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. 2014 Jul;4(14):2913-30.
doi: 10.1002/ece3.1136. Epub 2014 Jun 16.

Establishing macroecological trait datasets: digitalization, extrapolation, and validation of diet preferences in terrestrial mammals worldwide

Affiliations

Establishing macroecological trait datasets: digitalization, extrapolation, and validation of diet preferences in terrestrial mammals worldwide

Wilm Daniel Kissling et al. Ecol Evol. 2014 Jul.

Abstract

Ecological trait data are essential for understanding the broad-scale distribution of biodiversity and its response to global change. For animals, diet represents a fundamental aspect of species' evolutionary adaptations, ecological and functional roles, and trophic interactions. However, the importance of diet for macroevolutionary and macroecological dynamics remains little explored, partly because of the lack of comprehensive trait datasets. We compiled and evaluated a comprehensive global dataset of diet preferences of mammals ("MammalDIET"). Diet information was digitized from two global and cladewide data sources and errors of data entry by multiple data recorders were assessed. We then developed a hierarchical extrapolation procedure to fill-in diet information for species with missing information. Missing data were extrapolated with information from other taxonomic levels (genus, other species within the same genus, or family) and this extrapolation was subsequently validated both internally (with a jack-knife approach applied to the compiled species-level diet data) and externally (using independent species-level diet information from a comprehensive continentwide data source). Finally, we grouped mammal species into trophic levels and dietary guilds, and their species richness as well as their proportion of total richness were mapped at a global scale for those diet categories with good validation results. The success rate of correctly digitizing data was 94%, indicating that the consistency in data entry among multiple recorders was high. Data sources provided species-level diet information for a total of 2033 species (38% of all 5364 terrestrial mammal species, based on the IUCN taxonomy). For the remaining 3331 species, diet information was mostly extrapolated from genus-level diet information (48% of all terrestrial mammal species), and only rarely from other species within the same genus (6%) or from family level (8%). Internal and external validation showed that: (1) extrapolations were most reliable for primary food items; (2) several diet categories ("Animal", "Mammal", "Invertebrate", "Plant", "Seed", "Fruit", and "Leaf") had high proportions of correctly predicted diet ranks; and (3) the potential of correctly extrapolating specific diet categories varied both within and among clades. Global maps of species richness and proportion showed congruence among trophic levels, but also substantial discrepancies between dietary guilds. MammalDIET provides a comprehensive, unique and freely available dataset on diet preferences for all terrestrial mammals worldwide. It enables broad-scale analyses for specific trophic levels and dietary guilds, and a first assessment of trait conservatism in mammalian diet preferences at a global scale. The digitalization, extrapolation and validation procedures could be transferable to other trait data and taxa.

Keywords: Diet ecology; Mammalia; ecological trait data; feeding guild; phylogenetic conservatism; trophic structure.

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Figures

Figure 1
Figure 1
Four terrestrial mammal species representing different diet preferences. Upper left: African Elephant (Loxodonta africana), a typical herbivore. Upper right: the Gray Wolf (Canis lupus), a carnivore. Lower left: the Daubenton's Bat (Myotis daubentonii), an insectivore. Lower right: the European Badger (Meles meles), an omnivore. Photo credits: W. Daniel Kissling (elephant), Gary Kramer (wolf), Gilles San Martin (bat), Kókay Szabolcs (badger). The latter three were obtained from Wikimedia Commons (http://commons.wikimedia.org).
Figure 2
Figure 2
Schematic overview of how macroecological trait datasets can be established. Trait information from the literature is converted into a digital database (digitalization) and errors can be minimized and assessed during the digitalization process (calibration and quality check). Missing trait data are then filled from other taxonomic or phylogenetic levels (extrapolation). The performance of the extrapolation procedure can afterward be assessed internally and externally (validation). Finally, guilds or functional groups are classified (guild classification) and spatially mapped (spatial visualization). Grey boxes (middle and right) illustrate the key processes in the establishment of macroecological trait datasets, whereas white boxes (left) illustrate datasets that are additionally needed.
Figure 3
Figure 3
Hierarchical structure of the sixteen diet categories used for data entry into MammalDIET. Diet categories represent four different hierarchical levels of diet information. Detailed information about these diet categories is provided in Appendix Table S1.
Figure 4
Figure 4
Summary of (A) extrapolation, (B) internal validation, and (C) external validation of global diet knowledge in mammals. In (A) each boxplot summarizes the percentage of species within mammal families (n = 140) according to how extrapolation of diet information was done (FillCode = 0, 1, 2.1, 2.2 and 3). Extrapolation was not necessary for those species (n = 2033) for which species-level information was already available from the data sources (FillCode = 0). For the other species (n = 3331), extrapolation was performed from the genus level (FillCode = 1), from one other species in the genus (FillCode = 2.1), from more than one species in the genus (FillCode = 2.2), or from family level (FillCode = 3). Raw data are provided in Appendix Table S2. In (B) each boxplot summarizes the proportion of correctly predicted diet ranks for high (grey boxes) and low (white boxes) hierarchical levels (compare Fig. 3). High hierarchical levels include the diet categories “Animal”, “Plant”, “Vertebrate”, and “Invertebrate”, whereas the low hierarchical levels include all other diet categories. Information on ranks 1–3 is provided in Table 2. The “0″ indicates that a diet category was not used (i.e., assumed absence). In (C), extrapolated diet data are validated independently with an external validation dataset (Mammals of Africa, see text for details). The percentage of correctly predicted diet ranks is given for each of the sixteen diet categories for rank 1 data only (gray bars) and for rank 1 and 2 data combined (white bars). Numbers below diet categories give the sample size (number of species) for each validation. Boxes in (A) and (B) represent the interquartile range (IQR), horizontal lines within the boxes represent medians, whiskers extend to 1.5 times the IQR, and outliers are plotted as dots.
Figure 5
Figure 5
External validation of extrapolating diet knowledge for two mammal orders that contain species which use a broad range of either plant or animal diet categories. (A) Rodentia (here rodents such as African dormice, gerbils, mice, etc.) are predominantly herbivorous and insectivorous, but different species use different plant diet categories. (B) Carnivora (mostly represented here by genets and mongooses) predominantly feed on animal material, but the importance of different animal diet categories varies among species. Bars illustrate the percentage of correctly predicted diet ranks of each of the 16 diet categories for rank 1 data (gray bars) and for rank 1 and 2 data combined (white bars). Numbers below diet categories give the sample size (number of species) for each validation. The results of this external validation are based on the Mammals of Africa (see text for details).
Figure 6
Figure 6
External validation of extrapolating diet knowledge for three mammal orders that contain species which feed on a few diet categories. (A) Primates (here mostly monkeys and galagos) feed on “Plant” and “Fruit”, but the use of leaves and invertebrates varies among species. (B) Cetartiodactyla (here duikers, dik-diks, etc.) are herbivores with a specialization on fruits and leaves, but being a browser (“Woody” leaves) or a grazer (“Herbaceous” leaves) varies among species. (C) Eulipotyphla (shrews) are highly insectivorous (incl. invertebrates) with other food items being only eaten by a few species. Bars illustrate the percentage of correctly predicted diet ranks of each of the 16 diet categories for rank 1 data (gray bars) and for rank 1 and 2 data combined (white bars). Numbers below diet categories give the sample size (number of species) for each validation. The results of this external validation are based on the Mammals of Africa (see text for details).
Figure 7
Figure 7
Global species richness maps of trophic levels (A–C) and dietary guilds (D–H). For definition of trophic levels and dietary guilds see Table 5. The grids are in Behrmann projection (a cylindrical equal-area projection) with a resolution of 2° equivalents. Color ramps are in quantile classification, numbers beside color ramps refer to species richness. Grid cells with less than 50% land cover as well as those covering Antarctica are not included.
Figure 8
Figure 8
Proportions of trophic levels (A–C) and dietary guilds (D–H). For definition of trophic levels and dietary guilds see Table 5. The grids are in Behrmann projection (a cylindrical equal-area projection) with a resolution of 2° equivalents. Color ramps are in quantile classification. Grid cells with less than 50% land cover as well as those covering Antarctica are not included.

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