Abstract

This overview highlights the multiple pathways by which ungulates distribute diaspores across landscapes and over long distances. There are three primary and three secondary seed dispersal pathways where ungulate dispersal agents contribute to diaspore propagation, both with and without the intervention of other biotic (dung beetles) and abiotic (wind, gravity …) agents. These pathways may combine successive inter-dependent (diplochory) or totally isolated steps. Native, introduced and domestic ungulates co-occur in many ecosystems. They locally constitute diverse long-distance-vector assemblages for both native and exotic plants. However, ungulate taxonomic diversity conceals a much higher diversity in terms of the functional traits involved in ungulate-mediated dispersal: body morphology, body mass, feeding regime, digestive strategy, fur morphology, sociality, grooming behavior, habitat use. These traits may strongly affect one or more of the different phases in the dispersal loop (emigration, transfer and immigration), and consequently, impact the overall seed dispersal effectiveness, both quantitatively and qualitatively. In this study, we compare internal pathways, or endozoochory, where seeds have to survive digestive treatments (ingestion, ingestion and regurgitation), with external pathways, or epizoochory, where diaspores are carried on the outside of the vectors, including both primary epizoochory (direct adhesion to fur) and secondary epizoochory (diaspore-laden mud adhering to hooves or the body, transfer through contact with a conspecific). We further point out that it is not exceptional for diaspores from a given plant to be dispersed by a single vector through different processes, and for some to be moved through only a specific process but by a network of different vectors. Two parameters, however, remain obscure in this field of dispersal ecology: i) the relative proportion of all diaspores produced that are carried over long distances by large ungulates, and ii) the relative importance of ungulates on the whole as the main dispersal agent for plants. In a previous paper, we showed in particular that nearly half of the plants dispersed by fur-epizoochory did not feature any specific adaptations. We conclude by discussing methodological challenges and stressing research perspectives , like taking animal behavior and animal cognition into account, studying invasion of exotic plants and altitudinal plant dispersal.

Highlights

  • Contrasting with the defaunation process currently impacting large mammals in tropical forest ecosystems (Galetti and Dirzo, 2013), ungulate populations in temperate forests are rapidly increasing, sometimes locally reaching higher populations than their historic records

  • We propose to adapt the seminal conceptual framework of seed dispersal effectiveness for frugivory and endozoochory proposed by Schupp (1993) and revisited by Schupp et al (2010) to the two other ungulatemediated primary processes: regurgitation and fur-epizoochory

  • External mechanisms, where diaspores are carried on the outside of the vectors on various body parts, include primary fur-epizoochory and secondary epizoochorous processes: transfer through contact with a conspecific (Liehrmann et al, 2018) and diaspore-laden mud adhering to hooves (Schulze et al, 2014) or the body (Heinken and Raudnitschka, 2002)

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Summary

INTRODUCTION

Contrasting with the defaunation process currently impacting large mammals in tropical forest ecosystems (Galetti and Dirzo, 2013), ungulate populations in temperate forests are rapidly increasing, sometimes locally reaching higher populations than their historic records This phenomenon concerns overabundant native deer populations (Côté et al, 2004) and introduced species thriving in different parts of the world (e.g., Canada, Japan, Australia, New Zealand). Endozoochory, including frugivory, is the most widely studied ungulate-mediated plant dispersal mechanism (e.g., hoof- and fur-epizoochory concerns

Regurgitation or Partial Endozoochory
Endozoochory and Secondary Dispersal by Dung Beetles
Spatial Trajectory of Conveyed Diaspores
Number of Visits
Number of Diaspores Loaded Per Visit
Quality of the deposition site for germination and growth
Treatment Quality by the Vector
Deposit Quality of the Released Diaspores
Quality of the Deposition Site for Germination and Growth
Body mass
Transfer endo c
Total number of plant species dispersed
Endozoochory by at Least Two Ungulates
Methodological Challenges
Findings
Research Perspectives
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