A new method to determine the diet of pygmy hippopotamus in Taï National Park, Côte d’Ivoire
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.
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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.
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Claims the paper makes
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- 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
- Findingown data§ 3.2 Faecal samples · p. 8
In the HC tree, the clustering of plant groups (monocots, dicots and ferns) was represented along the two main branches as well as some plant families: Rapataceae, Marantaceae, Pteridaceae and Rubiaceae.
“In the HC tree, the clustering of the plant groups (monocots, dicots and ferns) was represented along the two main branches as well as some plant families: Rapataceae, Marantaceae, Pteridaceae and Rubiaceae.”
Quote in the paperNot a substantial claim - Findingown data§ 5 CONCLUSIONS · p. 13
Similar fragments of monocots (grasses), dicots (shrubs and tree leaves) and ferns were observed in almost all faeces analysed, collected from ten different individuals distributed over a combined home range area of about 50 km2.
“Indeed, we observed similar fragments of monocots (grasses), dicots (shrubs and tree leaves) and ferns in almost all faeces analysed, collected from ten different individuals distributed over a combined home range area of about 50km 2 .”
Quote in the paperSubstantial claim - Findingown data§ Note · p. 10
Nephrolepis bisserata could be identified in 7 of 10 faecal samples.
“For the ferns, Nephrolepis bisserata could be identified in 7 of 10 faecal samples (Appendix 3).”
Quote in the paperNot a substantial claim - Findingown data§ 3.2 Faecal samples · p. 8
After analysing each branch of the tree with corresponding pictures, the presence of seven species of plants from the reference list was confirmed.
“After analysing each branch of the tree (Figure S1) with the corresponding pictures, we were able to confirm the presence of seven species of plants from our reference list (Table 3).”
Quote in the paperSubstantial claim - Findingown data§ 4 DISCUSSION · p. 11
Species classified as favourites were difficult to find or absent in our area (e.g., Staurogyne paludosa, Justicia tenella, and Floscopa africana).
“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§ 4.1 Limitations · p. 12
The species identity of the seeds could not be determined.
“We were unable to determine the species identity of the seeds.”
Quote in the paperNot a substantial claim - Findingown data§ 3.1 Plant reference library · p. 7
The MCA did not allow gathering taxonomic ranks (species) inside families or at the generic level.
“There was no other structure apart from the distinction between these three groups. Indeed, the MCA did not allow us to gather taxonomic ranks (species) inside the families and even less at the generic level.”
Quote in the paperNot a substantial claim - Findingown data§ 4 DISCUSSION · p. 11
Centoteca lappaceae was found in all faecal samples.
“Centoteca lappaceae (grass, found in all samples)”
Quote in the paperNot a substantial claim - Findingown data§ 5 CONCLUSIONS · p. 13
The study compiled a tropical plant database consisting of 60 species.
“With this study, we also compiled a tropical plant database consisting of 60 species that may serve future faunistic studies in West African forests.”
Quote in the paperSubstantial claim - Findingown data§ 3.2 Faecal samples · p. 8
The reference library and the faeces library shared similar positions on the spatial projection.
“We noticed that the reference library and the faeces library shared similar positions on the spatial projection (Figure 2).”
Quote in the paperSubstantial claim - Interpretationcitation§ 4.1 Limitations · p. 12
The analyses could be compromised by the fact that male and female home ranges can overlap, suggesting that two different individuals could have contributed to each sampling location.
“Finally, these analyses could be compromised by the fact that male and female home ranges can overlap, suggesting that two different individuals could have contributed to each sampling location (Roth et al., 2004).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Roth et al., 2004 source
- Interpretationreasoning§ 4 DISCUSSION · p. 11
Pygmy hippos qualify as herbivorous generalists.
“This fact further supports our findings that pygmy hippos qualify as herbivorous generalists (Hentschel, 1990; Robinson et al., 2017), such that an individual's diet composition will largely depend on the local flora encountered during foraging.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Hentschel, 1990
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Robinson et al., 2017
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 1 INTRODUCTION · p. 2
Recent botanical studies have shown that microscopically determined foliar anatomy can produce reliable information for species identification, particularly at the family and group level.
“recent botanical studies have shown that the microscopically determined foliar anatomy can produce reliable information for species identification, particularly at the family and group level (Metcalfe and Chalk, 1950, 1957; Shah et al., 2018a, b; Ullah et al., 2018a, b).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Metcalfe and Chalk, 1950, 1957
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Shah et al., 2018a, b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Ullah et al., 2018a, b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationown data§ 4 DISCUSSION · p. 10
The method has potential as an interactive research tool for rapid diet identification in combination with established criteria, such as trichomes, silicates or scales.
“This analysis was based on only five categorical variables, thus demonstrating the potential of the method as an interactive research tool for rapid diet identification in combination with established criteria, such as trichomes, silicates or scales.”
Quote in the paperSubstantial claim - Premisecitation§ 4 DISCUSSION · p. 11
Dicot species have a thin cell wall.
“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 claim0 of 2 sources checked- cites Bodmer, 1990 source
- cites Shipley, 1999
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 4 DISCUSSION · p. 10
A generalist foraging strategy allows pygmy hippos to avoid over-ingestion of plant toxins.
“a foraging strategy that allows them to avoid over-ingestion of plant toxins (Freeland and Janzen, 1974).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Freeland and Janzen, 1974 source
- Study design and conductcitation§ 4.1 Limitations · p. 12
Less than 4% of the plant species of Taï National Park were part of the plant reference library (60 of 1356 documented species).
“Less than 4% of the plant species of TNP were part of the library (60 of 1356 documented species by Scouppe (2011)).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Scouppe (2011)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationown data§ ABSTRACT · p. 1
Pygmy hippopotami qualified as intermediate feeders.
“which revealed that pygmy hippopotami qualified as intermediate feeders”
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.
- Interpretationreasoning§ 4 DISCUSSION · p. 10
Pygmy hippos are to be classified as herbivore generalists.
“Hence, we can confirm that pygmy hippos are to be classified as herbivore generalists, a foraging strategy that allows them to avoid over-ingestion of plant toxins (Freeland and Janzen, 1974).”
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.
- Interpretationreasoning§ 4 DISCUSSION · p. 11
An individual pygmy hippo's diet composition will largely depend on the local flora encountered during foraging.
“This fact further supports our findings that pygmy hippos qualify as herbivorous generalists (Hentschel, 1990; Robinson et al., 2017), such that an individual's diet composition will largely depend on the local flora encountered during foraging.”
Quote in the paperSubstantial claim - Interpretationcitation§ 4 DISCUSSION · p. 11
Classifying pygmy hippos as intermediate feeders is supported by their gregarious and territorial behaviour, dentition, and other dietary studies.
“Classifying pygmy hippos as intermediate feeders is also supported by their gregarious and territorial behaviour, dentition (Lang, 1975), and other dietary studies (Bülow, 1987; Hentschel, 1990).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Lang, 1975
Couldn’t be matched to a paper in Crossref, so it can’t be 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.
- Interpretationreasoning§ 4 DISCUSSION · p. 11
The failure to find evidence for those dicots may be due to the study focusing on large fragments only.
“This may be due to the fact that we focussed on large fragments only (large particles ingested).”
Quote in the paperSubstantial claim - Interpretationown data§ 5 CONCLUSIONS · p. 13
Pygmy hippos should be classified as intermediate feeders.
“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 - Study design and conductcitation§ 2.3 Faecal sample library · p. 5
Selecting faecal samples at least 2 km apart ensures independent data points.
“To obtain independent data points, only samples that were at least 2 km apart were selected (home range estimates; females = 0.5 km 2 ; males = 1.5 km 2 ; Bülow, 1987; Hentschel, 1990) (Figure 1).”
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.
- Rests on an assumption: Pygmy hippo faeces are easy to locate in the natural habitat.
- May combine two claims. 23
- Classified as probably not a substantial claim. 9
May combine two claims.
Findingown data§ 5 CONCLUSIONS · p. 13C. 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 claimProbably not a substantial claim: may not pass S3 (load-bearing).
Findingown data§ 3.2 Faecal samples · p. 8Many 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 claimMay combine two claims.
Findingown data§ 4 DISCUSSION · p. 10At 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. 12It 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 claimProbably not a substantial claim: may not pass S3 (load-bearing).
Findingown data§ 4.1 Limitations · p. 12Matching 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 claimMay combine two claims.
Findingown data§ 4 DISCUSSION · p. 10The 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 claimMay combine two claims.
Findingown data§ Note · p. 10Marantaceae 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 claimMay combine two claims.
Findingown data§ Note · p. 108 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 claimMay combine two claims.
Findingown data§ 4 DISCUSSION · p. 11The 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 claimMay combine two claims.
Findingown data§ 4 DISCUSSION · p. 11No 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 claimMay combine two claims.
Findingown data§ ABSTRACT · p. 1Pygmy 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. 7Despite 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 claimMay combine two claims.
Findingown data§ 4 DISCUSSION · p. 10Free-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. 7The 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. 11Pygmy hippos are non-ruminant generalist intermediate feeders.
“All these observations support that pygmy hippos are non-ruminant generalist intermediate feeders.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductnone given§ 4.1 Limitations · p. 12Many 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 claimMay combine two claims.
Interpretationreasoning§ 5 CONCLUSIONS · p. 13Pygmy 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 claimClassified as probably not a substantial claim. May combine two claims.
Premisecitation§ 1 INTRODUCTION · p. 2In 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. 11Previous 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. 7The 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 claimMay combine two claims.
Interpretationreasoning§ 3.2 Faecal samples · p. 8The 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 claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ 4 DISCUSSION · p. 11Intermediate 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. 5Pygmy 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. 12This 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 claimMay combine two claims.
Interpretationown data§ 5 CONCLUSIONS · p. 13Pygmy 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 claimClassified as probably not a substantial claim. May combine two claims.
Study design and conductreasoning§ 2.3 Faecal sample library · p. 6n = 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. 11Thin 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 claimMay combine two claims.
Study design and conductreasoning§ 4.1 Limitations · p. 12The 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 claimMay combine two claims.
Interpretationown data§ 4 DISCUSSION · p. 11Pygmy 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.
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.”
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The grassy ecosystems of Madagascar in context: Ecology, evolution, and conservation
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“This includes the common hippo (Hippopotamus amphibious), pygmy hippo (Choeropsis liberiensis) [49], common tapir (Tapirus terrestris) [50], Malayan tapir (Tapirus indicus) [51], beaver (Castor fiber) [52], and European water vole (Arvicola amphibius) [53].”
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Kempkes, J. B., Si, Y., van ’t Zelfde, M., Digbeu, H. E. O., Bogui, E. B., Ouattara, K., Koné, I., & Paris, M. (2026) · Global Ecology and Conservation · doi:10.1016/j.gecco.2026.e04346
Self-citation
- 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
- ORCID
- 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
- Semantic Scholar
- a57847fb8c49adaa938393114254dd49cb3a9ce5
- 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
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