Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory

Rebecca A Nelson; Lauren L Sullivan; Erika I Hersch-Green; Eric W Seabloom; Elizabeth T Borer; Pedro M Tognetti; Peter B Adler; Lori Biederman; Miguel N Bugalho; Maria C Caldeira; Juan P Cancela; Luísa G Carvalheiro; Jane A Catford; Chris R Dickman; Aleksandra J Dolezal; Ian Donohue; Anne Ebeling; Nico Eisenhauer; Kenneth J Elgersma; Anu Eskelinen; Catalina Estrada; Magda Garbowski; Pamela Graff; Daniel S Gruner; Nicole Hagenah; Sylvia Haider; W Stanley Harpole; Yann Hautier; Anke Jentsch; Nicolina Johanson; Sally E Koerner; Lucíola S Lannes; Andrew S MacDougall; Holly Martinson; John W Morgan; Harry Olde Venterink; Devyn Orr; Brooke B Osborne; Pablo L Peri; Sally A Power; Xavier Raynaud; Anita C Risch; Mani Shrestha; Nicholas G Smith; Carly J Stevens; G F Ciska Veen; Risto Virtanen; Glenda M Wardle; Amelia A Wolf; Alyssa L Young; Susan P Harrison

doi:10.1038/s42003-025-07882-7

Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory

Rebecca A Nelson^*
, Lauren L Sullivan
, Erika I Hersch-Green
, Eric W Seabloom
, Elizabeth T Borer
, Pedro M Tognetti
, Peter B Adler
, Lori Biederman
, Miguel N Bugalho
, Maria C Caldeira
, Juan P Cancela
, Luísa G Carvalheiro
, Jane A Catford
, Chris R Dickman
, Aleksandra J Dolezal
, Ian Donohue
, Anne Ebeling
, Nico Eisenhauer
, Kenneth J Elgersma
, Anu Eskelinen

Catalina Estrada, Magda Garbowski, Pamela Graff, Daniel S Gruner, Nicole Hagenah, Sylvia Haider, W Stanley Harpole, Yann Hautier, Anke Jentsch, Nicolina Johanson, Sally E Koerner, Lucíola S Lannes, Andrew S MacDougall, Holly Martinson, John W Morgan, Harry Olde Venterink, Devyn Orr, Brooke B Osborne, Pablo L Peri, Sally A Power, Xavier Raynaud, Anita C Risch, Mani Shrestha, Nicholas G Smith, Carly J Stevens, G F Ciska Veen, Risto Virtanen, Glenda M Wardle, Amelia A Wolf, Alyssa L Young, Susan P Harrison

^*Corresponding author for this work

Research output: Contribution to journal › Article › Academic › peer-review

Abstract

Forbs ("wildflowers") are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects.

Original language	English
Article number	444
Journal	Communications Biology
Volume	8
Issue number	1
DOIs	https://doi.org/10.1038/s42003-025-07882-7
Publication status	Published - 15 Mar 2025

Bibliographical note

Publisher Copyright:
© The Author(s) 2025.

Funding

Author contributions, site acknowledgments, and funding acknowledgments are documented in Supplementary Data . The authors would like to thank Dr. Neal Williams and Dr. Fernanda Valdovinos for providing additional feedback on this manuscript as well as the editors and peer reviewers. The authors would like to thank the Dirzo Lab at Stanford University for providing input on this research. This work was generated using data from the Nutrient Network ( http://www.nutnet.org ) experiment, funded at the site-scale by individual researchers (see Supplemental Data ). Coordination and data management have been supported by funding to E. Borer and E. Seabloom from the National Science Foundation Research Coordination Network (NSF-DEB-1042132) and Long Term Ecological Research (NSF-DEB-1234162 and NSF-DEB-1831944 to Cedar Creek LTER) programs, and the Institute on the Environment (DG-0001-13). We also thank the Minnesota Supercomputer Institute for hosting project data and the Institute on the Environment for hosting Nutrient Network meetings.

Funders	Funder number
National Science Foundation	NSF-DEB-1042132
Long Term Ecological Research	NSF-DEB-1831944, NSF-DEB-1234162
Institute on the Environment	DG-0001-13

Keywords

Animals
Biodiversity
Grassland
Herbivory
Mammals/physiology
Nitrogen/metabolism
Nutrients
Poaceae

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Cite this

Nelson, R. A., Sullivan, L. L., Hersch-Green, E. I., Seabloom, E. W., Borer, E. T., Tognetti, P. M., Adler, P. B., Biederman, L., Bugalho, M. N., Caldeira, M. C., Cancela, J. P., Carvalheiro, L. G., Catford, J. A., Dickman, C. R., Dolezal, A. J., Donohue, I., Ebeling, A., Eisenhauer, N., Elgersma, K. J., ... Harrison, S. P. (2025). Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory. Communications Biology, 8(1), Article 444. https://doi.org/10.1038/s42003-025-07882-7

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title = "Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory",

abstract = "Forbs ({"}wildflowers{"}) are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects.",

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doi = "10.1038/s42003-025-07882-7",

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Nelson, RA, Sullivan, LL, Hersch-Green, EI, Seabloom, EW, Borer, ET, Tognetti, PM, Adler, PB, Biederman, L, Bugalho, MN, Caldeira, MC, Cancela, JP, Carvalheiro, LG, Catford, JA, Dickman, CR, Dolezal, AJ, Donohue, I, Ebeling, A, Eisenhauer, N, Elgersma, KJ, Eskelinen, A, Estrada, C, Garbowski, M, Graff, P, Gruner, DS, Hagenah, N, Haider, S, Harpole, WS, Hautier, Y, Jentsch, A, Johanson, N, Koerner, SE, Lannes, LS, MacDougall, AS, Martinson, H, Morgan, JW, Olde Venterink, H, Orr, D, Osborne, BB, Peri, PL, Power, SA, Raynaud, X, Risch, AC, Shrestha, M, Smith, NG, Stevens, CJ, Veen, GFC, Virtanen, R, Wardle, GM, Wolf, AA, Young, AL & Harrison, SP 2025, 'Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory', Communications Biology, vol. 8, no. 1, 444. https://doi.org/10.1038/s42003-025-07882-7

TY - JOUR

T1 - Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory

AU - Nelson, Rebecca A

AU - Sullivan, Lauren L

AU - Hersch-Green, Erika I

AU - Seabloom, Eric W

AU - Borer, Elizabeth T

AU - Tognetti, Pedro M

AU - Adler, Peter B

AU - Biederman, Lori

AU - Bugalho, Miguel N

AU - Caldeira, Maria C

AU - Cancela, Juan P

AU - Carvalheiro, Luísa G

AU - Catford, Jane A

AU - Dickman, Chris R

AU - Dolezal, Aleksandra J

AU - Donohue, Ian

AU - Ebeling, Anne

AU - Eisenhauer, Nico

AU - Elgersma, Kenneth J

AU - Eskelinen, Anu

AU - Estrada, Catalina

AU - Garbowski, Magda

AU - Graff, Pamela

AU - Gruner, Daniel S

AU - Hagenah, Nicole

AU - Haider, Sylvia

AU - Harpole, W Stanley

AU - Hautier, Yann

AU - Jentsch, Anke

AU - Johanson, Nicolina

AU - Koerner, Sally E

AU - Lannes, Lucíola S

AU - MacDougall, Andrew S

AU - Martinson, Holly

AU - Morgan, John W

AU - Olde Venterink, Harry

AU - Orr, Devyn

AU - Osborne, Brooke B

AU - Peri, Pablo L

AU - Power, Sally A

AU - Raynaud, Xavier

AU - Risch, Anita C

AU - Shrestha, Mani

AU - Smith, Nicholas G

AU - Stevens, Carly J

AU - Veen, G F Ciska

AU - Virtanen, Risto

AU - Wardle, Glenda M

AU - Wolf, Amelia A

AU - Young, Alyssa L

AU - Harrison, Susan P

PY - 2025/3/15

Y1 - 2025/3/15

N2 - Forbs ("wildflowers") are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects.

AB - Forbs ("wildflowers") are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects.

KW - Animals

KW - Biodiversity

KW - Grassland

KW - Herbivory

KW - Mammals/physiology

KW - Nitrogen/metabolism

KW - Nutrients

KW - Poaceae

UR - https://www.scopus.com/pages/publications/105000362524

U2 - 10.1038/s42003-025-07882-7

DO - 10.1038/s42003-025-07882-7

M3 - Article

C2 - 40089613

SN - 2399-3642

VL - 8

JO - Communications Biology

JF - Communications Biology

IS - 1

M1 - 444

ER -

Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory

Abstract

Bibliographical note

Funding

Keywords

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