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not have the desired effect because of their low colonization in the intestine. By contrast, L. equi and
L. animalis may be more effective in herbivores. Lactobacillus acidophilus (L. acidophilus), which is
frequently found in the intestines of humans, was not found in the feces of zoo animals. We did not
examine primates in the present study, but the lactic acid bacteria used as probiotics for humans may not be
suitable for the zoo animals examined in this study. Although Lactobacillus spp. were not found in the feces
of the piscivorous or marine animals in this study, L. salivarius has been isolated from a rectal swab from a
bottlenose dolphin . The types of bacteria comprising the intestinal flora of piscivorous or marine animals
[59]
differed from those in the carnivorous and herbivorous terrestrial animals at the phylum level. Thus, the
selection of lactobacilli used as probiotics should be optimized.

DECLARATIONS
Acknowledgments
We are extremely grateful to the Tobe Zoological Park of Ehime Prefecture (Tobe, Ehime, Japan) and the
New Yashima Aquarium (Takamatsu, Kagawa, Japan) for providing animal fecal samples. We thank
Suzanne Leech, Ph.D., for editing the draft of this manuscript.

Authors’ contributions
Made substantial contributions to the conception and design of the study and performed data analysis and
interpretation: Horie M, Ohno T, Umemura M
Contributed to supervision: Iwahashi H, Murotomi K

Availability of data and materials
The isolated lactic acid bacteria strain was deposited at Japan Collection of Microorganisms (JCM), Riken
BioResource Research Center. The 16S rRNA gene sequence has been submitted to DDBJ. The datasets
generated and analyzed during the current study are available from the corresponding author upon
reasonable request.

Financial support and sponsorship
None.

Conflicts of interest
All authors declared that there are no conflicts of interest.

Ethical approval and consent to participate
We confirmed the consent form for the collection of animal fecal samples.

Consent for publication
Not applicable.

Copyright
© The Author(s) 2024.

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