Cited by 7Wildlife Ecology and ConservationPrimate Behavior and EcologyIsotope Analysis in Ecology

A new method to determine the diet of pygmy hippopotamus in Taï National Park, Côte d’Ivoire

Hendier, Alba, Chatelain, Cyrille, Du Pasquier, Pierre‐Emmanuel, Paris, Monique, Ouattara, Karim, Koné, Inza, Croll, Daniel, Zuberbühler, Klaus·2021·African Journal of Ecology

Abstract

Diet determination of endangered species is an essential element in defining successful conservation strategies and optimising captive breeding programmes. In this study, we developed a new diet identification system, derived from standard faecal analysis, to determine the diet of an elusive and endangered herbivore, the pygmy hippopotamus (Choeropsis liberiensis). We collected faecal samples from 10 free‐ranging individuals covering a combined home range area of about 50 km 2 in Taï National Park, Côte d’Ivoire. In subsequent laboratory analyses, we extracted a large number of leaf epidermis fragments from spatially separated faecal samples and compared them with a reference plant database. Using Multiple Correspondence Analysis (MCA) of epidermis fragments combined with direct visual inspection, we identified the most frequently consumed plant species, which revealed that pygmy hippopotami qualified as intermediate feeders. Their diet was based on at least seven species of monocotyledonae, dicotyledonae and fern groups, with a preference for a small number of other plant species. We evaluate the merit of our method and discuss our findings for developing effective conservation and captive breeding strategies in an endangered species with a wild population of less than 2500 adult individuals.

AI snapshot

Written from the paper’s full text.

TL;DR

The paper develops a faecal epidermis identification method combining microscopy, Multiple Correspondence Analysis and visual inspection to infer the diet of free-ranging pygmy hippopotamus in Taï National Park. It identifies at least seven consumed plant species and classifies pygmy hippos as generalist intermediate feeders rather than browsers.

Key findings

  • From 10 faecal samples, 8 of 10 contained monocots, dicots and ferns; the two remaining samples (2 and 6) lacked ferns. At least seven plant species were identified from the reference list.
  • Frequently detected plants included *Nephrolepis bisserata* in 7 of 10 samples, Marantaceae species in 7 of 10 samples, and *Heritiera utilis* in 4 of 10 samples. *Centotheca lappacea* was reported as found in all samples, and *Streptogyna crinita* was also identified.
  • The authors classify pygmy hippos as herbivore generalists and non-ruminant generalist intermediate feeders, with suggested preference for *N. bisserata*, *Pteris burtonii*, Marantaceae species and *S. crinita*.
  • *Centotheca lappacea* and *Heritiera utilis* are presented as new additions to the known diet, and the importance of grasses is highlighted for captive and wild conservation strategies.

Methods

  • Fieldwork was conducted in Taï National Park, Côte d’Ivoire, from July to November 2017 in an area of about 50 km². The authors collected 70 faeces and 330 GPS location points, then selected 10 faecal samples at least 2 km apart, treated as 10 individuals.
  • A reference library was built from 60 collected plant species, with 56 usable species and 112 reference epidermis samples from both leaf sides. Faecal analysis used 2 g per sample, macroscopic sorting, and 48 randomly selected leaf fragments per sample, yielding 480 fragments and 2,880 pictures; 130 fragments were finally analysed.
  • Identification used five categorical epidermal features in Multiple Correspondence Analysis, Hierarchical Clustering with the Ward method, and visual inspection of targeted pictures. The MCA reference library had a total inertia of 2.4; the first two axes explained 35.2% and the first five axes 65.7% of variance.

Limitations

  • The paper states that the plant reference library covered less than 4% of Taï National Park’s documented flora (60 of 1,356 species), though the authors argue it may include about one third of relevant shrub and herbaceous species; they recommend expanding samples across areas and seasons and adding chemical analyses.
  • The paper states that MCA variables were limited, stomata could not be used due to slide quality, adaxial sides were less informative, only large plant fragments were analysed, fruits and seeds were not identified, and dry material may have been better boiled for rehydration.
  • The paper also notes that male and female home ranges can overlap, so two individuals could have contributed to one sampling location; camera-trap identification was largely unsuccessful, and DNA barcoding would require fresh faeces difficult to obtain in remote field conditions.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1111/aje.12888

Cites

47 of 47 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

0 of 22 cited sources checked · 0 supported · 0 flagged

  • Findingown data
    § Note · p. 10

    In the dicots, Herritiera utilis remnants were found four times.

    “Finally, in the dicots, we found four times Herritiera utilis remnants (Figure 3).”
    Quote in the paperNot a substantial claim
  • Findingown data
    § 4 DISCUSSION · p. 11

    Grasses are an integral part of the pygmy hippo diet.

    “Our data show that grasses are an integral part of the pygmy hippo diet, but it is unclear whether this is properly taken into account by captive facilities.”
    Quote in the paperSubstantial claim
  • Findingown data
    § 4 DISCUSSION · p. 11

    When comparing our data to the already published list, we could only confirm about 50% of species.

    “When comparing our data to the already published list, we could only confirm about 50% of species, while species classified as favourites were difficult to find or absent in our area (e.g., Staurogyne paludosa , Justicia tenella , and Floscopa africana ).”
    Quote in the paperSubstantial claim
  • Findingown data
    § Note · p. 10

    For the monocots of the Poaceae family, evidence was found for Centhoteca lappaceae and Streptogyna crinita.

    “For the monocots of the Poaceae family, we found evidence for Centhoteca lappaceae , and Streptogyna crinita (Appendix 4 and 5).”
    Quote in the paperNot a substantial claim
  • Findingown data
    § 4 DISCUSSION · p. 11

    Herritiera utilis was found in at least four of the ten samples.

    “Herritiera utilis (tree leaves, found in at least four of the ten samples)”
    Quote in the paperNot a substantial claim
  • May combine two claims. 23
  • Classified as probably not a substantial claim. 9
  • May combine two claims.

    Findingown data
    § 5 CONCLUSIONS · p. 13

    C. lappaceae was found in all samples and confirmed the importance of grasses in the diet of this species.

    “In addition, C . lappaceae (monocot) was found in all samples and confirmed the importance of grasses in the diet of this species.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data
    § 3.2 Faecal samples · p. 8

    Many plant species of the reference plant list shared the same position on the HC tree.

    “However, many plant species of the reference plant list shared the same position on the tree (Figure S1).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data
    § 4 DISCUSSION · p. 10

    At least seven plant species were identified, although several epidermis fragments remained unidentified.

    “Secondly, we were able to identify at least seven plant species, although several epidermis fragments remained unidentified (limitations below).”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: Diet determination of endangered species is an essential element in defining successful conservation strategies and optimising captive breeding programmes.
    • Rests on an assumption: There is little information from the wild on pygmy hippos.
    • Rests on an assumption: We are not aware of any systematic data on diet preference and composition in free-ranging pygmy hippos.
  • May combine two claims.

    Findingreasoning
    § 4.1 Limitations · p. 12

    It was almost impossible to carry out plant identification and recognise different individuals based on the camera trap videos.

    “However, it was almost impossible to carry plant identification, and recognise different individuals based on the videos.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data
    § 4.1 Limitations · p. 12

    Matching seeds were found across samples collected during the rainy season.

    “During the sorting of the faeces, we found matching seeds across samples collected during the rainy season (Appendix 8).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data
    § 4 DISCUSSION · p. 10

    The study analysed the diet of 10 individual pygmy hippos over a combined range of about 50 km2 and found evidence for commonly found plants in the faecal samples.

    “We analysed the diet of 10 individual pygmy hippos over a combined range of about 50 km 2 and found evidence for commonly found plants in the faecal samples.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data
    § Note · p. 10

    Marantaceae family was present in 7 of 10 samples, often Marantocloa purpurea.

    “For Marantaceae family, they were present in 7 of 10 samples (Appendix 6 and 7; often Marantocloa purpurea ).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data
    § Note · p. 10

    8 of 10 faecal samples contained monocots, dicots and fern species; the two remaining samples (2, 6) did not contain any fern.

    “Results showed that 8 of 10 faecal samples contained monocots, dicots and fern species. The two remaining samples (2, 6) did not contain any fern (Table 3).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data
    § 4 DISCUSSION · p. 11

    The seven plant species described in the results were frequently found in most faecal samples, particularly plants from the Poaceae family (grasses).

    “The seven plant species described in the results were frequently found in most faecal samples, particularly the plants from the Poaceae family (grasses).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data
    § 4 DISCUSSION · p. 11

    No evidence was found for Desmodium adscendens, Dissotis rotundifolia, Geophila afzelii, Geophila hirsuta, and Cercestis afzelii in the faecal samples.

    “However, we could not find any evidence for the following dicots: Desmodium adscendens , Dissotis rotundifolia , Geophila afzelii , Geophila hirsuta , and Cercestis afzelii .”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data
    § ABSTRACT · p. 1

    Pygmy hippo diet was based on at least seven species of monocotyledonae, dicotyledonae and fern groups, with a preference for a small number of other plant species.

    “Their diet was based on at least seven species of monocotyledonae, dicotyledonae and fern groups, with a preference for a small number of other plant species.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Diet determination of endangered species is an essential element in defining successful conservation strategies and optimising captive breeding programmes.
    • Rests on an assumption: There is little information from the wild on pygmy hippos.
    • Rests on an assumption: We are not aware of any systematic data on diet preference and composition in free-ranging pygmy hippos.
  • May combine two claims.

    Findingown data
    § 3.1 Plant reference library · p. 7

    Despite the horseshoe effect, three main groups were observed through the categories of the variables, corresponding mostly to the three major taxonomic groups: monocots, dicots and ferns.

    “Despite this effect, three main groups were observed through the categories of the variables (Appendix 2), which corresponded mostly to the three major taxonomic groups: monocots, dicots and ferns.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data
    § 4 DISCUSSION · p. 10

    Free-ranging pygmy hippos have a varied diet, which includes major plant groups including monocots, dicots and ferns.

    “With this method, we found that, firstly, free-ranging pygmy hippos have a varied diet, which includes major plant groups including monocots, dicots and ferns.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Diet determination of endangered species is an essential element in defining successful conservation strategies and optimising captive breeding programmes.
    • Rests on an assumption: There is little information from the wild on pygmy hippos.
    • Rests on an assumption: We are not aware of any systematic data on diet preference and composition in free-ranging pygmy hippos.
  • May combine two claims.

    Findingown data
    § 3.1 Plant reference library · p. 7

    The MCA of the plant reference library (n=112, 5 variables) had a total inertia of 2.4; first two axes explained 35.2%, first five axes 65.7% of total variance.

    “The MCA ( n = 112 individuals; 5 variables) had a total inertia of 2.4. The first two axes (F1 and F2) explained 35.2%, and the first five axes 65.7% of the total variance.”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: The two sides of the leaves can be difficult to identify in faeces.
  • May combine two claims.

    Interpretationreasoning
    § 4 DISCUSSION · p. 11

    Pygmy hippos are non-ruminant generalist intermediate feeders.

    “All these observations support that pygmy hippos are non-ruminant generalist intermediate feeders.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Study design and conductnone given
    § 4.1 Limitations · p. 12

    Many food item references share the same characteristics and it is sometimes difficult to distinguish the epidermis of different species without confirmation by picture.

    “Furthermore, many of the food item references share the same characteristics and, sometimes, it is difficult to distinguish the epidermis of different species without a confirmation by picture.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationreasoning
    § 5 CONCLUSIONS · p. 13

    Pygmy hippos in the Taï area appear to have a food preference for N. bisserata, S. crinita, Marantaceae species, C. lappaceae, and H. utilis.

    “Moreover, if these analyses were combined with additional evidence, then pygmy hippos in the Taï area appear to have a food preference for specific species, notably N . bisserata (fern), S . crinita (monocot), Marantaceae species (monocot), C . lappaceae (monocot), and H . utilis (dicot).”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation
    § 1 INTRODUCTION · p. 2

    In captivity, pygmy hippos suffer from persistent health problems, such as polycystic kidney disease and dental skin and foot problems, which may originate from inadequate diet.

    “In captivity, individuals suffer from persistent health problems, such as polycystic kidney disease and dental skin and foot problems, which may originate from inadequate diet (Flacke et al., 2017; von Houwald et al., 2007).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Flacke et al., 2017 source
    • cites von Houwald et al., 2007

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationcitation
    § 4 DISCUSSION · p. 11

    Previous studies were based on feeding trials, feeding signs and direct observations in a restricted study area, which could explain the new observation.

    “This new observation could be explained by the fact that previous studies were based on feeding trials, feeding signs and direct observations in a restricted study area (Hentschel, 1990).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Hentschel, 1990

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationown data
    § 3.1 Plant reference library · p. 7

    The light horseshoe shape (Guttman effect) in the MCA projection was due to two variables, most likely the macroscopic variable 'macro veins' and the 'microscopic layout'.

    “We observed a light horseshoe shape (Guttman effect) on the active individual's (reference plants; blue) projection in F1xF2 axes (Figure 2, Appendix 1 for the projection on F1×F3 axes). This effect was due to two variables that contributed in similar ways, most likely the macroscopic variable 'macro veins' and the 'microscopic layout'.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationreasoning
    § 3.2 Faecal samples · p. 8

    The HC tree helped to target similar epidermis, but visual inspection was still necessary to determine the species in the faeces fragments.

    “The HC tree helped to target similar epidermis, but visual inspection was still necessary to determine the species in the faeces fragments.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation
    § 4 DISCUSSION · p. 11

    Intermediate feeders are able to adapt their diets according to the availability of resources and the seasons.

    “Furthermore, intermediate feeders are able to adapt their diets according to the availability of resources and the seasons (Hofmann, 1989).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Hofmann, 1989 source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation
    § 2.3 Faecal sample library · p. 5

    Pygmy hippo faeces can be classified as either territorial or as litter droppings, depending on their consistency and shape.

    “Similar to common hippos ( Hippopotamus amphibius Linnaeus , 1758), pygmy hippo faeces can be classified as either territorial or as litter droppings (Robinson et al., 2017), depending on their consistency and shape (soft/shapeless or solid, respectively).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Robinson et al., 2017

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductnone given
    § 4.1 Limitations · p. 12

    This study focused only on large plant fragments.

    “A third limitation is that this study focused only on large plant fragments.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationown data
    § 5 CONCLUSIONS · p. 13

    Pygmy hippos are generalist herbivores with a wide range of plant species consumed, including grasses and shrubs.

    “Our results confirmed that pygmy hippos are generalist herbivores with a wide range of plant species consumed, including grasses and shrubs, suggesting they should be classified as intermediate feeders.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Study design and conductreasoning
    § 2.3 Faecal sample library · p. 6

    n = 48 leaf fragments per sample allowed a representative subsample and kept the same number of leaf fragments per faeces.

    “Some samples having more leaves than others, n = 48, allowed us to have a representative subsample and to keep the same number of leaf fragments per faeces.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Pygmy hippo faeces are easy to locate in the natural habitat.
  • May combine two claims.

    Interpretationcitation
    § 4 DISCUSSION · p. 11

    Thin cell walls suggest that dicot species are better absorbed and may have been overlooked by the method.

    “Dicot species have a thin cell wall (Bodmer, 1990; Shipley, 1999), suggesting that they are better absorbed and therefore may have been overlooked by our method.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Study design and conductreasoning
    § 4.1 Limitations · p. 12

    The study was limited in descriptions because fewer features are visible on the adaxial side, and cells looked very similar, making it difficult to distinguish species (e.g., adaxial side of Dialium aubrevillei and Napoleona leonensis).

    “Indeed, as there are fewer features visible on the adaxial side, we were limited in the descriptions. The cells looked very similar, and it was difficult to distinguish one from another (i.e., adaxial side of Dialium aubrevillei and Napoleona leonensis ).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationown data
    § 4 DISCUSSION · p. 11

    Pygmy hippos in Taï National Park have a preference for N. bisserata, Pteris burtonii, Marantaceae species and S. crinita.

    “More generally, the data suggest that pygmy hippos in TNP have a preference for N . bisserata , Pteris burtonii , Marantaceae species and S . crinita , in line with what has been proposed by Bülow (1987) and Hentschel (1990).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Bülow (1987)

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Hentschel (1990)

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

29 of 29
  • The paper relies on this without support.

    Diet determination of endangered species is an essential element in defining successful conservation strategies and optimising captive breeding programmes.

    “Diet determination of endangered species is an essential element in defining successful conservation strategies and optimising captive breeding programmes.”
  • The paper relies on this without support.

    There is little information from the wild on pygmy hippos.

    “There is little information from the wild, which is mainly due to the species' cryptic behaviour and difficulties accessing their natural habitat.”
  • The paper relies on this without support.

    We are not aware of any systematic data on diet preference and composition in free-ranging pygmy hippos.

    “but we are not aware of any systematic data on diet preference and composition in free-ranging animals.”
  • The paper relies on this without support.

    Pygmy hippo faeces are easy to locate in the natural habitat.

    “Pygmy hippo faeces are easy to locate in the natural habitat.”
  • The paper relies on this without support.

    The two sides of the leaves can be difficult to identify in faeces.

    “As the two sides of the leaves can be difficult to identify in faeces, we measured each side of the leaf in this library.”
  • The paper relies on this without support.

    Pygmy hippos are unlikely to target most of the plant species of Taï National Park as food items, but will focus on shrubs and herbaceous plants, which represent only between 10 and 15% of all plant species.

    “However, pygmy hippos are unlikely to target most of these plants as food items, but will focus on shrubs and herbaceous plants, which represent only between 10 and 15% of all plant species.”
  • The authors state this as an assumption.

    Almost one third of the plant species of Taï National Park were analysed in this study.

    “Therefore, we assume that almost one third of these plant species were analysed in this study.”
  • The paper relies on this without support.

    Fruits and seeds are part of the pygmy hippo diet.

    “In order to consider the complete diet of the pygmy hippos, one should look at smaller fragments and also include fruits and seeds, because fruits and seeds are also part of the pygmy hippo diet.”
8 of 8

Cites

Flagged first, then by citations

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Cited by

Sorted by citations

Authors
Hendier, A., Chatelain, C., Du Pasquier, P., Paris, M., Ouattara, K., Koné, I., Croll, D., & Zuberbühler, K.
Affiliations
  • Swiss Centre for Scientific Research (CI)
  • Jardin Botanique de Genève (CH)
  • Institute of Biology University of Neuchâtel Neuchâtel Switzerland
  • University of Pretoria (ZA)
  • University of Neuchâtel (CH)
  • University of St Andrews (GB)
Corresponding author
Hendier, Alba
Venue
African Journal of Ecology
Publisher
Wiley
Published
2021-06-09
Volume
59
Issue
4
Pages
809-825
ISSN
  • 0141-6707 (ISSN-L)
  • 1365-2028
OpenAlex
W3169232486
Cited by
7
References
45
Citations by year
2026: 1 · 2024: 3 · 2023: 3
FWCI
0.78
Type
journal article
Language
en
Open access
closed
Locations
  • African Journal of Ecology (publishedVersion)
  • St Andrews Research Repository (St Andrews Research Repository) (acceptedVersion)
  • UpSpace Institutional Repository (University of Pretoria) (submittedVersion)
Topics
  • Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
  • Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
  • Physical Sciences › Environmental Science › Ecology › Isotope Analysis in Ecology
Keywords
pygmy hippopotamus, Choeropsis liberiensis, fecal analysis, herbivore diet, endangered species conservation, captive breeding, plant species preference, Taï National Park

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-27.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-27.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

What counts as a claim ›How we measure confidence ›How we check this ›

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