Temperature controls phenology in continuously flowering Protea species of subtropical Africa
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Date
Authors
Daru, Barnabas H.
Kling, Matthew M.
Meineke, Emily K.
Van Wyk, Abraham Erasmus (Braam)
Journal Title
Journal ISSN
Volume Title
Publisher
Wiley Open Access
Abstract
PREMISE OF THE STUDY: Herbarium specimens are increasingly used as records of plant flowering
phenology. However, most herbarium-based
studies on plant phenology focus on taxa
from temperate regions. Here, we explore flowering phenologic responses to climate in the
subtropical plant genus Protea (Proteaceae), an iconic group of plants that flower year-round
and are endemic to subtropical Africa.
METHODS: We present a novel, circular sliding window approach to investigate phenological
patterns developed for species with year-round
flowering. We employ our method to evaluate
the extent to which site-to-
site
and year-to-
year
variation in temperature and precipitation
affect flowering dates using a database of 1727 herbarium records of 25 Protea species.
We also explore phylogenetic conservatism in flowering phenology.
RESULTS: We show that herbarium data combined with our sliding window approach
successfully captured independently reported flowering phenology patterns (r = 0.93).
Both warmer sites and warmer years were associated with earlier flowering of 3–5 days/°C,
whereas precipitation variation had no significant effect on flowering phenology. Although
species vary widely in phenological responsiveness, responses are phylogenetically
conserved, with closely related species tending to shift flowering similarly with increasing
temperature.
DISCUSSION: Our results point to climate-responsive
phenology for this important plant
genus and indicate that the subtropical, aseasonally flowering genus Protea has temperature-driven
flowering responses that are remarkably similar to those of better-studied
northern
temperate plant species, suggesting a generality across biomes that has not been described
elsewhere.
Description
APPENDIX S1. Specimen collection frequency across day of flowering
year (DOFY), a normalized version of the Julian day of year.
Red vertical dashed lines correspond to January 1.
APPENDIX S2. Comparison of species peak flowering season (in Julian days) recorded from herbarium specimen records versus the literature (Rebelo, 2001). Rebelo (2001) reports both a “long” season of increased flowering activity and a narrower “short” season of maximal flowering activity for each species, the centers of which are shown here relative to the peak flowering date we calculated from herbarium data as described in the text. Although the y-axis ranges from 0–365, the x-axis has a slightly broader range—given the circular nature of the calendar year, a given Julian date can take multiple values (e.g., 10 = 375), and the value that best communicates alignment with the field guide data set is shown.
APPENDIX S3. Parameters used to characterize phenologic responsiveness to climate in Protea species, estimated from the mixed effects model.
APPENDIX S4. Changes in flowering times of Protea species across South Africa in relation to anomalies in temperature. Statistical analysis based on mixed effects model using both spatial temperature variation (A) and temporal climate (year-to- year temperature variation) (B) as predictors, with species as random effect. Negative slopes indicate advancement of flowering with warming. Lines indicate fitted slopes for individual Protea species. Points indicate input specimen data, and have been truncated for visualization at the extremes of the y-axis range.
APPENDIX S5. Species-specific statistics generated by the sliding window phenology analysis and the mixed effects model (MEM) climate analysis for each of the 25 Protea species.
APPENDIX S6. Relationship between the aseasonality of species’ annual flowering phenology cycles (aseasonality index) and their estimated phenological responses to temperature variation across space and time (coefficients from the linear mixed effects model). Dashed lines show linear regressions with 95% confidence intervals shaded.
APPENDIX S7. Tests of phylogenetic signal in different dimensions of Protea flowering.
APPENDIX S2. Comparison of species peak flowering season (in Julian days) recorded from herbarium specimen records versus the literature (Rebelo, 2001). Rebelo (2001) reports both a “long” season of increased flowering activity and a narrower “short” season of maximal flowering activity for each species, the centers of which are shown here relative to the peak flowering date we calculated from herbarium data as described in the text. Although the y-axis ranges from 0–365, the x-axis has a slightly broader range—given the circular nature of the calendar year, a given Julian date can take multiple values (e.g., 10 = 375), and the value that best communicates alignment with the field guide data set is shown.
APPENDIX S3. Parameters used to characterize phenologic responsiveness to climate in Protea species, estimated from the mixed effects model.
APPENDIX S4. Changes in flowering times of Protea species across South Africa in relation to anomalies in temperature. Statistical analysis based on mixed effects model using both spatial temperature variation (A) and temporal climate (year-to- year temperature variation) (B) as predictors, with species as random effect. Negative slopes indicate advancement of flowering with warming. Lines indicate fitted slopes for individual Protea species. Points indicate input specimen data, and have been truncated for visualization at the extremes of the y-axis range.
APPENDIX S5. Species-specific statistics generated by the sliding window phenology analysis and the mixed effects model (MEM) climate analysis for each of the 25 Protea species.
APPENDIX S6. Relationship between the aseasonality of species’ annual flowering phenology cycles (aseasonality index) and their estimated phenological responses to temperature variation across space and time (coefficients from the linear mixed effects model). Dashed lines show linear regressions with 95% confidence intervals shaded.
APPENDIX S7. Tests of phylogenetic signal in different dimensions of Protea flowering.
Keywords
Cape Floristic Region, Day of flowering year, Flowering phenology, Global warming, Herbarium specimens, Protea
Sustainable Development Goals
Citation
Daru, B. H., M. M. Kling, E. K. Meineke, and A. E. van
Wyk. 2019. Temperature controls phenology in continuously
flowering Protea species of subtropical Africa. Applications in
Plant Sciences 7(3): e1232.