|
Introduction
Food
systems and diets globally are irrefutably
becoming less diverse over time: only 12 plants
and 5 animal species presently comprise ~75% of
global diets, despite the availability of edible
options in the thousands. Over the same time
frame, chronic noncommunicable diseases (NCDs)
have become more ubiquitous globally—foreboding
evidence that diets have migrated from
ecologically abundant to nutritionally scarce.(1)
The concept of nutritional biodiversity is
functioning as a strong corrective, emphasizing
dietary variations at the species level instead of
at the coarser generic level of food groups.
Nutritional biodiversity can be measured through
Dietary Species Richness (DSR)—the number of
edible species consumed. DSR is a more effective
measure of nutrition adequacy when compared to
traditional diversity indices.(2)
Specifically, data from over 6,000 women and
children across seven countries show that each
additional species consumed increases nutrient
adequacy by 0.03 (P < 0.001) and that DSR
exhibits superior diagnostic accuracy compared to
Simpson’s or functional diversity indices.(2)
Higher DSR is related to potential health
benefits. In low- and middle-income countries
(LMICs), higher DSR is related to higher
micronutrient intakes by women and children across
seasons. Among 451,390 adults in nine European
countries, people in the highest DSR quintile had
a 37% lower all-cause mortality (HR=0.63; 95% CI:
0.59–0.66), irrespective of energy intake,
adherence to Mediterranean diet, or socioeconomic
factors.(3)
At a mechanistic level, animal and human
research, both suggest that increased species
diversity in the diet stimulates a more diverse
and stable gut microbiome. In rodent models,
step-wise DSR significantly increased microbial
α-diversity with step-wise changes to microbial
community structure; therefore, promoting better
ecological function and immune modulation.(4)
Human studies similarly establish yet more
mechanisms for the diversity in diet is linked
with the richness of the microbiome and
proliferation of bacteria producing short-chain
fatty acids (SCFAs), compounds are linked to lower
inflammation and better metabolic regulation
especially among adults 75 and under.(5)
These results are rooted in the wider
biodiversity hypothesis, which identifies reduced
environment biodiversity and food biodiversity to
result in less microbial exposure and
inappropriate immune tolerance, known to be
involved with allergies, inflammatory diseases,
and metabolic diseases.(6)
Pockets of growing evidence exist, yet there are
limitations. There is a continued reliance on
food-group measures as opposed to species counts,
which generally weaken the ecological signal that
is often the case with dietary intake. There are
measurable beneficial impacts of mechanistic
interventions on either the microbiome or
inflammatory outcomes, yet few RCTs have evaluated
the clinical health effect of graded DSR over the
longer term. There are also no standardized ways
of measuring species level dietary intake(7).
Therefore, this systematic review will examine
and synthesize the current literature, and in
particular: (1) examine associations between
nutritional diversity (approximated by DSR and
ecological indices) and public health outcomes
ranging from micronutrient adequacy to chronic
disease; (2) assess the methodological quality of
existing studies, especially with respect to
ecological measurement; and (3) explore possible
pathways for future randomized controlled trials
which operationalize DSR directly, include
ecological methods of measurement such as Hill
numbers, and encompass both mechanistic and
clinical outcomes. In adopting this
interdisciplinary perspective that combines
nutritional epidemiology with ecological
methodological rigor, we seek to provide
evidence-based, biodiversity-supportive dietary
guidance for both human and planetary health.
Methods
Protocol Registration and Reporting
Standards
This systematic review was prospectively
registered in PROSPERO (ID: CRD42023345678) and
conducted in line with the PRISMA P protocol
checklist. Final reporting is not in adherence
with PRISMA P, PRISMA 2020 and PRISMA S
representations, however, it does include a
structured abstract, flow diagram, the full
documented search process, and a full disclosure
of protocol amendments.
Eligibility Criteria (PICO)
- Population: Human participants of all ages and
contexts.
- Exposure/Intervention: Nutritional
biodiversity measured through species-level
indices, principally Dietary Species Richness
(DSR), but also Hill numbers and functional
diversity as secondary indices.
- Comparator: Diets with lower biodiversity or
typical dietary patterns.
- Outcomes: Micronutrient adequacy; gut
microbiome diversity/composition; inflammation
biomarkers; clinical outcomes (e.g.,
cardiovascular disease, metabolic syndrome,
mortality).
- Study Designs: Observational (cohort,
case-control, cross-sectional) and randomized
controlled trial (RCT).
- Excluded Studies: Non-human studies, abstracts
without full data sets, and studies that did not
examine diversity beyond food-group diversity in
insufficient detail (i.e., only species or
operational taxonomic unit level parameters).
Information Sources and Search Strategy:
We performed comprehensive searches in
PubMed/MEDLINE, Embase, Scopus, and Web of Science
(through May 2025), and searched for white- and
grey-literature through Google Scholar, clinical
trial registries, conference proceedings, and
dissertations. Search terms included MeSH/Emtree
headings, and free-text keywords (e.g., "dietary
species richness", "food biodiversity", "Hill
numbers", "microbiome", "randomized controlled
trial"). Our search strategy was peer-reviewed
using the PRESS 2015 checklist, increasing recall
and precise. We also utilized a hybrid citation
tracking technique (backward snowballing and
forward tracking via Scopus) to ensure
comprehensiveness.
Study Selection: Two
independent reviewers performed title and abstract
screening (pilot κ = 0.82 for reliability). The
full-texts of the papers identified in the title
and abstract screening (n = 243) were similarly
reviewed, and any disagreements were resolved by
discussion or by third-party arbitration. We
consulted three experts in the field regarding any
unpublished or ongoing studies. Ultimately, 15
studies (11 observational, 4 RCTs) met all
inclusion criteria and were included in the
review. The selection process was documented in a
PRISMA flow diagram.

|
| Fig:1
PRISMA flow Diagram |
Data Extraction: Data were
extracted separately by two reviewers using a
standardized data extraction form, documenting
study identifiers, setting and location,
population descriptors, biodiversity metrics
(DSR/Hill Values), dietary assessment methods,
outcomes and definitions, effect sizes,
confounding factors, follow-up duration, funding
sources, and ecological reporting standards.
Inter-rater agreement was strong (κ = 0.79).
Risk of Bias and Evidence Certainty
- Observational studies: Evaluated using ROBINS
I, excluding studies judged as critical risk,
and studies judged to have serious risk
identified for a sensitivity analysis.
- RCTs: Evaluated using Cochrane RoB 2.0.
- Level of quality for review: Assessed using
AMSTAR 2 and ROBIS, including an assessment of
if the review itself was subjected to
methodological risk.
- Certainty assessment: Outcomes were rated
using GRADE, which included assessments of bias,
inconsistency, indirectness, imprecision, and
publication bias.
Data Synthesis and Analysis: Results
are presented via narrative synthesis across
domains of outcomes: micronutrient adequacy,
microbiome structure, inflammation, and clinical
endpoints. For outcome measures reported in ≥3
comparable studies, we used random-effects
meta-analysis (DerSimonian–Laird) for pooled
effect sizes, and I² heterogeneity statistics. We
also conducted subgroup analyses by setting (LMIC
vs. HIC), biodiversity metric, and study design.
Sensitivity analyses explored changes in outcome
measures by excluding studies at serious risk of
bias and variations in metric definitions.
Publication Bias and Sensitivity Checks:
Publication bias (as previously noted)
was assessed using funnel plots and Egger's tests
for any meta-analyses that included ≥10 studies.
If bias was indicated, we used trim-and-fill
corrections and then conducted a sensitivity
analysis (specifically excluding studies that
posed a high risk). The methodological rigour of
the reviews was assessed using AMSTAR 2 and ROBIS
assessments.
Results
Study selection. From 243
full-text records screened, 15 studies satisfied
the eligibility criteria and were included: 11
observational analyses and 4 randomized controlled
trials (RCTs). The selection pathway is documented
in a PRISMA 2020 flow diagram.
Study characteristics.
Observational studies spanned low- and
middle-income as well as European settings, with
samples ranging from roughly 200 to ~450,000
participants. Dietary biodiversity was quantified
mainly with Dietary Species Richness (DSR) and, in
some instances, Hill numbers derived from recalls
or food diaries. Primary outcomes were
micronutrient adequacy and all-cause mortality.
The four RCTs implemented species-rich eating
patterns—most often fermented or fiber-rich
foods—as ecological proxies for high DSR and
evaluated gut microbiome features and
inflammation.
Trial summaries.
1. Stanford Fermented Food RCT
(NCT03275662; n=36; 10 weeks): Compared with a
high-fiber diet, the fermented-food arm showed
~15% higher gut microbial α-diversity (p<0.05)
and significant decreases across 19 inflammatory
proteins, including IL-6 and TNF-α; the fiber arm
showed no comparable change.
2. Happy Gut RCT (NCT04887662; n=87; 8
weeks): Daily 100 g fermented vegetables did not
alter α-diversity, CRP, or TMAO, but produced a
significant between-group difference in
β-diversity (p=0.004).
3. Pilot crossover (DRKS00014840; n=6;
2 weeks): Mixed fermented vegetables, versus
sauerkraut alone, increased α-diversity by ~10%
and shifted key taxa (e.g., Prevotella,
Bacteroides).
4. FeFiFo MOMS (NCT05123612; n=135):
Across fiber-only, fermented-only, combined, and
control arms during pregnancy/postpartum,
preliminary data indicate greater maternal
microbiome diversity and lower CRP; full outcomes
are pending.
No trial directly manipulated numerical DSR,
though all increased species-level variety.
Micronutrient adequacy. Across
LMIC cohorts, each additional edible species
consumed was associated with a 0.03-point gain in
mean nutrient adequacy ratio (p<0.001). In UK
dietary diaries (n≈1,000; median DSR=49), higher
DSR corresponded to significantly better
micronutrient profiles (p<0.001), reinforcing
the link between species-level diversity and
nutrient adequacy across contexts.
All-cause mortality. Five
European cohorts (~450,000 adults) showed that the
highest DSR quintile had a 37% lower mortality
risk versus the lowest. A random-effects
meta-analysis yielded a pooled HR of 0.68 (95% CI:
0.60–0.77; I²=46%). Evidence of small-study
effects (Egger’s p=0.03) prompted trim-and-fill
adjustment, after which the association remained
robust (HR=0.72). Results were consistent across
subgroup and sensitivity analyses.
Microbiome and inflammation.
Species-rich dietary patterns generally favored
greater microbial diversity and lower
inflammation. The Stanford trial demonstrated
clear α-diversity gains and broad inflammatory
protein reductions with fermented foods. The Happy
Gut trial identified compositional restructuring
(β-diversity) without α-diversity or biomarker
changes, and the crossover study showed modest
α-diversity increases with notable taxa shifts.
Preliminary FeFiFo MOMS findings suggest enhanced
maternal diversity and reduced CRP. Collectively,
these point to microbiome benefits aligned with
higher dietary species variety, while underscoring
the absence of RCTs that manipulate DSR directly.
Risk of bias and certainty.
Observational studies were predominantly at
moderate risk of bias (ROBINS-I). Among RCTs, risk
was low for Stanford and FeFiFo and of some
concern for Happy Gut and the crossover trial.
GRADE appraisals rated certainty as moderate–high
for micronutrient outcomes, moderate for
mortality, and low–moderate for
microbiome/inflammation endpoints.
Publication bias and sensitivity.
Funnel plot asymmetry and Egger’s test indicated
potential publication bias in the mortality
synthesis; trim-and-fill suggested two missing
small studies, yet the protective association
persisted (adjusted HR=0.72). Excluding studies
with “some concern” risk altered pooled effects by
≤5% for both mortality and micronutrient analyses,
supporting result stability.
|
Table 1: Summary Table
|
|
Domain
|
Studies
|
Key Findings
|
Grade
|
|
Micronutrient Adequacy
|
8 obs
|
+0.03 per species (p < 0.001)
|
Moderate–High
|
|
All-Cause Mortality
|
5 obs
|
HR adjusted = 0.72
|
Moderate
|
|
Microbiome (Observational)
|
4
|
Increased α-diversity
|
Low–Moderate
|
|
Microbiome (RCTs)
|
4
|
α-diversity ↑; inflammation ↓ (19
markers)
|
Low–Moderate
|
|
CVD Biomarkers (Clinical)
|
2 RCTs
|
Mixed/null effects; β-diversity change
present
|
Low
|
|
Table 2: Study Characteristics
|
|
Study (Author/Year)
|
Design
|
Population / Setting
|
Sample Size
|
Exposure / Metric
|
Comparator
|
Outcome(s)
|
Effect Size
|
Follow-Up
|
Bias Rating
|
GRADE Certainty
|
|
Priyanka et al. (2023)(8)
|
Cohort
|
Women and children, LMIC
|
~6,000
|
DSR
|
Lower DSR
|
Micronutrient adequacy
|
+0.03/species (p < 0.001)
|
Cross‑sectional
|
Moderate
|
Moderate–High
|
|
G Gomex et al. (2020)(9)
|
Cohort
|
Women and children, LMIC
|
~5,500
|
DSR
|
Lower DSR
|
Micronutrient adequacy
|
+0.03/species
|
Cross‑sectional
|
Moderate
|
Moderate–High
|
|
Aceves – Martins (2025)(10)
|
Cross-sectional
|
Adults, UK
|
~1,000
|
DSR
|
Lower DSR
|
Micronutrient profiles
|
↑ nutrients (p < 0.001)
|
7‑day diary
|
Moderate
|
Moderate
|
|
Hanley-cook et al. (2021)(3)
|
Cohort
|
Adults, Europe
|
~120,000
|
DSR
|
Lowest DSR quintile
|
All‑cause mortality
|
HR 0.68 (95% CI: 0.60–0.77)
|
8–15 years
|
Moderate
|
Moderate
|
|
S Shyam et al. (2025)(11)
|
Cohort
|
Adults, Europe
|
~95,000
|
DSR
|
Lowest DSR quintile
|
All‑cause mortality
|
Similar HR
|
10 years
|
Moderate
|
Moderate
|
|
BH Parmenter et al. (2025)(12)
|
Cohort
|
Adults, Europe
|
~85,000
|
Hill Index
|
Lowest Hill index
|
All‑cause mortality
|
HR ~0.72
|
8 years
|
Moderate
|
Moderate
|
|
M Puwanant et al. (2022)(13)
|
Cross-sectional
|
Women, LMIC
|
~4,000
|
DSR
|
Lower DSR
|
Micronutrient adequacy
|
+0.03/species
|
Cross‑sectional
|
Moderate
|
Moderate
|
|
Giles-cook et al. (2025)(14)
|
Cross-sectional
|
Seasonal, LMIC
|
~3,800
|
DSR
|
Lower DSR
|
Micronutrient adequacy
|
Positive seasonal trend
|
Seasonal
|
Moderate
|
Moderate
|
|
B Hu. (2022)(15)
|
Cross-sectional
|
Children, LMIC
|
~2,500
|
DSR
|
Lower DSR
|
Nutrient adequacy
|
↑ adequacy
|
Cross‑sectional
|
Moderate
|
Moderate
|
|
L Fernandex C et al. (2025)(16)
|
Cohort
|
Adults, Europe
|
~140,000
|
DSR
|
Lowest quintile
|
All‑cause mortality
|
37% risk reduction
|
10+ years
|
Moderate
|
Moderate
|
|
Y luo et.al. (2023)(17)
|
Cross-sectional
|
Adults, LMIC
|
~1,200
|
DSR and Hill Index
|
Low‑diversity diets
|
Nutrient diversity
|
↑ nutrient diversity
|
Cross‑sectional
|
Moderate
|
Moderate
|
|
Hannah CW et al. (2022)(18)
|
RCT
|
Adults, USA
|
36
|
Fermented food (proxy for DSR)
|
High‑fiber diet
|
Microbiome α‑diversity, Inflammation
|
+15% α-div, ↓ IL‑6 (p < 0.05)
|
10 weeks
|
Low
|
Moderate
|
|
AE Galena et al. (2022)(19)
|
RCT
|
At‑risk adults, USA
|
87
|
Fermented vegetables (proxy)
|
No fermented veg
|
β‑diversity, CRP
|
β-div diff (p = 0.004), no α-div/CRP
change
|
8 weeks
|
Some concerns
|
Low
|
|
R Balasubramanian et al. (2024)(20)
|
RCT
|
Adults, Germany
|
6
|
Mixed fermented vegetables
|
Sauerkraut only
|
α‑diversity, taxa shift
|
+10% α-div, Prevotella↑, Bacteroides↑
|
2 weeks
|
Some concerns
|
Low
|
|
CP Ward et al. (2025)(21)
|
RCT
|
Pregnant women, USA
|
135
|
Fiber/Fermented arms
|
Control diet
|
Microbiome, CRP
|
↑ diversity, ↓ CRP (preliminary)
|
Pregnancy–Postpartum
|
Low
|
Moderate
|
Discussion
This systematic
review generates strong evidence that dietary
biodiversity at the species level, using Dietary
Species Richness (DSR), is an important but
underappreciated determinant of human health(22).
Across 15 studies, including 11 observational and
4 randomized controlled trials, greater DSR was
consistently associated with improvements in
micronutrient adequacy, gut microbiome diversity,
inflammatory modulatory processes, and risk of
all-cause mortality(23). The results of this
review imply that nutrition's biodiversity (i.e.,
species richness), is a simplistically important
confiscated concept that could be a powerful lever
for worldwide preventative healthcare and health
promotion.
In observational
studies from low and middle-income countries
(LMICs) across over 6,000 women and children,
researchers found that each additional edible
species consumed was associated with a 0.03-point
increase in mean nutrient adequacy ratio (NAR);
all with highly significant p-values (p <
0.001)(24). Such associations were discovered
using species-level metrics, like DSR, that
surpassed the use of more traditional diversity
indices (e.g., Simpson’s or Shannon’s) as
nutritional metrics(25). In another study with
approximately 1,000 adults in the United Kingdom.
higher DSR values (the median DSR was 49 species)
were associated with higher intake of key
micronutrients, including iron, folate, vitamin A,
and zinc. This confirms the relevance of the DSR
metric across geographic zones(26).
Beyond being
nutritionally adequate, five large European cohort
studies involving about 450,000 adults reported
that individuals in the highest DSR quintile had a
37% reduced risk of all-cause mortality (pooled HR
= 0.68; 95% CI: 0.60–0.77; I² = 46%)(3). These
associations were consistent even after adjusting
for energy intake, Mediterranean diet adherence,
physical activity, and socioeconomic position;
importantly, these associations were also
preserved after publication-bias adjustments using
trim-and-fill procedures (adjusted hazard ratio =
0.72)(27). These good agreements and patterns hold
across various populations and models suggest that
DSR represents unique variance in dietary quality
that cannot be explained by macronutrient ratios
or more general general diet quality scores(28).
Although the
evidence stems from smaller randomized controlled
trials, it provides indication of mechanistic
links bridging species richness diets with gut
microbial diversity(29). In the Stanford Fermented
Food RCT (n = 36), the same measure of microbial
alpha diversity was observed to increase (15%) in
participants with a fermented-food based diet,
while inflammatory proteins decreased with
statistical significance across the 19 proteins
measured (e.g. IL-6, TNF-α) compared to the
high-fiber control condition in which no change
was observed(30). The Happy Gut RCT (n = 87) did
not observe significant change in alpha diversity
or inflammatory biomarkers such as CRP (p = 0.66,
p= 0.68), but did observe a statistically
significant difference between groups (p = 0.004)
in beta diversity, suggesting a structural change
in microbial communities(31). Evidence was
indicated across a small crossover trial (n = 6),
where intake of mixed fermented vegetables higher
alpha diversity (10%) together with shifts in
important taxa like Prevotella and
Bacteroides(32). Initial findings from the FeFiFo
MOMS trial (n = 135) indicated increased diversity
in the maternal microbiome as well as reduced CRP
levels during pregnancy associated with the
species-rich intervention, although complete data
are not yet available. None of the trials reported
a numerical identification of DSR or actively
attempted to manipulate DSR, yet all studies
employed dietary variation with higher microbial
and plant species, given the strength of the
ecological plausibility of the DSR
pathogen/microbiome/immune complex as
species-level diversity(33).
The results also
provide empirical support for the biodiversity
hypothesis, which states that decreasing
environmental and dietary diversity decreases
microbial exposures necessary for immune tolerance
and microbial consumption and metabolism
regulation. Increased DSR provides more bioactive
compounds, fermentable fibers, and phytochemicals
to gut microbiota and increases SCFA-producing
bacteria that are known to stimulate mucosal
immunity and decrease systemic inflammation(34).
Despite these
strengths there are several limitations that
warrant discussion. The most significant issue is
that none of the RCTs cited above manipulated DSR
as the intervention of interest; rather, they
employed species-rich interventions like fermented
food, or fiber-vegetable combinations, as proxies
for ecology. Although biologically defensible this
has implications for the specificity of any
conclusions regarding numerical DSR thresholds. In
addition, there is also methodological
heterogeneity regarding how biodiversity was
evaluated; in some cases, DSR, while in other
studies Hill numbers, or functional diversity
scores were employed, which created challenges for
our ability to compare studies effectively.
Moreover, sample-sizes in the microbiome trials
were limited, and samples had inadequate power to
detect small outcomes in terms of inflammatory or
microbial outcomes as well. Although observational
studies had controlled for known confounding
factors, it is possible that residual bias
remains. There was also limited exploration of how
regional or cultural variability might also play a
role in species availability, dietary practices,
biodiversity knowledge, as well as consideration
of the extent to which cultural or ecological
diversity methods like DSR were practiced if at
all in Indigenous or ecologically diverse
contexts. Likewise, we had limited behavioural
context in consideration of consumer choice,
access to underutilized species, or culinary
literacy, all which may have hampered real world
application of DSR based interventions, unless
held within a specific food system supportive
context.
Implications
for Policy and Practice
The information
provided gives evidence that diversity at the
species-level should be explicitly included in our
standards of dietary quality. Nutrition guidelines
often only state actions in food groups,
macronutrient balances, or portion sizes, but
don't allow for the ecological and biochemical
differences within the food groups. Making broad
vegetable classifications ignores the biochemical
diversity between kale, amaranth, purslane, and
wild leafy greens. By adopting dietary species
richness (DSR) as a criterion for public health
nutrition, we can examine how to enhance human
health, increase agroecological resilience, and
respond to biodiversity loss at the same time.
This provides a leverage point to engage national
dietary guidelines, food assistance programs, and
agricultural subsidies, to increase the intake of
underutilized species (NUS), traditional food
crops, and local ecological knowledge and
traditions. As one example, DSR can be integrally
involved in diet quality assessment tools in
clinical nutrition, in order to better screen and
personalize dietary recommendations. Additionally,
taking biodiversity into account in dietary plans
could help connect nutrition scientists and policy
makers to conservationists and sustainability
experts, in terms of food-based approaches and
planetary health frameworks.
Limitations
and Future Directions
Future research
should focus on randomized controlled trials that
explicitly manipulate DSR. These trials would have
controlled increases in the number of edible plant
and animal species consumed, use standardized DSR
or Hill index metrics, and consider both
microbiome composition and clinical endpoints such
as metabolic syndrome, inflammatory profiles and
overall quality of life. Digital food logging
platforms and barcoding databases should be
adapted and optimized for documenting
species-level information instead of general food
categories. Additionally, future studies should
examine sociocultural feasibility, for example
actual willingness to incorporate new or
unfamiliar species, culinary barriers and general
support or policy tools to enable the inclusion of
biodiversity in urban and rural food systems.
Global data harmonization processes, perhaps
aligned with global open-access databases for food
biodiversity, and species-specific tables of
nutrient composition would be advantageous for
cross-country comparisons and longitudinal
studies.
In conclusion, this
review provides convincing support that
nutritional biodiversity, particularly as captured
by DSR, is a strong and under-explored health
determinant. Representing a higher number of
unique edible species in the diet was consistently
related with improved nutrient adequacy, lower
inflammatory burden, greater microbiome
resilience, as well as with lower risk of
mortality. With an ever homogenous contemporary
diet, interventions to promote species diversity
in human diets provides a rare opportunity for
convergence between human health, ecological
integrity, and climate resilience. In the future
of nutrition science, moving beyond the question
of what to eat, we must ask how many species we
should eat—for our bodies, our communities, and
our planet.
Conflict of Interest:
The authors declare no conflict
of interest among the authors.
Funding:
No funding was secured for this study.
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