Invaders break assembly rules to beat the natives: How cheatgrass cheats
Data files
Dec 04, 2024 version files 105.57 KB
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FIELD_DATA_FOR_DRYAD.xlsx
72.08 KB
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GREENHOUSE_DATA_FOR_DRYAD.xlsx
26.86 KB
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README.md
6.64 KB
Abstract
Understanding how some introduced plants achieve invasive status while most simply become naturalized is a fundamental question in invasion ecology. Traditional approaches comparing native and introduced plants have linked ruderal traits such as annual life history, high fecundity, and rapid growth rates to invasiveness. However, they do not explain why other introduced species bearing similar traits fail to become invasive, possibly because generic comparisons ignore local processes that drive community assembly. Herein, we contrasted native and introduced annuals in the context of local successional processes to elucidate how introduced annual bromes like cheatgrass (Bromus tectorum) overtake perennial grasslands in the intermountain western United States. We created disturbed plots and seeded them first with annuals representing natives, naturalized species, or invasive bromes. We then seeded plots with native perennial community dominants to examine how the different annuals influenced succession. Native annuals established transient populations that facilitated perennial establishment compared to unseeded controls, enabling the shift to perennial dominance. Naturalized annuals mirrored the natives, but invasive annuals maintained robust populations at high biomass that inhibited perennial establishment and impeded succession. Mechanistically, invasive annuals reduced soil moisture and elevated plant biomass, litter, and soil N. However, only litter abundance correlated with perennial seedling recruitment across treatments. Overall, litter showed a unimodal relationship wherein lower litter abundance associated with native and naturalized annuals appeared to facilitate perennial seedling establishment while higher litter levels generated by invasives appeared to suppress perennial establishment and inhibit succession. Additional experiments provided little support for the roles of pathogen spill-over or plant-soil feedbacks favoring the introduced bromes. The domination of perennial grasslands by annual bromes may be driven by litter buildup that allows these introduced plants to break local succession rules by acting as both early seral and climax species. Traits like litter accumulation may strongly influence invasion outcomes but are indistinguishable using trait comparisons lacking community context.
README: Invaders break assembly rules to beat the natives: How cheatgrass cheats
https://doi.org/10.5061/dryad.9ghx3fft6
Description of the data and file structure
These data assay plant community succession via field and greenhouse experiments to test whether processes diverge between communities initiated with invasive annual bromes or native and naturalized annuals. In the files described further below, note that cases where no data were collected are indicated with a period (".")
Files and variables
File: FIELD_DATA_FOR_DRYAD.xlsx
Description:
DATA FROM MAIN FIELD EXPERIMENT AND EMBEDDED SEED FATE EXPERIMENT
Variables
- FIELD EXP TAB
YEAR: SAMPLING YEAR, 2016-2019
PLOT: EXPERIMENTAL UNIT UNIQUELY LABELED
SPECIES: FOUR LETTER CODE REPRESENTING FOCAL ANNUAL SPECIES ORIGINALLY SEEDED INTO PLOT (SEE FIG. S1 FOR KEY)
TREAT: ANNUAL SPECIES CATEGORY REPRESENTED BY FOCAL SEEDED SPECIES, WITH CNTL=UNSEEDED CONTROL, INVA=INVASIVE, NATU=NATURALIZED, AND NATV = NATIVE
FOCAL: COVER (%) OF ANNUAL SPECIES SEEDED INTO PLOT (NOTE: TRACE LEVELS CODED AS 0.1% FOR ALL COVER VARIABLES, AND COVER OF BROMUS JAPONICUS AND B. TECTORUM LUMPED GIVEN DIFFICULTY OF SPLITTING THESE TAXA)
FOCAL_T: TOTAL COVER (%) OF FOCAL ANNUALS COMPRISED OF FOCAL SPECIES SEEDED INTO PLOT PLUS COVER OF FUNCTIONALLY SIMILAR FOCAL SPECIES COLONIZING FROM OTHER PLOTS
COLNZ: COVER (%) OF NATIVE AND NATURLIZED FOCAL ANNUALS COLONIZING FROM OTHER PLOTS
COLNZ_MEAN: MEAN COVER (%) PER COLONIZING NATIVE OR NATURALIZED FOCAL ANNUAL SPECIES
LITTER: LITTER COVER (%)
N_SEEDL: NUMBER OF SEEDLINGS OF SEEDED NATIVE PERENNIALS COUNTED IN THE FIRST GROWING SEASON AFTER SOWING, 2017 AND 2018
NH4: SOIL AMMONIUM MEASURED IN 2018
NO3: SOIL NITRATE MEASURED IN 2018
SOM: SOIL ORGANIC MATTER (LOI %) MEASURED IN 2018
PH: SOIL PH MEASURED IN 2018
P: SOIL PHOSPHORUS (PPM) MEASURED IN 2018
K: SOIL POTASSIUM (PPM) MEASURED IN 2018
S: SOIL SULFATE (PPM) MEASURED IN 2018
ZN: SOIL ZINC (PPM) MEASURED IN 2018
FE: SOIL IRON (PPM) MEASURED IN 2018
MN: SOIL MANGANESE (PPM) MEASURED IN 2018
CU: SOIL COPPER (PPM) MEASURED IN 2018
CA: SOIL CALCIUM (PPM) MEASURED IN 2018
MG: SOIL MAGNESIUM (PPM) MEASURED IN 2018
NA: SOIL SODIUM (PPM) MEASURED IN 2018
CEC: SOIL CATION EXCHANGE CAPACITY (ME/100 G) MEASURED IN 2018
VWC: SOIL VOLUMETRIC WATER CONTENT (%) MEASURED IN 2018
BM_FOCAL_19: ABOVEGROUND BIOMASS (G) OF FOCAL SEEDED ANNUAL SPECIES, AS HARVESTED IN 2019 (NOTE: B. JAPONICUS AND B. TECTORUM WERE DINSTIGUISHABLE AT HARVEST AND TREATED AS SEPARATE SPECIES FOR THIS METRIC)
BM_FOCAL_T_19: TOTAL ABOVEGROUND BIOMASS (G) OF FOCAL ANNUALS COMPRISED OF FOCAL SPECIES SEEDED INTO PLOT PLUS COVER OF FUNCTIONALLY SIMILAR FOCAL SPECIES COLONIZING FROM OTHER PLOTS, AS HARVESTED IN 2019
BM_PEREN_19: ABOVEGROUND BIOMASS (G) OF SEEDED PERENNIALS FROM 2018 SEEDLING COHORT, AS HARVESTED IN 2019
BM_LITTER_19: ABOVEGROUND BIOMASS (G) OF LITTER, AS HARVESTED IN 2019
LITTER_PC: PRIMARY PRINCIPLE COMPONENT BUILT FROM LITTER COVER IN THE SAMPLING YEAR (2017 OR 2018, RESPECTIVELY) AND LITTER BIOMASS FROM 2019
LITTER_PC_TX: LITTER_PC SCALED SO THAT MINIMUM NUMBER WAS >0 FOR DISPLAY PURPOSES ONLY
* PSSP SEED FATE TAB
PLOT: EXPERIMENTAL UNIT UNIQUELY LABELED (REPRESENTING SUBSET OF PLOTS FROM MAIN EXPERIMENT)
SPECIES: FOUR LETTER CODE REPRESENTING FOCAL ANNUAL SPECIES ORIGINALLY SEEDED INTO PLOT (SEE FIG. S1 FOR KEY)
TREAT: ANNUAL SPECIES CATEGORY REPRESENTED BY FOCAL SEEDED SPECIES, WITH CNTL=UNSEEDED CONTROL, INVA=INVASIVE, NATU=NATURALIZED, AND NATV = NATIVE
N_EMERG_F: NUMBER OF MARKED PSSP (PSEUDOROEGNERIA SPICATA) SEEDS WITH EVIDENCE OF EMERGENCE IN FALL 2018 COUNT
N_EMERG_SP: NUMBER OF MARKED PSSP SEEDS WITH EVIDENCE OF EMERGENCE IN SPRING 2019 COUNT (EXCLUDES THOSE SCORED AS EMERGING IN FALL)
N_EMERG: TOTAL NUMBER OF MARKED PSSP SEEDS WITH EVIDENCE OF EMERGENCE (FALL 2018 COUNT PLUS SPRING 2019 COUNT)
P_EMERG: PROPORTION OF TOTAL MARKED PSSP SEEDS (OF N=20 PER PLOT) WITH EVIDENCE OF EMERGENCE (FALL 2018 COUNT PLUS SPRING 2019 COUNT)
N_SURV_SP: NUMBER OF MARKED PSSP SEEDLINGS FROM FALL SURVING UNTIL SPRING 2019 COUNT
P_SURV_SP: PROPORTION OF MARKED PSSP SEEDLINGS FROM FALL SURVING UNTIL SPRING 2019 COUNT (N_SURV_SP/N_EMERG_F)
N_RECOV: NUMBER OF MARKED PSSP SEEDS WITH NO EVIDENCE OF EMERGENCE (FROM FALL OR SPRING) THAT WERE RECOVERED FOR LAB INSPECTION
N_PYREN: NUMBER OF RECOVERED PSSP SEEDS WITH SIGNS OF INFECTION BY THE PATHOGEN PYRENOPHORA SEMENIPERDA (NO RECOVERED SEEDS HAD SIGNS OF FUSARIUM INFECTION)
D_PYREN: FOR PLOTS WITH RECOVERED PSSP SEEDS, INDICATES DETECTION OF INFECTION BY THE PATHOGEN PYRENOPHORA SEMENIPERDA (1=DETECTED, 0=NOT DETECTED)
File: GREENHOUSE_DATA_FOR_DRYAD.xlsx
Description:
DATA FROM GREENHOUSE EXPERIMENTS FOCUSED ON PLANT-SOIL FEEDBACKS (PSF)
Variables
- PSF EXP1 TAB - FEEDBACKS OF FOCAL ANNUALS ON THEMSELVES
ROUND: FIRST OR SECOND GREENHOUSE GROWTH PERIOD (TRAINED SOIL FROM EACH ROUND 1 POT WAS USED TO GROW CONSPECIFICS IN THE SAME POT IN ROUND 2)
SPECIES: FOUR LETTER CODE REPRESENTING FOCAL ANNUAL SPECIES PLANTED IN POT (SEE FIG. S1 FOR KEY)
TREAT: ANNUAL SPECIES CATEGORY REPRESENTED BY PLANTED FOCAL SPECIES, WITH INVA=INVASIVE, NATU=NATURALIZED, AND NATV = NATIVE
SOIL_TYPE: LIVE=FIELD SOIL USED IN THE MAIN EXPERIMENT; INERT=POTTING SOIL CONTROL USED TO TEST FOR BROAD CHANGES IN GROWING CONDITIONS BETWEEN ROUNDS
POT_ID: UNIQUE ID FOR EACH POT
NPLANTS: NUMBER OF PLANTS SURVIVING UNTIL THE END OF THE ROUND
MASS_1: SHOOT BIOMASS (G) OF FIRST PLANT WEIGHED AT THE END OF THE ROUND
MASS_2: SHOOT BIOMASS (G) OF SECOND PLANT WEIGHED AT THE END OF THE ROUND
MASS_3: SHOOT BIOMASS (G) OF THIRD PLANT WEIGHED AT THE END OF THE ROUND
MASS_MEAN: MEAN SHOOT BIOMASS (G) PER PLANT AT THE END OF THE ROUND
* PSF EXP2 TAB - FEEDBACKS OF FOCAL ANNUALS ON NATIVE PERENNIALS
SPECIES1: FOUR LETTER CODE REPRESENTING FOCAL ANNUAL SPECIES GROWN IN POT IN ROUND 1 TO TRAIN SOIL (SEE FIG. S1 FOR KEY)
TREAT: ANNUAL SPECIES TYPE REPRESENTED BY FOCAL SPECIES PLANTED IN ROUND 1, WITH BRTE=BROMUS TECTORUM (INVASIVE) AND NOT=EITHER NATURALIZED OR NATIVE SPECIES
SPECIES2: FOUR LETTER CODE REPRESENTING NATIVE PERENNIAL SPECIES GROWN IN TRAINED SOIL IN ROUND 2 (SEE FIG. S4 FOR KEY)
POT_ID: UNIQUE ID FOR EACH POT
MASS2: SHOOT BIOMASS (G) OF NATIVE PERENNIAL PLANT WEIGHED AT THE END OF ROUND 2
Code/software
No specialized code/software needed.
Methods
Data for this study derive from a main field experiment, an embedded seed fate experiment, and two greenhouse experiments. In the field, we primarily measured response variables that assayed plant community development and associated changes in soil conditions. The greenhouse experiments involved the same focal annual and perennial species used in the field experiments, but with a focus on testing for plant-soil feedbacks. Please see README information (included in this archive and within data files) in addition to the published article for detailed methods.