pachystachya and the related species C. macloviana and C. preslii (Cyperaceae). Systematic Botany 13, 146–153.Google Scholar
Widén, B. (1993) Demographic and genetic effects on reproduction as related to population size in a rare perennial herb, Senecio integrifolius . Biological Journal of the Linnean Society 50, 179–195.Google Scholar
Willi, Y., Van Buskirk, J. and Fischer, M. (2005) A three-fold genetic allee effect: population size affects cross-compatibility, inbreeding depression and drift load in the self-incompatible Ranunculus reptans . Genetics 169, 2255–2265.Google Scholar
Willi, Y., Dietrich, S., van Kleunen, M. and Fischer, M. (2007) Inter-specific competitive stress does not affect the magnitude of inbreeding depression. Evolutionary Ecology Research 9, 959–974.Google Scholar
Willis, J.H. (1993a) Effects of different levels of inbreeding on fitness components in Mimulus guttatus . Evolution 47, 864–876.Google Scholar
Willis, J.H. (1993b) Partial self-fertilization and inbreeding depression in two populations of Mimulus guttatus . Heredity 71, 145–154.Google Scholar
Willis, J.H. (1999a) The role of genes of large effect on inbreeding depression in Mimulus guttatus . Evolution 53, 1678–1691.Google Scholar
Willis, J.H. (1999b) Inbreeding load, average dominance and the mutation rate for mildly deleterious alleles in Mimulus guttatus . Genetics 153, 1885–1898.Google Scholar
Winn, A.A., Elle, E., Kalisz, S., Cheptou, P.-O., Eckert, C.G., Goodwillie, C., Johnston, M.O., Moeller, D.A., Ree, R.H., Sargent, R.D. and Vallejo-Marin, M. (2011) Analysis of inbreeding depression in mixed-mating plants provides evidence for selective interference and stable mixed mating. Evolution 65, 3339–3359.Google Scholar
Wolfe, L.M. (1993) Inbreeding depression in Hydrophyllum appendiculatum: role of maternal effects, crowding, and parental mating history. Evolution 47, 374–386.Google Scholar
Wyatt, R. (1983) Reproductive biology of the granite outcrop endemic Sedum pusillum (Crassulaceae). Systematic Botany 8, 24–28.Google Scholar