HomeAll CategoriesSpecies Richness Calculator (Free, with Chao1) | StatsUnlock

Species Richness Calculator (Free, with Chao1) | StatsUnlock

Species Richness Calculator (Free, with Chao1) | StatsUnlock

Species Richness Calculator

A free online tool to measure species richness (S) and estimate true richness with Margalef, Menhinick, Chao1, and jackknife from your survey counts, with instant charts and report-ready results.

BiodiversitySpecies RichnessChao1EcologyFreeOnline

Last updated: 15 September 2026 · Built for ecologists and wildlife researchers by StatsUnlock

⚡ 0. Quick Answer

The species richness calculator counts how many species are in your sample and estimates how many you missed. Enter counts such as 45, 30, 15, 7, 3, 2, 2, 1, 1, 1 and the tool returns observed richness S = 10, Chao1 = 11.0, Margalef 1.926 and Menhinick 0.967, with sample completeness at 91 percent.

🌿 Key Takeaways

  • The species richness calculator counts how many species are in your sample and estimates how many you may have missed.
  • You only need the count of individuals per species. Paste them comma separated, upload a CSV file, or type them in.
  • It returns observed richness (S) plus Margalef and Menhinick indices, which adjust for sample size, and Chao1 and jackknife estimators of true richness.
  • A higher richness means more species; the estimators tell you whether more sampling would likely reveal even more species.
  • Typical field uses: camera trapping, point counts, vegetation plots, transects, and reef fish surveys.

🌱 1. What Is Species Richness?

Species richness is the simplest biodiversity measure there is: the number of different species in a defined area or sample, written S. It ignores abundance entirely. One elephant and one thousand ants count as two species, exactly the same as one thousand of each.

That simplicity is both its strength and its weakness. Everyone understands it, and it needs no maths. But it depends completely on how hard you looked. Survey a forest for one hour and you find 12 species; survey it for a week and you find 40. The number changed, the forest did not.

This is why richness estimators exist. Margalef and Menhinick scale S by sample size so sites surveyed with different effort can be compared. Chao1 and the jackknife estimator go further: they use the rare species in your sample, the ones seen only once or twice, to estimate how many species you never detected at all. If your sample holds many singletons, there are almost certainly more species out there.

observed S = 10 Chao1 estimate = 11.0 Sampling effort (individuals, trap nights, quadrats) Species found still climbing: keep sampling flattening: nearly complete
The species accumulation curve. Observed richness climbs with effort; the gap to the Chao1 line is the species you have not found yet.

1.1 The five numbers this tool reports

MeasureSymbolWhat it doesUnits
Observed richnessSCounts the species you actually recordedSpecies count
Margalef's indexDMgScales S by the log of sample sizeUnitless
Menhinick's indexDMnScales S by the square root of sample sizeUnitless
Chao1-Estimates true richness from singletons and doubletonsSpecies count
Jackknife 1-A second estimator of true richness, usually higher than Chao1Species count

What to conclude: report S first, because it is what readers expect. Report Chao1 beside it, because the gap between the two is the honest measure of how complete your survey was.

1.2 Reading sample completeness

S / Chao1ReadingWhat to do
Above 95 percentNearly complete GoodReport richness with confidence
85 to 95 percentGood GoodUsable. Note the estimate alongside S.
70 to 85 percentPartial WatchAdd effort, or compare sites by Chao1 rather than S
Below 70 percentUndersampled ConcernDo not compare raw S between sites. Rarefy or keep surveying.

What to conclude: the worked example on this page reaches 91 percent, which is good. A site at 60 percent with 20 species may well be richer than a site at 98 percent with 24.

📥 2. Set Up Your Data

0 values entered

Supports .csv and .txt, headers detected automatically.
SpeciesCount

🔎 4. Interpretation of Results, In Detail

▶ Run the analysis above to generate a detailed, plain-English interpretation of your species richness result.

✍️ 5. How to Write Your Results in Research

▶ Run the analysis above to auto-fill all six examples with your results.

🪧 5.1 Research Poster Panel

A print-ready poster layout that updates with your result. Copy the text or use the visual scaffold for A0 / A1 conference posters.

▶ Run the analysis above to build your poster panel.

🧮 6. Formulas Used

Every number this species richness calculator reports comes from one of the seven formulas below. Card 1 is richness itself, and each later card corrects it for effort or estimates what you missed.

1

Observed species richness

S = count of species with ni > 0
  • S observed richness, a whole number of species
  • ni count of individuals of species i
  • Range: 1 upward. There is no maximum.
  • Requires: a species list. Counts are optional for S, but needed for every card below.
  • Example: the worked dataset on this page gives S = 10.
2

Margalef's richness index

DMg = (S - 1) / ln N
  • N total individuals across all species
  • ln N natural logarithm of the total
  • Use for: comparing sites where N differs. The log dampens the effect of large samples.
  • Example: (10 - 1) / ln 107 = 9 / 4.673 = 1.926.
  • Note: undefined when N = 1, because ln 1 = 0.
3

Menhinick's richness index

DMn = S / √N
  • Use for: the same job as Margalef, with a stronger correction for sample size.
  • Example: 10 / √107 = 10 / 10.344 = 0.967.
  • Rule: pick one of Margalef or Menhinick and use it consistently. They do not agree, and mixing them across sites is meaningless.
4

Chao1 estimated richness (bias-corrected form)

Chao1 = S + [ F1(F1 - 1) ] / [ 2(F2 + 1) ]
  • F1 singletons, species seen exactly once
  • F2 doubletons, species seen exactly twice
  • Range: S upward. It can never be below observed richness.
  • Example: 10 + (3 × 2) / (2 × 3) = 10 + 1 = 11.0.
  • Note: this tool uses the bias-corrected form, which is safe when doubletons are few or absent.
5

Chao1, the classic form

Chao1 = S + F1² / (2 F2)
  • Use for: matching older papers that quote this version.
  • Example: 10 + 9 / 4 = 12.25, higher than the bias-corrected 11.0.
  • Rule: it breaks when F2 = 0, because you cannot divide by zero. That is exactly why the corrected form exists.
6

First-order jackknife estimate

Jack1 = S + F1 × (N - 1) / N
  • Use for: a second opinion on true richness. It leans entirely on singletons.
  • Example: 10 + 3 × (106 / 107) = 12.97.
  • Note: usually higher than Chao1. If the two disagree sharply, your sample is too small to trust either.
7

Sample completeness

Completeness = (S / Chao1) × 100
  • Range: 0 to 100 percent.
  • Example: (10 / 11.0) × 100 = 91 percent.
  • Use for: deciding whether you can compare raw richness between sites at all.
  • Rule: below 70 percent, compare estimated richness rather than observed.

🧭 7. How to Use This Calculator

Ten steps from a field sheet to a publication-ready species richness result.

  1. Collect standardised counts. Record every species and the number of individuals, using the same effort at every site.
  2. Record rare species carefully. Chao1 and the jackknife depend on species seen once or twice, so a missed singleton changes your estimate.
  3. Pick an input tab. Use Paste or Type for a quick run, Upload CSV for a spreadsheet, or Manual Table to key in rows.
  4. Enter species names and counts. In Column Entry mode, put the species name on the left and the count on the right.
  5. Or load a sample dataset. Choose one of the built-in field datasets to see how the tool behaves before you use your own data.
  6. Name your study area. Type the site name into the Study Area box so it appears in every report template and in the poster panel.
  7. Choose which estimator to highlight. Observed S is the default. Switch to Chao1 when your survey is incomplete.
  8. Click Run Analysis. The tool returns S, Margalef, Menhinick, Chao1, jackknife, completeness, charts and a results table.
  9. Check sample completeness first. If it is below 70 percent, add effort before you compare sites.
  10. Copy or download your result. Use the report templates, the poster panel, Download Doc or Download PDF for your thesis or paper.

📗 8. How to Calculate Species Richness in Excel

Excel, step by step, with the same worked example used everywhere on this page

The worked example below uses ten species with counts 45, 30, 15, 7, 3, 2, 2, 1, 1, 1 (N = 107, three singletons, two doubletons). Excel, R, Python and this calculator all return S = 10, Chao1 = 11.0, Margalef = 1.926 and Menhinick = 0.967 for it.

8.1 The Excel functions you need

FunctionWhat it doesExample
COUNTIFCounts species present, giving S, and counts singletons and doubletons=COUNTIF(B2:B11,">0")
SUMAdds your counts to get N=SUM(B2:B11)
LNNatural logarithm, the denominator of Margalef=(B12-1)/LN(B13)
SQRTSquare root, the denominator of Menhinick=B12/SQRT(B13)
COUNTACounts species names when you have a list but no counts=COUNTA(A2:A11)
COUNTIFSCounts species present at one site in a multi-site sheet=COUNTIFS(C:C,"Site A",B:B,">0")

What to conclude: COUNTIF does nearly all the work here. Species richness is the one biodiversity measure you can calculate without a single multiplication.

8.2 Step by step in the sheet

Step 1. Count the species that are actually present

=COUNTIF(B2:B11,">0")=SUM(B2:B11)
0") in cell B12">richness.xlsx - ExcelB12fx=COUNTIF(B2:B11,">0")ABCDESpeciesCountIs singleton?Is doubleton?12Bulbul453Babbler304Sunbird155Barbet76Drongo37Warbler28Thrush29Owlet110Pitta111Trogon112S (species)10

Species names go in column A, counts in column B. Cell B12 holds =COUNTIF(B2:B11,">0") and returns S = 10. Use COUNTIF rather than COUNTA, so a species recorded as zero is not counted as present.

Step 2. Flag the singletons

=IF(B2=1,1,0)fill down to C11
richness.xlsx - ExcelC2fx=IF(B2=1,1,0)ABCDESpeciesCountIs singleton?Is doubleton?12Bulbul4503Babbler3004Sunbird1505Barbet706Drongo307Warbler208Thrush209Owlet1110Pitta1111Trogon11

C2 holds =IF(B2=1,1,0). Three species in this dataset were seen exactly once, shown in red. These singletons are what Chao1 runs on.

Step 3. Total the singletons and doubletons

=SUM(C2:C11)=SUM(D2:D11)=COUNTIF(B2:B11,1)
richness.xlsx - ExcelC12fx=SUM(C2:C11)ABCDESpeciesCountIs singleton?Is doubleton?12Bulbul45003Babbler30004Sunbird15005Barbet7006Drongo3007Warbler2018Thrush2019Owlet11010Pitta11011Trogon11012F1 singletons3

C12 holds =SUM(C2:C11) and returns F1 = 3. Column D does the same for doubletons, giving F2 = 2. The shortcut is =COUNTIF(B2:B11,1), which skips the helper column entirely.

Step 4. Build Chao1, Margalef and Menhinick

=B12+(C12*(C12-1))/(2*(D12+1))=(B12-1)/LN(B13)=B12/SQRT(B13)=B12/E12*100
richness.xlsx - ExcelE12fx=B12+(C12*(C12-1))/(2*(D12+1))ABCDESpeciesCountIs singleton?Is doubleton?12Bulbul45003Babbler30004Sunbird15005Barbet7006Drongo3007Warbler2018Thrush2019Owlet11010Pitta11011Trogon11012Chao111.00

The Chao1 cell returns 11.00. Margalef is =(B12-1)/LN(B13) = 1.926 and Menhinick is =B12/SQRT(B13) = 0.967. Completeness is =B12/E12*100 = 91 percent.

Tip: you can skip the helper columns completely. =COUNTIF(B2:B11,1) gives F1 and =COUNTIF(B2:B11,2) gives F2, so the whole Chao1 calculation fits in three cells.
Warning: COUNTA counts every non-empty cell, including species you recorded as absent with a 0 and any stray note. It will report a richness higher than the truth. Use COUNTIF with a ">0" condition instead, every time.

8.3 The awkward cases

CaseWhat breaksFix
No doubletons at all (F2 = 0)The classic Chao1 divides by zeroUse the bias-corrected form, which adds 1 to F2 in the denominator
No singletons at all (F1 = 0)Chao1 equals S exactlyCorrect, and it means your survey looks complete
Duplicate species rowsThe same species is counted twice, inflating SUse a PivotTable to sum counts by species name first
Morphospecies labels that differ by a space"Warbler sp1" and "Warbler sp 1" count as two speciesRun TRIM and a sort to catch near-duplicates before counting
Species recorded as present with a tick, not a numberYou get S but Margalef, Menhinick and Chao1 are impossibleGo back for counts, or report observed richness only

What to conclude: almost every wrong richness value in Excel is a data-entry problem rather than a formula problem. Sort your species column and read it before you count it.

8.4 Why answers differ between Excel, R and Python

Observed richness never differs; it is a count. The estimators do. R's vegan estimateR() returns the bias-corrected Chao1 by default, which is what this page and this calculator use, giving 11.0. Older papers and some software quote the classic form, F1 squared over 2F2, which gives 12.25 on the same data. Neither is wrong, but they are not interchangeable, so name the form in your methods. The jackknife also comes in individual-based and sample-based versions: this tool uses the individual-based form, while vegan's specpool() uses the sample-based one and needs a site-by-species matrix rather than a single vector.

8.5 Charting the result in Excel

  1. Species accumulation curve. Add a running count of new species as each sample is added, then Insert, Line chart. If it is still climbing at the right-hand edge, keep surveying.
  2. Observed versus estimated bars. Chart S, Chao1 and Jackknife 1 side by side. The gap is the story.
  3. Site comparison. One Margalef value per site in a clustered bar chart, with error bars if you have replicates.
  4. Label the axis honestly. Write "Species richness (S, observed)" so nobody mistakes an observed count for an estimate.

8.6 Excel error messages and what they mean

MessageCauseFix
#DIV/0!N is 1, so LN(N) is zero in the Margalef cellMargalef needs at least two individuals
#VALUE!Text inside a numeric rangeClear notes such as "present" from the count column
#NUM!SQRT received a negative numberCounts cannot be negative. Recheck data entry.
Chao1 below SA sign error in the formulaChao1 can never be below observed richness
S is larger than the species listCOUNTA was used instead of COUNTIFSwitch to COUNTIF with ">0"
Completeness above 100 percentS and Chao1 were swapped in the divisionCompleteness is S divided by Chao1, not the reverse

What to conclude: a Chao1 below S, or a completeness above 100 percent, is arithmetically impossible. Both point at the same swapped-cell mistake.

📘 9. How to Calculate Species Richness in R

R, one complete script you can run top to bottom

9.1 The complete script

Copy this into RStudio and press Run. It needs no packages. Change only the counts line. It prints S, Margalef, Menhinick, Chao1, jackknife and completeness, then saves one 300 dpi figure.

# ------------------------------------------------------------------
# Species Richness and Richness Estimators in R
# StatsUnlock.com - free ecology and wildlife analysis tools
# ------------------------------------------------------------------
set.seed(42)

# 1. YOUR DATA -- change only this line
counts <- c(45, 30, 15, 7, 3, 2, 2, 1, 1, 1)
species <- c("Bulbul", "Babbler", "Sunbird", "Barbet", "Drongo",
             "Warbler", "Thrush", "Owlet", "Pitta", "Trogon")

# Reading from a file instead? Uncomment the two lines below.
# dat <- read.csv("my_survey.csv")          # columns: species, count
# counts <- dat$count; species <- dat$species

# 2. CLEAN THE DATA
counts[is.na(counts)] <- 0        # missing values become zero
keep <- counts > 0                # a species recorded as zero is not present
counts <- counts[keep]
species <- species[keep]

# 3. CALCULATE
S <- length(counts)                       # observed species richness
N <- sum(counts)                          # total individuals
F1 <- sum(counts == 1)                    # singletons
F2 <- sum(counts == 2)                    # doubletons
margalef <- (S - 1) / log(N)              # Margalef's index
menhinick <- S / sqrt(N)                  # Menhinick's index
chao1 <- S + (F1 * (F1 - 1)) / (2 * (F2 + 1))   # bias-corrected Chao1
chao1_classic <- if (F2 > 0) S + F1^2 / (2 * F2) else NA   # classic form
jack1 <- S + F1 * ((N - 1) / N)           # first-order jackknife
completeness <- 100 * S / chao1           # percent of species detected

# Same answers using the vegan package:
# install.packages("vegan")
# library(vegan)
# vegan::estimateR(counts)        # returns S.obs, S.chao1, S.ACE

# 4. PRINT A PLAIN-ENGLISH VERDICT
verdict <- if (completeness >= 95) "survey nearly complete" else
           if (completeness >= 85) "survey good" else
           if (completeness >= 70) "survey partial" else "undersampled, keep surveying"
cat("Observed richness S     :", S, "\n")
cat("Total individuals N     :", N, "\n")
cat("Singletons F1           :", F1, "\n")
cat("Doubletons F2           :", F2, "\n")
cat("Margalef index          :", round(margalef, 3), "\n")
cat("Menhinick index         :", round(menhinick, 3), "\n")
cat("Chao1 (bias-corrected)  :", round(chao1, 2), "\n")
cat("Jackknife 1             :", round(jack1, 2), "\n")
cat("Sample completeness %   :", round(completeness, 1), "\n")
cat("Verdict                 :", verdict, "\n")

# 5. SAVE ONE 300 dpi FIGURE
png("richness_observed_vs_estimated.png", width = 1800, height = 1200, res = 300)
vals <- c(S, chao1, jack1)
bp <- barplot(vals,
              names.arg = c("Observed S", "Chao1", "Jackknife 1"),
              col = c("#27a558", "#2b7fd4", "#8b5cf6"), border = NA,
              ylim = c(0, max(vals) * 1.2),
              ylab = "Number of species",
              main = paste0("Richness, ", round(completeness, 0), "% detected"))
text(bp, vals, labels = round(vals, 1), pos = 3, cex = 0.9)
dev.off()
cat("Figure saved as richness_observed_vs_estimated.png\n")

9.2 What each line does

LineWhat it does
counts <- c(45, 30, ...)Your species counts. This is the only line most users change.
keep <- counts > 0Drops species recorded as absent. Leaving them in inflates S.
S <- length(counts)Observed species richness, after the zero filter.
F1 <- sum(counts == 1)Counts singletons. Every estimator below depends on this number.
F2 <- sum(counts == 2)Counts doubletons.
margalef <- (S - 1) / log(N)Margalef's index. R's log() is the natural log.
menhinick <- S / sqrt(N)Menhinick's index.
chao1 <- S + (F1*(F1-1)) / (2*(F2+1))Bias-corrected Chao1. Safe even when F2 is zero.
jack1 <- S + F1 * ((N-1)/N)First-order jackknife, the individual-based form.
vegan::estimateR(counts)The package version. It returns S.obs, S.chao1 and S.ACE in one call.
dev.off()Closes the device and writes the file. Forget this and the PNG stays empty.

9.3 What the figure shows

The script saves three bars: observed richness, Chao1 and the jackknife, each labelled with its value and the detection percentage in the title. This is the most useful figure for a richness result, because the height difference between the first bar and the other two is your undersampling, made visible. If the three bars are almost level, your species list is close to complete.

9.4 Problems and fixes

ProblemCauseFix
Chao1 equals S exactlyNo singletons in the sampleCorrect behaviour, and usually a sign of good sampling
estimateR gives a different Chao1vegan needs integer counts and uses its own correctionPass as.integer(counts), and compare the bias-corrected values
Error: could not find function "estimateR"vegan is not loadedRun install.packages("vegan") then library(vegan)
specpool() returns an errorIt needs a site-by-species matrix, not one vectorUse estimateR() for a single sample
Margalef is Inf or NaNN is 1, so log(N) is zeroMargalef needs at least two individuals
Jackknife far above Chao1Many singletons, few doubletonsBoth estimators are unstable here. Add survey effort.

9.5 One-liners worth keeping

CodeReturns
vegan::specnumber(x)Observed richness S from a vector or a matrix
vegan::estimateR(x)S.obs, Chao1 and ACE with standard errors
vegan::specpool(matrix)Chao, jackknife 1 and 2, and bootstrap for a set of sites
vegan::specaccum(matrix)A species accumulation curve you can plot directly
vegan::rarefy(x, sample = 100)Richness standardised to 100 individuals
iNEXT::iNEXT(x, q = 0)Rarefaction and extrapolation curves for richness

🐍 10. How to Calculate Species Richness in Python

Python, one complete script you can run top to bottom

Packages: numpy does the arithmetic and matplotlib draws and saves the figure. Install them once with pip install numpy matplotlib. scikit-bio is optional and ships chao1() and osd() for richness work. pandas is only needed if you read a CSV file.

10.1 The complete script

Save this as richness.py and run it. Change only the counts line. It prints every richness measure and saves one 300 dpi figure.

# ------------------------------------------------------------------
# Species Richness and Richness Estimators in Python
# StatsUnlock.com - free ecology and wildlife analysis tools
# ------------------------------------------------------------------
import numpy as np
import matplotlib
matplotlib.use("Agg")
import matplotlib.pyplot as plt

np.random.seed(42)

# 1. YOUR DATA -- change only this line
counts = np.array([45, 30, 15, 7, 3, 2, 2, 1, 1, 1], dtype=float)
species = ["Bulbul", "Babbler", "Sunbird", "Barbet", "Drongo",
           "Warbler", "Thrush", "Owlet", "Pitta", "Trogon"]

# Reading from a file instead? Uncomment the three lines below.
# import pandas as pd
# dat = pd.read_csv("my_survey.csv")          # columns: species, count
# counts, species = dat["count"].to_numpy(float), dat["species"].tolist()

# 2. CLEAN THE DATA
counts = np.nan_to_num(counts, nan=0.0)       # missing values become zero
keep = counts > 0                             # a species recorded as zero is absent
species = [s for s, k in zip(species, keep) if k]
counts = counts[keep]

# 3. CALCULATE
S = int(counts.size)                          # observed species richness
N = float(counts.sum())                       # total individuals
F1 = int(np.sum(counts == 1))                 # singletons
F2 = int(np.sum(counts == 2))                 # doubletons
margalef = (S - 1) / np.log(N)                # Margalef's index
menhinick = S / np.sqrt(N)                    # Menhinick's index
chao1 = S + (F1 * (F1 - 1)) / (2 * (F2 + 1))  # bias-corrected Chao1
chao1_classic = S + F1 ** 2 / (2 * F2) if F2 > 0 else float("nan")
jack1 = S + F1 * ((N - 1) / N)                # first-order jackknife
completeness = 100 * S / chao1                # percent of species detected

# Same answer using scikit-bio:
# from skbio.diversity.alpha import chao1 as sk_chao1
# sk_chao1(counts.astype(int))

# 4. PRINT A PLAIN-ENGLISH VERDICT
if completeness >= 95:
    verdict = "survey nearly complete"
elif completeness >= 85:
    verdict = "survey good"
elif completeness >= 70:
    verdict = "survey partial"
else:
    verdict = "undersampled, keep surveying"
print(f"Observed richness S     : {S}")
print(f"Total individuals N     : {N:.0f}")
print(f"Singletons F1           : {F1}")
print(f"Doubletons F2           : {F2}")
print(f"Margalef index          : {margalef:.3f}")
print(f"Menhinick index         : {menhinick:.3f}")
print(f"Chao1 (bias-corrected)  : {chao1:.2f}")
print(f"Jackknife 1             : {jack1:.2f}")
print(f"Sample completeness %   : {completeness:.1f}")
print(f"Verdict                 : {verdict}")

# 5. SAVE ONE 300 dpi FIGURE
labels = ["Observed S", "Chao1", "Jackknife 1"]
vals = [S, chao1, jack1]
fig, ax = plt.subplots(figsize=(6, 4))
bars = ax.bar(labels, vals, color=["#27a558", "#2b7fd4", "#8b5cf6"])
ax.bar_label(bars, fmt="%.1f", padding=3)
ax.set_ylim(0, max(vals) * 1.2)
ax.set_ylabel("Number of species")
ax.set_title(f"Richness, {completeness:.0f}% detected")
plt.tight_layout()
plt.savefig("richness_observed_vs_estimated.png", dpi=300)
print("Figure saved as richness_observed_vs_estimated.png")

10.2 What each line does

LineWhat it does
matplotlib.use("Agg")Draws without a screen, so the script runs on a server or in a notebook.
counts = np.array([...])Your species counts. The only line most users change.
keep = counts > 0Drops species recorded as absent, which would otherwise inflate S.
S = int(counts.size)Observed species richness after the filter.
F1 = np.sum(counts == 1)Counts singletons, the engine behind every estimator here.
margalef = (S - 1) / np.log(N)Margalef's index. np.log is the natural log.
menhinick = S / np.sqrt(N)Menhinick's index.
chao1 = S + (F1*(F1-1))/(2*(F2+1))Bias-corrected Chao1, safe when F2 is zero.
jack1 = S + F1 * ((N-1)/N)First-order jackknife, individual-based form.
ax.bar_label(bars, fmt="%.1f")Prints the value on top of each bar so the figure reads without a table.
plt.savefig(..., dpi=300)Writes the 300 dpi figure most journals ask for.

10.3 What the figure shows

The saved PNG shows observed richness beside the two estimators, with the detection percentage in the title, identical in meaning to the R figure. For a species accumulation curve instead, shuffle your individual records and plot the running count of unique species against sample number, repeating the shuffle 100 times and averaging.

10.4 Problems and fixes

ProblemCauseFix
skbio chao1 differs from this scriptscikit-bio applies its own bias correction ruleRead the docstring, then compare like with like
ZeroDivisionError on completenessChao1 came out as zero, which needs an empty sampleCheck the array is not empty after the keep filter
F1 is always 0Counts stored as floats, so 1.0 == 1 comparison surprisesThe == 1 test works on floats, but round your data first if it came from a model
RuntimeWarning: divide by zeroN is 1, so np.log(N) is zero in the Margalef lineMargalef needs at least two individuals
ModuleNotFoundError: numpyThe package is not installed in this environmentRun pip install numpy matplotlib
Figure window never opensThe Agg backend has no screen outputThat is expected. Open the saved PNG file.

10.5 One-liners worth keeping

CodeReturns
(c > 0).sum()Observed species richness S
np.sum(c == 1)Singletons F1
skbio.diversity.alpha.chao1(c)Chao1 estimated richness from integer counts
skbio.diversity.alpha.osd(c)Observed richness, singletons and doubletons in one tuple
(len(c)-1)/np.log(c.sum())Margalef's index
len(c)/np.sqrt(c.sum())Menhinick's index

📋 11. Reference Tables

Six lookup tables for reading, estimating and sanity-checking a species richness result.

Table 11.1 Typical species richness by habitat and taxon

HabitatTaxonTypical STypical Margalef
Lowland tropical rainforestBirds40 to 1206.0 to 14.0
Dry deciduous forestBirds25 to 504.0 to 8.0
Temperate woodlandBirds15 to 352.5 to 5.5
Grassland, managedPlants10 to 302.0 to 5.0
Wetland, freshwaterWaterbirds12 to 402.0 to 6.0
Coral reef transectFish30 to 805.0 to 11.0
Camera trap gridLarge mammals6 to 201.0 to 3.0
Urban parkBirds8 to 201.5 to 3.5
Intensive croplandArthropods5 to 201.0 to 3.5
Polluted streamMacroinvertebrates3 to 100.5 to 2.0

What to conclude: raw S is only comparable within a row. Twelve species is excellent for a camera trap grid and poor for a rainforest bird survey.

Table 11.2 What the singleton pattern tells you

F1 as share of SChao1 behaviourWhat it means
0 percent (no singletons)Chao1 = SSurvey looks complete Good
Under 10 percentChao1 barely above SVery well sampled Good
10 to 25 percentChao1 modestly above SNormal for a good survey Good
25 to 50 percentChao1 clearly above SMore species remain undetected Watch
Above 50 percentChao1 far above S, unstableBadly undersampled Concern

What to conclude: the singleton share is the fastest quality check on any species list. Count it before you interpret anything else.

Table 11.3 Margalef and Menhinick at a glance

SNMargalef (S-1)/ln NMenhinick S/√N
5501.020.71
101071.930.97
105001.450.45
202003.591.41
2010002.750.63
5010007.091.58
100500011.631.41

What to conclude: compare rows 2 and 3. Same richness, five times the effort, and Menhinick halves while Margalef falls only a quarter. Menhinick corrects harder, so pick one and stay with it.

Table 11.4 Choosing a richness estimator

EstimatorRuns onBest whenWatch out for
Chao1 (bias-corrected)Singletons and doubletonsAbundance data from one sampleUnstable when singletons dominate
Chao1 (classic)Singletons and doubletonsMatching older published valuesUndefined when F2 = 0
Chao2Presence across samplesMultiple sites or replicate quadratsNeeds a site-by-species matrix
Jackknife 1Singletons onlyA quick second opinionUsually the highest estimate
ACEAll rare species up to 10Large samples with a long rare tailSensitive to the rare-species cutoff
BootstrapAll speciesConservative estimatesTends to underestimate

What to conclude: report Chao1 for single-sample abundance data, Chao2 when you have replicate samples. Stating which one you used is not optional.

Table 11.5 Minimum sampling effort by survey type

Survey typeMinimum effort for a usable SEffort for a stable species list
Bird point counts10 points across 1 visit30 points across 3 visits and 2 seasons
Line transects2 km walked10 km, repeated in 2 seasons
Camera trapping500 trap nights2,000 or more trap nights
Vegetation quadrats15 quadrats of 1 m²50 quadrats, stratified by habitat
Pitfall traps30 trap days150 trap days across 5 stations
Reef fish belt transects6 transects of 50 m20 transects across 3 depths
Light trap insects5 trap nights20 trap nights across the season

What to conclude: richness needs more effort than any other diversity measure, because rare species are exactly the ones you find last. These minimums are higher than the equivalents for Shannon or Simpson for that reason.

Table 11.6 Which diversity measure to report

MeasureWeightsRangeBest used when
Species richness SAll species equally1 upwardYou have a species list and effort is standardised
Chao1Rare speciesS upwardYour survey is incomplete and you need the true total
Margalef / MenhinickRichness, effort corrected0 upwardSample sizes differ between sites
Shannon Wiener H'Rare species0 to ~4.6You want richness and evenness in one number
Simpson's 1 - DCommon species0 to 1Dominance is the story, or the sample is small
Pielou's J'Evenness only0 to 1Comparing communities with different richness
Hill numbersTunable by q1 upwardYou need richness and diversity on one scale. S is Hill q = 0.

What to conclude: richness answers "how many", the diversity indices answer "how evenly". Most reviewers want both, so report S with Shannon or Simpson beside it.

📊 12. Example Results

Scenario 1: High-richness tropical forest

A bird survey records 28 species across 412 individuals. Chao1 estimates about 31 species, so the survey captured roughly 90% of the community, a rich, well-sampled site.

Scenario 2: Low-richness degraded site

A logged fragment holds only 6 species across 200 individuals. Margalef = 0.94, low for the region, pointing to a species-poor, disturbed community.

Scenario 3: Same richness, different effort

Two sites both record 15 species, but Site A sampled 100 individuals and Site B sampled 500. Menhinick corrects for this: Site A = 1.50, Site B = 0.67, showing Site A is richer per unit effort.

Scenario 4: Undersampling flagged by Chao1

A survey finds 20 species but many singletons. Chao1 estimates 34 species, a big gap that warns you to keep sampling before reporting richness.

Scenario 5: Camera-trap mammal survey

Across 1,400 trap nights, 8 mammal species are recorded with few rare species. Chao1 = 8.5, so the community is almost fully sampled.

Scenario 6: Edge case / caution

Only 3 individuals across 3 species gives S = 3, but the sample is far too small to trust. Always build a species accumulation curve before reporting richness.

Scenario 7: Same species count, different completeness

Two wetlands each record 18 species. Site A has 2 singletons, so Chao1 = 18.3 and completeness is 98 percent. Site B has 9 singletons, so Chao1 = 21.6 and completeness is 83 percent. Site B is probably the richer wetland, and the raw count hides it completely.

Scenario 8: The worked example used in Sections 8, 9 and 10

Ten species with counts 45, 30, 15, 7, 3, 2, 2, 1, 1, 1 give N = 107, S = 10, F1 = 3 singletons, F2 = 2 doubletons, Margalef 1.926, Menhinick 0.967, Chao1 11.0, Jackknife 1 12.97 and 91 percent completeness. Excel, R, Python and this calculator all agree, so use this dataset to check your own setup.

🥾 13. How to Collect Raw Data in the Field

Species richness is only as good as the survey behind it. Follow these ready-to-use steps in the field to collect clean, comparable species data. Print this list and take it with you.

start end Line Transect Walk the line, record every species
Fig 1. Line transect. Walk a fixed straight line at steady pace and record every species seen or heard, noting new species as you go.
observer radius r Point Count Stand still, list every species in radius
Fig 2. Point count. Stand at a fixed station for a set time (e.g. 10 min) and list all species within a fixed radius; repeat at several stations to build richness.
Quadrat List all species inside a fixed frame 1 m × 1 m frame
Fig 3. Quadrat. For plants or sessile species, place a fixed frame (e.g. 1 m × 1 m) and record every species inside it; repeat across many frames.
camera Camera Trap Motion-triggered photos over trap nights
Fig 4. Camera trap. For shy mammals, fix motion-triggered cameras, run them for equal trap nights per site, and count each new species detected.
FIELD DATASHEET · Site: Pine Ridge SpeciesCountDetect Robin|||| |||seen Blue Tit|||| |heard Wren||||heard Chaffinch|||seen Nuthatch||seen Date 14/07 Observer RP 0630–0700 Clear
Fig 5. Field datasheet. One row per species: note each new species and its count, plus date, observer, time, and weather. Enter the per-species counts into the calculator.
Sampling effort (sites / days) Species found estimated true richness Species Accumulation Curve
Fig 6. Accumulation curve. Plot species found against effort. When the curve flattens toward the estimated true richness line, you have sampled enough.
  1. Define your question first. Decide what you are comparing (site vs site, season vs season, before vs after) before you collect any data.
  2. Pick one taxonomic group and stick to it. Count only birds, only trees, or only mammals. Mixing groups makes richness meaningless.
  3. Choose a standard sampling method and use it everywhere: point counts, line transects, quadrats, camera traps, pitfall traps, or mist nets.
  4. Fix your sampling effort. Keep the same transect length, plot size, count duration, or number of trap nights at every site.
  5. Sample enough to reach a plateau. Richness keeps rising with effort, so keep sampling until few new species appear.
  6. Use replicates. Place at least 3 stations, plots, or transects per site so you can build accumulation curves.
  7. Mark station locations with GPS so the exact spot can be resurveyed in future years.
  8. Record every species, and count individuals too. Counts let you calculate Margalef, Menhinick, and Chao1, not just observed richness.
  9. Note singletons and doubletons carefully. Chao1 relies on species seen once or twice, so record rare species accurately.
  10. Avoid double-counting. Move in one direction and use a set detection radius so the same species is logged once per station.
  11. Log detection method (seen, heard, trapped, photographed) for later corrections.
  12. Identify to species where possible. If you cannot, use a consistent morphospecies label and keep it identical across sheets.
  13. Use a prepared datasheet. Columns: Date, Site, Station, Observer, Species, Count, Detection type, Notes.
  14. Record effort metadata on every sheet: start and end time, weather, observer name, and equipment used.
  15. Photograph or voucher tricky species to settle identification disputes back at the desk.
  16. Enter counts as whole numbers, not percentages or densities.
  17. Back up your data the same day by photographing paper sheets or typing counts into a spreadsheet.
  18. Keep a field notebook for disturbance, human activity, unexpected species, or equipment problems.
  19. Combine species across stations before entry, then paste the per-species totals into this calculator (for example 52, 48, 55, 61, 47).
  20. Never invent or round zeros away. A species absent from the sample is simply not listed.

🎯 14. When to Use This Tool

14.1 Quick decision checklist

  • ✅ You have a species list from a defined area or a standardised amount of effort.
  • ✅ Your question is "how many species are here", not "how evenly are they spread".
  • ✅ You have counts as well as names, so the estimators can run.
  • ✅ You need to know whether your survey is complete before you report anything.
  • ❌ Effort differs a lot between sites and you plan to compare raw S. Use Margalef, or rarefy.
  • ❌ Your data is percent cover for plants with no individual counts. Report S only, with no estimators.
  • ❌ One species dominates and that is your real question. Use Simpson's index instead.

14.2 Four real situations

SituationWhat you doWhat richness tells you
Baseline survey for an environmental impact assessmentStandardised transects across the development footprintThe species list of record, with Chao1 showing how much you may have missed
Comparing a reserve with adjacent farmlandSame point count protocol at both, same seasonWhether protection holds more species, after correcting for effort
Deciding when to stop surveyingPlot the accumulation curve as data comes inWhen the curve flattens and Chao1 closes on S, you can stop
A student thesis on habitat fragmentationQuadrats or traps in fragments of different sizesWhether smaller fragments hold fewer species, the classic test

14.3 Sampling design guidance

  1. Fix effort first. Richness is more sensitive to effort than any other biodiversity measure, so decide trap nights, transect length or quadrat count before you start.
  2. Sample until the curve flattens. A rising accumulation curve means your S is a floor, not an estimate.
  3. Record rare species with care. A singleton missed is an estimator changed. Photograph anything uncertain.
  4. Spread effort across habitats. Richness rises fastest when you add a new habitat type, so stratify rather than clustering stations.
  5. Repeat across seasons. Migrants, flowering plants and emerging insects mean a single season always undercounts.

14.4 Which metric next

  • Want evenness as well as counts? Go to the Shannon Wiener index.
  • Want to know if one species dominates? Use Simpson's index or Berger-Parker.
  • Have replicate samples rather than one pooled count? Use Chao2 and the sample-based jackknife.
  • Comparing sites with very different effort? Use rarefaction before anything else.
  • Want richness and diversity on one scale? Use Hill numbers, where S is q = 0.

🛠️ 15. Troubleshooting and Common Errors

Ten problems users hit with species richness, and how to fix each one.

Chao1 came out exactly equal to S

Cause: your sample has no singletons. Fix: nothing to fix. When every species was seen at least twice, Chao1 concludes you have found them all. It is the best result you can get.

Chao1 is far higher than S

Cause: many species seen only once. Fix: this is a warning, not an error. Your survey is incomplete. Add effort, or report Chao1 rather than S when comparing sites.

My richness rises every time I add a survey day

Cause: expected. Richness climbs with effort until it plateaus. Fix: plot a species accumulation curve. Keep sampling until it flattens, or rarefy all sites to the smallest sample size.

Two sites have the same S but very different effort

Cause: raw richness is not comparable across unequal effort. Fix: use Margalef or Menhinick, which scale S by sample size, or rarefy both to the same number of individuals.

The tool reports more species than I recorded

Cause: duplicate species rows, usually a name typed two slightly different ways. Fix: sort your species column and read it. "Warbler sp1" and "Warbler sp 1" count as two species.

Margalef returns an error or infinity

Cause: N is 1, so ln N is zero and the division fails. Fix: Margalef needs at least two individuals. With a single individual, report observed richness only.

My Chao1 does not match the value in a published paper

Cause: two Chao1 forms exist. This tool uses the bias-corrected version; older papers often quote the classic F1 squared over 2F2. Fix: on the worked example those give 11.0 and 12.25. Name the form you used.

Jackknife and Chao1 disagree sharply

Cause: many singletons and few doubletons make both estimators unstable. Fix: treat the gap as a signal that your sample is too small for either estimate to be trusted. Add effort.

Can I calculate richness without counts?

Cause: presence-only data. Fix: yes for observed S, which is just a count of species. No for Margalef, Menhinick, Chao1 and the jackknife, all of which need abundances or replicate samples.

Charts do not appear after I click Run

Cause: the Chart.js library was blocked by a network filter or an offline device. Fix: reload the page on a normal connection. The numbers and the results table still work without charts.

⚠️ 16. Assumptions and Limitations

Species richness is the most intuitive biodiversity measure and the most easily misread. Its whole weakness is that it depends on how hard you looked.

16.1 What the measure assumes

  • Survey effort is standardised across every site being compared.
  • All species had a fair chance of being detected. Cryptic and nocturnal species break this badly.
  • Every individual was identified to the same taxonomic level.
  • Species names are used consistently, with no accidental duplicates or splits.
  • For the estimators, individuals were sampled at random and independently.
  • For Chao1 specifically, rare species were recorded accurately, because singletons drive the estimate.

16.2 What it cannot tell you

  • How abundant anything is. One individual and ten thousand count the same.
  • Whether the community is balanced or dominated by one species. Use Shannon or Simpson for that.
  • Whether the species are native, rare, threatened, or invasive. Richness treats all species as equal, so replacing ten forest specialists with ten weeds leaves it unchanged.
  • Whether a difference between sites is statistically significant. That needs replicates and a test.
  • Ecosystem function or habitat quality on its own.

16.3 The four limitations reviewers raise most

LimitationWhy it mattersWhat to do
Effort dependenceS rises with sampling and has no natural ceilingReport effort explicitly, rarefy, and show the accumulation curve
Estimator instabilityChao1 and the jackknife swing widely when singletons dominateReport both, plus the singleton count, so readers can judge
Blind to identityLosing endemics and gaining weeds leaves S unchangedPublish the species list and conservation status alongside
Detection biasCryptic species are systematically missedUse occupancy models, or state the bias openly

What to conclude: never report a richness value without the effort that produced it. S = 24 means nothing on its own; S = 24 from 1,400 trap nights with Chao1 = 26 means something.

🔍 17. Conclusion

▶ Run the analysis above to generate a personalised conclusion for your dataset.

❓ 18. Frequently Asked Questions

What is species richness and how is it measured?

Species richness is the number of different species in a defined area or sample, written S. It is measured by counting species, not individuals. One elephant and a thousand ants count as two species. Survey effort must be standardised before richness can be compared between sites.

How do you calculate species richness step by step?

Count the species with at least one individual recorded. That is observed richness S. For Margalef, subtract 1 and divide by the natural log of the total individuals. For Chao1, add a correction built from the number of species seen once and twice.

What is the Excel formula for species richness?

Use =COUNTIF(B2:B11,">0") to count species present. Do not use COUNTA, which counts every non-empty cell including zeros and notes. For Chao1 use =S+(F1*(F1-1))/(2*(F2+1)), where F1 and F2 come from =COUNTIF(B2:B11,1) and =COUNTIF(B2:B11,2).

How do I calculate species richness in R?

Observed richness is length(counts[counts > 0]), or vegan::specnumber(x). For estimators use vegan::estimateR(counts), which returns observed richness, Chao1 and ACE together. Use vegan::specpool() when you have a site-by-species matrix rather than one sample.

Which Python package calculates species richness?

scikit-bio is the main one: skbio.diversity.alpha.chao1() for the estimator and osd() for observed richness, singletons and doubletons in one call. NumPy alone is enough for the basics, since observed richness is just (counts > 0).sum().

What is the Chao1 estimator and why does it matter?

Chao1 estimates how many species you did not detect, using the number seen once and twice. If many species appeared only once, more are almost certainly out there. It matters because observed richness is always a floor, never the true total.

What are singletons and doubletons?

Singletons are species represented by exactly one individual in your sample, and doubletons by exactly two. Chao1 and the jackknife estimator are built entirely from these two counts, which is why rare species must be recorded accurately in the field.

What is the difference between Margalef and Menhinick?

Both scale richness by sample size so unequal effort can be compared. Margalef divides by the natural log of N, Menhinick by the square root of N. Menhinick corrects harder. Pick one and use it consistently, because they do not agree.

How much sampling do I need for reliable species richness?

More than any other diversity measure needs, because rare species are found last. Camera traps usually need 2,000 trap nights for a stable list, bird point counts 30 points across three visits. Stop when the species accumulation curve flattens.

What is a species accumulation curve?

A plot of species found against sampling effort. It rises steeply at first, then flattens as new species become rare. If your curve is still climbing at the end of the survey, your richness value is an underestimate, not a result.

Can I compare species richness between sites?

Only when effort is identical, or after correcting for it. With unequal effort, use Margalef or Menhinick, or rarefy every site to the smallest sample size. Comparing raw S across different effort is the single most common richness error.

What is sample completeness and how do I report it?

Sample completeness is observed richness divided by estimated richness, as a percentage. Above 95 percent your list is nearly complete; below 70 percent you are undersampled. Report it beside S so readers can judge the survey, not just the number.

Why is my Chao1 value different from a published paper?

Two forms exist. The bias-corrected form used here gives 11.0 on this page's worked example, while the classic F1 squared over 2F2 gives 12.25. The classic form also breaks when there are no doubletons. Always name the form you used.

Does species richness tell me if a habitat is healthy?

Not on its own. Richness treats every species as equal, so a site that loses ten forest specialists and gains ten weeds scores exactly the same. Report a species list and conservation status alongside the number.

How do I report species richness in a journal or report?

Give the number, the effort, and the estimate, for example: S = 10 species from 107 individuals (Chao1 = 11.0, 91 percent completeness). State which estimator form you used and how effort was standardised across sites.

What is the difference between species richness and species diversity?

Richness counts species and ignores abundance. Diversity indices such as Shannon and Simpson combine richness with evenness, so they also reflect how individuals are spread among species. Two sites with identical richness can have very different diversity.

Can I calculate species richness without counting individuals?

Yes for observed richness, which is simply a count of species. No for Margalef, Menhinick, Chao1 and the jackknife, which all need abundances. With presence-only data across replicate samples, use the Chao2 estimator instead.

What is a good species richness value?

There is no universal figure, because it depends entirely on habitat, taxon and effort. Twelve species is excellent for a camera trap grid and poor for a rainforest bird survey. Always compare against a reference site sampled the same way.

What units does species richness use?

Observed richness is a plain count with no units. Margalef and Menhinick are dimensionless. Chao1 and the jackknife are in species units. Species density, a related measure, is species per unit area such as species per hectare.

Is this species richness calculator free and accurate?

Yes. It is completely free, runs entirely in your browser, stores no data, and uses the standard published formulas for Margalef, Menhinick, bias-corrected Chao1 and the first-order jackknife. Results are cross-checked against R and Python.

📝 19. Cite This Tool

Use one of these three formats when you mention this species richness calculator in a thesis, report, or paper.

APA 7th edition

StatsUnlock. (2026). Species richness calculator: Free online ecology tool with Chao1 and Margalef [Computer software]. https://statsunlock.com/species-richness-calculator/

BibTeX

@misc{statsunlock_richness_2026,
  title        = {Species Richness Calculator: Free Online Ecology Tool with Chao1 and Margalef},
  author       = {{StatsUnlock}},
  year         = {2026},
  howpublished = {\url{https://statsunlock.com/species-richness-calculator/}},
  note         = {Accessed: 2026-09-15}
}

Methods paragraph

Species richness was measured as the observed number of species (S) and standardised for sampling effort using Margalef's index, DMg = (S - 1) / ln N (Margalef, 1958). True richness was estimated with the bias-corrected Chao1 estimator and the first-order jackknife (Chao, 1984; Colwell & Coddington, 1994), and sample completeness was calculated as S / Chao1. All values were computed with the StatsUnlock species richness calculator (https://statsunlock.com/species-richness-calculator/) and verified in R with vegan::estimateR().

🔗 20. Related Tools

Free ecology and wildlife calculators that pair well with species richness.

Shannon Wiener Index Calculator

Adds evenness to your species count. Richness says how many, Shannon says how evenly.

Simpson's Diversity Index Calculator

Answers the dominance question that richness cannot: is one species taking over?

Hill Numbers Calculator

Puts richness, Shannon and Simpson on one scale. Observed richness is Hill q = 0.

Evenness Index Calculator (Pielou's J')

Separates evenness from richness when comparing sites with different species counts.

Fisher's Alpha Diversity Calculator

A richness measure designed to stay stable as sample size changes.

Camera Trap Analysis Calculator

Turns photo detections and trap nights into the standardised species counts this tool needs.

📖 21. Glossary of Terms

ACE Abundance-based Coverage Estimator, a richness estimator that uses all species with ten or fewer individuals.
Alpha diversity Diversity within a single site or sample. Species richness is the simplest alpha measure.
Asymptote The level a species accumulation curve flattens toward. Reaching it means sampling is adequate.
Beta diversity The change in species composition between sites.
Chao1 An estimator of true richness built from the number of singletons and doubletons.
Chao2 The presence-absence version of Chao1, for replicate samples rather than one pooled count.
Detection probability The chance a species present at a site is actually recorded during a survey.
Doubleton (F2) A species represented by exactly two individuals in the sample.
Effort The amount of survey work done: trap nights, transect kilometres, point counts, or quadrats.
Jackknife estimator A richness estimator that adds a correction based on the number of rare species.
Margalef's index Richness scaled by the natural log of sample size, (S - 1) / ln N.
Menhinick's index Richness scaled by the square root of sample size, S divided by the square root of N.
Morphospecies A consistent label for an unidentified species, such as "Warbler sp. 1", used across all data sheets.
Observed richness (S) The number of species actually recorded in a sample.
Quadrat A fixed-size sampling frame, usually square, placed on the ground to count plants or slow-moving animals.
Rarefaction Standardising samples to the same number of individuals so richness can be compared fairly.
Sample completeness Observed richness divided by estimated richness, as a percentage.
Singleton (F1) A species represented by exactly one individual. Chao1 and the jackknife run on these.
Species accumulation curve A plot of species found against effort. When it flattens, sampling is adequate.
Species density Species per unit area, such as species per hectare, rather than per individual sampled.
Trap night One camera trap active for 24 hours. The standard unit of camera trap effort.
Transect A fixed line along which observations are recorded at a steady speed and effort.
True richness The real number of species in the community, including those never detected.
Undersampling Stopping a survey before the accumulation curve flattens, so richness is underestimated.
📚 22. References (20 Peer-Reviewed Sources)

The following references support the ecological methods used in this species richness calculator, covering biodiversity measurement, species diversity, and best practices in ecological sampling and wildlife analysis.

  1. Magurran, A. E. (2004). Measuring biological diversity. Blackwell Publishing.
  2. Margalef, R. (1958). Information theory in ecology. General Systems, 3, 36–71.
  3. Menhinick, E. F. (1964). A comparison of some species-individuals diversity indices. Ecology, 45(4), 859–861. doi.org/10.2307/1934933
  4. Chao, A. (1984). Nonparametric estimation of the number of classes in a population. Scandinavian Journal of Statistics, 11(4), 265–270.
  5. Colwell, R. K., & Coddington, J. A. (1994). Estimating terrestrial biodiversity through extrapolation. Phil. Trans. R. Soc. B, 345(1311), 101–118. doi.org/10.1098/rstb.1994.0091
  6. Gotelli, N. J., & Colwell, R. K. (2001). Quantifying biodiversity: Procedures and pitfalls. Ecology Letters, 4(4), 379–391. doi.org/10.1046/j.1461-0248.2001.00230.x
  7. Chao, A., et al. (2014). Rarefaction and extrapolation with Hill numbers. Ecological Monographs, 84(1), 45–67. doi.org/10.1890/13-0133.1
  8. Hsieh, T. C., Ma, K. H., & Chao, A. (2016). iNEXT: rarefaction and extrapolation of Hill numbers. Methods Ecol. Evol., 7(12), 1451–1456. doi.org/10.1111/2041-210X.12613
  9. Colwell, R. K. (2013). EstimateS: Statistical estimation of species richness (v9). purl.oclc.org/estimates
  10. Oksanen, J., et al. (2022). vegan: Community ecology package. R package v2.6-4. CRAN.R-project.org/package=vegan
  11. Krebs, C. J. (1999). Ecological methodology (2nd ed.). Benjamin Cummings.
  12. Whittaker, R. H. (1972). Evolution and measurement of species diversity. Taxon, 21(2/3), 213–251. doi.org/10.2307/1218190
  13. Gotelli, N. J., & Chao, A. (2013). Measuring and estimating species richness. In Encyclopedia of Biodiversity (2nd ed., pp. 195–211). Academic Press.
  14. Bibby, C. J., Burgess, N. D., Hill, D. A., & Mustoe, S. H. (2000). Bird census techniques (2nd ed.). Academic Press.
  15. Ahumada, J. A., et al. (2011). Community structure and diversity of tropical forest mammals. Phil. Trans. R. Soc. B, 366(1578), 2703–2711. doi.org/10.1098/rstb.2011.0115
  16. Niedballa, J., et al. (2016). camtrapR: An R package for camera trap data. Methods Ecol. Evol., 7(12), 1457–1462. doi.org/10.1111/2041-210X.12600
  17. Roswell, M., Dushoff, J., & Winfree, R. (2021). A conceptual guide to measuring species diversity. Oikos, 130(3), 321–338. doi.org/10.1111/oik.07202
  18. Peet, R. K. (1974). The measurement of species diversity. Annual Review of Ecology and Systematics, 5, 285–307. doi.org/10.1146/annurev.es.05.110174.001441
  19. R Core Team. (2024). R: A language and environment for statistical computing. R-project.org
  20. Chao, A., & Chiu, C. H. (2016). Species richness: Estimation and comparison. Wiley StatsRef: Statistics Reference Online, 1–26. doi.org/10.1002/9781118445112.stat03432.pub2

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