Influence of group size and floral display on pollinator behaviour in Moricandia arvensis
DOI:
https://doi.org/10.26786/1920-7603(2026)882Keywords:
Brassicaceae, floral density, neighbourhood, pollinator sharing, Moricandia, pollinator foraging behaviourAbstract
Optimal foraging theory predicts that pollinators will visit dense floral patches over sparse ones. Understanding how the local abundance of floral resources influences pollinator behaviour is crucial for assessing the effect of floral traits on plant reproduction. In this study, we experimentally investigate the role of plant group size and floral display on pollinator foraging behaviour visiting Moricandia arvensis (Brassicaceae). We performed two field experiments: the first manipulated the number of plants per group (group size), and the second manipulated both the total number of flowers per group (group floral display) and group size. We then recorded the foraging behaviour of pollinators using probability of approaching the group, number of plants visited per group, bout length and floral visitation rate. Group floral display was the primary factor influencing pollinator foraging behaviour, except for the probability of approaching the group, where a significant interaction between group floral display and group size was observed. Specifically, the effect of larger floral displays on attracting pollinators increased disproportionately in larger groups. Because the increase in floral visitation rate due to larger displays was less than the increase in the floral display itself, visitation rate per flower was lower in groups with larger displays than in those with smaller displays. Although pollinator foraging behaviour depended largely on local floral resources, the number of plants per group partially shaped the effect of the group floral display on pollinator attraction and visitation rate. This interaction indicates complex effects of neighbouring floral traits on plant-pollinator interactions, which may have important consequences for mating patterns and plant reproductive success.
References
Barbot E, Dufaÿ M, Tonnabel J, Godé C, De Cauwer I (2022) On the function of flower number: disentangling fertility from pollinator-mediated selection. Proceedings of the Royal Society B, 289:20221987. DOI: https://doi.org/10.1098/rspb.2022.1987
Bates D, Mächler M, Bolker B, Walker S (2015) Fitting Linear Mixed-Effects Models Using lme4. Journal of Statistical Software 67:51. DOI: https://doi.org/10.18637/jss.v067.i01
Biernaskie JM, Walker SC, Gegear RJ (2009) Bumblebees learn to forage like Bayesians. The American Naturalist 174:413–423. DOI: https://doi.org/10.1086/603629
Dauber J, Biesmeijer JC, Gabriel D, Kunin WE, Lamborn E, Meyer B, Nielsen A, Potts SG, Roberts SPM, Sõber V, Settele J, Steffan-Dewenter I, Stout JC, Teder T, Tscheulin T, Vivarelli D, Petanidou T (2010) Effects of patch size and density on flower visitation and seed set of wild plants: a pan-European approach. Journal of Ecology 98:188–196. DOI: https://doi.org/10.1111/j.1365-2745.2009.01590.x
Essenberg CJ (2012) Explaining Variation in the Effect of Floral Density on Pollinator Visitation. The American Naturalist 180:153–166. DOI: https://doi.org/10.1086/666610
Essenberg CJ (2013) Scale-dependent shifts in the species composition of flower visitors with changing floral density. Oecologia 171:187–196. DOI: https://doi.org/10.1007/s00442-012-2391-z
Feldman TS (2006) Pollinator aggregative and functional responses to flower density: does pollinator response to patches of plants accelerate at low-densities? Oikos 115:128–140. DOI: https://doi.org/10.1111/j.2006.0030-1299.14493.x
Ghazoul J (2005) Pollen and seed dispersal among dispersed plants. Biological Reviews 80:413–43. DOI: https://doi.org/10.1017/S1464793105006731
Gómez JM, González-Megías A, Narbona E, Navarro L, Perfectti F, Armas C (2022) Phenotypic plasticity guides Moricandia arvensis divergence and convergence across the Brassicaceae floral morphospace. New Phytologist 233:1479–1493. DOI: https://doi.org/10.1111/nph.17807
Gómez JM, Perfectti F, Armas C, Narbona E, González-Megías A, Navarro L, DeSoto L, Torices R (2020) Within-individual phenotypic plasticity in flowers fosters pollination niche shift. Nature Communications 11:4019. DOI: https://doi.org/10.1038/s41467-020-17875-1
Gómez JM, Perfectti F, Bosch J, Camacho JPM (2009) A geographic selection mosaic in a generalized plant–pollinator–herbivore system. Ecological Monographs, 79: 245-263. DOI: https://doi.org/10.1890/08-0511.1
Gómez JM, Torices R, Lorite J, Klingenberg CP, Perfectti F (2016) The role of pollinators in the evolution of corolla shape variation, disparity and integration in a highly diversified plant family with a conserved floral Bauplan. Annals of Botany 117:889–904. DOI: https://doi.org/10.1093/aob/mcv194
Goulson D (2000) Why do pollinators visit proportionally fewer flowers in large patches? Oikos 91:485–492. DOI: https://doi.org/10.1034/j.1600-0706.2000.910309.x
Grindeland JM, Sletvold N, Ims RA. (2005) Effects of floral display size and plant density on pollinator visitation rate in a natural population of Digitalis purpurea. Functional Ecology 19:383–390. DOI: https://doi.org/10.1111/j.1365-2435.2005.00988.x
Hegland SJ (2014) Floral neighbourhood effects on pollination success in red clover are scale-dependent. Functional Ecology 28:561–568. DOI: https://doi.org/10.1111/1365-2435.12223
Hegland SJ, Boeke L (2006) Relationships between the density and diversity of floral resources and flower visitor activity in a temperate grassland community. Ecological Entomology 31:532–538. DOI: https://doi.org/10.1111/j.1365-2311.2006.00812.x
Heinrich B, Raven PH (1972) Energetics and Pollination Ecology. Science 176:597–602. DOI: https://doi.org/10.1126/science.176.4035.597
Janovský Z, Mikát M, Hadrava J, Horčičková E, Kmecová K, Požárová D, Smyčka J, Herben T (2013) Conspecific and heterospecific plant densities at small-scale can drive plant-pollinator interactions. PloS one 8:e77361. DOI: https://doi.org/10.1371/journal.pone.0077361
Klinkhamer PGL, De Jong TJ (1990) Effects of Plant Size, Plant Density and Sex Differential Nectar Reward on Pollinator Visitation in the Protandrous Echium vulgare (Boraginaceae). Oikos 57:399. DOI: https://doi.org/10.2307/3565970
Klinkhamer PGL, De Jong TJ, Linnebank LA (2001) Small-scale spatial patterns determine ecological relationships: An experimental example using nectar production rates. Ecology Letters 4:559–567. DOI: https://doi.org/10.1046/j.1461-0248.2001.00267.x
Knight TM (2003) Floral density, pollen limitation, and reproductive success in Trillium grandiflorum. Oecologia 137:557–63. DOI: https://doi.org/10.1007/s00442-003-1371-8
Kunin WE (1993) Sex and the Single Mustard: Population Density and Pollinator Behavior Effects on Seed-Set. Ecology 74:2145–2160. DOI: https://doi.org/10.2307/1940859
Kunin WE (1997) Population Size and Density Effects in Pollination - Pollinator Foraging and Plant Reproductive Success in Experimental Arrays of Brassica kaber. Journal of Ecology 85:225–234. DOI: https://doi.org/10.2307/2960653
Lenth RV (2016) Least-Squares Means: The {R} Package {lsmeans}. Journal of Statistical Software 69:1–33. DOI: https://doi.org/10.18637/jss.v069.i01
Lewis I, Friedman J (2025) Kin discrimination causes plastic responses in floral and clonal allocation. Proceedings of the Royal Society B: Biological Sciences 292:20242387. DOI: https://doi.org/10.1098/rspb.2024.2387
Makino TT, Ohashi K, Sakai S (2007) How do floral display size and the density of surrounding flowers influence the likelihood of bumble bee revisitation to a plant? Functional Ecology 21:87–95. DOI: https://doi.org/10.1111/j.1365-2435.2006.01211.x
Martén-Rodríguez S, Quesada M, Momen B (2012) Effects of Local Plant Density and Incomplete Dichogamy on the Reproductive Success of the Rare Neotropical Palm Geonoma epetiolata. Biotropica 44:680–688. DOI: https://doi.org/10.1111/j.1744-7429.2012.00860.x
Metcalfe DB, Kunin WE (2005) The effects of plant density upon pollination success, reproductive effort and fruit parasitism in Cistus ladanifer L. (Cistaceae). Plant Ecology 185:41–47. DOI: https://doi.org/10.1007/s11258-005-9082-3
Moeller DA (2004) Facilitative interactions among plants via shared pollinators. Ecology 85:3289–3301. DOI: https://doi.org/10.1890/03-0810
Nielsen A, Dauber J, Kunin WE, Lamborn E, Jauker B, Moora M, Potts SG, Reitan T, Roberts S, Sõber V, Settele J, Steffan-Dewenter I, Stout JC, Tscheulin T, Vaitis M, Vivarelli D, Biesmeijer JC, Petanidou T (2012) Pollinator community responses to the spatial population structure of wild plants: A pan-European approach. Basic and Applied Ecology 13:489–499. DOI: https://doi.org/10.1016/j.baae.2012.08.008
Nottebrock H, Schmid B, Mayer K, Devaux C, Esler KJ, Böhning-Gaese K, Schleuning M, Pagel J, Schurr FM (2017) Sugar landscapes and pollinator-mediated interactions in plant communities. Ecography 40:1129–1138. DOI: https://doi.org/10.1111/ecog.02441
Ohashi K, Yahara T (2002) Visit larger displays but probe proportionally fewer flowers: counterintuitive behaviour of nectar-collecting bumble bees achieves an ideal free distribution. Functional Ecology 16:492–503. DOI: https://doi.org/10.1046/j.1365-2435.2002.00644.x
Parachnowitsch AL, Kessler A (2010) Pollinators exert natural selection on flower size and floral display in Penstemon digitalis. New Phytologist 188: 393-402. DOI: https://doi.org/10.1111/j.1469-8137.2010.03410.x
Seifan M, Hoch E-M, Hanoteaux S, Tielbörger K (2014) The outcome of shared pollination services is affected by the density and spatial pattern of an attractive neighbour. Journal of Ecology 102:953–962. DOI: https://doi.org/10.1111/1365-2745.12256
Sih A, Baltus M-S (1987) Patch Size, Pollinator Behavior, and Pollinator Limitation in Catnip. Ecology 68:1679–1690. DOI: https://doi.org/10.2307/1939860
Sletvold N, Grindeland JM, Ågren J (2010) Pollinator‐mediated selection on floral display, spur length and flowering phenology in the deceptive orchid Dactylorhiza lapponica. New Phytologist 188: 385-392. DOI: https://doi.org/10.1111/j.1469-8137.2010.03296.x
Stout JC, Allen JA, Goulson D (1998) The influence of relative plant density and floral morphological complexity on the behaviour of bumblebees. Oecologia 117:543–550. DOI: https://doi.org/10.1007/s004420050691
Sun S, Broom M, Johanis M, Rychtář J (2021) A mathematical model of kin selection in floral displays. Journal of Theoretical Biology 509:110470. DOI: https://doi.org/10.1016/j.jtbi.2020.110470
Takagi K, Ohashi K (2025) Realized flower constancy in bumble bees: Optimal foraging strategy balancing cognitive and travel costs and its possible consequences for floral diversity. Functional Ecology 39:863–875. DOI: https://doi.org/10.1111/1365-2435.70008
Thomson JD (1981) Spatial and Temporal Components of Resource Assessment by Flower-Feeding Insects. Journal of Animal Ecology 50:49. DOI: https://doi.org/10.2307/4030
Torices R, DeSoto L, Narbona E, Gómez JM (2021) Effects of the Relatedness of Neighbours on Floral Colour. Frontiers in Ecology and Evolution 9:1–11. DOI: https://doi.org/10.3389/fevo.2021.589781
Torices R, Gómez JM, Pannell JR (2018) Kin discrimination allows plants to modify investment towards pollinator attraction. Nature Communications 9:2018. DOI: https://doi.org/10.1038/s41467-018-04378-3
Underwood N, Hambäck PA, Inouye BD (2020) Pollinators, herbivores, and plant neighborhood effects. Quarterly Review of Biology 95:37–57. DOI: https://doi.org/10.1086/707863
Veddeler D, Klein AM, Tscharntke T (2006) Contrasting responses of bee communities to coffee flowering at different spatial scales. Oikos 112:594–601. DOI: https://doi.org/10.1111/j.0030-1299.2006.14111.x
Worley AC, Barrett SC (2000) Evolution of floral display in Eichhornia paniculata (Pontederiaceae): direct and correlated responses to selection on flower size and number. Evolution 54: 1533-1545. DOI: https://doi.org/10.1111/j.0014-3820.2000.tb00699.x
Downloads
Published
How to Cite
Issue
Section
License
Copyright (c) 2026 Rubén Torices, Adela González-Mejías, José María Gómez

This work is licensed under a Creative Commons Attribution 4.0 International License.



