Abstract
Nonnative feral pigs (Sus scrofa) are recognized throughout the New World as a highly significant introduced species in terms of ecosystem alteration. Similarly, nonnative soil macroinvertebrates (e.g. earthworms, ground beetles) invade and alter the structure and function of native habitats globally. However, the relationship between feral pigs and soil macroinvertebrates remains largely unknown. This study analyzed relationships between these taxa using nine sites located inside and outside of feral pig management units representing a ~ 25 year chronosequence of removal in tropical montane wet forests in Hawai‘i. Soil macroinvertebrates were sampled from plots categorized as: actively trampled by feral pigs, actively rooted by feral pigs, feral pigs present with no signs of recent activity, or feral pigs removed over time. In total, we found 13 families of primarily nonnative soil macroinvertebrates. Plots with active trampling correlated with lower total macroinvertebrate abundance, biomass, and family richness. Plots with active rooting were correlated with higher abundance of nonnative earthworms (Lumbricidae and Megascolicidae) and ground beetles (Carabidae). The abundance, biomass, and biodiversity of macroinvertebrates did not vary with time since feral pig removal. Collectively, these results indicate: (1) trampling by feral pigs negatively influences soil macroinvertebrates; (2) feral pigs either modify habitats while rooting thereby facilitating earthworm and ground beetle habitat use or selectively seek out target prey species of soil macroinvertebrates; and (3) removal of feral pigs has minimal impacts on soil macroinvertebrates over time. These results are important globally due to the broadly overlapping ranges of S. scrofa and nonnative macroinvertebrates.

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Acknowledgements
We would like to thank the following people for their valuable contributions to this project: Dr. J. Crumsey and K. Laushman for study design; K. Francisco, Dr. J. Friday, and M. Pina for logistical support; Dr. C. Ritzenthaler for specimen identification; N. Babila for data collection and entry; E. Barrido, D. Christensen, N. Friday, S. Rivera, L. Sletton, and T. Tomita for fieldwork assistance; and, Dr. C. Giardina, Dr. G. Metzler, and three for helpful feedback on the manuscript. This work was supported by: the Watson T. Yoshimoto Fellowship distributed by the Ecology, Evolution, and Conservation Biology Graduate Specialization Program at the University of Hawai‘i at Mānoa; the College of Tropical Agriculture and Human Resources at the University of Hawai‘i at Mānoa via the U.S. Department of Agriculture National Institute of Food and Agriculture Hatch (HAW01127H) and McIntyre Stennis (HAW01123 M) Programs; the Department of Defense Strategic Environmental Research and Development Program (RC-2433); and the Invasive Species Program of the U.S. Geological Survey. We would like to thank the State of Hawai‘i Department of Land and Natural Resources, Division of Forestry and Wildlife and Hawai‘i Volcanoes National Park for access to research areas. Additionally, this article was, in part, created by a U.S. government employee and is in the public ___domain. Public ___domain information may be freely distributed and copied, but it is requested that any subsequent use be given appropriate acknowledgement. Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the U.S. Government.
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Wehr, N.H., Litton, C.M., Lincoln, N.K. et al. Relationships between soil macroinvertebrates and nonnative feral pigs (Sus scrofa) in Hawaiian tropical montane wet forests. Biol Invasions 22, 577–586 (2020). https://doi.org/10.1007/s10530-019-02117-3
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DOI: https://doi.org/10.1007/s10530-019-02117-3