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Germination niche of a pioneer woody species (Manihot grahamii hook.): a strategy of seed heat stimulation to cope with disturbance in dry subtropical forests

Published online by Cambridge University Press:  10 November 2023

Sebastián R. Zeballos Open the ORCID record for Sebastián R. Zeballos [Opens in a new window] ,
Paula Venier ,
Mariana Pereyra Open the ORCID record for Mariana Pereyra [Opens in a new window] ,
Denise Simian  and
Guillermo Funes
Sebastián R. Zeballos*
Affiliation:
Instituto Multidisciplinario de Biología Vegetal (FCEFyN, CONICET-UNC), Córdoba, Argentina
Paula Venier
Affiliation:
Instituto Multidisciplinario de Biología Vegetal (FCEFyN, CONICET-UNC), Córdoba, Argentina
Mariana Pereyra
Affiliation:
Instituto Multidisciplinario de Biología Vegetal (FCEFyN, CONICET-UNC), Córdoba, Argentina
Denise Simian
Affiliation:
Centro de Recursos Naturales Renovables de la Zona Semiárida (CONICET-UNS), Bahía Blanca, Provincia de Buenos Aires, Argentina
Guillermo Funes
Affiliation:
Instituto Multidisciplinario de Biología Vegetal (FCEFyN, CONICET-UNC), Córdoba, Argentina Facultad de Ciencias Exactas, Físicas y Naturales, Universidad Nacional de Córdoba, Córdoba, Argentina
*
Corresponding author: Sebastián R. Zeballos; Email: sebazeba@hotmail.com
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Abstract

Light and temperature conditions trigger germination in specific temporal windows and microhabitats, thus determining the germination niche of plant species. Manihot species grow in fire-prone ecosystems and their seeds show heat tolerance. Successful establishment in disturbed environments might be related to seed attributes that allow seeds to cope with or avoid stressful environments. We studied some characteristics of the germination niche of Manihot grahamii, a pioneer woody species present in dry subtropical forests of central Argentina, to understand its successful establishment in disturbed environments. We evaluated the germination ecology of the seeds of M. grahamii with the aims to (1) characterize seed traits (viability, mass and moisture content); (2) determine whether it has dormancy and if it is physical or physiological; (3) evaluate the effect of several pre-treatments (gibberellic acid, after ripening, dry prechilling and dry prechilling + warm) on seed dormancy; and (4) assess the effect of different environmental events of high temperatures on the germination process simulating two treatments: fire intensities (with three levels of heat shock) and a gap temperature. M. grahamii seeds have large mass (0.24 g), low moisture content (8%), physiological dormancy, negative photoblastic behaviour and high heat tolerance. Dormancy was alleviated and seeds became insensitive to light when they were exposed to pre-treatments of dry prechilling + warm and high-temperature treatments. This germination strategy promotes secure germination timing into the rainy season on undisturbed habitats as well as a cue for competition-released gaps which in turn favour recruitment in open, disturbed and dry habitats, respectively. In the context of global change, with an increasing habitat fragmentation and fire frequency, M. grahamii could become more abundant and extend its geographic distributional range in central Argentina.

Keywords

Chaco ForestDormancyEspinal ForestFireGermination timingPhotoblastismSeed functional traits
Type
Research Paper
Information
Seed Science Research , Volume 33 , Issue 3 , September 2023 , pp. 182 - 192
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Copyright
Copyright © The Author(s), 2023. Published by Cambridge University Press

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References

Adjata, KD, Tchaniley, L, Banla, E, Tchansi, KK and Gumedzoe, YMD (2013) Study of germination conditions of cassava (Manihot esculenta Crantz) seeds obtained by genetic selection. Scientific Research and Essays 8, 2138–2143.CrossRefGoogle Scholar
Arcamone, JR and Jaureguiberry, P (2018) Germination response of common annual and perennial forbs to heat shock and smoke treatments in the Chaco Serrano, central Argentina. Austral Ecology 43, 567–577.CrossRefGoogle Scholar
Argañaraz, JP, Radeloff, V, Bar-Massada, A, Gavier, G, Scavuzzo, M and Bellis, LM (2015) Mapeo y evaluación del riesgo de incendio en la interfase urbano-rural de las sierras de Córdoba. V Jornadas y II Congreso Argentino de Ecología de Paisajes Azul, May 2015, Buenos Aires, Argentina.Google Scholar
Argañaraz, JP, Cingolani, AM, Bellis, LM and Giorgis, MA (2020) Fire incidence along an elevation gradient in the mountains of central Argentina. Ecología Austral 30, 268–281.CrossRefGoogle Scholar
Baskin, CC and Baskin, JM (2004) Germinating seeds of wildflowers, an ecological perspective. HortTechnology 14, 467–473.CrossRefGoogle Scholar
Baskin, CC and Baskin, JM (2014) Seeds: Ecology, Biogeography, and Evolution of Dormancy and Germination, 2nd Edn. San Diego, CA, Elsevier.Google Scholar
Baskin, CC and Baskin, JM (2022) Plant Regeneration from Seeds: A Global Warming Perspective. San Diego, CA, Academic Press.Google Scholar
Baskin, JM, Davis, BH, Baskin, CC, Gleason, SM and Cordell, S (2004) Physical dormancy in seeds of Dodonaea viscosa (Sapindales, Sapindaceae) from Hawaii. Seed Science Research 14, 81–90.CrossRefGoogle Scholar
Baskin, CC, Thompson, K and Baskin, JM (2006) Mistakes in germination ecology and how to avoid them. Seed Science Research 16, 165–168.CrossRefGoogle Scholar
Bazzaz, FA and Pickett, STA (1980) Physiological ecology of tropical succession: a comparative review. Annual Review of Ecology and Systematics 11, 287–310.CrossRefGoogle Scholar
Becwar, MR, Stanwood, PC and Leonhardt, KW (1983) Dehydration effects on freezing characteristics and survival in liquid nitrogen of desiccation-tolerant and desiccation-sensitive seeds. Journal of the American Society for Horticultural Science 108, 613–618.CrossRefGoogle Scholar
Bell, DT, Plummer, JA and Taylor, SK (1993) Seed germination ecology in southwestern Western Australia. The Botanical Review 59, 24–73.CrossRefGoogle Scholar
Bewley, JD, Bradford, K and Hilhorst, HWM (2013) Seeds: Physiology of Development, Germination and Dormancy. New York, Springer Science & Business Media.CrossRefGoogle Scholar
Can, L and Küçüker, O (2015) Seed morphology and surface microstructure of some Euphorbia (Euphorbiaceae) taxa distributed in Turkey-in-Europe. Turkish Journal of Botany 39, 449–457.CrossRefGoogle Scholar
Carta, A, Skourti, E, Mattana, E, Vandelook, F and Thanos, CA (2017) Photoinhibition of seed germination: occurrence, ecology and phylogeny. Seed Science Research 27, 131–153.CrossRefGoogle Scholar
Chambers, JC, MacMahon, JA and Haefner, JH (1991) Seed entrapment in alpine ecosystems: effects of soil particle size and diaspore morphology. Ecology 72, 1668–1677.CrossRefGoogle Scholar
Chazdon, RL and Fetcher, N (1984) Light environments of tropical forests, pp. 27–36 in Medina, E; Mooney, HA; Vázquez-Yánes, C (eds), Physiological Ecology of Plants of the Wet Tropics, Dordrecht, Springer.CrossRefGoogle Scholar
Cochrane, A (2019) Effects of temperature on germination in eight Western Australian herbaceous species. Folia Geobotanica 54, 29–42.CrossRefGoogle Scholar
Cochrane, A, Hoyle, GL, Yates, CJ, Wood, J and Nicotra, AB (2014) Predicting the impact of increasing temperatures on seed germination among populations of Western Australian Banksia (Proteaceae). Seed Science Research 24, 195–205.CrossRefGoogle Scholar
D'Agostino, AB, Gurvich, DE, Ferrero, MC, Zeballos, SR and Funes, G (2012) Requerimientos germinativos de enredaderas características del Chaco Serrano de Córdoba, Argentina. Revista de Biología Tropical 60, 1513–1523.Google Scholar
Daws, MI, Crabtree, LM, Dalling, JW, Mullins, CE and Burslem, DF (2008) Germination responses to water potential in neotropical pioneers suggest large-seeded species take more risks. Annals of Botany 102, 945–951.CrossRefGoogle ScholarPubMed
De Fina, AL (1992) Aptitud agroclimática de la República Argentina. Academia Nacional de Agronomía y Veterinaria. Buenos Aires, AR.Google Scholar
Donohue, K, Rubio de Casas, R, Burghardt, L, Kovach, K and Willis, CG (2010) Germination, postgermination adaptation, and species ecological ranges. Annual Review of Ecology, Evolution, and Systematics 41, 293–319.CrossRefGoogle Scholar
Drennan, PM and Van Staden, J (1992) The effect of ethrel and temperature on the germination of Manihot glaziovii Muell. Arg. seeds. Plant Growth Regulation 11, 273–275.CrossRefGoogle Scholar
Duputié, A, Salick, J and McKey, D (2011) Evolutionary biogeography of Manihot (Euphorbiaceae), a rapidly radiating Neotropical genus restricted to dry environments. Journal of Biogeography 38, 1033–1043.CrossRefGoogle Scholar
Elias, M and McKey, D (2000) The unmanaged reproductive ecology of domesticated plants in traditional agroecosystems: An example involving cassava and a call for data. Acta Oecologica 21, 223–230.CrossRefGoogle Scholar
Ellis, RH, Hong, TD and Roberts, EH (1981) The influence of desiccation on cassava seed germination and longevity. Annals of Botany 47, 173–175.CrossRefGoogle Scholar
Espadaler, XT and Gómez, C (1996) Seed production, predation and dispersal in the Mediterranean myrmecochore Euphorbia characias (Euphorbiaceae). Ecography 19, 7–15.CrossRefGoogle Scholar
Essl, F, Dullinger, S, Genovesi, P, Hulme, PE, Jeschke, JM, Katsanevakis, S, Kühn, I, Lenzner, B, Pauchard, A, Pyšek, P, Rabitsch, W, Richardson, DM, Seebens, H, van Kleunen, M, van Der Putten, WH, Vilà, M and Bacher, S (2019) A conceptual framework for range-expanding species that track human-induced environmental change. BioScience 69, 908–919.CrossRefGoogle Scholar
Fehlenberg, V, Baumann, M, Gasparri, NI, Piquer-Rodriguez, M, Gavier-Pizarro, G and Kuemmerle, T (2017) The role of soybean production as an underlying driver of deforestation in the South American Chaco. Global Environmental Change 45, 24–34.CrossRefGoogle Scholar
Ferreras, AE, Funes, G and Galetto, L (2015) The role of seed germination in the invasion process of Honey locust (Gleditsia triacanthos L., Fabaceae): comparison with a native confamilial. Plant Species Biology 30, 126–136.CrossRefGoogle Scholar
Funes, G, Díaz, S and Venier, P (2009) La temperatura como principal determinante de la germinación en especies del Chaco seco de Argentina. Ecología Austral 19, 129–138.Google Scholar
Giorgis, M and Tecco, PA (2014) Árboles y arbustos invasores de la Provincia de Córdoba (Argentina): una contribución a la sistematización de bases de datos globales. Boletín de la Sociedad Argentina de Botánica 49, 681–603.CrossRefGoogle Scholar
Gómez-González, S and Cavieres, LA (2009) Litter burning does not equally affect seedling emergence of native and alien species of the Mediterranean-type Chilean matorral. International Journal of Wildland Fire 18, 213–21.CrossRefGoogle Scholar
Grubb, PJ (1977) The maintenance of species-richness in plant communities: the importance of the regeneration niche. Biological Reviews 52, 107–145.CrossRefGoogle Scholar
International Seed Testing Association (2014) International Rules for Seed Testing, 2014 edition. Zürich, Switzerland, ISTA.Google Scholar
Jaureguiberry, P and Díaz, S (2015) Post-burning regeneration of the Chaco seasonally dry forest: germination response of dominant species to experimental heat shock. Oecologia 177, 689–699.CrossRefGoogle ScholarPubMed
Kazancı, DD and Tavşanoğlu, Ç (2019) Heat shock-stimulated germination in Mediterranean Basin plants in relation to growth form, dormancy type and distributional range. Folia Geobotanica 54, 85–98.CrossRefGoogle Scholar
Kottek, M, Grieser, J, Beck, C, Rudolf, B and Rubel, F (2006) World map of the Köppen-Geiger climate classification updated. Meteorologische Zeitschrift 15, 259–263.CrossRefGoogle Scholar
Lagoa, AMMA and Pereira, MFA (1987) The role of the caruncle in the germination of seeds of Ricinus communis. Plant Physiology and Biochemistry 25, 125–128.Google Scholar
Lan, QY, Xia, K, Wang, XF, Liu, JW, Zhao, J and Tan, YH (2014) Seed storage behaviour of 101 woody species from the tropical rainforest of southern China: a test of the seed-coat ratio–seed mass (SCR–SM) model for determination of desiccation sensitivity. Australian Journal of Botany 62, 305–311.CrossRefGoogle Scholar
Larson, JE and Funk, JL (2016) Regeneration: an overlooked aspect of trait-based plant community assembly models. Journal of Ecology 104, 1284–1298.CrossRefGoogle Scholar
Le Fer, D and Parker, VT (2005) The effect of seasonality of burn on seed germination in chaparral: the role of soil moisture. Madroño 52, 166–174.CrossRefGoogle Scholar
Leopold, AC and Vertucci, CW (1989) Moisture as a regulator of physiological reaction in seeds. Seed Moisture 14, 51–67.Google Scholar
Lipoma, ML, Funes, G and Díaz, S (2018) Fire effects on the soil seed bank and post-fire resilience of a semi-arid shrubland in central Argentina. Austral Ecology 43, 46–55.CrossRefGoogle Scholar
Long, RL, Gorecki, MJ, Renton, M, Scott, JK, Colville, L, Goggin, DE, Commander, LE, Westcott, DA, Cherry, H and Finch-Savage, WE (2015) The ecophysiology of seed persistence: a mechanistic view of the journey to germination or demise. Biological Reviews 90, 31–59.CrossRefGoogle ScholarPubMed
Loudermilk, EL, O'Brien, JJ, Mitchell, RJ, Cropper, WP Jr, Hiers, JK, Grunwald, S, Grego, J and Fernandez-Diaz, JC (2012) Linking complex forest fuel structure and fire behavior at fine scales. International Journal of Wildland Fire 21, 882–93.CrossRefGoogle Scholar
Luna, B and Moreno, J (2010) Range-size, local abundance and germination niche-breadth in Mediterranean plants of two lifeforms. Plant Ecology 210, 85–95.CrossRefGoogle Scholar
Luna, B, Moreno, JM, Cruz, A and Fernandez-Gonzalez, F (2007) Heatshock and seed germination of a group of Mediterranean plant species growing in a burned area: an approach based on plant functional types. Environmental and Experimental Botany 60, 324–333.CrossRefGoogle Scholar
Marques, AR, Atman, AP, Silveira, FA and de Lemos-Filho, JP (2014) Are seed germination and ecological breadth associated? Testing the regeneration niche hypothesis with bromeliads in a heterogeneous neotropical montane vegetation. Plant Ecology 215, 517–529.CrossRefGoogle Scholar
Milberg, P, Andersson, L and Thompson, K (2000) Large-seeded spices are less dependent on light for germination than small-seeded ones. Seed Science Research 10, 99–104.CrossRefGoogle Scholar
Moreschi, EG, Funes, G, Zeballos, SR and Tecco, PA (2019) Post-burning germination responses of woody invaders in a fire-prone ecosystem. Austral Ecology 44, 1163–1173.CrossRefGoogle Scholar
Moyo, M, Kulkarni, MG, Finnie, JF and Van Staden, J (2009) After-ripening, light conditions, and cold stratification influence germination of marula [Sclerocarya birrea (A. Rich.) Hochst. subsp. caffra (Sond.) Kokwaro] seeds. HortScience 44, 119–124.CrossRefGoogle Scholar
Muñoz Garachana, D, Aragón, R and Baldi, G (2018) Estructura espacial de remanentes de bosque nativo en el Chaco Seco y el Espinal. Ecología Austral 28, 553–564.CrossRefGoogle Scholar
Nartey, F, Møller, BL and Andersen, MR (1974) Changes in the major constituents of Manihot esculenta seeds during germination and growth. Economic Botany 28, 145–154.CrossRefGoogle Scholar
Nassar, NMA and Teixeira, RP (1983) A quebra da dorméncia da semente das espécies selvagens da mandioca. Manihot spp. Ciéncia e Cultura 35, 630–632.Google Scholar
Nassar, NM, Hashimoto, DYC and Fernandes, SDC (2008) Wild Manihot species: botanical aspects, geographic distribution and economic value. Genetics and Molecular Research 7, 16–28.CrossRefGoogle ScholarPubMed
Ooi, MK, Denham, AJ, Santana, VM and Auld, TD (2014) Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change. Ecology and Evolution 4, 656–671.CrossRefGoogle ScholarPubMed
Ospina, J, Gómez, CLG and Orozco, MSS (2000) Latencia en semillas de Manihot y Ricinus (Euphorbiaceae). Acta Agronómica 50, 30–37.Google Scholar
Pacini, E (1990) Mercurialis annua L. (Euphorbiaceae) seed interactions with the ant Messor structor (Latr.), Hymenoptera: Formicidae. Acta Botanica Neerlandica 39, 253–262.CrossRefGoogle Scholar
Paula, S and Pausas, JG (2008) Burning seeds: germinative response to heat treatments in relation to resprouting ability. Journal of Ecology 96, 543–552.CrossRefGoogle Scholar
Pearson, TRH, Burslem, DFRP, Mullins, CE and Dalling, JW (2002) Germination ecology of neotropical pioneers: interacting effects of environmental conditions and seed size. Ecology 83, 2798–2807.CrossRefGoogle Scholar
Pujol, B, Gigot, G, Laurent, G, Pinheiro-Kluppel, M, Elias, M, Hossaert-McKey, M and McKey, D (2002) Germination ecology of cassava (Manihot esculenta Crantz, Euphorbiaceae) in traditional agroecosystems: seed and seedling biology of a vegetatively propagated domesticated plant. Economic Botany 56, 366–379.CrossRefGoogle Scholar
Ranal, MA and Santana, DGD (2006) How and why to measure the germination process? Brazilian Journal of Botany 29, 1–11.CrossRefGoogle Scholar
R Core Team (2022) R: A language and environment for statistical computing. Vienna, Austria, R Foundation for Statistical Computing. Available at https://www.R-project.org/.Google Scholar
Renard, D, Schatz, B and McKey, DB (2010) Ant nest architecture and seed burial depth: implications for seed fate and germination success in a myrmecochorous savanna shrub. Écoscience 17, 194–202.CrossRefGoogle Scholar
Rezvani, M, Zaefarian, F and Amini, V (2014) Effects of chemical treatments and environmental factors on seed dormancy and germination of shepherd's purse (Capsella bursa-pastoris (L.) Medic.). Acta Botanica Brasilica 28, 495–501.CrossRefGoogle Scholar
Rival, L and McKey, D (2008) Domestication and diversity in manioc (Manihot esculenta Crantz ssp. esculenta, Euphorbiaceae). Current Anthropology 49, 1119–1128.CrossRefGoogle Scholar
Roca, J, Jaureguiberry, P and Gurvich, DE (2021) Are wildfires affecting seed germination in cactus? An experimental assessment. Austral Ecology 46, 818–832.CrossRefGoogle Scholar
Rogers, DJ and Appan, SG (1973) Manihot and Manihotoides (Euphorbiaceae): A computer-assisted study. Flora Neotropica 13, 1–272.Google Scholar
Ruprecht, E, Lukács, K, Domokos, P, Kuhn, T and Fenesi, A (2016) Hydration status influences seed fire tolerance in temperate European herbaceous species. Plant Biology 18, 295–300.CrossRefGoogle ScholarPubMed
Saatkamp, A, Cochrane, A, Commander, L, Guja, LK, Jimenez-Alfaro, B, Larson, J, Nicotra, A, Poschlod, P, Silveira, FAO, Cross, AT, Dalziell, EL, Dickie, J, Erickson, TD, Fidelis, A, Fuchs, A, Golos, PJ, Hope, M, Lewandrowski, W, Merritt, DJ, Miller, BP, Miller, RG, Offord, CA, Ooi, MKJ, Satyanti, A, Sommerville, KD, Tangney, R, Tomlinson, S, Turner, S and Walck, JL (2019) A research agenda for seed-trait functional ecology. New Phytologist 221, 1764–1775.CrossRefGoogle ScholarPubMed
Sala, OE, Stuart Chapin, FIII, Armesto, JJ, Berlow, E, Bloomfield, J, Dirzo, R, Huber-Sanwald, E, Huenneke, LF, Jackson, RB, Kinzig, A, Leemans, R, Lodge, DM, Mooney, HA, Oesterheld, M, Poff, NL, Sykes, MT, Walker, BH, Walker, M and Wall, DH (2000) Global biodiversity scenarios for the year 2100. Science 287, 1770–1774.CrossRefGoogle ScholarPubMed
Schofield, CJ and Bucher, EH (1986) Industrial contributions to desertification in South America. Trends in Ecology & Evolution 1, 78–80.CrossRefGoogle ScholarPubMed
Soltani, E, Baskin, CC and Baskin, JM (2017) A graphical method for identifying the six types of non-deep physiological dormancy in seeds. Plant Biology 19, 673–682.CrossRefGoogle ScholarPubMed
Suda, CN and Giorgini, JF (2000) Seed reserve composition and mobilization during germination and initial seedling development of Euphorbia heterophylla. Revista Brasileira de Fisiologia Vegetal 12, 226–245.CrossRefGoogle Scholar
Takaki, M (2001) New proposal of classification of seeds based on forms of phytochrome instead of photoblastism. Revista Brasileira de Fisiologia Vegetal 13, 104–108.CrossRefGoogle Scholar
Tangney, R, Merritt, DJ, Fontaine, JB and Miller, BP (2019) Seed moisture content as a primary trait regulating the lethal temperature thresholds of seeds. Journal of Ecology 107, 1093–1105.CrossRefGoogle Scholar
Tangney, R, Merritt, DJ, Callow, JN, Fontaine, JB and Miller, BP (2020) Seed traits determine species’ responses to fire under varying soil heating scenarios. Functional Ecology 34, 1967–1978.CrossRefGoogle Scholar
Thanos, CA, Georghiou, K, Douma, DJ and Marangaki, CJ (1991) Photoinhibition of seed germination in Mediterranean maritime plants. Annals of Botany 68, 469–475.CrossRefGoogle Scholar
Tweddle, JC, Dickie, JB, Baskin, CC and Baskin, JM (2003) Ecological aspects of seed desiccation sensitivity. Journal of Ecology 91, 294–304.CrossRefGoogle Scholar
Vander Wall, SB and Longland, WS (2004) Diplochory: are two seed dispersers better than one? Trends in Ecology & Evolution 19, 155–161.CrossRefGoogle ScholarPubMed
van Tooren, BF (1988) The fate of seeds after dispersal in chalk grassland: the role of the bryophyte layer. Oikos 53, 41–48.CrossRefGoogle Scholar
Venier, P, Cabido, M and Funes, G (2017) Germination characteristics of five coexisting neotropical species of Acacia in seasonally dry Chaco forests in Argentina. Plant Species Biology 32, 134–146.CrossRefGoogle Scholar
Walck, JL, Hidayati, SN, Dixon, KW, Thompson, KEN and Poschlod, P (2011) Climate change and plant regeneration from seed. Global Change Biology 17, 2145–2161.CrossRefGoogle Scholar
Whitworth-Hulse, JI, Magliano, PN, Zeballos, SR, Gurvich, DE, Spalazzi, F and Kowaljow, E (2020) Advantages of rainfall partitioning by the global invader Ligustrum lucidum over the dominant native Lithraea molleoides in a dry forest. Agricultural and Forest Meteorology 290, 108013.CrossRefGoogle Scholar
Zeballos, SR, Giorgis, MA, Cabido, MR, Acosta, ATR, Iglesias, MDR and Cantero, JJ (2020) The lowland seasonally dry subtropical forests in central Argentina: vegetation types and a call for conservation. Vegetation Classification and Survey 1, 87–102.CrossRefGoogle Scholar
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