Harder & Johnson S1
Table S1 Studies included in the survey of phenotypic selection. These studies were identified by
searching the Web of Science for papers that reported ‘phenotypic selection’ and
‘flower*’ or ‘inflorescence*’: we also supplemented this collection with other known
examples that the search overlooked. To be included in this survey, a study must have
measured some component of pollination dispersal and/or fertility, so studies that
considered only pollinator visitation were excluded. We focused on studies that
considered floral and/or inflorescence traits of open-pollinated plants and recorded
selection on phenology only if it was measured in conjunction with floral and/or
inflorescence traits. Only records for open-pollinated, unmanipulated plants, including
observations for planted populations, were included.
Alexandersson R, Johnson SD. 2002. Pollinator-mediated selection on flower-tube length in a
hawkmoth-pollinated Gladiolus (Iridaceae). Proceedings of the Royal Society of London
Series B-Biological Sciences 269: 631-636.
Andersson S. 1992. Phenotypic selection in a population of Crepis tectorum ssp. pumila
(Asteraceae). Canadian Journal of Botany 70: 89-95.
Andersson S. 1996. Floral variation in Saxifraga granulata: phenotypic selection, quantitative
genetics and predicted response to selection. Heredity 77: 217-223.
Andersson S, Widén B. 1993. Pollinator-mediated selection on floral traits in a synthetic
population of Senecio integrifolius (Asteraceae). Oikos 66: 72-79.
Arista M, Ortiz PL. 2007. Differential gender selection on floral size: an experimental approach
using Cistus salvifolius. Journal of Ecology 95: 973-982.
Ashman T, Diefenderfer C. 2001. Sex ratio represents a unique context for selection on
attractive traits: consequences for the evolution of sexual dimorphism. American
Naturalist 157: 334-347.
Harder & Johnson S2
Benitez-Vieyra S, Medina AM, Glinos E, Cocucci AA. 2006. Pollinator-mediated selection on
floral traits and size of floral display in Cyclopogon elatus, a sweat bee-pollinated orchid.
Functional Ecology 20: 948-957.
Campbell DR. 1989. Measurements of selection in a hermaphroditic plant: variation in male and
female pollination success. Evolution 43: 318-334.
Campbell DR. 1991. Effects of floral traits on sequential components of fitness in Ipomopsis
aggregata. American Naturalist 137: 713-737.
Campbell DR, Waser NM, Price MV. 1996. Mechanisms of hummingbird-mediated selection
for flower width in Ipomopsis aggregata. Ecology 77: 1463-1472.
Caruso CM. 2000. Competition for pollination influences selection on floral traits of Ipomopsis
aggregata. Evolution 54: 1546-1557.
Caruso CM. 2001. Differential selection on floral traits of Ipomopsis aggregata growing in
contrasting environments. Oikos 94: 295-302.
Caruso CM, Peterson SB, Ridley CE. 2003. Natural selection on floral traits of Lobelia
(Lobeliaceae): spatial and temporal variation. American Journal of Botany 90: 13331340.
Caruso CM, Remington DLD, Ostergren KE. 2005. Variation in resource limitation of plant
reproduction influences natural selection on floral traits of Asclepias syriaca. Oecologia
146: 68-76.
Caruso CM, Yakobowski SJ. 2008. Selection on floral and carbon uptake traits of Lobelia
siphilitica is similar in females and hermaphrodites. Journal of Evolutionary Biology 21:
1514-1523.
Conner JK, Rice AM, Stewart C, Morgan MT. 2003. Patterns and mechanisms of selection on
a family-diagnostic trait: evidence from experimental manipulation and lifetime fitness
selection gradients. Evolution 57: 480-486.
Harder & Johnson S3
Dieringer G, Cabrera L. 1994. Sexual selection of anther trichomes and sexual dimorphism in
Ibervillea lindheimeri (Cucurbitaceae, Melothrieae). American Journal of Botany 81:
111-118.
Eckhart VM. 1993. Do hermaphrodites of gynodioecious Phacelia linearis (Hydrophyllaceae)
trade off seed production to attract pollinators? Biological Journal of the Linnean Society
50: 47-63.
Fenster CB, Ritland K. 1994. Evidence for natural selection on mating system in Mimulus
(Scrophulariaceae). International Journal of Plant Sciences 155: 588-596.
Fishman L, Willis JH. 2008. Pollen limitation and natural selection on floral characters in the
yellow monkeyflower, Mimulus guttatus. New Phytologist 177: 802-810.
Frey FM. 2004. Opposing natural selection from herbivores and pathogens may maintain floralcolor variation in Claytonia virginica (Portulacaceae). Evolution 58: 2426-2437.
Galen C. 1989. Measuring pollinator-mediated slection on morphometric floral traits bumblebees and the alpin sky pilot, Polemonium viscosum. Evolution 43: 882-890.
Gilbert F, Willmer P, Semida F, Ghazoul J, Zalat S. 1996. Spatial variation in selection in a
plant-pollinator system in the wadis of Sinai, Egypt. Oecologia 108: 479-487.
Giles BE, Pettersson TM, Carlsson-Graner U, Ingvarsson PK. 2006. Natural selection on
floral traits of female Silene dioica by a sexually transmitted disease. New Phytologist
169: 729-739.
Gómez JM. 2000. Phenotypic selection and response to selection in Lobularia maritima:
importance of direct and correlational components of natural selection. Journal of
Evolutionary Biology 13: 689-699.
Gómez JM. 2003. Herbivory reduces the strength of pollinator-mediated selection in the
Mediterranean herb Erysimum mediohispanicum: consequences for plant specialization.
American Naturalist 162: 242-256.
Harder & Johnson S4
Gómez JM, Perfectti F, Camacho JPM. 2006. Natural selection on Erysimum
mediohispanicum flower shape: Insights into the evolution of zygomorphy. American
Naturalist 168: 531-545.
Gómez JM, Bosch J, Perfectti F, Fernández JD, Abdelaziz M, Camacho JPM. 2008. Spatial
variation in selection on corolla shape in a generalist plant is promoted by the preference
patterns of its local pollinators. Proceedings of the Royal Society of London, Series B
275: 2241-2249.
Hall MC, Willis JH. 2006. Divergent selection on flowering time contributes to local adaptation
in Mimulus guttatus populations. Evolution 60: 2466-2477.
Herrera CM. 1993. Selection on floral morphology and environmental determinants of
fecundity in a hawk moth-pollinated violet. Ecological Monographs 63: 251-275.
Hodgins KA, Barrett SCH. 2008. Natural selection on floral traits through male and female
function in wild populations of the heterostylous daffodil Narcissus triandrus. Evolution
62: 1751-1763.
Holland JN, Chamberlain SA. 2007. Ecological and evolutionary mechanisms for low seed :
ovule ratios: need for a pluralistic approach? Ecology 88: 706-715.
Irwin RE. 2000. Morphological variation and female reproductive success in two sympatric
Trillium species: evidence for phenotypic selection in Trillium erectum and Trillium
grandiflorum (Liliaceae). American Journal of Botany 87: 205-214.
Johnston MO. 1991. Natural selection on floral traits in two species of Lobelia with different
pollinators. Evolution 45: 1468-1479.
Little KJ, Dieringer G, Romano M. 2005. Pollination ecology, genetic diversity and selection
on nectar spur length in Platanthera lacera (Orchidaceae). Plant Species Biology 20:
183-190.
Maad J. 2000. Phenotypic selection in hawkmoth-pollinated Platanthera bifolia: targets and
fitness surfaces. Evolution 54: 112-123.
Harder & Johnson S5
Maad J, Alexandersson R. 2004. Variable selection in Platanthera bifolia (Orchidaceae):
phenotypic selection differed between sex functions in a drought year. Journal of
Evolutionary Biology 17: 642-650.
Medel R, Botto-Mahan C, Kalin-Arroyo M. 2003. Pollinator-mediated selection on the nectar
guide phenotype in the Andean monkey flower, Mimulus luteus. Ecology 84: 1721-1732.
Mitchell RJ, Shaw RG, Waser NM. 1998. Pollinator selection, quantitative genetics, and
predicted evolutionary responses of floral traits in Penstemon centranthifolius
(Scrophulariaceae). International Journal of Plant Sciences 159: 331-337.
Moeller DA, Geber MA. 2005. Ecological context of the evolution of self-pollination in Clarkia
xantiana: population size, plant communities, and reproductive assurance. Evolution 59:
786-799.
Morgan MT, Schoen DJ. 1997. Selection on reproductive characters: floral morphology in
Asclepias syriaca. Heredity 79: 433-441.
Nuismer SL, Cunningham BM. 2005. Selection for phenotypic divergence between diploid and
autotetraploid Heuchera grossulariifolia. Evolution 59: 1928-1935.
O'Connell LM, Johnston MO. 1998. Male and female pollination success in a deceptive
orchid, a selection study. Ecology 79: 1246-1260.
Parachnowitsch AL, Caruso CM. 2008. Predispersal seed herbivores, not pollinators, exert
selection on floral traits via female fitness. Ecology 89: 1802-1810.
Parra-Tabla V, Bullock SH. 2005. Ecological and selective effects of stigma-anther separation
in the self-incompatible tropical tree Ipomoea wolcottiana (Convolvulaceae). Plant
Systematics and Evolution 252: 85-95.
Petit C, Thompson JD. 1998. Phenotypic selection and population differentiation in relation to
habitat heterogeneity in Arrhenatherum elatius (Poaceae). Journal of Ecology 86: 829840.
Pilson D. 2000. Herbivory and natural selection on flowering phenology in wild sunflower,
Helianthus annuus. Oecologia 122: 72-82.
Harder & Johnson S6
Sandring S, Riihimäki MA, Savolainen O, Ågren J. 2007. Selection on flowering time and
floral display in an alpine and a lowland population of Arabidopsis lyrata. Journal of
Evolutionary Biology 20: 558-567.
Schemske DW, Horvitz CC. 1989. Temporal variation in selection on a floral character.
Evolution 43: 461-465.
Schueller SK. 2007. Island-mainland difference in Nicotiana glauca (Solanaceae) corolla
length: a product of pollinator-mediated selection? Evolutionary Ecology 21: 81-98.
Totland Ø. 1999. Effects of temperature on performance and phenotypic selection on plant traits
in alpine Ranunculus acris. Oecologia 120: 242-251.
Totland Ø. 2001. Environment-dependent pollen limitation and selection on floral traits in an
alpine species. Ecology 82: 2233-2244.
Totland Ø, Andersen HL, Bjelland T, Dahl V, Eide W, Houge S, Pedersen TR, Vie EU.
1998. Variation in pollen limitation among plants and phenotypic selection on floral traits
in an early-spring flowering herb. Oikos 82(3): 491-501.
van Kleunen M, Burczyk J. 2008. Selection on floral traits through male fertility in natural
plant population. Evolutionary Ecology 22: 39-54.
Wilson P. 1995. Selection for pollination success and the mechanical fit of Impatiens flowers
aroung bumblebee bodies. Biological Journal of the Linnean Society 55: 355-383.
Wright JW, Meagher TR. 2004. Selection on floral characters in natural Spanish populations of
Silene latifolia. Journal of Evolutionary Biology 17: 382-395.
Harder & Johnson S7
Table S2 Categorization of traits considered in the survey of phenotypic selection. The
frequencies represent the number of measurements of a trait per study, species, fitness
measure, population, year, morph, and/or cytotype. PC = principal component; RW =
relative warp.
Trait class
Trait
Attraction
nectar guide shape
2
0.19
nectar guide size
1
0.09
anther dimorphism
3
0.28
anther exsertion
3
0.28
anther height
4
0.37
anther trichomes
2
0.19
carpel length
4
0.37
corolla tube width
6
0.56
corpusculum length
5
0.47
flower diameter
30
2.8
flower length
17
1.59
flower width
56
5.22
hood separation
5
0.47
hood width
5
0.47
horn length
5
0.47
30
2.8
stamen length
4
0.37
stigmatic slit length
2
0.19
stigmatic slit width
5
0.47
stigma exsertion
31
2.89
stigma height
10
0.93
Contact
porch petal height
Frequency
Percent
Harder & Johnson S8
Flower color
Flower production
Flower shape
Flower size
stigma size
1
0.09
style length
2
0.19
tube diameter
1
0.09
fluctuating asymmetry
1
0.09
flower color
4
0.37
number of red dots
1
0.09
optical density
2
0.19
pigment
2
0.19
target
2
0.19
visual rank
2
0.19
96
8.96
head number
2
0.19
head size
1
0.09
head width
1
0.09
inflorescence number
6
0.56
flower shape PC
4
0.37
flower shape RW1
6
0.56
flower shape RW2
6
0.56
flower shape RW3
5
0.47
flower shape RW4
5
0.47
width:length
1
0.09
calyx diameter
5
0.47
corolla diameter
5
0.47
corolla lobe length
9
0.84
corolla lobe width
8
0.75
corolla size
1
0.09
corolla width
3
0.28
flower number
Harder & Johnson S9
flower area
10
0.93
flower circumference
30
2.8
flower diameter
15
1.4
flower height
4
0.37
flower length:width
1
0.09
flower length
1
0.09
flower mass
1
0.09
flower size
15
1.4
flower size PC
6
0.56
flower size PC1
5
0.47
flower size PC2
5
0.47
flower size PC3
5
0.47
flower size composite
2
0.19
16
1.49
labellum length
6
0.56
labellum width
6
0.56
lip size
4
0.37
petal length
6
0.56
18
1.68
petal width
5
0.47
sepal length
4
0.37
sepal width
4
0.37
28
2.61
flower density
2
0.19
head diameter
1
0.09
inflorescence display
13
1.21
inflorescence height
40
3.73
flower width
petal size
Inflorescence display
% flowers female phase
Harder & Johnson S10
Nectar depth
pedicel length
4
0.37
plant size PC
2
0.19
claw length
5
0.47
corolla tube length
9
0.84
10
0.93
hood height
2
0.19
hood length
5
0.47
spur length
18
1.68
tube length
54
5.04
1
0.09
36
3.36
ovary length
1
0.09
ovule number
3
0.28
flowering date
39
3.64
flowering date mean
6
0.56
flowering date median
9
0.84
flowering duration
7
0.65
flowering date final
4
0.37
flowering period
3
0.28
flowering synchrony
4
0.37
anther length
1
0.09
anther number
1
0.09
% non-viable pollen
1
0.09
protandry
9
0.84
flower length
Nectar production
nectar production rate
nectar volume
Ovule production
Phenology
Pollen production
Protandry
Harder & Johnson S11
Table S3 Experimental studies surveyed for functional evidence of floral and inflorescence
adaptation. The three papers describing results from an array experiment with Mimulus
ringens (Karron et al., 2004; Mitchell et al., 2004, 2005) were considered to be part of a
single study.
Andersson S. 1994. Floral stability, pollination efficiency, and experimental manipulation of the
corolla phenotype in Nemophila menziesii (Hydrophyllaceae). American Journal of
Botany 81: 1397-1402.
Andersson S. 1996. Floral display and pollination success in Senecio jacobaea (Asteraceae):
interactive effects of the head and corymb size. American Journal of Botany 83: 71-75.
Andersson S. 2008. Pollinator and nonpollinator selection on ray morphology in Leucanthemum
vulgare (oxeye daisy, Asteraceae). American Journal of Botany 95: 1072-1078.
Arista M, Ortiz PL. 2007. Differential gender selection on floral size: an experimental approach
using Cistus salvifolius. Journal of Ecology 95: 973-982.
Armbruster WS, Antonsen L, Pélabon C. 2005. Phenotypic selection on Dalechampia
blossoms: honest signaling affects pollination success. Ecology 86: 3323-3333.
Campbell DR. 1996. Evolution of floral traits in a hermaphroditic plant: field measurements of
heritabilitles and genetic correlations. Evolution 50(4): 1442-1453.
Campbell DR, Waser NM, Price MV. 1994. Indirect selection on stigma position in Ipomopsis
aggregata via a genetically correlated trait. Evolution 48: 55-68.
Carlson JE. 2008. Hummingbird responses to gender-biased nectar production: are nectar biases
maintained by natural or sexual selection? Proceedings of the Royal Society of London, B
275: 1717-1726.
Conner JK, Rice AM, Stewart C, Morgan MT. 2003. Patterns and mechanisms of selection on
a family-diagnostic trait: evidence from experimental manipulation and lifetime fitness
selection gradients. Evolution 57: 480-486.
Harder & Johnson S12
Cresswell JE. 2000. Manipulation of female architecture in flowers reveals a narrow optimum
for pollen deposition. Ecology 81: 3244-3249.
Friedman J, Barrett SCH. 2008. A phylogenetic analysis of the evolution of wind pollination
in the angiosperms. International Journal of Plant Sciences 169: 49-58.
Golding YC, Sullivan MS, Sutherland JP. 1999. Visits to manipulated flowers by Episyrphus
balteatus (Diptera: Syrphidae): partitioning the signals of petals and anthers. Journal of
Insect Behavior 12: 39-45.
Harder LD, Aizen MA. 2004. The functional significance of synchronous protandry in
Alstroemeria aurea. Functional Ecology 18: 467-474.
Harder LD, Johnson SD. 2005. Adaptive plasticity of floral display size in animal-pollinated
plants. Proceedings of the Royal Society of London, B 272: 2651-2657.
Herrera CM. 2001. Deconstructing a floral phenotype: do pollinators select for corolla
integration in Lavandula latifolia? Journal of Evolutionary Biology 14: 574-584.
Ishii HS, Harder LD. 2006. The size of individual Delphinium flowers and the opportunity for
geitonogamous pollination. Functional Ecology 20: 1115-1123.
Jersáková J, Johnson SD. 2006. Lack of floral nectar reduces self-pollination in a fly-pollinated
orchid. Oecologia 147: 60-68.
Jersáková J, Johnson SD. 2007. Protandry promotes male pollination success in a mothpollinated orchid. Functional Ecology 21: 496-504.
Johnson SD, Andersson S. 2002. A simple field method for manipulating ultraviolet reflectance
of flowers. Canadian Journal of Botany 80: 1325-1328.
Johnson SD, Steiner KE. 1997. Long-tongued fly pollination and evolution of floral spur length
in the Disa draconis complex (Orchidaceae). Evolution 51: 45-53.
Karron JD, Mitchell RJ, Holmquist KG, Bell JM, Funk B. 2004. The influence of floral
display size on selfing rates in Mimulus ringens. Heredity 92: 242-248.
Kessler D, Gase K, Baldwin IT. 2008. Field experiments with transformed plants reveal the
sense of floral scents. Science 321: 1200-1202.
Harder & Johnson S13
Kohn JR, Barrett CH. 1992. Experimental studies on the functional significance of heterostyly.
Evolution 46: 43-55.
Lamborn E, Ollerton J. 2000. Experimental assessment of the functional morphology of
inflorescences of Daucus carota (Apiaceae): testing the 'fly catcher effect'. Functional
Ecology 14: 445-454.
Mitchell RJ, Karron JD, Holmquist KG, Bell JM. 2004. The influence of Mimulus ringens
floral display size on pollinator visitation patterns. Functional Ecology 18: 116-124.
Mitchell RJ, Karron JD, Holmquist KG, Bell JM. 2005. Patterns of multiple paternity in fruits
of Mimulus ringens (Phrymaceae). American Journal of Botany 92: 885-890.
Muchhala N. 2007. Adaptive trade-off in floral morphology mediates specialization for flowers
pollinated by bats and hummingbirds. American Naturalist 169: 494-504.
Nilsson LA. 1988. The evolution of flowers with deep corolla tubes. Nature 334: 147-149.
Peter CI, Johnson SD. 2008. Mimics and magnets: the importance of color and ecological
facilitation in floral deception. Ecology 89: 1583-1595.
Schemske DW, Bradshaw HD, Jr. 1999. Pollinator preference and the evolution of floral traits
in monkeyflowers (Mimulus). Proceedings of the National Academy of Sciences, U.S.A.
96: 11910-11915.
Tadey M, Aizen MA. 2001. Why do flowers of a hummingbird-pollinated mistletoe face down?
Functional Ecology 15: 782-790.
Ushimaru A, Kawase D, Imamura A. 2006. Flowers adaptively face down-slope in 10 forestfloor herbs. Functional Ecology 20: 585-591.
Waelti MO, Muhlemann JK, Widmer A, Schiestl FP. 2008. Floral odour and reproductive
isolation in two species of Silene. Journal of Evolutionary Biology 21: 111-121.
Walker-Larsen J, Harder LD. 2001. Vestigial organs as opportunities for functional
innovation: the example of the Penstemon staminode. Evolution 55: 477-487.
Waser NM, Price MV. 1983. Pollinator behaviour and natural selection for flower colour in
Delphinium nelsonii. Nature 302: 422-424.
Harder & Johnson S14
Weeks EL, Frey FM. 2007. Seed production and insect visitation rates in Hesperis matronalis
are not affected by floral symmetry. International Journal of Plant Sciences 168: 611617.
Wilson P. 1995. Selection for pollination success and the mechanical fit of Impatiens flowers
around bumblebee bodies. Biological Journal of the Linnean Society 55: 355-383.
Wilson P. 1995. Variation in the intensity of pollination in Drosera tracyi: selection is strongest
when resources are intermediate. Evolutionary Ecology 9: 382-396.
Zhao ZG, He YL, Wang MT, Du GZ. 2007. Variations of flower size and reproductive traits in
self-incompatible Trollius ranunculoides (Ranunculaceae) among local habitats at Alpine
Meadow. Plant Ecology 193: 241-251.