DOI: 10.5061/dryad.3bk3j9ksb
title: Data from: seed dispersal by frugivores without seed swallowing: evaluating the contributions of stomatochoric seed dispersers

description: The process of seed dispersal that underpins ecosystem maintenance is
 performed by diverse arrays of fruit-eating animals. However, seed
 dispersal studies are primarily focused on a subset of these animal
 communities that disperse seeds by endozoochory. Stomatochory (seed
 dispersal in which seeds are carried externally and are not swallowed) is
 rarely considered to be effective, despite an increasing number of
 taxa-focused studies that indicate otherwise. We collated the available
 information on stomatochory to provide a quantitative overview of the
 dispersal mechanism, including plant-types and fruit-traits dispersed,
 dispersal distances and germination potential for all available taxa. We
 compared seed sizes dispersed, dispersal distances, and germination
 potential with corresponding data on endozoochory for bats and primates.
 We also identified the main taxa dispersing seeds by stomatochory and
 assessed what factors influenced the distances that they carried seeds to.
 Stomatochoric dispersers can displace large quantities of seeds, including
 large seeds and those of large fruits, and over short to long distances
 (&gt; 1 km). Compared to similar-sized endozoochoric dispersers, they
 can disperse larger seeds, but over shorter distances, on average. Similar
 to endozoochory, seed handling by stomatochory also improves the
 germination potential of dispersed seeds. Dispersal distances achieved by
 stomatochory were influenced by body mass, daily path length, seed width,
 fruit type and seed handling techniques. Five main taxonomic groups of
 stomatochoric dispersers were identified: bats, parrots, squirrels,
 corvids and Old World monkeys (cercopithecines). Parrots perform dispersal
 services for the largest fruits and over the longest distances. However,
 given the lack of research on stomatochory, it is likely other taxa are
 also important stomatochoric dispersers but have not been identified yet.
 More research attention must be directed towards seed dispersal services
 that are not provided by endozoochory. Many stomatochoric dispersers are
 common animals within communities and could be playing dominant seed
 dispersal roles, even without swallowing seeds. Community-wide studies
 should incorporate all seed dispersal interactions, rather than focusing
 solely on endozoochory. This will ensure a more robust understanding of
 community-wide patterns.
description: We searched for published articles reporting stomatochory
 observations using Google Scholar, Web of Science and Scopus. Searches
 included the words “seed carrying” or “pulp predators” or “pulp consumers”
 or “pulp feeders” or "stomatochory". We scanned the titles and
 abstracts of the papers to determine if they were likely to include
 information on stomatochory and extracted the appropriate information from
 the full paper. We also used these papers to identify taxa that regularly
 consumed pulp-only or often carried seeds away from parent crowns and
 conducted more intensive searches on these taxa (mainly primates, rodents,
 bats, parrots, corvids). We excluded papers on synzoochory and only
 included papers documenting carrying of fruits for the immediate
 consumption of pulp (e.g., by squirrels), or seeds (e.g., husking stations
 of rats). Ruminants are also identified as “seed spitters”, but in these
 cases seeds are transported in the rumen and are a form of endozoochory so
 are not considered here (Delibes et al. 2019). For each
 stomatochory interaction between animal species and plant species we
 recorded details on the type of stomatochory behaviour: (i) spat (seed
 stored within the mouth or cheek pouches, and then spat out once pulp is
 removed. The seed/s might be spat with mouthfuls of fruit fibre) (Aziz et
 al., 2021; Lambert, 1999), (ii) dropped (fruit is carried in the mouth,
 beak or by the feet (Tella et al. 2020) and pulp is removed externally,
 before the seed is dropped, or the seed might be dropped incidentally
 while traveling (Aziz et al. 2021, Blanco et al. 2015). We only recorded
 an interaction when the animal had been recorded to remove a seed from the
 vicinity of the parent plant on at least one occasion. The searches
 generated many papers of pulp feeding of birds, but only rarely was seed
 removal confirmed to occur; hence, these interactions were not included in
 the database.  When available, we also recorded details
 on intact rate (how many handled seeds were deposited undamaged),
 proportion of seeds removed beyond the canopy area, seed dispersal
 distances, seed germination, fruit and seed size, plant life-form and
 fruit-type; the trait data for plants was supplemented by searching for
 information from other sources. When a choice of data summary was
 available (e.g., for dispersal distance) we preferentially used medians,
 but some values represent mean since this was the only value available. We
 also collected data on germination tests for endozoochoric interactions
 involving cercopithecine primates and bats, as a comparison to the
 stomatochory data.  Literature cited
 Aziz, S.A., McConkey, K.R., Tanalgo, K., Sritongchuay, T., Low,
 M.-R., Yong, J.Y., Mildenstein, T.L., Nuevo-Diego, C.E., Lim, V.-C.,
 Racey, P.A., 2021. The Critical Importance of Old World Fruit Bats for
 Healthy Ecosystems and Economies. Frontiers in Ecology and Evolution 9.
 https://doi.org/10.3389/fevo.2021.641411 Blanco, G., Hiraldo, F., Rojas, A., Denes, F. V., Tella, J. L. 2015. Parrots as key multilinkers in ecosystem structure and functioning. Ecology and Evolution 5: 4141-4160. https://doi.org/10.1002/ece3.1663 Delibes M., Castaneda I, Fedriani J. M. 2019. Spitting seeds from the cud: a review of an endozoochory exclusive to ruminants. Frontiers in Ecology and Evolution 7 https://doi.org/10.3389/fevo.2019.00265 Lambert, J.E., 1999. Seed handling in chimpanzees (<em>Pan troglodytes</em>) and redtail monkeys (<em>Cercopithecus ascanius</em>): Implications for understanding hominoid and cercopithecine fruit‐processing strategies and seed dispersal. American Journal of Physical Anthropology 109, 365–386. https://doi.org/10.1002/(SICI)1096-8644(199907)109:365 Tella, J.L., Hiraldo, F., Pacífico, E., Díaz-Luque, J.A., Dénes, F.V., Fontoura, F.M., Guedes, N., Blanco, G., 2020. Conserving the diversity of ecological interactions: The role of two threatened macaw species as legitimate dispersers of “megafaunal” fruits. Diversity 12, 45. https://doi.org/10.3390/d12020045
description: # Seed dispersal by frugivores without seed swallowing: evaluating the
 contributions of stomatochoric seed dispersers
 The database collates
 information on stomatochoric seed dispersal interactions from articles
 published until 2022. All vertebrate seed dispersers were covered in the
 initial searches, but the final database consists of birds and mammals
 only, from across all habitat types and regions (for which data were
 available). Stomatochoric seed dispersal interactions are defined as
 dispersal occurring when seeds are transported in the mouth or limbs of
 the animal, without being swallowed. The database also contains data on
 endozoochoric interactions for cercopithecine monkeys and bats. ##
 Description of the data and file structure The stomatochory database is an
 excel file with four worksheets. In all datasheets NA refers to missing
 data. `Main stomatochory dataset` has 20 columns; each row refers to a
 specific animal-plant interaction: Animal family Animal genus Animal
 species Plant family Plant genus Plant species Life form (of the plant
 species) Habitat (in which the interaction occurred) Location of study
 Country Mode of dispersal (whether seed is 'dropped' or
 'spat' from the mouth) Average dispersal distance (as reported
 in the study using median or mean as reported) Maximum dispersal distance
 (as reported in the study) Percent seeds dispersed under the crown (under
 the crown of the source tree) Fruit width (in mm) Seed length (in mm) Seed
 width (in mm) Fruit-type (as described in the literature) Intact rate
 (percent seeds dispersed without being damaged) References
 `Stomat_germ_ratio_alltaxa` has 14 columns; each row refers to a specific
 animal-plant interaction. This dataset collates the results of germination
 tests from published studies on stomatochory. It includes all animal taxa
 for which data could be found. Animal family Animal genus Animal species
 Plant family Plant genus Plant species Does it germinate? (yes/no answer
 as to whether the tested seeds germinated or not) Germination percent for
 stomatochory (percent of seeds tested that germinated) Number of
 stomatochory seeds that germinated (sample size that germinated) Total
 number of stomatochory seeds tested (sample size that were tested)
 Germination percent for control seeds (percent of control seeds that
 germinated) Number of control seeds that germinated (sample size for
 control that germinated) Total number of control seeds tested (sample size
 for control that were tested) `endo_germ-ratio_cerco&bats` has 13
 columns; each row refers to a specific animal-plant interaction. This
 dataset collates the results of germination tests from published studies
 on endozoochory for cercopithecine monkeys and bats only. Animal family
 Animal genus Animal species Plant family Plant genus Plant species
 Germination percent for ingested (percent of seeds tested that germinated)
 Number of ingested seeds that germinated (sample size that germinated)
 Total number of ingested seeds tested (sample size that were tested)
 Germination percent for control seeds (percent of control seeds that
 germinated) Number of control seeds that germinated (sample size for
 control that germinated) Total number of control seeds tested (sample size
 for control that were tested) `References` has a single column listing the
 references used in all four worksheets. The references are arranged
 alphabetically.