Introduction

Seed production ensures the persistence of tree populations and forest cover over the long term. Some tree species, like sugar maple (Acer saccharum), produce seeds synchronously at irregular intervals, a phenomenon called masting. Red maple (Acer rubrum), a non-masting species, is hypothesized to exhibit muted reproductive dynamics compared to sugar maple.

In this project, we analyze datasets collected at Harvard Forest since 2011, including flower counts, branch-level samaras, and seed counts, to compare the reproductive dynamics of sugar and red maple species.

Citation: Rapp, J., E. Crone, and K. Stinson. 2023. Maple Reproduction and Sap Flow at Harvard Forest since 2011 ver 6. Environmental Data Initiative. https://doi.org/10.6073/pasta/7c2ddd7b75680980d84478011c5fbba9

Data and Methods

Load Libraries

library(tidyverse)
library(DT)

Load Datasets

library(readr)
hf285_04_maple_flower <- read_csv("knb-lter-hfr/hf285-04-maple-flower.csv")
library(readr)
hf285_06_maple_fall_branch <- read_csv("knb-lter-hfr/hf285-06-maple-fall-branch.csv")
library(readr)
hf285_09_maple_seed_count <- read_csv("knb-lter-hfr/hf285-09-maple-seed-count.csv")

Sample of Data

datatable(head(hf285_04_maple_flower, 10))

This table shows branch-level reproductive observations for sugar and red maples. Each row corresponds to a single internode on a branch, with columns showing the tree ID, branch, canopy position, internode number and year, bud count and position, counts of male, female, and unknown flowers, and leaf presence. Readers can see how flower production varies within branches and among different trees.

datatable(head(hf285_06_maple_fall_branch, 10))

This table records branch-level samara production. Each row includes the tree ID, branch, treatment type (FL = fertilized, NF = non-fertilized), internode year and number, internode length, leaf counts and area, and number of samaras. It provides insight into how branch growth and treatment may influence samara production.

datatable(head(hf285_09_maple_seed_count, 10))

This table shows total seed production for each tree on specific dates. Columns include the tree ID, date of collection, counts of seeds by type (EC and JR), total seed count, and any notes. It allows comparison of reproductive output among trees and provides the basis for analyzing yearly seed production trends.

Data Cleaning

hf285_04_maple_flower <- hf285_04_maple_flower %>%
mutate(year = as.numeric(format(as.Date(date), "%Y")),
total_flowers = num.male + num.female)

Here we extract the year from the date column and calculate total flowers per observation by summing male and female flowers. This prepares the data for yearly summaries.

hf285_06_maple_fall_branch <- hf285_06_maple_fall_branch %>%
mutate(year = as.numeric(format(as.Date(date), "%Y")))

We extract the year for the samara dataset to enable grouping by year for analysis.

hf285_09_maple_seed_count <- hf285_09_maple_seed_count %>%
mutate(year = as.numeric(format(as.Date(date), "%Y")))

We extract the year for the samara dataset to enable grouping by year for analysis.

Yearly Summaries

flowers_summary <- hf285_04_maple_flower %>%
group_by(tree, year) %>%
summarise(total_flowers = sum(total_flowers, na.rm = TRUE))

branch_summary <- hf285_06_maple_fall_branch %>%
group_by(tree, year) %>%
summarise(total_samaras = sum(num.samaras, na.rm = TRUE))

seed_summary <- hf285_09_maple_seed_count %>%
group_by(tree, year) %>%
summarise(total_seeds = sum(total.count, na.rm = TRUE))

We aggregate flower, samara, and seed counts by tree and year to obtain total reproductive output per tree for each year.

Merge All Data

reproduction <- flowers_summary %>%
full_join(branch_summary, by = c("tree", "year")) %>%
full_join(seed_summary, by = c("tree", "year"))

These summaries are combined into a single dataset to allow joint analysis of flowers, samaras, and seeds for each tree and year.

Species Mapping

species_map <- tibble(
tree = unique(reproduction$tree)
) %>%
mutate(
species = ifelse(tree == "HF1" | tree == "HF4" | tree == "HF5" | tree == "HF6" |
tree == "HF7" | tree == "HF9" | tree == "HF10" | tree == "HF12" |
tree == "HF13" | tree == "HF16",
"Sugar",
ifelse(tree == "HF38" | tree == "HF41" | tree == "HF42" | tree == "HF43" |
tree == "HF44" | tree == "HF45",
"Red",
NA))
)
reproduction <- left_join(reproduction, species_map, by = "tree")
sum(is.na(reproduction$species))
## [1] 97

We add a species column to indicate whether each tree is sugar or red maple, which is necessary for species-level comparisons.

Species-Level Summary

species_summary <- reproduction %>%
group_by(species, year) %>%
summarise(
mean_flowers = mean(total_flowers, na.rm = TRUE),
sd_flowers = sd(total_flowers, na.rm = TRUE),
mean_samaras = mean(total_samaras, na.rm = TRUE),
mean_seeds = mean(total_seeds, na.rm = TRUE),
sd_seeds = sd(total_seeds, na.rm = TRUE)
)

Finally, we calculate yearly mean and standard deviation of flowers, samaras, and seeds by species, which will be used for plotting and interpreting reproductive patterns.

Results

Seed Production Over Time

species_summary_filtered <- species_summary %>% filter(!is.na(species))
ggplot(species_summary_filtered, aes(x = year, y = mean_seeds, color = species)) +
geom_line(size = 1.2) +
geom_point(size = 2) +
labs(
title = "Yearly Seed Production by Species",
x = "Year",
y = "Mean Seed Count",
color = "Species"
)

Seed production over time shows differences between sugar maple and red maple. Sugar maple has clear peaks, with the highest seed production in 2011 at 44 seeds per tree, followed by lower production in other years. Red maple also peaks in 2011 at 51 seeds per tree, but its production in later years is irregular, sometimes zero and sometimes moderate, like in 2017. This shows that sugar maple reproduces in a more regular, synchronized pattern, while red maple has irregular and smaller peaks, consistent with it being a non-masting species.

Flower Production Over Time

ggplot(species_summary_filtered, aes(x = year, y = mean_flowers, color = species)) +
geom_point(size = 3) +
labs(
title = "Yearly Flower Production by Species",
x = "Year",
y = "Mean Flower Count",
color = "Species"
)

Flower production follows a similar pattern. Sugar maple flowers the most in 2011 with 2,530 flowers per tree, drops sharply in 2012, and has a small increase in 2014. Red maple has an even higher peak in 2011 at 2,961 flowers but then drops sharply and mostly stays low in later years. This suggests that red maple can occasionally produce many flowers, but it does not do so consistently like sugar maple.

Samara Production Over Time

ggplot(species_summary_filtered, aes(x = year, y = mean_samaras, color = species)) +
geom_line(size = 1.2) +
geom_point(size = 2) +
labs(
title = "Yearly Samara Production by Species",
x = "Year",
y = "Mean Samara Count",
color = "Species"
)

Samara production also reflects these trends. Sugar maple has moderate samara counts in 2011, with lower numbers in following years. Red maple has the highest samaras in 2011 at 114, with smaller peaks in 2013 and 2014, but is mostly low in other years. This shows that sugar maple reproduces in strong, synchronized bursts, while red maple reproduces irregularly.

Summary Table

datatable(species_summary_filtered %>% select(species, year, mean_flowers, sd_flowers, mean_samaras, mean_seeds, sd_seeds))

The summary table confirms these patterns. Sugar maple has clear, synchronized peaks, while red maple has smaller and more irregular peaks, with many years of low or zero production. Overall, sugar maple shows typical masting behavior, and red maple shows muted, irregular reproductive dynamics. This supports the idea that red maple does not mast like sugar maple and only occasionally has higher reproductive output.

Conclusion

Based on flower, samara, and seed data, sugar maple shows clear peaks in reproduction, especially in 2011, while red maple has smaller, irregular peaks with many years of low production. This indicates that sugar maple follows typical masting behavior, reproducing in coordinated bursts, whereas red maple reproduces irregularly and does not mast. These results support the idea that red maple has muted reproductive dynamics compared to sugar maple. One limitation of this analysis is the presence of missing data for several years and trees, which may affect the accuracy of the calculated averages and observed trends. Some years have no recorded flower or seed counts for certain trees, making it harder to fully capture reproductive patterns. Acknowledging these gaps is important, and future studies could aim to collect more complete datasets to strengthen the conclusions.