Articulo 2

TAXONOMIC DESCRIPTION
Zheng et al., Int. J. Syst. Evol. Microbiol. 2020;70:2782–2858

DOI 10.1099/ijsem.0.004107
A taxonomic note on the genus Lactobacillus: Description

of 23 novel genera, emended description of the genus
Lactobacillus Beijerinck 1901, and union of Lactobacillaceae
and Leuconostocaceae
Jinshui Zheng1†, Stijn Wittouck2†, Elisa Salvetti3†, Charles M.A.P. Franz4, Hugh M.B. Harris5, Paola Mattarelli6, Paul
W. O’Toole5, Bruno Pot7, Peter Vandamme8, Jens Walter9,10, Koichi Watanabe11,12, Sander Wuyts2, Giovanna E. Felis3,*,†,
Michael G. Gänzle9,13,*,† and Sarah Lebeer2†
Abstract
The genus Lactobacillus comprises 261 species (at March 2020) that are extremely diverse at phenotypic, ecological and gen-
otypic levels. This study evaluated the taxonomy of Lactobacillaceae and Leuconostocaceae on the basis of whole genome
sequences. Parameters that were evaluated included core genome phylogeny, (conserved) pairwise average amino acid identity,
clade-specific signature genes, physiological criteria and the ecology of the organisms. Based on this polyphasic approach, we
propose reclassification of the genus Lactobacillus into 25 genera including the emended genus Lactobacillus, which includes
host-adapted organisms that have been referred to as the Lactobacillus delbrueckii group, Paralactobacillus and 23 novel genera
for which the names Holzapfelia, Amylolactobacillus, Bombilactobacillus, Companilactobacillus, Lapidilactobacillus, Agrilactobacil-
lus, Schleiferilactobacillus, Loigolactobacilus, Lacticaseibacillus, Latilactobacillus, Dellaglioa, Liquorilactobacillus, Ligilactobacillus,
Lactiplantibacillus, Furfurilactobacillus, Paucilactobacillus, Limosilactobacillus, Fructilactobacillus, Acetilactobacillus, Apilactobacil-
lus, Levilactobacillus, Secundilactobacillus and Lentilactobacillus are proposed. We also propose to emend the description of
the family Lactobacillaceae to include all genera that were previously included in families Lactobacillaceae and Leuconosto-
caceae. The generic term ‘lactobacilli’ will remain useful to designate all organisms that were classified as Lactobacillaceae until
2020. This reclassification reflects the phylogenetic position of the micro-organisms, and groups lactobacilli into robust clades
with shared ecological and metabolic properties, as exemplified for the emended genus Lactobacillus encompassing species
adapted to vertebrates (such as Lactobacillus delbrueckii, Lactobacillus iners, Lactobacillus crispatus, Lactobacillus jensensii,
Lactobacillus johnsonii and Lactobacillus acidophilus) or invertebrates (such as Lactobacillus apis and Lactobacillus bombicola).
Introduction anaerobic and non-spore-forming micro-organisms. The

genus is classified in the phylum Firmicutes, class Bacilli, order
The genus Lactobacillus was proposed by Beijerinck in 1901 Lactobacillales, family Lactobacillaceae, which contains the
and includes Gram-
positive, fermentative, facultatively genera Lactobacillus, Paralactobacillus, and Pediococcus. The
Author affiliations: 1Huazhong Agricultural University, State Key Laboratory of Agricultural Microbiology, Hubei Key Laboratory of Agricultural
Bioinformatics, Wuhan, Hubei, PR China; 2Research Group Environmental Ecology and Applied Microbiology, Department of Bioscience Engineering,
University of Antwerp, Antwerp, Belgium; 3Dept. of Biotechnology, University of Verona, Verona, Italy; 4Max Rubner-Institut, Department of Microbiology
and Biotechnology, Kiel, Germany; 5School of Microbiology & APC Microbiome Ireland, University College Cork, Co. Cork, Ireland; 6University of
Bologna, Dept. of Agricultural and Food Sciences, Bologna, Italy; 7Research Group of Industrial Microbiology and Food Biotechnology (IMDO), Vrije
Universiteit Brussel, Brussels, Belgium; 8Laboratory of Microbiology, Department of Biochemistry and Microbiology, Ghent University, Ghent, Belgium;
9
Department of Agricultural, Food & Nutritional Science, University of Alberta, Edmonton, Canada; 10Department of Biological Sciences, University of
Alberta, Edmonton, Canada; 11National Taiwan University, Dept. of Animal Science and Technology, Taipei, Taiwan, ROC; 12Food Industry Research and
Development Institute, Bioresource Collection and Research Center, Hsinchu, Taiwan, ROC; 13Hubei University of Technology, College of Bioengineering
and Food Science, Wuhan, Hubei, PR China.
*Correspondence: Giovanna E. Felis, giovanna.felis@univr.it; Michael G. Gänzle, mgaenzle@ualberta.ca
Keywords: Lactobacillus; Pediococcus; Lactobacillaceae; Leuconostocaceae; revised taxonomy.
Abbreviations: AAI, pairwise amino acid identity; ANI, average nucleotide identity; cAAI, pairwise amino acid identity of conserved genes.
†These authors also contributed equally to this work
†These authors share senior authorship.
Six supplementary tables and seven supplementary figures are available with the online version of this article.
004107 © 2020 The Authors

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Leuconostocaceae, including the genera Convivina, Fructoba-

cillus, Leuconostoc, Oenococcus and Weissella, are the closest
relatives at the family level [1].
The early taxonomy of lactobacilli was based on phenotypic
traits including optimal growth temperature, sugar utilization
and spectrum of metabolites produced [2]. Later in the 20th
century, genotypic and chemotaxonomic criteria including
DNA–DNA hybridization, the mol% G+C content and the
chemical structure of the peptidoglycan were used for the
delineation of new bacterial species. Since 1983, the similarity
of 16S rRNA genes has been used in bacterial taxonomy to
provide phylogenetic schemes as backbones for classifica-
tion and nomenclature [3]. Within the last 15 years, the
sequencing of whole bacterial genomes became widely avail-
able and average nucleotide identity (ANI) values of genes
shared between two bacterial genomes was introduced as the
gold standard for delineation of new bacterial species [4, 5].
Fig. 1. Cumulative number of species described in the genera
Although proposals have been made to apply a method-free
Pediococcus and Lactobacillus until January 2020. The species
species concept that is based on cohesive evolutionary forces description in January 2020 is included in the 2010–2019 count.
[6], an ANI value of 95 % (94–96 %) has been almost consist-
ently used in recent years to describe new bacterial species
[7, 8] and was suggested to reflect the biological significance
of the prokaryotic species concept [9]. Published species of It has been increasingly recognized that the genus Lactoba-
Lactobacillaceae and Leuconostocaceae were recently evalu- cillus as currently defined displays a level of genetic diversity
ated based on single-copy core genes and a fixed genome- that by far exceeds what is generally found in bacterial genera
genome similarity cutoff; the species were exclusive and and even bacterial families. The different phylogroups within
discontinuous [10]. the genus, however, are composed of species with a phylo-
genetic and physiological diversity that match the diversity
The introduction of genotypic methods such as 16S rRNA- of other bacterial genera [14]. Since 2015, several large-scale
based phylogeny already revealed the extensive diversity of phylogenetic analyses based on core genome phylogeny of
the genus Lactobacillus, which includes the genus Pedio- a comprehensive representation of species of the genera
coccus as an integral part [11–13]. Phylogenetic trees on Lactobacillus and Pediococcus have revealed the discrepancy
the basis of single genes including the 16S rRNA gene allow between the taxonomy of these genera and other bacterial
grouping of species to phylogenetic clades; however, they genera [14, 16, 17]. The continuous species descriptions in
do not have sufficient resolution to inform on the phyloge- the last years – 80 since 2015 – has brought the number of
netic relationships of different clades [13, 14]. Phylogenetic validly named species of Lactobacillus and Pediococcus to 272
studies of lactobacilli and pediococci on the basis of core (Fig. 1 and Table S1, available in the online version of this
genome phylogeny confirmed this phylogenetic heteroge- article), thus further increasing the diversity that is associ-
neity; in addition, the greater resolution of core genome ated with the genus Lactobacillus. The current taxonomy,
phylogeny established the presence of consistent clades or although widely accepted by medical community, food
phylogroups that are characterized by common phenotypic and health-related industries, by lay-persons, and used in
and ecological traits [12–17]. Physiological and phyloge- national and international regulations, impedes research
netic considerations, and genomic analyses also replaced the aimed at understanding the ecology, physiology, evolution
earlier differentiation between ‘obligate homofermentative’, and applications of this important group of organisms. This
‘facultative heterofermentative’ and ‘obligate heterofermen- is because micro-organisms that are genetically very distinct
tative’ lactic acid bacteria with a simpler differentiation and metabolically, ecologically and functionally very diverse
of homofermentative lactic acid bacteria that metabolize are grouped within the same genus. The lack of refined taxo-
hexoses via the Embden–Meyerhoff pathway to pyruvate nomic structure, moreover, encourages the addition of new
as the key metabolic intermediate, and heterofermentative species to a genus that has already been shown to be overly
lactic acid bacteria, that metabolize hexoses via the phos- heterogeneous and prevents the detection and description of
phoketolase pathway to pyruvate and acetyl-phosphate as functional properties or other communalities shared between
key intermediates [14, 18]. Within the genus Lactobacillus, members of the subgroups. However, a proposal for a formal
homofermentative and heterofermentative lactobacilli form re-evaluation of the taxonomy of the genera Lactobacillus and
distinct phylogenetic clades [14, 16], while homolactic or Pediococcus has not been made so far.
heterolactic metabolism is conserved in other lactic acid Here we re-evaluate the genetic relatedness and phylogeny
bacteria at the family level. Pentose metabolism, however, of the species within the present genus Lactobacillus and
is variable at the species or strain level [14]. its sister taxa in the Lactobacillaceae and Leuconostocaceae
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through a polyphasic approach [19]. For this, we considered multiple outgroups were added. To select outgroups, we used
average nucleotide identity (ANI), average amino acid iden- the Genome Taxonomy Database [27] to filter for the best
tity (AAI), core-gene average amino acid identity (cAAI), quality genome for each of the 50 genera belonging to the
core genome phylogeny, signature genes, and metabolic or order Lactobacillales (excluding species of Lactobacillaceae
ecological criteria. Within the Lactobacillaceae, 26 lineages and Leuconostocaceae), as evaluated by CheckM completeness
were identified that are reliably separated and are character- values. Potential outgroups with more than 5 % estimated
ized by conserved phenotypes and clade-specific signature contamination were excluded. The list of genomes used in
genes. Twenty-three of these clades are described here as new the analysis is shown in Table S2. Genes were predicted
genera, the description of the genera Lactobacillus and Para- for ingroup and outgroup genomes with Prodigal version
lactobacillus is emended while the description of the genus 2.6.3 [21]. Single-copy core genes were extracted using
Pediococcus remains unchanged. Furthermore, data of the progenomics version 0.1.0 [28] with the following options:
present study demonstrate that the family Leuconostocaceae 30 seed genomes, minimum presence in 25 seed genomes,
Schleifer 2010 should be considered a later synonym of the and required single-copy presence in 99 % of all genomes.
family Lactobacillaceae Winslow et al. 1917 (Approved Lists Core genes were aligned using mafft version 7.407 [29].
1980). The aligned core genes were then concatenated into a protein
supermatrix, and columns with more than 10 % gaps were
removed using trimal version 1.4.rev15 [24]. A phylogeny
Methods was inferred on the supermatrix with iq-tree version 1.6.11
Phylogenomic re-evaluation with RAxML and [30], using the LG+G+F protein substitution model. Branch
genome sequences of type strains support was calculated using 1000 bootstrap trees with the
UFBoot2 algorithm [31]. Finally, the full tree was rooted using
Genome sequences for type strains of all species in the
the clade (Listeria, Listeria_A, Brochothrix) as an outgroup.
families Lactobacillaceae and Leuconostocaceae which were
The tree was visualized using ggtree version 1.16.0 [32].
available on 19 August 2019 were obtained from GenBank
(genome set 1; Table S1). A listing of all species in the
Lactobacillaceae is also available on https://site.unibo.it/ Calculation of average amino acid identity (AAI) and
subcommittee-lactobacillus-bifidobacterium/en and http:// average amino acid identity of core proteins (cAAI)
lactobacillus.ualberta.ca; the file hosted on the websites will AAI was calculated between each pair of type strain genome
be periodically updated by adding new species that have been sequences (genome set 1) using CompareM [33]. cAAI values
validly published. All genomes were reannotated by Prokka were additionally calculated to minimize the impact of hori-
[20], which uses Prodigal [21] for gene prediction. Protein zontal gene transfer on pairwise AAI values. Acquisition of
sequences for each genome were extracted for gene clus- genes by lateral gene transfer substantially contributes to
tering. FastOrtho was used to generate gene families based genetic diversity in bacteria [34]. The extent of gene acquisi-
on protein sequences. First, an all-against-all alignment was tion by lateral gene transfer depends on the lifestyle of the
performed by blastP[22] with an E-value cutoff of 10−10. micro-organism [35]. Lateral gene transfer distorts the molec-
Then, ortholog groups were created with the MCL algorithm ular clock of bacterial evolution and the inferred phylogenetic
with an inflation value of 2. The clustering results were filtered relationship between different species [36], particularly for
by a local Perl script as previously described [14] to reduce micro-organisms that share a habitat and evolve by acquisi-
the influence of the fragmented protein sequences. All of the tion of similar genes. To minimize the impact of lateral gene
114 single-copy core gene families of Lactobacillaceae and transfer, we defined the cAAI based on the protein sequences
Leuconostocaceae were used for phylogenetic analysis. Protein of core gene families. We used soft core gene families that are
sequences of each family were aligned by muscle [23]. All shared by more than 90 % of the genomes studied. Protein
the 114 alignments were trimmed with TrimAl [24] and were sequences belonging to the soft-core gene families of each
concatenated into a new alignment through a local Perl script. genome were used to calculate cAAI values by CompareM.
RAxML [25] was used for phylogenetic analysis based on the Differences between cAAI and AAI indicate divergent or
core gene alignment with PROTGAMMAILGF (LG+I+G+F) convergent evolution through acquisition of diverse or
as the substitution model and 500 bootstrap samplings. The common genes, respectively. In addition, habitat adaptation
phylogenetic tree was visualized with iTOL [26]. by specific phylogenetic clades was assessed by analysis of
clade-specific genes as described below.
Phylogenomic re-evaluation with iq-tree and best
quality genomes Identification of genes that are exclusive to specific
A second tree of Lactobacillaceae and Leuconostocaceae phylogenetic clades (signature genes)
species was calculated by using one representative genome The complete pangenome of all best- quality genomes
per species (genome set 2) as described earlier [10]. Of the (genome set 2) was inferred using OrthoFinder 2.3.3 [37] with
2459 high-quality genomes analysed, 16 genomes could not MMseqs2 version d36de [38] as the sequence search program.
be assigned to a known species owing to low whole-genome For the analysis of gene family presence/absence and signa-
similarity or low 16S rRNA gene similarity to type strains; ture genes, the outgroup genomes were removed from the
we labelled these species as ‘unassigned species’. In addition, tree and the pangenome dataset. The gene family presence/
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absence plot was created by grouping together gene fami- between different habitats [17]. For example, the L. mali
lies with the same presence/absence pattern across species, group that predominantly includes micro-organisms with a
leaving out two types of trivial patterns: gene family presence free-living lifestyle clusters separately from the L. salivarius
in a single species and gene family presence in all species. The group, which predominantly includes micro-organisms with a
patterns with a frequency (number of gene families) of four or vertebrate host-adapted lifestyle [17, 27](Fig. 2). New species
more were then visualized in descending order of frequency. that were described since 2017 all conform to the lifestyle that
Signature genes were defined as gene families present in all was assigned to other species in the same phylogenetic tree
genomes of a clade (subtree) and absent in all other genomes. (Fig. 2) [17].
Phylogroups were defined by their type species; species were
assigned to the phylogroup of the type species with whom Evaluation on basis of cAAI and AAI values
they shared the most recent common ancestor.
To validate the assumption that each of these 26 phylogenetic
For all analysis involving the core genome phylogenetic tree groups in the Lactobacillaceae encompass a phylogenetic
and the gene family presence/absence patterns, R version diversity that is commonly assigned to a bacterial genus, all 38
3.6.1 (www.R-project.org) was used for data processing 364 pairwise cAAI and AAI values for the Leuconostocaceae
and visualization. R packages that were crucial included and Lactobacillaceae species for which genome sequences of
the tidyverse version 1.2.1 (https://CRAN.R-project.org/ the type strains were available were calculated (Table S3 and
package=tidyverse) and tidygenomes version 0.1.2 [39]. The S4). The intra-family cAAI and AAI values for Lactobacil-
code used for these analyses is deposited in two repositories laceae, which are largely shaped by intra-genus cAAI values
on GitHub [40]. of Lactobacillus species, overlap with the inter-family cAAI
and AAI values for Lactobacillaceae and Leuconostocaceae,
confirming the phylogenetic heterogeneity of Lactobacillus as
Results well as the lack of a clear separetion between Lactobacillaceae
Phylogenomic evaluation of Lactobacillaceae and Leuconostocaceae (Fig. 3 and Fig. S1). If intra-group
cAAI and AAI values are calculated for the 26 phylogenetic
A core genome phylogenetic tree of all Lactobacillaceae type
groups of Lactobacillaceae excluding Pediococcus species, the
strains for which genome sequence data was available in
distribution of intra-group cAAI and AAI values matched
August 2019 is shown in Fig. 2. The Lactobacillaceae consist-
the intra-genus distribution observed in Pediococcus, Weis-
ently form 26 phylogenetic groups that match all prior core-
sella and Leuconostoc (Fig. 4 and Fig. S2). Inter-genus cAAI
genome phylogenetic analyses of the family although more
values of more than 70 % are observed for the L. brevis and L.
than 60 genomes were added since 2015 [13, 14, 16, 17] and
collinoides groups. Despite these relatively high cAAI values,
are supported with 100 % bootstrap values, indicating they
the assignment of the L. brevis group and the L. collinoides
have been shaped by cohesive evolutionary forces. Below,
group is justified on the basis of the consistent clustering in
we propose genus status for each of these 26 phylogenetic
phylogenetic trees (Fig. 2) and the distinct metabolism and
groups, which include an emended genus Lactobacillus (for
ecology of species in the L. brevis and L. collinoides groups.
the phylogenetic group that includes L. delbrueckii, the type
Likewise, the description of the genus Convivina despite high
species of Lactobacillus), Paralactobacillus, Pediococcus and
AAI values to Fructobacillus was based on ecological and
23 new genera consisting of species previously assigned to
metabolic criteria Fructobacillus [42].
the genus Lactobacillus.
The intra-group distribution of cAAI and AAI values is shown
The L. delbrueckii group that contains the type strain of the
in Fig. 5 and Fig. S3. With the exception of the L. brevis, L.
genus forms a monophyletic clade with L. floricola and L.
collinoides, L. reuteri and L. salivarius groups, all groups are
amylophilus. Heterofermentative lactobacilli together with
exclusive, i.e. the lowest intra-group cAAI values are higher
pediococci and the L. plantarum group also form a mono-
than the highest inter-group cAAI values for any species in
phyletic clade; the remaining homofermentative lactobacilli,
the group (indicated in red in Fig. 5; data provided in Table
however, are not monophyletic (Fig. 2). Virtually all of the
S3). The L. delbrueckii group, the L. casei group and the L. sali-
Lactobacillus species that were described over the past 2 years
varius group are among the most diverse groups. The diver-
cluster within one of the phylogenetic groups identified in
sity within the L. casei and the L. salivarius groups relative
2017 (Fig. 2) [17]. Of the recently described species, only
to other groups is higher on the basis of AAI values (Figure
the candidate species Lactobacillus jinshani [41] can not be
S3) than when assessed on the basis of cAAI values (Fig. 5).
assigned to one of the established phylogenetic groups and
This implies that the diversification of the species in these
forms a separate lineage that is currently represented by only
groups is mediated by horizontal gene transfer and acquisi-
this candidate species.
tion of lifestyle-associated genes (see below). This observa-
We have previously shown that the lifestyle of species within tion matches the diverse lifestyles in the L. salivarius group,
groups, as inferred from isolation source, genomic, func- which contains free-living species or species with unknown
tional, ecological and physiological information, is remark- lifestyle in addition to host-adapted species. Information on
ably conserved within these groups and can be assigned to the ecology of the divergent L. pantheris/L. sharpeae clade
insect or flower-associated habitats, vertebrate hosts, to a within the L. casei group is currently too limited to conclude
free-living lifestyle, or to a nomadic lifestyle that transitions on the lifestyle of these organisms.
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Fig. 2. Core genome phylogenetic tree of Lactobacillaceae. The phylogenomic analysis is based on the concatenated alignment of protein
sequences for the 114 single-copy core genes. The maximum likelihood tree was inferred by RAxML as described previously [14] using
the 244 Lactobacillus and Pediococcus species for which genome sequence data was available on the NCBI database on 19 August 2019.
The tree was rooted via midpoint rooting. Bootstrap support values were calculated from 500 replicates, and only values above 90 %
were labelled. Members of the same phylogenetic group that are the basis for the proposed taxonomy are indicated by the same colour
for branches, and the type strain of each group is printed in bold. Outer rings provide information on genomic features and the inferred
lifestyle of the species. The colour gradient in red represents the GC content of each genome sequence; higher GC contents are indicated
by darker shading. The solid circles in brown represent genome sizes; the area of the circle correlates with the genome size. The second
ring indicates the inferred natural habitats of the species as vertebrate host-adapted (red), insect-adapted (orange), nomadic (green),
free-living (blue) or unassigned (white). This assignment of species to lifestyle was based on [17].
Identification of genes that are exclusive to specific properties. Signature genes therefore allow inferences about
phylogenetic clades (signature genes) the evolutionary forces that likely shaped the cluster.
To determine whether the 26 phylogenetic groups can be Most of the proposed new genera are supported by signature
characterized by clade-specific genes, we analysed the pres- genes. This is especially the case for the L. delbrueckii, L.
ence/absence of gene families within all species of the families alimentarius, L. mellifer, L. concavus, L. perolens, L. sakei, L.
Lactobacillaceae and Leuconostocaceae [43]. Gene families coryniformis, L. casei, L. plantarum, L. rossiae, L. vaccinos-
that are present in all best-quality genomes of a certain clade tercus, L. kunkeei, L. fructivorans and L. brevis groups. Each of
but are absent in all other genomes were termed signature these phylogroups contain at least four signature genes (Fig. 6,
genes. Signature genes reflect a common evolutionary history Table S5). The genera Leuconostoc, Fructobacillus, Oenococcus,
of a phylogenetic clade or result from horizontal gene transfer Weissella and Pediococcus also show a relatively large number
and relate to common lifestyles, ecologies and physiological of signature genes. Of the remaining groups, signature genes
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only one genome, making it impossible to assess their

harbouring of signature genes. Although phylogroups with
a smaller number of species obviously show larger numbers
of signature genes, the impact of phylogroup sample size is
limited. Even large phylogroups such as the L. delbrueckii
and L. alimentarius groups show a relatively large number
of signature genes. The identification of signature genes thus
conforms to the phylogenetic analyses (Fig. 2) and cAAI
values (Fig. 5), likely reflecting a common evolutionary
history of phylogroups/genera, and is therefore suitable to
provide additional information on the delineation of bacterial
genera. The current signature genes analysis supports most of
the new genera proposed in this work, particularly the separa-
tion of the L. brevis and L. collinoides groups, which were
combined in earlier studies [16, 27], and the separation of the
L. salivarius and L. mali groups, which were also assigned to a
single phylogenetic group in the past [11, 13, 14].
Phylogenomic evaluation of Lactobacillaceae and

Fig. 3. Box plot of the intra-family and inter-family cAAI values in the Leuconostocaceae
families Leuconostocaceae and Lactobacillaceae. The reference line is We further used the phylogenetic trees and cAAI values to
drawn at a cAAI of 59 % in both panels. explore the relationships of Lactobacillaceae and Leuconos-
tocaceae. To identify clades and nodes that are consistently
identified by different approaches, we deliberately used two
can be identified if the most distant member(s) of the group different datasets, type strain genomes and best-quality
is (are) omitted (Fig. 2). The L. buchneri group contains genomes, and different bioinformatics pipelines. The overall
signature genes if L. senioris is omitted and the L. collinoides topology of the phylogenetic trees was remarkably consistent.
group contains signature genes if the L. malefermentans/L. However, the core genome phylogenetic tree generated using
oryzae outgroup is omitted. The L. reuteri, L. salivarius and the type strains of the 26 phylogenetic groups of the Lacto-
L. mali groups do not contain signature genes and can also bacillaceae, 31 type strains of other genera in the Lactobacil-
not be divided into sub-clades that contain signature genes lales and eight type strains from Bacillales as outgroup placed
(Fig. 6). Of note, the L. reuteri group and the L. salivarius the Leuconostocaceae as a monophyletic cluster within the
group were also identified as non-exclusive on the basis of Lactobacillaceae that shares the root with heterofermentative
cAAI values (Fig. 5). The remaining six phylogroups contain lactobacilli, the genus Pediococcus and the L. plantarum group
Fig. 4. Frequency distribution of pairwise cAAI in Lactobacillaceae and Leuconostocaceae. Inter-genus cAAI values are shown in descending
order; Intra-genus cAAI values are shown in ascending order for the current taxonomy (dotted lines) and for the proposed taxonomy
that divides the genus Lactobacillus in 25 genera (solid lines). Genera are colour coded; Lactobacillus; Pediococcus, Leuconostoc and
Weissella. The number of pairwise cAAI values that is represented by the lines is indicated in the figure legend. Owing to the low number
of values, intra-genus cAAI values for Fructobacillus, Oenococcus and Convivina are not shown. The data used for the graph is provided
in Table S3.
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Fig. 5. Range of pairwise intra-genus cAAI values for all current and proposed genera in the families Lactobacillaceae and Leuconostocaceae.
The dotted line designates the lowest cAAI value for current genera excluding Lactobacillus. Bars and symbols are coloured gray if the
genus is exclusive, i.e. if the lowest intra-genus cAAI is higher than any inter-genus cAAI of species in that genus. Bars and symbols
are coloured red and green, respectively, if the genus is non-exclusive, i.e. if the lowest intra-genus cAAI is lower than the highest
inter-genus cAAI of species in that genus. Trivial values (100 % for self to self comparison) are shown only for genera that include one
species. The data used for the graph is provided in Table S3. The y-axis label uses the proposed taxonomy as follows: Lentilactobacillus
– L. buchneri group; Secundilactobacillus – L. collinoides group; Levilactobacillus – L. brevis group; Fructilactobacillus – L. fructivorans
group; Acetilactobacillus, no prior designation; Apilactobacillus – L. kunkeei group; Limosilactobacillus – L. reuteri group; Paucilactobacillus
– L. vaccinostercus group; Furfurilactobacillus – L. rossiae group; Lactiplantibacillus – L. plantarum group; Ligilactobacillus – part of
L. salivarius group; Liquorilactobacillus - part of L. salivarius group; Dellaglioa – L. algidus; Loigolactobacillus – L. coryniformis group;
Paralactobacillus – L. selangorensis; Latilactobacillus – L. sakei group; Lacticaseibacillus – L. casei group; Agrilactobacillus – L. composti;
Schleiferilactobacillus – L. perolens group; Lapidilactobacillus – L. dextrinicus / convacus; Companilactobacillus – L. alimentarius group;
Bombilactobacillus – L. mellifer/mellis group; Lactobacillus – L. delbrueckii group; Amylolactobacillus – L. amylophilus group; Holzapfelia
– L. florum.
(Fig. 7a). The core genome tree based on the genome set used between heterofermentative lactobacilli and the L. delbrueckii
for the signature gene analysis (best-quality genomes) placed group (54–58 %). These findings do not support a taxonomic
the Leuconostocaceae as a monophyletic cluster within the separation of the two families.
heterofermentative lactobacilli, sharing the root with the L.
rossiae, L. reuteri and L. vaccinostercus groups (Fig. 7b and
Fig. S4). This means that the family Lactobacillaceae is not Discussion
monophyletic unless Leuconostocaceae are included. The
Family-level considerations
close relationship of Leuconostocaceae and Lactobacillaceae
is also supported by the cAAI values (Fig. 8, Table S3). Inter- The genus Pediococcus has consistently been recognized as
family cAAI and AAI values between Leuconostocaceae and an integral part of the genus Lactobacillus. Some studies
Lactobacillaceae overlap with the intra-family cAAI values. also suggested that the family Leuconostocaceae, which was
Moreover, the cAAI values between Leuconostocaceae and described in in 2010 on the basis of 16S rRNA sequence
heterofermentative lactobacilli (Table S3) range from 53 similarities [1, 44], are, on the contrary, an integral part of
to 60% and are thus in the same range as the cAAI values the lactobacilli [15, 16]. The phylogenetic and comparative
2788
Fig. 6. Gene family presence/absence patterns in Lactobacillaceae and Leuconostocaceae. Each column represents a gene family
presence/absence pattern in species of Lactobacillaceae and Leuconostoaceae, where presence is indicated with a dot. The absolute
number of gene families that conform to each pattern is visualized in the marginal bar plot at the top. Separations between phylogroups
are indicated with horizontal black lines. We defined genes that were present in all genomes of a clade and in none of the genomes
outside of that clade as ‘signature genes’ (dark blue); other genes are shown in light blue. Only presence/absence patterns followed by
four or more gene families are shown. Patterns of presence in a single species or all species are not shown. Unassigned species are
clusters of closely related genomes which could not be assigned to a known species due to low whole-genome similarity to a type strain
and/or low 16S rRNA similarity to a type strain.
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Fig. 7. (a) Core genome phylogenetic tree with the type strains of 25 phylogenetic groups of the genus Lactobacillus, 31 type strains of
other genera in the Lactobacillales and eight type strains from Bacillales as an outgroup. The phylogenomic analysis is based on the
concatenated protein sequences of 170 single copy core genes. The maximum-likelihood tree was inferred by RAxML as described
previously [5, 12]. Bootstrap support values were calculated from 500 replicates and only values of >80 % are labelled. Members of the
same bacterial families are colour coded. (b) Overview tree of phylogroups of Lactobacillaceae and Leuconostocaceae. Subtree of the tree
shown in Fig. S4; only the branches corresponding to type species of phylogroups of Lactobacillaceae or genera of Leuconostocaceae
are shown.
genomic analyses conducted here confirmed that Leuconos- the most distantly related members of Lactobacillaceae and
tocaceae and Lactobacillaceae are closely related and inter- Leuconostocaceae, respectively, with respect to their cAAI
twined. Moreover, two different datasets and bio-informatic values (50.58 %). The uncertainty with respect to the last
approaches placed the Leuconostocaceae as a monophyletic common ancestor of Leuconostocaceae and Lactobacillaceae
cluster within the Lactobacillaceae. This result is also in line has no implications for their taxonomic assignment into a
with the Genome Taxonomy Database (https://gtdb.ecog- single family. As each node separating homofermentative
enomic.org/; [24]) in which the members of the Leuconosto- from heterofermentative lactic acid bacteria represents a
caceae were considered as members of the Lactobacillaceae. switch of a micro-organism from homofermentation to
heterofermentation, however, it relates to the molecular and
The current analyses also indicate that heterofermentative
ecological evolution of lactic acid bacteria and may thus be
lactobacilli are more closely related to Leuconostoc and
of interest for future studies.
Weissella than they are to the L. delbrueckii group (Fig. 8,
Table S2 and S3). This confirms that the two fermentation Within Lactobacillaceae, the L. delbrueckii group forms a
types, homofermentative and heterofermentative, are closely robust monophyletic clade with the L. floricola group and
linked to the phylogeny of lactobacilli. The genetic related- the L. amylophilus group. Species in this clade also differ from
ness between heterofermentative lactobacilli and Leuconos- other lactobacilli with regards to physiological criteria, e.g.
tocaceae as assessed on the basis of cAAI values is closer than vancomycin sensitivity related to the inter-peptide of the
the relationship between heterofermentative lactobacilli and peptidoglycan, the absence of pyruvate formate lyase, and the
the L. delbrueckii group. Exceptions are Oenococcus species, absence of the pentose–phosphate pathway enabling homo-
which share less than 54.5 % cAAI to any current member fermentative pentose metabolism [14, 45]. The L. mellifer
of the Lactobacillaceae. Despite the overall congruent group and the L. alimentarius group represent the link
topology, the phylogenetic trees generated in this study are between the L. delbrueckii group and other lactobacilli with
inconsistent with respect to the position of Leuconostocaceae respect to phylogeny and phenotypes; Pediococcus and the
within the Lactobacillaceae. Depending on the dataset used L. plantarum group represent the evolutionary link between
for calculation of the tree, the root of Leuconostocaceae is homofermentative and heterofermentative lactic acid
shared with all heterofermentative lactobacilli, Pediococcus bacteria. Although pediococci and the L. plantarum group
and the L. plantarum group (Fig. 7A), or shared with the L. obviously share major metabolic features with homofermen-
reuteri, L. vaccinostercus and the L. rossiae group only (Fig. 8 tative lactobacilli [18], they are phylogenetically related to
and Fig. S4, and Fig. S5) [16]. All of these trees support heterofermentative lactobacilli and Leuconostocaceae (this
the observation that L. iners and Oenococcus species are study).
2790
Fig. 8. Heat map depicting the cAAI values of the 38 782 pairwise comparisons of all species in the families Leuconostocaceae and
Lactobacillaceae for which genome sequence data was available in August 2019. The source file is provided as Table S3.
Criteria for delineation of new genera of nucleotide identities [10]. Threshold values for the deline-
Considerations of the different forces that shape bacterial ation of new bacterial genera on the basis of AAI or cAAI
evolution led to the proposal of a method-free concept in values have not been consistently used [43] but the transition
bacterial taxonomy that considers ecological divergence rather zones for intra-genus and inter-genus AAI and cAAI values
than numerical thresholds [6]. For species level taxonomy, the can be derived by comparison with other bacterial genera,
ANI value between two genomes has been consistently used or alternative taxonomic approaches [27]. A framework for
to delineate new taxa, however, the phylogenetic signal of ANI taxonomy of isolates with unknown ecology including uncul-
values is lost for ANI values below 70 %, i.e. ANI is not a tured organisms was previously established on the basis of
useful metrics for genus level taxonomy. This was particularly the relative evolutionary distance of bacteria as determined
noted for the taxonomy of genus Lactobacillus [14, 16]. The by core genome phylogeny [27]. The Genome Taxonomy
pairwise average amino acid identity (AAI) and the conserved Database classified species of the genus Lactobacillus into 18
proteins (POCP) have been proposed for classification at the phylogenetic clades equivalent to genera which overlap with
genus level [5, 46]; of these two, the AAI is the more powerful the 24 phylogenetic clades that were proposed on the basis
metrics because it is based on the proteins sequences and of ecological and phylogenetic considerations [14, 17]. An
does not only account for the presence or absence of protein. approach focusing on ecology and evolution has also been
Although formal thresholds for genus-level AAI values have applied to lactobacilli [17] and demonstrated a remarkable
not been established, the vast majority bacterial intra-genus overlap of their phylogeny and ecology. A method-free
AAI values is higher than 68 % [5]. Moreover, AAI values can approach, however, relies on information on the ecology of
be calculated on the basis of core proteins to exclude proteins a multiple isolates of a specific bacterial taxon. Currently,
that were acquired by lateral gene transfer (this study), an the ecology and evolution are well-studied for only a small
approach that was also employed for comparison on the basis subset of species in the genus Lactobacillus, while the ecology
2791
of a large number of species and several phylogenetic groups the genus level using an AAI threshold of 68 % to the type
remains unclear because of the lack of data or the low number strain of the most closely related genus [5] (Fig. S3 and S7).
of species in the group [17]. Building on previous studies Values below that level require a more detailed analysis that
on the taxonomy, ecology and phylogeny of Lactobacillus, also includes core genome phylogeny and cAAI values, and
we based the delineation of novel genera on the following may justify the description of novel genera when phylogenetic
criteria, listed in decreasing order of priority: (i) the novel placement and AAI and cAAI values are supported by meta-
genera represent monophyletic phylogenetic groups; (ii) bolic and ecological criteria. We believe that this proposed
intra-genus AAI and cAAI values are higher than inter-genus taxonomy will facilitate further research and discovery. The
AAI and cAAI values, and these show limited overlap; (iii) diversity of lactobacilli – the generic term remains useful to
species in the proposed genera have common characteristics designate organisms in all 25 genera that are currently clas-
with regard to ecology and physiology that differentiate the sified as Lactobacillus species – is likely much greater than
proposed genus from other genera, and this distinction is currently known as a majority of strains were isolated from
generally supported by the presence signature genes [17, 43]; humans, domesticated animals, and food. Sampling of more
and (iv) the proposed genera are largely consistent with the diverse plant or environmental sources, and of wild animals
previously established phylogenetic groups [13, 14, 17] and will likely extend our perspective on the phylogenetic and
with the classification based on relative evolutionary distance metabolic diversity of Lactobacillaceae.
[27].
Classification of the Lactobacillaceae species into 26 genera
comprising an emended genus Lactobacillus (i.e. one confined Species-level considerations
to the L. delbrueckii group), Pediococcus, Paralactobacillus Taxonomic implications of the study
and 23 new genera that correspond to the phylogenetic
Principle 8 of the International Code of Nomenclature of
groups shown in Fig. 2 fit these four criteria best. The
Prokaryotes states that each order or taxon of a lower rank
proposed genera Lacticaseibacillus (the L. casei group) and
with a given circumscription, position and rank can bear
Ligilactobacillus (the L. salivarius group) remain relatively
only one correct name, i.e. the earliest that is in accordance
heterogeneous, particularly with respect to their AAI values,
with the Rules of this Code. Based on the data discussed
indicating that specific subgroups in these genera currently
above, we consider the family Leuconostocaceae Schleifer
adapt to different ecological habitats. This was proposed for
2010, which was circumscribed on the basis of phylogenetic
the L. casei group with respect to oxidative stress tolerance
analyses of 16S rRNA sequences a later synonym of the family
[47] and for the L. salivarius group where megaplasmids and
specific exopolysaccharide clusters may relate to human host Lactobacillaceae Winslow et al. 1917 (Approved Lists 1980).
adaptation [48]. Conversely, the proposed genus Secundilac- Below we propose an emended description of the family
tobacillus (the L. collinoides group) and the proposed genus Lactobacillaceae.
Levilactobacillus (the L. brevis group) are relatively homoge- We further propose that the 23 phylogenetic groups and
neous with respect to their cAAI values, but they are separated single lines of descent (Fig. 2) represent novel genera. The
by phylogenetic analysis, their metabolism, and their ecology. new taxa are described below and an emended description of
The L. collinoides group is differentiated from L. brevis by the genera Lactobacillus and Paralactobacillus are proposed.
adaptation to hexose-depleted habitats, which is reflected by The new genera are described in an order that reflects their
lack of mannitol-dehydratase, diol-dehydratase activity, and a phylogenetic relationships and recognizes the distinct phylo-
metabolic focus on pentoses. These metabolic and ecological genetic position of homofermentative and heterofermentative
differences are also supported by the more divergent AAI lactobacilli (Figs 2 and 5–8).
values and the identification of signature genes.
The genome-based analyses confirmed that the only strain of
Overall, the available evidence is highly consistent in its P. lolii that has been described, P. lolii DSM 19927T, is a strain
support with the proposed taxonomic classification. Moreover, of P. acidilactici, with cAAI and AAI values to the P. acidilac-
16S rRNA gene sequence identities and AAI values readily tici type strain of 99.4 and 98.1 %, respectively [50] (Tables
allow new species to be assigned to one of the 26 genera of S3 and S4). P. lolii is thus a later synonym of P. acidilactici.
Lactobacillaceae. Species that exhibit a 16S rRNA gene identity
that is greater than 94.5 % to the type species of a genus [49] The genome-based analyses also suggested that the two previ-
and cluster integral to that genus when using 16S rRNA gene ously described subspecies of L. aviarius should be elevated
phylogeny are generally assigned correctly even in the absence to species level. Lactobacillus aviarius was isolated from
of further analyses (Fig. S6). However, 16S rRNA gene iden- the intestine of chickens in 1984 and two subspecies were
tity and 16S rRNA phylogeny should be complemented with recognised based on different sugar fermentation profiles:
core genome phylogeny in combination with whole-genome L. aviarius subsp. aviarius does not ferment trehalose and
similarity metrics to validate the taxonomy at the genus level cellobiose (while L. aviarius subsp. araffinosus does) but
[4]. We compared the performance of several whole-genome metabolizes galactose, lactose, melibiose and raffinose (while
metrics, i.e. ANI, CNI, AAI and cAAI. Of these four, ANI L. aviarius subsp. araffinosus does not) [51]. The availability of
and AAI can easily be computed using publicly available the genome sequences of the type strains of both subspecies
tools. Novel species of Lactobacillaceae can be classified on revealed that they are only distantly related, which support
2792
that these two subspecies should be recognised as different Lactobacillaceae, which reflects its strict adaptation to the
species. human vagina.
In addition to their relevance in intestinal and vaginal ecosys-
Emended description of the family Lactobacillaceae tems, Lactobacillus species frequently occur in dairy and
The emended family Lactobacillaceae is circumscribed on the cereal fermentations and are widely used as starter cultures
basis of phylogenomic analyses reported in the present paper, for production of fermented dairy products [58, 59].
and includes all the genera previously included in families
Lactobacillaceae Winslow et al. 1917 (Approved Lists 1980) A phylogenetic tree of all species in the genus Lactobacillus is
and Leuconostocaceae Schleifer 2010, i.e. Convivina, Fructo- provided in Fig. S6A.
bacillus, Lactobacillus, Leuconostoc, Oenococcus, Paralactoba- The type species of the genus Lactobacillus is L. delbrueckii.
cillus, Pediococcus and Weissella. Although the nomenclature of species in the emended genus
Cells are Gram-positive, non-spore-forming facultative or Lactobacillus remains unchanged, a list of species and a list of
strict anaerobic bacteria. Cells are coccoid or rod-shaped, their properties is provided below.
which may form chains, pairs or tetrads (genus Pediococcus).
Main product of the fermentative metabolism is lactate, and Lactobacillus delbrueckii
other products may be acetate, ethanol, CO2, formate or Lactobacillus delbrueckii (Bacillus delbrücki) (del.brueck’i.i.
succinate. Complex nutritional requirements for amino acids, N.L. gen. n. delbrueckii, of Delbrück, named for M. Delbrück,
peptides, nucleic acid derivatives, vitamins, salts, fatty acids or a German bacteriologist).
fatty acid esters, and fermentable carbohydrates. Lactobacil-
laceae are the only family in the Lactobacillales that includes Type species of the genus Lactobacillus. Because of the high
homofermentative and heterofermentative micro-organisms. phenotypic and genotypic similarities between L. delbrueckii,
L. leichmannii, L. lactis and L. bulgaricus, only L. delbrueckii is
Type genus: Lactobacillus Beijerinck 1901 212 (Approved retained as a separate species. Both L. lactis and L. leichmannii
Lists). are treated as L. delbrueckii subsp. lactis and L. bulgaricus as
L. delbrueckii subsp. bulgaricus [60]. All strains produce d(−)-
lactic acid.
Homofermentative Lactobacillaceae
Emended description of Lactobacillus
Lactobacillus delbrueckii subsp. bulgaricus
Lactobacillus species are Gram-positive, homofermentative,
Lactobacillus delbrueckii subsp. bulgaricus (bul.ga’ri.cus. N.L.
thermophilic and non- spore-forming rods. Most Lacto-
masc. adj. bulgaricus, Bulgarian).
bacillus species do not ferment pentoses and none of the
organisms encode genes for the pentose–phosphate pathway Strains of this subspecies were previously referred to as L.
or pyruvate formate lyase. The emended description of the bulgaricus. L. delbrueckii subsp. bulgaricus ferment glucose
genus includes all organisms that were previously assigned to d(−)-lactic acid and also metabolise fructose, mannose
to the L. delbrueckii group [17]. Lactobacillus species are and lactose, but not sucrose [60, 61]. L. delbrueckii subsp.
host-adapted; the Lactobacillus melliventris clade (previously bulgaricus undergoes genome reduction which particularly
termed the Firm-5 clade) is adapted to social bees [52] while relates to loss or silencing of genes coding for metabolism of
all other Lactobacillus species are adapted to vertebrate hosts. carbohydrates [57]. The genome size is 1.76 Mbp and the G+C
Lactobacillus species ferment a relatively broad spectrum of content of DNA is 49.9 mol%.
carbohydrates and have the strain-specific ability to ferment
extracellular fructans, starch, or glycogen [53, 54]. The L. Isolated from yoghurt and cheese and an important starter
melliventris clade species also ferment a wider range of carbo- cultures in the production of these foods [58, 59] but also
hydrates when compared to insect-adapted species in the detected by culture-independent methodology in the intes-
genera Apilactobacillus and Bombilactobacillus. In intestinal tinal microbiota of suckling piglets [54].
habitats, Lactobacillus species are generally associated with The type strain is ATCC 11842T=CCUG 41390T=CIP
heterofermentative lactobacilli. For specific examples, it was 101027T= DSM 20081T=IFO (now NBRC) 13953T= JCM
shown that co-habitation of Lactobacillus species with hetero- 1002T=LMG 6901T= LMG 13551T=NCTC 12712T=VKM
fermentative lactobacilli is based on long-term evolutionary B-1923T.
relationships in biofilms [55] and a complementary prefer-
ence for carbon sources [18, 56]. Many Lactobacillus species Genome sequence accession number: JQAV00000000.
are able to ferment mannitol, which also reflects co-habitation 16S rRNA gene accession number: CR954253.
with heterofermenters. The metabolic focus of L. delbrueckii
on lactose [57] explains its dominance in yoghurt and cheese
fermentations but also relates to its presence in the intestine Lactobacillus delbrueckii subsp. delbrueckii
of suckling piglets [54]. The genus Lactobacillus remains a Lactobacillus delbrueckii ( del.
bruec’ki.i. N.L. gen. n.
relatively heterogenous genus with L. iners as the most distant delbrueckii, of Delbrück, named after M. Delbrück, a German
member. L. iners has the smallest genome size among all bacteriologist).
2793
L. delbrueckii subsp. delbrueckii ferments glucose to d(−)-lactic ferments glucose to d(−)-lactic acid and also metabolises
acid and also metabolises sucrose, fructose and mannose, sucrose, fructose and mannose, maltose and trehalose [60].
but not lactose. Characteristics of the species/subspecies are The genome size is 1.87 Mbp and the mol% G+C content of
provided by [61–65]. The genome size is 1.75 Mbp and the DNA is 49.9 .
mol% G+C content of DNA is 49.9.
Isolated from milk, cheese, compressed yeasts and grain mash.
Isolated from vegetable source, sour grain mash and
The type strain is L 110T=ATCC 12315T=CCUG 31454T=CIP
fermented grains.
101028T=DSM 20072T = JCM 1248T=LMG 7942T=NBRC
The type strain is ATCC 9649T=CCUG 34222T=CIP 102622T=NRRL B-4525T.
57.8T=DSM 20074T=IFO (now NBRC) 3202T=JCM
Genome sequence accession number: AZDE00000000.
1012T=LMG 6412T=NCIMB 8130T (formerly NCDO
213)=NRRL B-763T=VKM B-1596T. 16S rRNA gene accession number: AY050173.
Genome sequence accession number: AZCR00000000.
Lactobacillus delbrueckii subsp. sunkii
16S rRNA gene accession number: AY773949.
Lactobacillus delbrueckii subsp. sunkii (sun’ki.i. N.L. gen.
n. sunkii, of sunki, the name of the non-salted, traditional,
Lactobacillus delbrueckii subsp. indicus Japanese pickle from which the type strain was isolated).
Lactobacillus delbrueckii subsp. indicus (in’di.cus. L. masc. adj.
L. delbrueckii subsp. sunkii ferments glucose to d(−)-lactic
indicus, of India, Indian, referring to the geographical origin
acid and also metabolises sucrose, fructose and mannose,
of the strains).
maltose but not lactose [68]. The genome size is 2.0 Mbp and
Lactobacillus delbrueckii subsp. indicus metabolises sucrose, the mol% G+C content of DNA is 50.1.
fructose, lactose and mannose, but is unable to ferment
Isolated from a traditionally fermented Japanese red turnip.
maltose and trehalose [66]. The genome size is 1.88 Mbp and
the mol% G+C content of DNA is 49.5. The type strain is YIT 11221T=DSM 24966T=JCM 17838T.
Isolated from a fermented dairy product dahi from India. Genome sequence accession number: CP018217.
The type strain is NCC725 =CCUG 51390 =DSM
T T
16S rRNA gene accession number: AB641833.
15996T=JCM 15610T=LMG 22083T.
Genome sequence accession number: AZFL00000000. Lactobacillus acetotolerans
Lactobacillus acetotolerans (a.ce.to.to’le.rans. L. neut. n.
acetum vinegar; L. pres. part. tolerans tolerating, enduring;
N.L. part. adj. acetotolerans vinegar tolerating).
Lactobacillus delbrueckii subsp. jakobsenii
Lactobacillus acetotolerans produces dl-lactic acid from
Lactobacillus delbrueckii subsp. jakobsenii (ja.kob.se’ni.i. N.L.
glucose, mannose, fructose and trehalose but not from
gen. n. jakobsenii, of Jakobsen, named after Mogens Jakobsen
arabinose, sucrose, lactose, melibiose and raffinose [69]. The
for his significant contributions in the field of food microbi-
genome size is 1.59 Mbp and the mol% G+C content of DNA
ology including biodiversity, production and safety of African
is 36.3.
fermented foods and beverages).
The species is part of the core microbiome of mash fermenta-
Lactobacillus delbrueckii subsp. jakobsenii metabolises
tions for production of grain liquor and vinegar in China and
sucrose, maltose and trehalose but is unable to ferment lactose
Japan [58]; it was also isolated from other plant fermentations,
[67]. The genome size is 1.75 Mbp and the mol% G+C content
silage, and from the intestine of swine, ducks, and cattle.
of DNA is 50.3.
The type strain is ATCC 43578T=CCUG 32229T=CIP
Isolated from dolo wort used in the production of the
103180T=DSM 20749T=JCM 3825T=LMG 10751T=NBI 3014T.
fermented African beverge dolo in Burkina Faso.
Genome sequence accession number: AYZC00000000.
The type strain is ZN7a-9T=DSM 26046T=LMG 27067T.
16S rRNA gene accession number: M58801.
Genome sequence accession number: JQCG00000000.
16S rRNA gene accession number: ALPY00000000. Lactobacillus acidophilus
Lactobacillus acidophilus (a.ci.do’phi.lus. N.L. neut. n. acidum
Lactobacillus delbrueckii subsp. lactis acid; Gr. masc. adj. philos loving; N.L. masc. adj. acidophilus
Lactobacillus delbrueckii subsp. lactis (lac’tis. L. gen. n. lactis, acid-loving).
of milk).
Lactobacillus acidophilus produces dl- lactic acid from
L. delbrueckii subsp. lactis includes strains previously desig- cellobiose, galactose, lactose, maltose, mannose, sucrose and
nated as L. lactis and L. leichmannii; L. delbrueckii subsp. lactis trehalose, but not from mannitol [70–72]. Specific strains
2794
have been used as probiotics. The genome size of the type Genome sequence accession number: AZCM00000000.
strain is 1.95 Mbp and the mol% G+C content of DNA is 34.6.
Isolated from the intestinal tract of humans and animals,
human mouth, human vagina, sourdough and wine. Lactobacillus apis
The type strain is ATCC 4356 =CCUG 5917 =CIPT T
Lactobacillus apis (a’pis. L. gen. n. apis of/from a honey bee,
76.13T=DSM 20079T=IFO (now NBRC) 13951T=JCM the genus name of the true honey bee Apis mellifera L., refer-
1132T=LMG 9433T= LMG 13550T=NCTC 12980T=NRRL ring to the insect host of the first strains).
B-4495T=VKM B-1660T.
Lactobacillus apis cells produce l-lactic acid from glucose.
Genome sequence accession number: AZCS00000000. Growth is found under strictly anaerobic and microaerophilic
16S rRNA gene accession number: AY773947. conditions. Produces acid from glucose, fructose, mannose
and salicin, but not from galactose, maltose, raffinose, cello-
Lactobacillus amylolyticus biose, sucrose and lactose [77]. The genome size is 1.70 Mbp
and the mol% G+C content of DNA is 36.9.
Lactobacillus amylolyticus (a.my.lo.ly'ti.cus. Gr. neut. n. amylon
starch; N.L. masc. adj. lyticus (from Gr. masc. adj. lytikos) able Isolated from the stomach contents of honeybees (Apis
to loosen; N.L. masc. adj. amylolyticus starch-digesting). mellifera L.).
Lactobacillus amylolyticus can grow up to 52 °C with an The type strain is R4BT=CCM 8403T=LMG 26964T.
optimum growth temperature between 45 °C and 48 °C. No Genome sequence accession number: PDKP00000000.
growth occurs at 20 °C. These bacteria produce dl-lactic acid
from glucose, fructose, galactose, glucose, maltose, mannose, 16S rRNA gene accession number: KF386017.
sucrose raffinose and melibiose. Neither acid nor gas are
produced from arabinose, cellobiose, lactose, mannitol, Lactobacillus bombicola
rhamnose, ribose and trehalose [73]. The genome size is 1.54
Lactobacillus bombicola [bom.bi’co.la. L. masc. n. bombus a
Mbp and the mol% G+C content of DNA is 38.2.
boom, a deep hollow noise, buzzing, also the zoological genus
Isolated from malt, mash and unhopped wort in breweries but name of the bumble bee; L. suf. –cola (derived from L. masc.
was also identified in sourdough and tofu whey. or fem. n. incola incola, inhabitant) dwelling, occurring in;
N.L. masc. n. bombicola occurring in Bombus].
The type strain is LA 5T=CCUG 39901T=DSM 11664T=JCM
12529T=LMG 18796T. Lactobacillus bombicola produces only d(−)-lactic acid from
Genome sequence accession number: AZEP00000000. glucose. It is capable of utilizing cellobiose, ribose, glucose,
fructose, mannose and trehalose for growth an acid produc-
16S rRNA gene accession number: FR683095. tion, but not arabinose, maltose, lactose, melibiose and
sucrose [42]. The genome size is 1.64 Mbp and the mol% G+C
Lactobacillus amylovorus content of DNA is 34.6.
Lactobacillus amylovorus (a.my.lo.vo’rus. Gr. neut. n. amylon Isolated from the gut of a Bombus lapidarius bumble bee.
starch, L. v. vorare to devour, N.L. masc. adj. amylovorus
starch-devouring). The type strain is H70-3T=LMG 28288T=DSM 28793T.
The species includes strains that were previously designated Genome sequence accession number: NPNG00000000.
as Lactobacillus sobrius [74]. 16S rRNA gene accession number: LK054485.
Lactobacillus amylovorus produces both d(−)- and l(+)-lactic
acid and acid is produced from cellobiose, fructose, galactose, Lactobacillus colini
glucose, maltose, mannose, sucrose, starch and trehalose. Lactobacillus colini (co.li’ni. N.L. gen. n. colini of Colinus,
Arabinose, melezitose, melibiose, raffinose, rhamnose and scientific name of bobwhites).
ribose are not fermented. The ability to ferment starch and
the presence of an extracellular amylolytic enzyme are distin- L. colini ferments a broad range of hexoses and disaccharides
guishing characteristics [75]. Growth maximum is 45 to 48 but pentoses are not fermented. Optimum growth is observed
°C. The genome size of the type strain is 2.02 Mbp and the at 37 °C and between pH 5.0 and 5.5 [78]. The mol% G+C
mol% G+C content of DNA is 37.8. content of DNA is 35.6.
The microorganism is a characteristic representative of swine Isolated from bobwhites, a ground-dwelling bird [78].
intestinal microbiota [54, 76]; also isolated from sourdough
The type strain is 111144 L1T=DSM 101872T=KCTC 21086T.
and cattle waste-corn fermentation.
Genome accession number: not available at time of
The type strain is ATCC 33620T=CCUG 27201T=CIP
publication.
102989T=DSM 20531T=JCM 1126T=LMG 9496T=NCAIM
B.01458T=NRRL B-4540T. 16S rRNA gene accession number: KU161105.
2795
Lactobacillus crispatus Lactobacillus gallinarum

Lactobacillus crispatus (cris.pa’tus. L. part. adj. crispatus Lactobacillus gallinarum (gal.li.na'rum. L. gen. pl. n. galli-
curled, crisped, referring to morphology observed original narum of hens).
in broth media).
Lactobacillus gallinarum produces dl-lactic acid. Acid is
Lactobacillus crispatus is synonymous with Lactobacillus produced from glucose, mannose, galactose, sucrose, fruc-
acidophilus group A2 of Johnson et al. [72]. It produces tose, cellobiose, melibiose and raffinose. No acid formation
dl-lactic acid from glucose, cellobiose, fructose, galactose, from arabinose, rhamnose, ribose, trehalose and mannitol.
lactose, maltose, mannose and sucrose [79–81]. Arabinose, Synonymous with Lactobacillus acidophilus group A4 of
mannitol, melezitose or rhamnose are not fermented. Grows Johnson et al. [72]. Strains are tolerant to 4.0 % NaCl [72, 84].
equally well at 37 and 45 °C. The genome size is 2.06 Mbp and The genome size of the type strain is 1.94 Mbp and the mol%
the mol% G+C content of DNA is 36.6. G+C content of DNA is 36.5.
One of the predominant species found in the human female Isolated from the chicken intestine.
lower genital tract. Isolated from human faeces, vagina and
The type strain is L917, ScavT=ATCC 33199T=CCUG
buccal cavities, from crops and caeca of chicken and from
30724T=CIP 103611T=DSM 10532T= JCM 2011T= LMG
patients with purulent pleurisy, leucorrhea and urinary tract 9435T=VPI 1294T.
infections. Also isolated from type II sourdoughs.
Genome sequence accession number: AZEL00000000.
The type strain is VPI 3199T=ATCC 33820T=CCUG
30722T=CIP 102990T=CIPP IIT=DSM 20584T= JCM 16S rRNA gene accession number: AJ417737.
1185T=LMG 9479T.
Genome sequence accession number: AZCW00000000. Lactobacillus gasseri
Lactobacillus gasseri (gas’se.ri. N.L. gen. n. gasseri, of Gasser,
16S rRNA gene accession number: AF257097. named for F. Gasser, a French bacteriologist).
Lactobacillus gasseri strains are all positive for glucose,
Lactobacillus equicursoris
mannose, galactose, maltose, sucrose and fructose fermenta-
Lactobacillus equicursoris (e.qui.cur’so.ris. L. masc. n. equus tion and produce dl-lactic acid. Strains do not ferment ribose,
a horse; L. gen. n. cursoris of a runner or racer; N.L. gen. n. mannitol and raffinose. Further characteristics are provided
equicursoris of a racing horse, referring to the isolation of the by [84–86]. The genome size is 1.89 Mbp and the mol% G+C
type strain from faeces of a thoroughbred racehorse). content of DNA is 35.3.
L. equicursoris produces d(−)-lactic acid from glucose. It One of the predominant species in the human female lower
utilises glucose, fructose, galactose, mannose, cellobiose, genital tract. Also isolated from the human mouth, intes-
lactose and mannitol for growth and acid production [82]. tinal tract, and from the intestine of animals. Also found in
The genome size is 2.05 Mbp and the mol% G+C content of wounds, urine, blood, carious dentine, and pus of patients
DNA is 47.7. suffering from septic infections.
Isolated from a thoroughbred racehorse. The type strain is 63 AMT of Gasser=ATCC 33323T=CCUG
The type strain is DI70T=DSM 19284T=JCM 14600T. 31451T=CIP 102991T=DSM 20243T=JCM 1131T=LMG
9203T=NRRL B-14168T=NRRL B-4240T.
Genome sequence accession number: AZDU00000000.
Genome sequence accession number: NC_008530.
16S rRNA gene accession number: AF519171.
Lactobacillus fornicalis
Lactobacillus gigeriorum
Lactobacillus fornicalis (for.ni.ca ́lis. N.L. masc. adj. fornicalis,
Lactobacillus gigeriorum (gi.ge.ri.o‘rum. L. gen. pl. n. gigeri-
of the posterior fornix).
orum of or from the entrails or gizzards of poultry).
L. fornicalis produces dl-lactic acid from glucose. Acid is
Lactobacillus gigeriorum produces mainly d-lactic acid from
produced from ribose, fructose, galactose, glucose, mannose,
glucose, fructose, mannose, maltose, sucrose, galactose and
maltose, cellobiose, sucrose, trehalose, melezitose, mannitol
cellobiose. Ribose, mannitol, lactose and raffinose are not
and sorbitol [83]. The mol% G+C content of DNA is 37.
fermented [87].
Isolated from the human posterior vaginal fornix.
The genome size is 1.91 Mbp and the mol% G+C content of
The type strain is TV 1018T=DSM 13171T=ATCC 700934T. DNA is 36.9.
Genome sequence accession number: not available at the time Isolated from a crop of a chicken.
of publication.
The type strain is 202T=LEM 202T=CRBIP 24.85T=DSM
16S rRNA gene accession number: Y18654. 23908T.
2796
Genome sequence accession number: AYZO00000000. 15019T=JCM 1120T=LMG 6413T=LMG 13555T=NRRL

B-4526T.
16S rRNA gene accession number: FR681899.
Genome sequence accession number: AZEK00000000.
Lactobacillus hamsteri 16S rRNA gene accession number: AM113779.
Lactobacillus hamsteri (ham’ste.ri. N.L. gen. n. hamster of the
hamster from which the isolate was derived). Lactobacillus hominis
Lactobacillus hamsteri produces dl-lactic acid from glucose, Lactobacillus hominis (ho’mi.nis. L. gen. n. hominis of a
cellobiose, sucrose, mannitol, raffinose and ribose. No acid is human being).
produced from melizitose [88]. The genome size is 1.84 Mbp
Lactobacillus hominis produces acid from galactose, glucose,
and the mol% G+C content of DNA is 35.1.
fructose, mannose, cellobiose, maltose, lactose, sucrose,
Isolated from the intestine of a hamster. trehalose and raffinose. Mainly d-lactic acid (about 90 %) is
The type strain is Ha5F1T=ATCC 43851T= CCUG 30726T=CIP produced from glucose [93]. The genome size is 1.93 Mbp and
103365T=DSM 5661T=JCM 6256T=LMG 10754T. the mol% G+C content of DNA is 35.2.
Genome sequence accession number: AZGI00000000. Isolated from the human intestine.
16S rRNA gene accession number: AJ306298. The type strain is 61DT=CRBIP 24.179T=DSM 23910T.
Genome sequence accession number: AYZP00000000.
Lactobacillus helsingborgensis 16S rRNA gene accession number: FR681902.
Lactobacillus helsingborgensis (hel.sing.bor.gen’sis. N.L. masc.
adj. helsingborgensis pertaining to Helsingborg, the site of Lactobacillus iners
Lund University, Campus Helsingborg, Sweden, where the
type strain was characterized). Lactobacillus iners (in’ers. L. masc. adj. iners, inactive, lazy).
Lactobacillus helsingborgensis growth occurs at 15–50 °C Lactobacillus iners grows on blood agar but not on MRS
and d(−)-lactic acid is produced as the end product from or Rogosa agar, reflecting a strict adaptation to the human
hexose fermentation. Acid is produced during fermentation vagina. l(+)-Lactic acid is produced from glucose, some
of glucose, mannose, sorbose, sorbitol and sucrose, but not strains produce acid from maltose but acid is not produced
from maltose or lactose [89]. The genome size is 2.02 Mbp from lactose, mannitol, raffinose, ribose and sucrose [94]. The
and the mol% G+C content of DNA is 36.4. genome size is 1.27 Mbp and the mol% G+C content of DNA
is 32.5.
Isolated from the honey stomach of the honeybee A. mellifera
mellifera, and from alfalfa silage. One of the predominant species found in the human female
lower genital tract; also occurs on human skin [95]
The type strain is Bma5NT=DSM 26265T=CCUG 63301T.
The type strain is CCUG 28746T=CIP 105923T=DSM
Genome sequence accession number: JXJR00000000. 13335T=JCM 12513T=LMG 18914T.
16S rRNA gene accession number: JX099553. Genome sequence accession number: AZET00000000.
16S rRNA gene accession number: Y16329.
Lactobacillus helveticus
Lactobacillus helveticus [hel.ve’ti.cus. L. masc. adj. helveticus
Swiss, referring to the isolation of the type strain from Lactobacillus intestinalis
Emmental (Swiss) cheese]. Lactobacillus intestinalis (in.tes.tina’lis. N.L. masc. adj. intes-
tinalis, pertaining to the intestine).
Lactobacillus helveticus produces dl-lactic acid from glucose
galactose, lactose, mannose and trehalose but not from cello- Lactobacillus intestinalis produces dl-lactic acid from glucose,
biose, mannitol, raffinose and sucrose. L. helveticus [2, 90] is mannose, fructose, galactose, sucrose and mannitol, but not
an earlier heterotypic synonym of Lactobacillus suntoryeus from arabinose, xylose, rhamnose and trehalose [96]. The
[91]. The genome size of the type strain is 1.83 Mbp and the genome size is 2.01 Mbp and the mol% G+C content of DNA
mol% G+C content of DNA is 36.8. is 35.4.
Part of the core microbiome of chicken [92] but it was also Isolated from the intestines of rats, mice and pigs.
isolated from sour milk, cheese starter cultures and cheese,
The type strain is Th4T=ATCC 49335T=CCUG 30727T=CIP
particularly Emmental and Gruyère cheeses, and in tomato
104793T=DSM 6629T=JCM 7548T=LMG 14196T.
pomace and silage.
Genome sequence accession number: AZGN00000000.
The type strain is 12, Lh12T=ATCC 15009T=CCUG
30139T=CIP 103146T=DSM 20075T=IFO (now NBRC) 16S rRNA gene accession number: AJ306299.
2797
Lactobacillus jensenii Lactobacillus kefiranofaciens

Lactobacillus jensenii (jen.se’ni.i. N.L. gen. n. jensenii, of Lactobacillus kefiranofaciens (ke.fi.ra.no.fa'ci.ens. N.L. neut. n.
Jensen, named for S. Orla-Jensen, a Danish microbiologist). kefiranum, a polysaccharide of kefir grain, kefiran; L. v. facio,
produce; N.L. part. adj. kefiranofaciens, kefiran producing).
Lactobacillus jensenii produces only the d(−)-lactic enanti-
omer. Cellobiose, sucrose and maltose fermented; lactose, Lactobacillus kefiranofaciens produces dl-lactic acid with an
mannitol and arabinose not fermented [97]. The genome excess of d(−)-lactic acid from glucose, fructose, galactose,
size is 1.61 Mbp and the mol% G+C content of DNA is sucrose, maltose, lactose and raffinose, but not from arab-
34.33. inose, ribose, cellobiose or trehalose [99].
One of the predominant species in the human female lower Two subspecies are recognised.
genital tract.
Lactobacillus kefiranofaciens subsp. kefiranofaciens
The type strain is 62GT=ATCC 25258T=CCUG 21961T=CCUG
Lactobacillus kefiranofaciens subsp. kefiranofaciens (ke.fi.
35572T=CIP 69.17T=DSM 20557T=JCM 1146T=JCM
rano. fa'ci.ens. L. n. kefiran, a polysaccharide of kefir grain,
15953T=LMG 6414T=NRRL B-4550T.
kefiran; L. v. facio, produce; N.L. part. adj. kefiranofaciens,
Genome sequence accession number: AYYU00000000. kefiran producing).
16S rRNA gene accession number: AF243176. The genome size of the type strain is 2.26 Mbp and the mol%
G+C content of DNA is 37.2.
Lactobacillus johnsonii Part of the microbiota of kefir grains [99, 100] but also isolated
Lactobacillus johnsonii (john.so'ni.i. N.L. gen. n. johnsonii of from other fermented dairy products.
Johnson; named for J. L. Johnson, an American microbiolo- The type strain is WT-2BT=ATCC 43761T=CCUG 32248T=CIP
gist). Synonymous with L. acidophilus group B2 of Johnson 103307T=DSM 5016T=JCM 6985T=LMG 19149T.
et al. [72].
Genome sequence accession number: AZGG00000000.
Lactobacillus johnsonii strains all ferment glucose, mannose,
galactose, maltose, sucrose and fructose and produce dl- 16S rRNA gene accession number: AM113781.
lactic acid. L. johnsonii does not ferment mannitol or ribose
[84]. The genome size is 1.77 Mbp and the mol% G+C content Lactobacillus kefiranofaciens subsp. kefirgranum
of DNA is 34.4. Lactobacillus kefiranofaciens subsp. kefirgranum (ke.fir.
gra’num. Turkish n. kefir, Caucasian sour milk; L. n. granum
Isolated from humans (gut, vagina) and the faeces of birds, grain; N.L. neut. n. kefirgranum kefir grain).
rodents, calves and pigs, and from type II sourdoughs.
The genome size of the type strain is 2.10 Mbp and the mol%
The type strain is ATCC 33200T=CCUG 30725T=CIP G+C content of DNA is 37.5.
103620T=DSM 10533T=JCM 2012T=VPI 7960T.
Part of the core microbiome of kefir grains [100, 101].
Genome sequence accession number: AZCY00000000.
The type strain is GCL 1701T=ATCC 51647T=CCUG
16S rRNA gene accession number: AJ002515. 39467T=CIP 104241T=DSM 10550T=JCM 8572T=LMG
15132T.
Lactobacillus kalixensis Genome sequence accession number: AZEM00000000.
Lactobacillus kalixensis (ka.lix.en‘sis. N.L. masc. adj. kalixensis
pertaining to Kalix, a town in northern Sweden, where the 16S rRNA gene accession number: AM113782.
gastric biopsies were sampled).
Lactobacillus kimbladii
Lactobacillus kalixensis produces both d(−)- and l(+)-lactate Lactobacillus kimbladii (kim.bla‘di.i. N.L. gen. n. kimbladii
from galactose, glucose, fructose, mannose, cellobiose, named after beekeeper Tage Kimblad, for his significant
maltose, lactose, sucrose, trehalose and raffinose. Acid is not contributions to the discovery of the LAB microbiota in the
produced arabinose, ribose, xylose, rhamnose and mannitol honey stomach of honeybees).
[98]. The genome size is 2.08 Mbp and the mol% G+C content
of DNA is 36.1. Lactobacillus kimbladii grows at 15 to 50°C and d(−)-lactate
from glucose, fructose, mannose and tagatose, but not from
Isolated from a biopsy of the healthy human gastric mucosa. ribose, sorbitol, cellobiose and lactose [89]. The genome size
The type strain is Kx127A2T=CCUG 48459T=DSM is 2.19 Mbp and the mol% G+C content of DNA is 36.0.
16043T=JCM 15954T=LMG 22115T. Isolated from the honey stomach of the honeybee A. mellifera.
Genome sequence accession number: AZFM00000000. The type strain is Hma2NT=DSM 26263T=CCUG 63633T.
16S rRNA gene accession number: AY253657. Genome sequence accession number: JXLH00000000.
2798
16S rRNA gene accession number: JX099549. Lactobacillus mulieris

Lactobacillus mulieris (mu.li’.e.ris. L. gen. n. mulieris of a
Lactobacillus kitasatonis woman, from where the bacterium was isolated).
Lactobacillus kitasatonis (ki.ta.sa.to‘nis. L. gen. n. kitasatonis Lactobacillus mulieris grows in the pH range of 5.0–8.5 and
referring to Shibasaburo Kitasato, the founder of Kitasato from 30 to 45 °C; optimum growth is observed at 37 °C [103].
Institute, the father of Japanese bacteriology). Lactic acid is produced from glucose, fructose, mannose,
Lactobacillus kitasatonis produces dl- lactic acid from N-acetylglucosamine, several disaccharides and starch but
mannose, maltose, galactose, sucrose and fructose while not from pentoses [103]. The genome size is 1.66 Mbp and
acid is not produced from arabinose, xylose, ribose, trehalose, the mol% G+C content of DNA is 34.2.
melibiose and raffinose [102]. The genome size of the type
Isolated from the urine of a woman; several strains previ-
strain is 1.91 Mbp and the mol% G+C content of DNA is 37.5.
ously classified as L. jensenii should be classified as L. mulieris
Isolated from the intestine of animals including chicken and [103].
swine.
The type strain is c10Ua161MT=CECT 9755T=DSM 108704T.
The type strain is T. Mitsuoka Ch-J-2–1T=JCM 1039T=KCTC
3155T. Genome sequence accession number: SDGL00000000.
Genome sequence accession number: AZFU00000000. 16S rRNA gene accession number: MK775269.

Lactobacillus panisapium
Lactobacillus panisapium (pa.nis.a’pi.um. L. masc. n. panis
Lactobacillus kullabergensis
bread; L. fem. n apis bee; N.L. gen. n. panisapium of bee
Lactobacillus kullabergensis (kul.la.ber.gen‘sis. N.L. masc. adj. bread).
kullabergensis of or belonging to the nature reserve Kullaberg,
where the discovery of these strains was made in 2005). Lactobacillus panisapium is facultatively anaerobic but
optimal growth is observed under anaerobic conditions.
Lactobacillus kullabergensis grows at 15–50 °C; d(−)-lactate is Growth occurs from 15 up to 55 °C (optimum, 37 °C) and it
produced as the end product from hexose fermentation. Acid produces acid from cellobiose, fructose, galactose, glucose,
is produced from glucose, fructose, trehalose and gentiobiose, melibiose, ribose, sucrose, mannose and raffinose [104]. The
but not from ribose, galactose, mannose, maltose, lactose and
genome size is 1.83 Mbp and the mol% G+C content of DNA
sucrose [89]. The genome size is 2.12 Mbp and the mol% G+C
is 37.4.
content of DNA is 35.8.
Isolated from bee bread of Apis cerana.
Isolated from the honey stomach of the honeybee A. mellifera
mellifera. The type strain is Bb 2–3T=DSM 102188T=ACCC 19955T.
The type strain is Biut2NT=DSM 26262T=CCUG 63631T. Genome sequence accession number: NPNH01000000.
Genome sequence accession number: JXBY00000000. 16S rRNA gene accession number: KX447147.
16S rRNA gene accession number: JX099550.
Lactobacillus paragasseri
Lactobacillus melliventris Lactobacillus paragasseri (pa.ra.gas’se.ri. Gr. prep. para,
Lactobacillus melliventris (mel.li.ven‘tris. L. neut. n. mel, mellis resembling; N.L. gen. n. gasseri, a species epithet; N.L. gen.
honey; L. masc. n. venter, -tris belly, stomach; N.L. gen. n. n. paragasseri, resembling Lactobacillus gasseri).
melliventris of the honey stomach, referring to the isolation Lactobacillus paragasseri produces both d(−)- and l(+)-lactic
of the first strains from the honey stomach of honeybees). acid as end products of glucose fermentation (75:25 ratio).
Lactobacillus melliventris grows from 15 to 50 °C. Produce Also ferments maltose, lactose, sucrose and trehalose, but not
acid from glucose, fructose, mannose and gentiobiose, but ribose and raffinose [105]. The genome size is 1.94 Mbp and
not from galactose, ribose, sucrose, maltose and lactose [89]. the mol% G+C content of DNA is 34.9.
The genome size is 2.12 Mbp and the mol% G+C content of Occurs in the human gastrointestinal tract.
DNA is 35.9.
The type strain is M. Rogosa 208XRT=JCM 5343T = ATCC
Isolated from the homey stomach of honeybees. 4963T=LMG 11478T=NCFB 1375T=KCTC 3172T=NCIMB
The type strain is Hma8NT=DSM 26256T=CCUG 63629T. 8931T=VPI 0334T.
Genome sequence accession number: JXLI00000000. Genome sequence accession number: AP018549.
16S rRNA gene accession number: JX099551. 16S rRNA gene accession number: LC374363.
2799
Lactobacillus pasteurii The type strain is MYMRS/TLU1T = DSM 24759T=CCM

Lactobacillus pasteurii (pas.teu‘ri.i. N.L. gen. masc. n. pasteurii 7945T.
of Pasteur). Genome sequence accession number: BFBY00000000.
Lactobacillus pasteurii produces mainly d(−)-lactic acid from 16S rRNA gene accession number: HQ851022.
glucose; acid is also produced from ribose, fructose, mannose,
cellobiose, maltose, lactose and sucrose [93]. The genome size
Lactobacillus taiwanensis
is 1.87 Mbp and the mol% G+C content of DNA is 38.5.
Lactobacillus taiwanensis (
tai.
wa.
nen’sis. N.L. masc. adj.
Isolated from the human intestine. taiwanensis of Taiwan, referring to the origin of the type
The type strain is 1517T=CRBIP 24.76T=DSM 23907T. strain).
Genome sequence accession number: AYZN00000000. L. taiwanensis produces both d- and l-lactic acid enantiomers
from glucose fermentation. Fructose, mannose, maltose,
16S rRNA gene accession number: FR681901. lactose, sucrose and trehalose are also fermented. No acid is
produced from raffinose or ribose [109]. The genome size is
Lactobacillus porci 1.88 Mbp and the mol% G+C content of DNA is 34.0.
Lactobacillus porci (por’ci. L. gen. n. porci of a pig, referring to Occurs in the mouse gastrointestinal tract [55] but also
the isolation of the type strain from small intestine of a pig). isolated from silage cattle feed.
Lactobacillus porci grows between 15 and 50 °C and produces The type strain is FIRDI 006T=BCRC 17755T=DSM 21401T.
d(−)-lactic acid from glucose; acid is also produced from
mannose, cellobiose, maltose, lactose, trehalose, raffinose Genome sequence accession number: AYZG00000000.
and sucrose [106]. The mol% G+C content of DNA is 51.5. 16S rRNA gene accession number: EU487512.
Isolated from the swine intestine.
The type strain is SG816T=KCTC 21090T=NBRC 112917T. Lactobacillus ultunensis
Lactobacillus ultunensis (ul.tun.en’sis. N.L. masc. adj. ultun-
Genome sequence accession number: not available at the time ensis pertaining to Ultuna, the site of Swedish University of
of publication. Agricultural Sciences in Uppsala, Sweden).
16S rRNA gene accession number: MF346092 Lactobacillus ultunensis produces both d- and l-lactate from
glucose. Acid is produced from galactose, fructose, mannose,
Lactobacillus psittaci cellobiose, maltose, lactose, sucrose, and trehalose. Acid is not
Lactobacillus psittaci (psit.ta‘ci. L. gen. masc. n. psittaci of the produced from arabinose, ribose, rhamnose and melezitose.
parrot, from which the organism was first isolated). Growth occurs at 42 but not at 45 °C [98]. The genome size
of the type strain is 2.16 Mbp and the mol% G+C content of
Lactobacillus psittaci produces acid from glucose, raffinose DNA is 36.0.
and sucrose, but not from lactose, mannitol and trehalose
[107]. The genome size is 1.54 Mbp and the mol% G+C Isolated from a biopsy of a healthy human gastric mucosa.
content of DNA is 35.6. The type strain is Kx146C1T = CCUG 48460T=DSM
Isolated from a hyacinth macaw. 16047T=JCM 16177T=LMG 22117T.
The type strain is B 1491/99T = CCUG 42378T=CIP Genome sequence accession number: AZFO00000000.
106492T=DSM 15354T=JCM 11552T. 16S rRNA gene accession number: AY253660.
Genome sequence accession number: AZFB00000000.
16S rRNA gene accession number: AJ272391. Lactobacillus xujianguonis
Lactobacillus xujianguonis (xu.jian.guo'nis. N.L. gen. masc. n.
Lactobacillus rodentium xujianguonis of Xu Jianguo, recognized for contributions to
medical microbiology).
Lactobacillus rodentium (ro.den‘ti.um. L. gen. pl. n. rodentium
of gnawers, referring to the fact that the micro-organism was Lactobacillus xujianguonis grows optimally at 37–40°C and at
isolated from wild rodents). neutral pH [110]. dl-Lactic acid is produced from pentoses,
hexoses, disaccharides, sorbitol and mannitol. The genome
Lactobacillus rodentium utilizes glucose, fructose, maltose,
size of the type strain is 2.22 Mbp and the mol% G+C content
melibiose and sucrose, but not ribose, mannitol or trehalose
of DNA is 38.8.
[108]. The genome size is 1.52 Mbp and the mol% G+C
content of DNA is 34.0. Isolated from a Himalayan marmot (Marmota himalayana).
Isolated from the colon of wild mice (Mus musculus) and The type strain is HT111-2T=CGMCC 1.13855T=KCTC
other rodent species [55, 108]. 15803T.
2800
Genome sequence accession number: RXIA00000000. The classification of A. amylotrophicus as separate species
was disputed as the first published genome sequence of the
16S rRNA gene accession number: MK294239.
type strain of A. amylotrophicus was virtually identical to
the sequence of A. amylophilus [14, 16]. The 16S rRNA,
Description of Amylolactobacillus pheS and rpoA sequences provided with the new descrip-
tion, however, clearly differentiate A. amylotrophicus from
gen. nov.
A. amylophilus.
Amylolactobacillus (A.my.lo.lac.to.ba.cil’lus Gr. neut. n.
amylon starch; N.L. masc. n. Lactobacillus a bacterial genus; Phenotypic properties that differentiate A. amylotrophicus
N.L. masc. n. Amylolactobacillus, a lactobacillus that ferments from A. amylophilus include the production of acid from
starch). cellobiose and salicin [112]. The mol% G+C content of DNA
is 43.5.
Strains of Amylolactobacillus are thin rods, (0.5–0.9 µm wide
and 1.2–3 µm long) occurring singly and in short chains, non- Isolated from corn silage.
motile, Gram-positive, catalase- negative, oxidase-negative, The type strain is LMG 11400T=DSM 20534T=JCM
non-spore-forming. Amylolactobacillus species are homo-
1124T=NRRL B-4436T.
fermentative and display extracellular amylolytic enzyme
activity. The mol% G+C content is between 44 and 46. Genome sequence accession number: not available at the time
of publication.
The type species is Amylolactobacillus amylophilus; Amylolac-
tobacillus was previously referred to as L. amylophilus group. 16S rRNA gene accession number: AM236149.
Description of Amylolactobacillus Description of Holzapfelia gen. nov.

amylophilus comb. nov. Holzapfelia (Holz.ap.fe’li.a. N.L. fem. n. Holzapfelia, named
Amylolactobacillus amylophilus (a.my.lo’phi.lus. Gr. neut. n. after Wilhelm Holzapfel, a microbiologist and taxonomist
amylon starch; Gr. masc. adj. philos loving; N.L. masc. adj. who made significant contributions to the understanding of
amylophilus starch-loving). the physiology and applications of lactic acid bacteria).
Basonym: Lactobacillus amylophilus Nakamura and Crowell Strains of Holzapfelia are Gram positive, rod- shaped,
1981, 216VP (Effective publication: Nakamura and Crowell catalase negative, homofermentative and aerotolerant. The
1979, 539). type strain of the single species in the genus was isolated
A. amylophilus strains ferment starch to l(+)-lactic acid, they from mountain flowers, other strains were identified in bee
also metabolise fructose, galactose, glucose, mannose and larvae. H. floricola grows at 15 but not at 37 °C; glucose and
maltose [111]. The genome size is 1.56 Mbp. The mol% G+C fructose are the only carbohydrates that are fermented. The
content of DNA is 43.6 [111]. sources of isolation as well as genomic and physiological
properties of the organism indicate an insect- and flower-
Isolated from swine waste-corn fermentation, corn-starch associated lifestyle of the genus.
processing industrial wastes and kocho (Ensete ventricosum)
bread. The type species of Holzapfelia is H. floricola.
The type strain is ATCC 49845T=LMG 6900T=DSM
20533T=CCUG 30137T=CIP 102988T=IFO (now
NBRC) 15881 =JCM 1125 =NCAIM B.01457T=NRRL
T T
Description of Holzapfelia floricola
B-4437T=NRRL B-4476T. comb. nov.
Genome sequence accession number: AYYS00000000. Holzapfelia floricola (flo.ri’co.la. L. fem. n. flos flower, blossom;
16S rRNA gene accession number: M58806. L. suff. –cola from L. masc. or fem. n. incola dweller; N.L. fem.
n. floricola flower-dweller).
Basonym: Lactobacillus floricola Kawasaki et al. 2011, 1358VP.
Description of Amylolactobacillus
amylotrophicus comb. nov. The description of the species is identical of that of the genus
. The genome size is 1.29 Mbp, the mol% G+C content is 34.5.
Amylolactobacillus amylotrophicus (a.my.lo.tro’phi.cus.
Gr. neut. n. amylon starch; N.L. masc. adj. trophicus from Isolated from flowers.
Gr. masc. adj. trophikos nursing, tending; N.L. masc. adj. The type strain is Ryu1-2T=DSM 23037T=JCM 16512T=NRIC
amylotrophicus growing on starch, pertaining to the ability 0774T.
to ferment starch).
Genome sequence accession number: AYZL00000000.
Basonym: Lactobacillus amylotrophicus Naser et al. 2006,
2526VP. 16S rRNA gene accession number: AB523780.
2801
Description of Bombilactobacillus Isolated from the digestive tract of laboratory-

reared
gen. nov. bumblebee queens (Bombus terrestris).
Bombilactobacillus (Bom.bi.lac.to.ba.cil’lus. L. masc. n. The type strain is BTLCH M1/2T=DSM 26517T=CCM 8440T.
bombus, buzzing of bees and bumblebees, N.L. masc. n. Lacto- Genome sequence accession number: CP031513 (strain
bacillus a bacterial genus; N.L. masc. n. Bombilactobacillus, a BI-2.5; the genome sequence for the type strain is not
lactobacillus from bees and bumble bees). available).
Bombilactobacillus species have been isolated from the 16 rRNA gene sequence accession number: KJ078643.
stomach and the hindgut of honey bees (Apis mellifera) and
bumble bees where they are associated with the heterofer-
mentative sister genus Apilactobacillus. Bombilactobacillus
Description of Bombilactobacillus
species have genome sizes which range from 1.81 to 1.84 Mbp
and a mol% G+C content ranging from 34.7 to 39.5. They mellis comb. nov
are homofermentative and thermophilic; comparable to other Bombilactobacillus mellis (mel’lis. L. gen. n. mellis, of honey,
insect-associated lactobacilli, bombilactobacilli ferment only referring to the isolation from the digestive tract of A.
a narrow spectrum of carbohydrates. mellifera)
A phylogenetic tree on the basis of 16S rRNA genes of all Basonym: Lactobacillus mellis Olofsson et al. 2014, 3115VP
species in the genus Bombilactobacillus is provided in Fig. The characteristics as provided by [89] are comparable to B.
S6B. mellifer. The genome size is 1.81 Mbp. The mol% G+C content
The type species is Bombilactobacillus mellifer comb. nov; of DNA is 36.4.
Bombilactobacillus was previously referred to as L. mellifer Isolated from the honey stomach of the honeybee Apis
group. mellifera.
The type strain is Hon2NT=DSM 26255T=CCUG 63289T.
Description of Bombilactobacillus
Genome sequence accession number: JXBZ00000000.
mellifer comb. nov.
Bombilactobacillus mellifer (mel’li.fer. L. masc. adj. mellifer, 16 rRNA gene sequence accession number: JX099545.
honey-producing, referring to the origin from the stomach
and digestive tract of honey bees Apis mellifera).
Description of Companilactobacillus
Basonym: Lactobacillus mellifer Olofsson et al. 2014, 3113VP
gen. nov.
Description provided by [89]. B. mellifer grows over a wide Companilactobacillus (Com.pani.lac.to.ba.cil’lus. M. L.
temperature range (15–50 °C) and pH values (pH 3–12) but n. companio friend, partner; N.L. masc. n. Lactobacillus
ferments only few hexoses, sucrose and raffinose [89]. The a bacterial genus; N.L. masc. n.; Companilactobacillus,
genome size is 1.82 Mbp. The mol% G+C content of DNA Companion-lactobacillus, the name refers to the associa-
is 39.5. tion of Companilactobacillus species with other lactobacilli,
Isolated from isolated from the honey stomach of the particularly heterofermentative organisms, in cereal and
honeybee Apis mellifera. vegetable fermentations [114]).
The type strain is Bin4NT=DSM 26254T=CCUG 63291T. Companilactobacillus species are Gram-positive, homofer-
mentative and non-spore-forming rods. Strains of Compa-
Genome sequence accession number: JXJQ00000000. nilactobacillus were isolated from fermented vegetables,
16 rRNA gene sequence accession number: JX099543. particularly fermented mustard or onion greens, and fruits
(17 type strains; five of these from mustard or onion fermen-
tations), sourdough or related cereal fermentations (7 type
Description of Bombilactobacillus strains), sausages or meat (seven type strains) or other plant
bombi comb. nov. sources; one isolates was obtained from a fermented dairy
product. Companilactobacillus appears not to be dominant
Bombilactobacillus bombi (bom’bi. L. masc. n. bombus,
in any of these fermentations but occurs consistently (type I
buzzing of bees and Bombus, N.L. gen. n. bombi of Bombus,
sourdoughs and fermented mustard or onion greens) or occa-
of a bumblebee).
sionally (meat, salami). Companilactobacillus metriopterae
Basonym: Lactobacillus bombi Killer et al. 2014, 2615VP was isolated from a grasshopper and forms and outgroup that
differs with respect to genome size (C. metriopterae 1.50 Mbp;
Description provided by [113]; compared to other bombilac-
other species range from 2.17 to 2.9 Mbp); mol% G+C content
tobacilli, B. bombi ferments a wider range of carbohydrates.
of DNA (C. metriopterae 32.8; other species range from 34.1 to
The genome size is 1.84 Mbp. The mol% G+C content of DNA 38.6) and physiological properties. The temperature require-
is 34.7. ments for growth are inconsistent; all strains grow between
2802
25 and 30 °C, most but not all species grow at 15 °C and Description of Companilactobacillus
some species grow at 45 °C. The fermentation of pentoses baiquanensis comb. nov.
by Companilactobacillus is species- or strain-specific. Infor-
Companilactobacillus baiquanensis (bai.quan.en´sis. N.L.
mation on the ecology or lifestyle of companilactobacilli is
masc. adj. baiquanensis, pertaining to Baiquan, a county in
scarce.
the Heilongjiang province of China).
A phylogenetic tree on the basis of 16S rRNA genes of all
Basonym: Lactobacillus baiquanensis Wei and Gu 2019, 3192VP
species in the genus Companilactobacillus is provided in Fig.
S6C. The type strain grows at 15 and 37 but not at 45 °C; dl-lactic
acid is produced from ribose, hexoses and maltose [118]. The
The type species is Companilactobacillus alimentarius comb.
genome size is 2.24 Mbp; the mol% G+C content is 34.4.
nov.; Companilactobacillus was previously referred to as L.
alimentarius group. Isolated from fermented Chinese cabbage.
The type strain is 184–8T=LMG 31050T=NCIMB 15152T=CCM
8895T=KCTC 21131T.
Genome sequence accession number: RHOQ00000000.
alimentarius comb. nov.
Companilactobacillus alimentarius (a.li.men.ta'ri.us. L. masc. 16S RNA gene sequence accession number: MK110828.
adj. alimentarius related to food).
Basonym: Lactobacillus alimentarius (ex Reuter 1970) Reuter Description of Companilactobacillus
1983a, 672VP
bobalius comb. nov.
Growth is observed between 15 and 37°C and with pentoses, Companilactobacillus bobalius (bo.ba’li.us. N.L. masc. adj.
hexoses and disaccharides as carbon sources [115]. The bobalius pertaining to the grape variety Bobal).
genome size is 2.34 Mbp. The mol% G+C content of DNA
is 35.4. Basonym: Lactobacillus bobalius Mañes-Lázaro et al. 2008,
2702VP; the validity of the species was questioned but
Isolated as spoilage organisms from marinated fish products, re-established on the basis of ANI values [119].
as fermentation organisms in fermented sausages, as spoilage
Growth occurs in the range of 15 to 40°C and with pentoses,
organism in ready-to-eat meats. A strain of the species had
hexoses, and disaccharides as carbon sources [120]. The
been used as biopreservatives culture [116]. It occurs in asso-
ciation with Fructilactobacillus sanfranciscensis in type I sour-
is 35.3.
doughs [114] and was isolated in other plant fermentations.
Isolated from grape must and forage.
The type strain is R13T=ATCC 29643T=CCUG 30672T=CIP
102986T=DSM 20249T=JCM 1095T=LMG 9187T. The type strain is 203T=CECT 7310T=DSM 19674T=JCM
16180T.
Genome sequence accession number: AZDQ00000000.
Genome sequence accession number: AZDY00000000.
16S RNA gene sequence accession number: M58804.
16S RNA gene sequence accession number: AY681134.
allii comb. nov. Description of Companilactobacillus
Companilactobacillus allii (al’li.i. L. gen. n. allii of Egyptian
crustorum comb. nov.
onions (Allium proliferum), the source of fermented onions, Companilactobacillus crustorum (crus.tor’um. L. gen. pl. n.
pa-kimchi, from which the type strain was isolated). crustorum, of baked goods or cakes).
Basonym: Lactobacillus allii Jung et al. 2017, 4939VP Basonym: Lactobacillus crustorum Scheirlinck et al. 2007,
1466VP
Growth is observed at 25 and 37 °C; hexoses and disaccha-
rides but not pentoses are fermented [117]. The genome size Growth is observed at 15 and at 45 °C; pentoses are not
is 2.51 Mbp. The mol% G+C content of DNA is 35.3. fermented; disaccharide fermentation is strain dependent
[121]. The genome size is 2.22 Mbp. The mol% G+C content
Isolated from scallion kimchi. of DNA is 35.
The type strain is WiKim39T=KCTC 21077T=JCM 31938T. Isolated from sourdough, dairy products and forages. Some
Genome sequence accession numbers: CP019323 (chromo- strains degrade saponins by β-glucuronidase activity [122].
some) and CP019324 (plasmid). The type strain is CCUG 53174T=JCM 15951T=LMG 23699T.
16S RNA gene sequence accession number: NR_159082. Genome sequence accession number: AZDB00000000.
2803
16S RNA gene sequence accession number: AM285450. Genome sequence accession number: RHNT00000000.
16S RNA gene sequence accession number: AB910349.
farciminis comb. nov. Description of Companilactobacillus
Companilactobacillus farciminis (far.ci'mi.nis. L. gen. n. farci- futsaii comb. nov.
minis of sausage).
Companilactobacillus futsaii (fut.sai’i. N.L. gen. n. of futsai,
Basonym: Lactobacillus farciminis Reuter 1983, 672VP the Taiwanese fermented mustard product from which the
Growth is observed between 15 and 42°C and with hexoses type strain was isolated).
and disaccharides but not with pentoses as carbon source Basonym: Lactobacillus futsaii Chao et al. 2012, 489VP. Two
[115]. The genome size is 2.48 Mbp. The mol% G+C content subspecies of C. futsaii, ‘C. futsaii ssp. futsaii’ and ‘C. futsaii ssp.
of DNA is 36.4. chongqingii’ were described [125] but have not yet been included
Isolated from meat products, sourdough, fermentend fish, on the Validation Lists. The two subspecies are highly similar
cold-smoked salmon, soy sauce mash, dairy products, table with respect to DNA–DNA hybridization values, 16rRNA and
olives and fermented vegetables and corn silage. recA gene sequence similarity and were established on the basis
of a divergent pheS sequence, requiring further confirmation.
The type strain is Rv4 naT=ATCC 29644T=DSM 20184T=JCM
1097T=LMG 9200T=NRRL B-4566T. Growth is observed between 15 and 30 °C; hexoses and disac-
charides but not pentoses support acid formation [126]. The
Genome sequence accession number: AZDR00000000. genome size is 2.53 Mbp. The mol% G+C content of DNA
16S RNA gene sequence accession number: M58817. is 35.6.
Isolated from traditional fermented mustard products, fu-tsai
and suan-tsai; it has been used experimentally for fermenta-
Description of Companilactobacillus tion of shrimp waste [127].
formosensis comb. nov.
The type strain is YM 0097T=BCRC 80278T=JCM 17355T.
Companilactobacillus formosensis (for.mo.sen’sis. N.L. masc.
adj. formosensis of or pertaining to Formosa [Taiwan] where Genome sequence accession number: AZDO00000000
the type strain was isolated).
16S RNA gene sequence accession number: HQ322270.S
Basonym: Lactobacillus formosensis Chang et al. 2015, 105VP
Growth is observed between 25 and 45°C but not at 10 °C; Description of Companilactobacillus
hexoses and disaccharides are fermented; pentoses are not
fermented [123]. The genome size is 2.47 Mbp. The mol% ginsenosidimutans sp. nov.
G+C content of DNA is 35.5. Companilactobacillus ginsenosidimutans (gin.se.no.si.di.
mu’tans N.L. neut. n. ginsenosidium, ginsenoside, L. pres.
Isolated from fermented soybean meal for use as poultry feed; part. mutans transforming, N.L. part. adj. ginsenosidimutans,
it has been used experimentally as silage inoculant. ginsenoside converting).
The type strain is S215T=NBRC 109509T=BCRC 80582T. Effective publication Jung et al. 2013 as Lactobacillus ginse-
Genome sequence accession number: QWDI00000000. nosidimutans. C. ginsenosidimutans is a kimchi isolate that
converts ginsenosides by β-glucosidase activity [128]. The
16S RNA gene sequence accession number: AB794060. genome size is 2.59 Mbp. The mol% G+C content of DNA
is 36.7.
Description of Companilactobacillus Isolated from kimchi.

furfuricola comb. nov. The type strain is EMML 3041T=KACC 15420T=LMG
Companilactobacillus furfuricola (fur.fu.ri’co.la. L. masc. n. 31607T=DSM 24154T.
furfur rice bran; L. suff. –cola from L. masc. or fem. n. incola Genome sequence accession number: CP012034.
inhabitant; N.L. masc. n. furfuricola rice bran-inhabitant).
16S RNA gene sequence accession number: HQ389549.
Basonym: Lactobacillus furfuricola Irisawa et al. 2014, 2904VP
Growth is observed between 20 and 37 °C [124]. The genome
size is 2.64 Mbp. The mol% G+C content of DNA is 38.6.
halodurans sp. nov.
Isolated from rice bran paste for Japanese pickles.
Companilactobacillus halodurans (ha.lo.du’rans. Gr. n. hals,
The type strain is Nu 27T=JCM 18764T=NRIC 0900T=DSM halos salt; L. pres. part. durans enduring; N.L. part. adj. halo-
27174T. durans salt-enduring, resisting).
2804
Effective publication Schuster et al., 2019, as Lactobacillus Description of Companilactobacillus

halodurans. C. halodurans grows between 10 and 37 °C and hulinensis comb. nov.
between pH 4.0 and 8.0, and at NaCl concentrations of up to
14 % [129]. Acid is produced from ribose, hexoses and some Companilactobacillus hulinensis (hu.lin.en´sis. N.L. masc. adj.
disaccharides. The genome size is 2.84 Mbp. The mol% G+C hulinensis, pertaining to Hulin, a county in the Heilongjiang
content of DNA is 35.8. province of China).
Isolated from spoiled fermented sausage. Basonym: Lactobacillus hulinensis Wei and Gu, 2019, 10VP
The type strain is TMW1.2172T=DSM 109452T=LMG31402T. Growth is observed at 15 and 37 °C but not at 45 °C; dl-
lactic acid is produced from hexoses and pentoses [118]. The
Genome sequence accession number: VDFP00000000. genome size is 2.35 Mbp, the mol% G+C content is 36.7.
16S RNA gene sequence accession number MK968448. Isolated from fermented Chinese cabbage.
The type strain is 8-1(1)T=LMG 31047T=NCIMB
Description of Companilactobacillus 15156T=CCM 8898T=KCTC 21115T.
heilongjiangensis comb. nov. Genome sequence accession number: RHOO00000000.
Companilactobacillus heilongjiangensis (hei.long.ji.ang.en’sis.
16S RNA gene sequence accession number: MK110830.
N.L. masc. adj. heilongjiangensis pertaining to Heilongjiang,
a river flowing through the province of China where the
bacterium was isolated). Description of Companilactobacillus
Basonym: Lactobacillus heilongjiangensis Gu et al. 2013, insicii comb. nov.
4098VP Companilactobacillus insicii (in.si’ci.i. L. gen. n. insicii from
Hexoses and disaccharides but not pentoses are fermented minced meat).
[130]. The genome size is 2.79 Mbp. The mol% G+C content Basonym: Lactobacillus insicii Ehrmann et al. 2016, 241VP
of DNA is 37.5.
Growth is observed in the range of 8–45 °C [132]; the genome
Isolated from fermented vegetables and type I sourdough. size is 2.54 Mbp. The mol% G+C content of DNA is 34.9.
The type strain is S4-3T=LMG 26166T=DSM 28069T=NCIMB Isolated from pork salami.
14701T.
The type strain is TMW 1.2011T=CECT 8802T=DSM 29801T.
Genome sequence accession number: CP012559.
Genome sequence accession number: RHNU00000000.
16S RNA gene sequence accession number: JF411966.
16S RNA gene sequence accession number: KP677494.
huachuanensis comb. nov.
jidongensis comb. nov.
Companilactobacillus huachuanensis (hua.chuan.en´sis. N.L.
masc. adj. huachuanensis pertaining to Huachuan County, a Companilactobacillus jidongensis (ji.dong.en´sis. N.L.
county in the Heilongjiang province of China). masc. adj. jidongensis, pertaining to Jidong, a county in the
Heilongjiang province of China).
Basonym: Lactobacillus huachuanensis Fu and Gu 2019,
2812VP Basonym: Lactobacillus jidongensis Wei and Gu 2019, 3192VP
The physiological properties of C. huachuanensis are compa- Growth is observed at 10 °C but not at 37 °C; dl-lactic acid
rable to C. heilongjiangensis [131]. The genome size is 3.02 is produced from hexoses but not from pentoses [118]. The
Mbp. The mol% G+C content of DNA is 36.4. genome size is 2.71 Mbp. The mol% G+C content of DNA
is 36.5.
Isolated from fermented Chinese cabbage.
The type strain is 395–6.2T=CCM 8927T=NCIMB15188T=L
MG31179T. The type strain is 204–8T=LMG31054T=NCIMB15159T=CC
M8900T=KCTC21133T=LMG31054T.
Genome sequence accession number: BJDF00000000.
Genome sequence accession number: RHOP00000000.
16S RNA gene sequence accession number: LC438522.
2805
Description of Companilactobacillus Description of Companilactobacillus

kedongensis comb. nov. kimchii comb. nov.
Companilactobacillus kedongensis (ke.dong.en´sis. N.L. Companilactobacillus kimchii (kim’chi.i. N.L. gen. n. kimchii
masc. adj. kedongensis, pertaining to Kedong, a county in the from kimchi, a Korean fermented-vegetable food).
Basonym: Lactobacillus kimchii Yoon et al. 2000, 1794VP; the
Basonym: Lactobacillus kedongensis Wei and Gu 2019, 3191 VP
classification of this taxon as a separate species has been ques-
tioned but was confirmed on the basis of ANI values [119].
Characteristics of C. kedongensis are generally similar to C.
jidongensis [118]. The genome size is 2.27 Mbp, the mol% Growth occurs in the range of 10 to 40°C and with pentoses,
G+C content is 36.2. hexoses, and disaccharides as carbon sources [134]. The
is 35.1.
The type strain is 116–2T=LMG31051T=NCIMB 15158T=CCM
Isolated from kimchi.
8899T=KCTC 21124T.
The type strain is MT-1077T=ATCC BAA-131T=DSM
Genome sequence accession number: RHOR00000000.
13961T=JCM 10707T=KCTC 8903PT.
Genome sequence accession number: AZDH00000000.
16S RNA gene sequence accession number: AF183558.
keshanensis comb. nov.
Companilactobacillus keshanensis (ke.shan.en´sis. N.L. masc.
adj. keshanensis, pertaining to a county in the Heilongjiang Description of Companilactobacillus
province of China). metriopterae comb. nov.
Basonym: Lactobacillus keshanensis Wei and Gu 2019, 3191VP Companilactobacillus metriopterae (me.tri.o.pte’rae. N.L. fem.
gen. n. metriopterae of Metrioptera, the genus of grasshoppers
Characteristics of C. keshanensis are similar to C. jidongensis from which the type strain was isolated).
[118]. The genome size is 2.30 Mbp. The mol% G+C content
of DNA is 35.6. Basonym: Lactobacillus metriopterae Chiba et al. 2018, 1486VP;
the species includes strains previously described as Lactoba-
Isolated from fermented Chinese cabbage. cillus terrae Kim et al. 2018 [135, 136]
The type strain is 33–7T=LMG 31166T=NCIMB 15153T=CCM C. metriopterae forms an outgroup of Companilactobacillus
8936T=KCTC 21118T=LMG 31166T. and has the smallest genome size (1.50 Mbp) and mol% G+C
Genome sequence accession number: RHOS00000000. content (32.8); it seems to transition to an insect associated
lifestyle. C. metriopterae is pigmented and ferments fewer
16S RNA gene sequence accession number: MK110816. carbohydrates than other companilactobacilli.
Isolated from the gut of grasshopper Metrioptera engelhardti,
Description of Companilactobacillus other isolates were obtained from soil.
kimchiensis comb. nov. The type strain is Hime 5-1T=JCM 31635T=DSM 103730T.
Companilactobacillus kimchiensis (kim.chi.en’sis. N.L. gen.
n. kimchiensis from kimchi, a Korean fermented-vegetable Genome sequence accession number: RSTD00000000.
food). 16S RNA gene sequence accession number: LC190736.
Basonym: Lactobacillus kimchiensis Kim et al. 2013, 1358VP.
Hexoses and disaccharides but not pentoses are fermented; Description of Companilactobacillus
growth is observed at 15–37 °C [133]. The genome size is 2.70
Mbp. The mol% G+C content of DNA is 35.5. mindensis comb. nov.
Companilactobacillus mindensis (min.den’sis. N.L. adj. mind-
Isolated from kimchi. ensis from Minden, Germany, where the type strain was
The type strain is L133T=DSM 24716T=JCM 17702T=KACC isolated).
15533T. Basonym: Lactobacillus mindensis Ehrmann et al. 2003, 11VP
Genome sequence accession number: JQCF00000000. Growth is observed between 15 and 30 °C. Pentoses are not
16S RNA gene sequence accession number: HQ906500. fermented [137]. The genome size is 2.34 Mbp. The mol%
Isolated from a type I sourdough.
2806
The type strain is TMW 1.80T=CCUG 48642T=DSM The type strain is LP33T=TMW 1.1265T=CIP 108546T=DSM
14500T=JCM 12532T=LMG 21508T. 16982T=JCM 16171T.
Genome sequence accession number: AZEZ00000000. Genome sequence accession number: AZFV00000000.
16S RNA gene sequence accession number: AJ313530. 16S RNA gene sequence accession number: AY690834.

mishanensis comb. nov. nodensis comb. nov.
Companilactobacillus mishanensis (mi.shan.en´sis. N.L. Companilactobacillus nodensis (no.den’sis. N.L. masc. adj.
masc. adj. mishanensis, pertaining to Mishan, a county in the nodensis referring to Noda, the Japanese city in which the
Heilongjiang province of China). type strain was isolated).
Basonym: Lactobacillus mishanensis Wei and Gu 2019, 3193VP Basonym: Lactobacillus nodensis Kashiwagi et al. 2009, 64VP
Characteristics are similar to C. hulinensis [118]. The genome Growth is observed between 15 and 37 °C and with pentoses
size is 2.47 Mbp, the mol% G+C is 36.6. and hexoses but not disaccharides as carbon sources [140].
The genome size is 2.68 Mbp. The mol% G+C content of DNA
Isolated from fermented Chinese cabbage. is 37.6.
The type strain is 256–3T=LMG 31048T=NCIMB 15160T=CCM Isolated from fermented rice bran paste and has been used
8901T=LMG 31048T. experimentally as adjunct culture in cheese [141].
Genome sequence accession number: RHON00000000. The type strain is iz4b-1T=DSM 19682T=JCM 14932T.
16S RNA gene sequence accession number: MK110831. Genome sequence accession number: AZDZ00000000.
musae comb. nov. Description of Companilactobacillus
Companilactobacillus musae (mu’sae. N.L. gen. n. musae of nuruki comb. nov.
Musa; pertaining to banana fruit from Musa paradisiaca var. Companilactobacillus nuruki (nu.ru’ki. N.L. gen. n. nuruki of
sapientum). Nuruk, a Korean traditional rice fermentation starter).
Basonym: Lactobacillus musae Chen et al. 2017, 5148VP Basonym: Lactobacillus nuruki Heo et al. 2018, 3277VP
Growth is observed between 25 and 37°C and with hexoses Growth is observed between 4 and 40 °C; a wide spectrum of
and disaccharides but not with pentoses as carbon sources carbohydrates including pentoses, hexoses and disaccharides
[138]. The genome size is 2.35 Mbp. The mol% G+C content is fermented [142].
of DNA is 34.6.
The genome size is 2.58 Mbp. The mol% G+C content of DNA
Isolated from banana fruits. is 34.2.
The type strain is 313T=NBRC 112868T=BCRC 81020T. Isolated from Nuruk, a rice bran fermentation starter.
Genome sequence accession number: BIFW00000000. The type strain is SYF10-1aT=KACC 18726T=NBRC 112011T.
16S RNA gene sequence accession number: LC184607. Genome sequence accession number: NIPR00000000.
16S RNA gene sequence accession number: MG786754.
nantensis comb. nov. Description of Companilactobacillus
Companilactobacillus nantensis (nan.ten’sis. N.L. masc. adj. paralimentarius comb. nov.
nantensis pertaining to Nantes, from where the first stain of
Companilactobacillus paralimentarius (par.a.li.men.ta’ri.us.
this species was isolated).
Gr. pref. para beside; L. masc. adj. alimentarius, a species
Basonym: Lactobacillus nantensis Valcheva et al. 2006, 589VP epithet; N.L. masc. adj. paralimentarius beside C. alimenta-
rius, referring to the close relationship with this organism).
Growth occurs at 15 but not at 45 °C; a wide range of carbo-
hydrates including ribose, hexoses and disaccharides are Basonym: Lactobacillus paralimentarius Cai et al. 1999, 1455VP
fermented [139]. The genome size is 2.91 Mbp. The mol%
The physiological properties of C. paralimentarius as described
G+C content of DNA 36.2.
[143] are similar to C. alimentarius. The genome size is 2.55
Isolated from a type I sourdough. Mbp. The mol% G+C content of DNA is 35.1.
2807
Isolated from sourdough, other cereal fermentations, and The type strain grows at 15 and 37 but not at 45 °C [145]. The
from poultry meat. genome size is 2.17 Mbp. The mol% G+C content of DNA
is 34.1.
The type strain is TB 1T=CCUG 43349T=CIP 106794T=DSM
13238T=JCM 10415T=LMG 19152T. Isolated from sausage.
Genome sequence accession number: AZES00000000. The type strain is R 19cT=CECT 5920T=DSM 20183T=JCM
18037T.
Genome sequence accession number: AZDG00000000.
16S RNA gene sequence accession number: AJ576006.
salsicarnum sp. nov.
Companilactobacillus salsicarnum (sal.si.carn´um. L. adj. Description of Companilactobacillus
salsus, salted; L. gen.n. carnis of meat; N.L. gen. n. salsi- versmoldensis comb. nov.
carnum of salted meat, referring to salami as the source of Companilactobacillus versmoldensis (vers.mold.en’sis. N.L.
isolation). masc. adj. versmoldensis is pertaining to Versmold, the town
in Germany where the strains were isolated).
Effective publication Schuster et al. 2019 as Lactobacillus
salsicarnum. C. salsicarnum grows between 10 and 37 °C and Basonym: Lactobacillus versmoldensis Kröckel et al. 2003,
between pH 4.5 and 8.5, and at NaCl concentrations of up to 8 516VP
% [129]. Acid is produced from ribose and arabinose, hexoses,
Description is provided by [146]. The genome size is 2.37
and maltose and trehalose. The genome size is 2.43 Mbp. The
Mbp. The mol% G+C content of DNA is 38.3.
Isolated from poultry salami.
Isolated from spoiled fermented sausage.
The type strain is KU-3T=ATCC BAA-478T=DSM
The type strain is TMW 1.2098T=DSM 109451T=LMG 31401T. 14857T=JCM 16174T=NCCB 100034T.
Genome sequence accession number: VDFN00000000. Genome sequence accession number: AZFA00000000.
16S RNA gene sequence accession number MK968446. 16S RNA gene sequence accession number: AJ496791.

suantsaicola comb. nov. zhachilii comb. nov.
Companilactobacillus suantsaicola (suan.tsai'co.la. N.L. neut. Companilactobacillus zhachilii ( zha.
chi’li.i. N.L. gen. n.
n. suantsaium, from suan-tsai, fermented mustard greens; L. zhachilii of zha-chili, a Chinese fermented mustard green
suff. –cola from L. masc. or fem. n. incola inhabitant; N.L. product).
masc. n. suantsaicola occurring in suan-tsai.)
Basonym: Lactobacillus zhachilii Zhang et al., 2019, 2199VP
Basonym: Lactobacillus suantsaicola Lin et al. 2019, 8 VP
Growth is observed between 15 and 45°C and with a wide
Growth is observed between 20 and 37°C and at pH 4.0 to range of hexoses and disaccharides but not with pentoses
10.0 [144]. The genome size is 2.60 Mbp. The mol% G+C [147]. The genome size is 2.71 Mbp. The mol% G+C content
content of DNA is 36.9. of DNA is 36.3.
Isolated from suan-tsai, a traditional fermented mustard Isolated from Zha-Chili, a traditional fermented product
green product of Taiwan. from mustard greens.
The type strain is R7T=BCRC 81127T=NBRC 113530T. The type strain is HBUAS52074T=GDMCC1.1417T=KCTC
21106T.
Genome sequence accession number: RKLY00000000.
Genome sequence accession number: CP031933.
16S RNA gene sequence accession number: MH810311.
16S RNA gene sequence accession number: MH392835.
tucceti comb. nov.
zhongbaensis comb. nov.
Companilactobacillus tucceti (tuc.ce’ti. L. gen. n. tucceti, from
Companilactobacillus zhongbaensis (zhong.ba.en´sis. N.L.
a sausage, referring to the origin of the type strain).
masc. adj. zhongbaensis from Zhongba, a county in Tibet,
Basonym: Lactobacillus tucceti Chenoll et al. 2009, 925VL China, where the type strain was isolated).
2808
Basonym: Lactobacillus zhongbaensis Wei and Gu 2019, 16S rRNA gene accession number: AY683322.
3193VP
Growth is observed at 15 and 37 °C but not at 45 °C; dl-lactic Description of Lapidilactobacillus
acid is produced from a broad range of pentoses, hexoses and
disaccharides [118]. The genome size is 3.04 Mbp, the mol% bayanensis comb. nov.
G+C content is 39.4. Lapidilactobacillus bayanensis (ba.yan.en´sis. N.L. masc. adj.
bayanensis pertaining to Bayan, a county in the Heilongjiang
Isolated from a fermented dairy product. province of China).
The type strain is M1575T=LMG 31045T=NCIMB Basonym: Lactobacillus bayanensis Wei and Gu 2019, 3191VP
15149T=CCM 8892T.
Cells grow at 15 °C but not at 37 °C. l(+)-Lactic acid and d(−)-
Genome sequence accession number: RHOM00000000. lactic acid are produced from glucose, fructose, mannose and
16S RNA gene sequence accession number: MK110832. maltose; pentoses or other disaccharides are not fermented
[118]. The genome size of the type strain is 2.43 Mbp. The
Description of Lapidilactobacillus Isolated from fermented Chinese cabbage.
gen. nov. The type strain 54–5T=LMG 31166T=NCIMB 15153T=CCM
Lapidilactobacillus [La.pi.di.lac.to.ba.cil’lus L. masc. n. lapis, 8936T=KCTC 21118T.
stone, referring to the isolation source of the type species
Lapidilactobacillus concavus; N.L. masc. n. Lactobacillus a Genome sequence accession number: RHOX00000000.
bacterial genus; N.L. masc. n. Lapidilactobacillus, a lactoba- 16S rRNA gene accession number: MK110807.
cillus isolated from stone (walls)].
The cells are Gram-positive rods or cocci, homofermentative,
non motile, non-spore-forming, facultatively anaerobes, and
Description of Lapidilactobacillus
catalase-negative. Pentose utilization and the pH range for dextrinicus comb. nov.
growth are strain dependent, the optimum pH is between Lapidilactobacillus dextrinicus (dex.tri′ni.cus. N.L. neut. n.
6.0 and 7.0. They are homofermentative and mainly produce dextrinum, dextrin; L. suff. -icus, suffix used with the sense of
l-lactic acid. The optimum temperature for growth is 30–37 belonging to; N.L. masc. adj. dextrinicus, related to dextrin).
°C. No ammonia is produced from arginine. The mol% G+C Basonym: Lactobacillus dextrinicus (Coster and White 1964)
content of DNA is between 38.1 and 49.0. Haakensen et al. 2009, 620VP.
A phylogenetic tree on the basis of 16S rRNA genes of all Strains of L. dextrinicus were previously classified as
species in the genus Lapidiactobacillus is provided in Fig. S6D. Pediococcus dextrinicus (Coster and White 1964) Back
The type species is Lapidilactobacillus concavus comb. 1978 (Approved Lists 1980) or Pediococcus cerevisiae subsp.
nov.; Lapidilactobacillus was previously referred to as L. dextrinicus. Cells are spherical, and may occur singly, in
concavus/dextrinicus group. pairs or clusters and rarely in chains. Clusters of four may be
observed, but they do not divide in two perpendicular direc-
tions at right angles. Only l(+)-lactic acid is produced from
Description of Lapidilactobacillus maltose, dextrin and starch [149]. The genome size of the type
strain is 1.81 Mbp. The mol% G+C content of DNA is 38.1.
concavus comb. nov.
Lapidilactobacillus concavus ( con.
ca’vus. L. masc. adj. Isolated from silage, fermenting vegetables, beer and sliced
concavus, curved, referring to the curved shape of the cells). vacuum-packed cooked sausage.
Basonym: Lactobacillus concavus Tong and Dong 2005, 2201VP The type strain is ATCC 33087=CCUG 18834T=CIP
103407T=DSM 20335T=JCM 5887T=LMG 11485T=NCIMB
Cells of L. concavus are non-motile curved rods; the optimum 701561T=VKM B-1603T.
pH for growth is 6.0–6.4; they can also grow at 8 % (v/v)
ethanol but not in 6.5 % (w/v) NaCl [148]. The genome size Genome sequence accession number: AYYK00000000.
of the type strain is 1.90 Mbp. The mol% G+C content of 16S rRNA gene accession number: D87679.
DNA is 43.3.
Isolated from the walls of a distilled-spirit-fermenting cellar
in China.
Description of Agrilactobacillus gen.
nov.
The type strain is C-5–1T=DSM 17758T=AS 1.5017T=JCM
Agrilactobacillus (A.gri.lac.to.ba.cil’lus. L. adj.masc. n. ager, a
14153T=LMG 22739T.
field; N.L. masc. n. Lactobacillus, a bacterial genus; Agrilac-
Genome sequence accession number: AZFX00000000. tobacillus, a lactobacillus from fields or soil).
2809
Gram positive, rod-shaped, catalase negative, homofermen- Gram-positive, rod-shaped, catalase-negative, homofermen-
tative and aerotolerant. The genome size ranges from 2.33 tative and aerotolerant. Strains of the genus were isolated
to 3.47 Mbp; the mol% G+C content of DNA ranges from from spoiled beverages including beer and fermented dairy
42.1 to 44.0. Origin, physiological properties and genome beverages, fermented vegetables, and fermented cereals.
characteristics suggest a free-living lifestyle of the genus. Growth is observed in the range of 15–42 °C; a wide range
Isolated from compost that included shochu mash and from of carbohydrates including pentoses, hexoses and oligosac-
vegetables (mustard) in Indonesia and China. charides are fermented. The genome size ranges from 3.14
The type species of the genus is A. composti. to 3.32 Mbp; the mol% G+C content of DNA ranges from
49.1 to 56.3.
Description of Agrilactobacillus species in the genus Schleiferilactobacillus is provided in Fig.
composti comb. nov. S6E.
Agrilactobacillus composti (com.pos’ti. N.L. gen. n. composti The type species of the genus is S. perolens comb. nov.; Schleif-
of compost, the source of the type strain). erilactobacillus was previously referred to as L. perolens group.
Basonym: Lactobacillus composti Endo and Okada 2007, 872VP
The type strain grows over a relatively wide pH range and Description of Schleiferilactobacillus
in the range of 10–37°C but not at 45 °C [150]. The genome
size is 3.47 Mbp and the mol% G+C content of DNA is 44.0. perolens comb. nov.
Schleiferilactobacillus perolens (per.o'lens, Latin preposition
Isolated from compost from shochu mash solids and from per through, penetrating; L. pres. part. olens having an odor;
pulque, a Mexican alcoholic beverage [151].
N.L. part. adj. perolens smelling, referring to the copious
The type strain is NRIC 0689T=DSM 18527T=JCM 14202T. diacetyl production by strains of the species).
Genome sequence accession number: AZGA00000000. Basonym: Lactobacillus perolens Back et al. 2000, 3VP
16S rRNA gene accession number: AB268118. Description of the species is as provided by [152]. Spoilage
was attributed to copious amounts of diacetyl produced by
S. perolens. The genome size is 3.31 Mbp and the mol% G+C
Description of Agrilactobacillus content of DNA is 49.1.
yilanensis comb. nov Isolated from spoiled soft drinks and brewery environments.
Agrilactobacillus yilanensis (yi.lan.en´sis. N.L. masc. adj.
The type strain is L 533T=DSM 12744T=JCM 12534T=LMG
yilanensis pertaining to Yilan, a county in the Heilongjiang
province of China). 18936T.
Basonym: Lactobacillus yilanensis Wei and Gu 2019, 3189VP Genome sequence accession number: AZEC00000000.
The type strain grows at 15 but not at 45 °C and ferments 16S rRNA gene accession number: Y19167.
a wide range of pentoses, hexoses and disaccharides [118].
The genome size is 2.33 Mbp and the mol% G+C content of
DNA is 41.7. Description of Schleiferilactobacillus
Isolated from a Chinese cabbage fermentation.
harbinensis comb. nov.
Schleiferilactobacillus harbinensis (har.bi.nen’sis. N.L. masc.
The type strain is 54–2T=LMG 31058T=NCIMB 15154T=CCM adj. harbinensis pertaining to Harbin, a city in northeastern
8896T=KCTC 21120T. China where the type strain was isolated).
Genome sequence accession number: RHOY00000000. Basonym: Lactobacillus harbinensis Miyamoto et al. 2006, 2VL
16S rRNA gene accession number: MK110806. Description of the species is as provided by [153]. The genome
size is 3.14 Mbp and the mol% G+C content of DNA is 53.8.
Description of Schleiferilactobacillus Isolated from fermented vegetables ‘Suan Cai’, the brewery
gen. nov. environment, fermented cereals and tomato pomace, and
spoiled soft drinks.
Schleiferilactobacillus (Schlei.fer.i.lac.to.ba.cil’lus. L. masc.
noun Schleiferi, of (Karl-Heinz) Schleifer, a German micro- The type strain is AHU 1762T=DSM 16991T=JCM
biologist and taxonomist who made seminal contributions 16178T=NBRC 100982T=SBT 10908T.
to bacterial taxonomy. N.L.masc. n. Lactobacillus a bacterial
Genome sequence accession number: AZFW00000000.
genus; N.L. masc. n. Schleiferilactobacillus, a lactobacillus
named after Karl-Heinz Schleifer). 16S rRNA gene accession number: AB196123.
2810
Description of Schleiferilactobacillus Isolates were obtained from diverse sources including chinese
shenzhenensis comb. nov. traditional pickle, infant faeces, corn liquor, oat silage,
commercial dietary supplements, sputum, nasopharynx
Schleiferilactobacillus shenzhenensis (shen.zhen.en’sis. N.L.
[47]. Information on the lifestyle of L. casei is counded by
masc. adj. shenzhenensis of Shenzhen, the city in the South
the unclear taxonomy over the past decades; most genomes
of China where the type strain was isolated).
of designated as L. casei in the NCBI database should be clas-
Basonym: Lactobacillus shenzhenensis Zou et al. 2013, 1821VP sified as L. paracasei instead [10, 47].
Growth is observed at 15 and 45 °C [154]. The genome size is The type strain is 03 [7, IAM 12473, Orland L-323, R.P. Titt-
3.27 Mbp and the mol% G+C content of DNA is 56.3. sler 303]T=ATCC 393T=BCRC 10697T=CCUG 21451T=CECT
Isolated from a fermented dairy beverage. 475T=CIP 103137T=DSM 20011T=IAM 12473T=NBRC
15883T=JCM 1134T=KCTC 3109T=LMG 6904T=NCIMB
The type strain is LY-73T=CCTCC M 2011481T=KACC 11970T=NCIMB 11970T=NRRL B-1922T.
16878T.
Genome sequence accession number: BALS00000000.
Genome sequence accession number: AVAA00000000.
Description of Lacticaseibacillus
Description of Lacticaseibacillus gen.
baoqingensis comb. nov.
nov.
Lacticaseibacillus baoqingensis (boa.qing.en’sis. N.L. masc.
Lacticaseibacillus (Lac.ti.ca.se.i.ba.cil'lus. L. neut. n. lac milk; adj. boaqingensis of Boaqing, from where the type strain was
L. n. caseus cheese, referring to the casei-group lactobacilli; isolated).
L. masc. n. bacillus a rod; N.L. masc. n. Lacticaseibacillus a
milk-derived rodlet from the [Lactobacillus] casei group). Basonym: Lactobacillus baoqingensis Long and Gu 2019,
2347VP
Strains of Lacticaseibacillus are homofermentative; some but
not all species metabolise pentoses via the phosphoketolase Original characteristics of L. baoqingensis strains are as
pathway. The mol % G+C content of DNA is between 46 and provided by Long and Gu [158].
58.0. The genome size ranges from 1.93 to 3.14 Mbp. Strains
The genome size of the type strain is 2.86 Mbp. The mol%
are non-motile, oxidase negative, often producing d(−)- and
l(+)-lactic acid from glucose. The temperature range for
growth is variable, but never below 10 °C and never above Isolated from a vegetable fermentation.
45 °C. One subspecies survives 70 °C for 40 s. Lys–d-Asp
The type strain is 47–3T=NCIMB 15165T=CCM 8903T=LMG
is the most common type of the peptidoglycan. The genus
31064T.
has considerable economic importance as it harbours several
species that are used as starter cultures in dairy fermentations Genome sequence accession number: RHOI00000000.
and as probiotics [58, 59].
species in the genus Lacticaseibacillus is provided in Fig. S6F.
The type species is Lacticaseibacillus casei comb. nov.; Lactica- Description of Lacticaseibacillus
seibacillus was previously referred to as L. casei group. brantae comb. nov.
Lacticaseibacillus brantae (bran′tae. N.L. gen. n. brantae, of
Branta, isolated from Branta canadensis, Canada goose).
Basonym: Lactobacillus brantae Volokhov et al. 2012, 2068VP
casei comb. nov.
Lacticaseibacillus casei (ca’se.i. L. gen. n. casei of cheese). Original characteristics of L. brantae strains are provided by
Basonym: Lactobacillus casei (Orla Jensen 1916, Hansen and mol% G+C content of DNA is 47.5.
Lessel 1971, 71 (Approved Lists) (Streptobacterium casei Orla-
Jensen 1919, 166). Isolated from the faeces of wild Canada goose (Branta
canadensis) and from experimental sourdoughs.
The species includes strains previously classified as Lactoba-
cillus zeae [155]. Original characteristics of L. casei strains The type strain is SL1108T=ATCC BAA-2142T=DSM
are provided by [2, 156, 157]. L. casei is differentiated from 23927T=LMG 26001T.
most other lactobacilli by catalase activity [47]. The genome
Genome sequence accession number: AYZQ00000000.
size of the type strain is 2.83 Mbp. The mol% G+C content
of DNA is 46.5. 16S rRNA gene accession number: HQ022861.
2811
Description of Lacticaseibacillus Description of Lacticaseibacillus

camelliae comb. nov. jixianensis comb. nov.
Lacticaseibacillus camelliae (ca.mel′li.ae. N.L. gen. n. camel- Lacticaseibacillus jixianensis (ji.xian.en’sis. N.L. masc. adj.
liae, of Camellia sinensis fermented tea leaves). jixianensis pertaining to Jixian, a county in the Heilongjiang
province of China).
Basonym: Lactobacillus camelliae Tanasupawat et al. 2007,
1371VL [160] Basonym: Lactobacillus jixianensis Long and Gu, 2019, 2346VP
Original characteristics of L. camelliae strains are as provided Original characteristics of L. jixianensis strains are provided
by [161] .The genome size of the type strain is 2.57 Mbp. The by [158]. The genome size of the type strain is 2.47 Mbp. The
mol% G+C content of DNA is 55.4. mol% G+C content of DNA is 58.3.
Isolated from fermented tea (Camellia sinensis) leaves and Isolated from fermented Chinese cabbage.
fermented tomato pomace.
The type strain is 159–4T=CCM 8911 T=NCIMB 15175 T.
The type strain is MCH3-1 =BCC 21233 =DSM 22697 =JCM
T T T
Genome sequence accession number: RHOJ00000000.
13995T=NRIC 0672T.
Genome sequence accession number: AYZJ00000000.
manihotivorans comb. nov.
Description of Lacticaseibacillus Lacticaseibacillus manihotivorans (ma.ni.ho.ti.vo’rans. N.L.
chiayiensis comb. nov. neut. n. manihotum cassava; L. v. vorare to devour; N.L. part.
Lacticaseibacillus chiayiensis (chia.yi.en’sis. N.L. masc. adj. adj. manihotivorans cassava-devouring).
chiayiensis of Chiayi, from where the type strain was isolated).
Basonym: Lactobacillus manihotivorans Morlon-Guyot et al.
Basonym: Lactobacillus chiayiensis effective publication 1998, 1107VP
Huang et al. 2018, 3382VL
Strains of L. manihotivorans express extracellular amylases
Original characteristics of L. chiayiensis strains are provided [165]. The genome size of the type strain is 3.14 Mbp. The
by [162, 163]. The genome size of the type strain is 2.87 Mbp. mol% G+C content of DNA is 47.7.
The mol% G+C content of DNA is 47.1.
Isolated from sour cassava starch fermentation and from
Isolated from cow manure. tomato pomace silage.
The type strain is NCYUAST=BCRC 81062T=NBRC 112906T. The type strain is OND 32T=CCUG 42894T=CIP
105851T=DSM 13343T=JCM 12514T = LMG 18010T.
Genome sequence accession number: MSSM00000000.
Genome sequence accession number: AZEU00000000.
16S rRNA gene accession number: MF446960.
hulanensis comb. nov. Description of Lacticaseibacillus
Lacticaseibacillus hulanensis (hu.lan.en’sis. N.L. masc. adj. nasuensis comb. nov.
hulanensis pertaining to Hulan, a district of Harbin city in Lacticaseibacillus nasuensis (na.su.en′sis. N.L. masc. adj.
China). nasuensis, named after Nasu-shiobara City, where strains
were first isolated).
Basonym: Lactobacillus hulanensis Zhao and Gu et al. 2019,
5VP Basonym: Lactobacillus nasuensis Cai et al. 2012, 1143VP
Original characteristics of L. hulanensis strains are provided Original characteristics of L. nasuensis strains are provided
[164]. The genome size of the type strain is 2.36 Mbp. The [166]. The genome size of the type strain is 2.28 Mbp. The
mol% G+C content of DNA is 52.7. mol% G+C content of DNA is 57.0.
Isolated from suancai, fermented Chinese cabbage Isolated from a Sudan grass [Sorghum sudanense (Piper)
Stapf.] silage.
The type strain is ZW163T=NCIMB15193T =
CCM8926T=CCTCC AB 2019015T. The type strain is SU 18T=JCM 17158T=CGMCC 1.10801T.
Genome sequence accession number: RRYD00000000. Genome sequence accession number: AZDJ00000000.
16S rRNA gene accession number: LC436604. 16S rRNA gene accession number: AB608051.
2812
Description of Lacticaseibacillus Description of Lacticaseibacillus

pantheris comb. nov. paracasei subsp. paracasei comb. nov.
Lacticaseibacillus pantheris (pan’ther.is. N.L. gen. n. pantheris Lacticaseibacillus paracasei subsp. paracasei (pa.ra.ca’se.i. Gr.
of the panther, referring to the isolation of the strains from prep. para resembling; N.L. gen. n. casei a species epithet; N.L.
jaguar faeces). gen. n. paracasei resembling Lactobacillus casei).
Basonym: Lactobacillus pantheris Liu and Dong 2002, 1747VP Basonym Lactobacillus paracasei subsp. paracasei Collins et
al. 1989, 107VP
Characteristics of the species are described in [167]. L.
pantheris together with L. thailandensis, L. sharpie, L. The genome size of the type strain is 2.88 Mbp. The mol%
songhuajiangensis and L. hulanensis forms a monophyletic G+C content of DNA is 46.5.
outgroup to Lacticaseibacillus; however, physiological char-
Isolated from dairy products, sewage, silage, humans and
acteristics and their ecology, as far as data are available, are
clinical sources.
similar to other Lacticaseibacillus species. The genome size
of the type strain is 2.55 Mbp. The mol% G+C content of The type strain is DSM 5622T=ATCC 25302T=AS 1.2826T=
DNA is 52.9. JCM 8130T=LMG 13087T.
Isolated from the faeces of a jaguar in Beijing Zoo and from Genome sequence accession number: AZGH00000000.
fermented vegetables.
16S rRNA gene accession number: D79212.
The type strain is A24-2-1T=DSM 15945T=AS 1.2826T=JCM
12539T=LMG 21017T.
Genome sequence accession number: AZFJ00000000. Description of Lacticaseibacillus
paracasei subsp. tolerans comb. nov.
Lacticaseibacillus paracasei subsp. tolerans (to.le’rans. L. masc.
part. tolerans tolerating, enduring; means survival during the
pasteurization of milk).
paracasei comb. nov. Basonym Lactobacillus paracasei subsp. tolerans Abo-Elnaga
and Kandler 1965, Collins et al. 1989, 108VP
Lacticaseibacillus paracasei (pa.ra.ca’se.i. Gr. prep. para resem-
bling; N.L. gen. n. casei a species epithet; N.L. gen. n. paracasei Some strains of the subspecies were initially described as
resembling Lactobacillus casei). L. casei subsp. tolerans [168]. Original characteristics of L.
paracasei subsp. tolerans strains are described in [170]. The
Basonym: Lactobacillus paracasei Collins et al. 1989, 107VP
genome size of the type strain is 2.38 Mbp. The mol% G+C
Strains of this species including strains previously referred content of DNA is 46.4.
to as L. casei subsp. alactosus, L. casei subsp. pseudoplan-
Isolated from dairy products and tomato pomace silage.
tarum and L. casei subsp. tolerans were transferred from L.
casei [168]. Original characteristics of L. paracasei strains The type strain is 7/74T=ATCC 25599T=CCUG 34829T=CIP
are described in [168]. Cells have superoxide dismutase 102994T=CIP 103024T=DSM 20258T=NBRC 15906T=JCM
activity [47], are rods (0.8–1.0×2.0–4.0 µm), often with 1171T=LMG 9191T=NCIMB 9709 T.
square ends, occurring singly or in chains. Growth is Genome sequence accession number: AYYJ00000000.
observed between 10 and 40 °C. Some strains grow at 5 and
45 °C. Two subspecies are validly published. The genome 16S rRNA gene accession number of the type strain:
size of the type strain is 2.88 Mbp. The mol% G+C content AB181950.
of DNA is 46.5.
Strains of this species have a nomadic lifestyle and were
isolated from a variety of courses including the human oral
cavity [169], fermented cereals, vegetables, meats, dairy prod- porcinae comb. nov.
ucts and in invertebrate hosts. Lacticaseibacillus porcinae (por.ci’nae. L. gen. n. porcinae of
pork).
The type strain is DSM 5622T=ATCC 25302T=AS 1.2826T=JCM
8130T=LMG 13087T. Basonym Lactobacillus porcinae Nguyen et al. 2013, 1758VP
Genome sequence accession number: AZGH00000000. Original characteristics of L. porcinae strains are described
in [171]. The genome size of the type strain is 2.84 Mbp. The
16S rRNA gene accession number of the type strain: D79212.
Two subspecies are recognized: Lacticaseibacillus paracasei
Isolate from nem chua (fermented beef) in northern Vietnam.
subsp. paracasei and Lacticaseibacillus paracasei subsp.
tolerans. The type strain is R-42633T=CCUG 62266T=LMG 26767T.
2813
Genome sequence accession number: RHNS00000000. Original characteristics of L. sharpeae strains are described
in [174, 175]. The genome size of the type strain is 2.45 Mbp.
16S rRNA gene accession number: HE616585.
Isolated from municipal sewage and spoiled meat.
Description of Lacticaseibacillus The type strain is 71T=ATCC 49974T=CIP 101266T=DSM
rhamnosus comb. nov. 20505T=JCM 1186T=LMG 9214T=NRRL B-14855T.
Lacticaseibacillus rhamnosus (rham.no’sus. N.L. masc. adj. Genome sequence accession number: AYYO00000000.
rhamnosus pertaining to rhamnose).
Basonym Lactobacillus rhamnosus (Hansen 1968, Collins et
al. 1989, 108VP (Lactobacillus casei subsp. rhamnosus Hansen
1968 [172], Description of Lacticaseibacillus songhuajiangensis
comb. nov.
Original characteristics of L. rhamnosus strains are described
Lacticaseibacillus songhuajiangensis (song.hua.ji.ang.en’sis.
in by [172]. The genome size of the type strain is 2.95 Mbp.
N.L. masc. adj. songhuajiangensis pertaining to the Songhua-
jiang River, a river flowing through Heilongjiang Province
The species has a nomadic lifestyle and was isolated from a of China).
broad range of habitats including dairy products, fermented
Basonym Lactobacillus songhuajiangensis Gu et al. 2013,
meat, fish, vegetables and cereals, sewage, humans (oral,
4698VP.
vaginal and intestinal), invertebrate hosts and clinical sources
[17, 169]. Original characteristics of L. songhuajiangensis strains are
described in [176]. The genome size of the type strain is 2.61
The type strain is ATCC 7469 =CCUG 21452 =CIP
T T
A157T=DSM 20021T=NBRC 3425T=JCM 1136T=LMG
6400T=NCAIM B.01147=NCCB 46033T=NCIMB Isolated from traditional sourdough.
6375 =NCTC 12953 = NRRL B-442 =VKM B-574T.
T T T
The type strain is 7–19T=LMG 27191T=NCIMB
Genome sequence accession number: AZCQ00000000. 14832T=CCUG 62990T.
16S rRNA gene accession number: D16552. Genome sequence accession number: RHNR00000000.
16S rRNA gene accession number: HF679038.
Description of Lacticaseibacillus thailandensis
saniviri comb. nov. comb. nov.
Lactobacillus saniviri (sa.ni.vi′ri. L. masc. adj. sanus healthy;
Lacticaseibacillus thailandensis (thai.lan.den’sis. N.L. masc.
L. gen. n. viri, of a man; N.L. gen. n. saniviri, of a healthy
adj. thailandensis, pertaining to Thailand, where the type
man).
strain was isolated).
Basonym Lactobacillus saniviri Oki et al. 2012, 605VP
Basonym Lactobacillus thailandensis Tanasupawat et al. 2007,
Original characteristics of L. saniviri strains are described in 1371VL
Original characteristics of L. thailandensis strains are
described in [161]. The genome size of the type strain is 2.06
Isolated from the faeces of a healthy man and from fermented Mbp. The mol% G+C content of DNA is 53.5.
rice and fish.
Isolated from fermented fish (pla-ra) in Thailand.
The type strain is YIT 12363 T=JCM 17471 T=DSM 24301T.
The type strain is MCH5-2T=BCC 21235T=DSM 22698T=JCM
Genome sequence accession number: JQCE00000000. 13996T=NRIC 0671T.
16S rRNA gene accession number: AB602569. Genome sequence accession number: AYZK00000000.
sharpeae comb. nov. Emended description of
Lacticaseibacillus sharpeae (shar’pe.ae. N.L. gen. n. sharpeae Paralactobacillus
of Sharpe; named for M. Elisabeth Sharpe, an English
Paralactobacillus (Pa.ra.lac.to.ba.cil’lus. Gr. prep. para resem-
bacteriologist).
bling; N.L. masc. n. Lactobacillus a bacterial genus; Paralac-
Basonym Lactobacillus sharpeae Weiss et al. 1981, 266VP tobacillus resembling the genus Lactobacillus).
2814
Cell are Gram-positive, homofermentative, non-motile, non- Basonym: Lactobacillus sakei corrig. Katagiri, Kitahara and
spore-forming rods, usually occurring singly or as pairs. They Fukami 1934, 157 (Approved Lists); emend. Klein et al. 1996
produce both d(−)- and l(+)- lactic acid from glucose. They
L. sakei strains are frequently slightly curved and irregular
produce acid from mannose and salicin but not from lactose,
cells, especially during stationary growth phase. Many of the
melibiose, raffinose, ribose or xylose. They grow at 15 °C but
strains grow also at 2–4 °C, the majority of them produce
not at 45 °C and with 6.5 % NaCl.
l(+)-lactic acid in MRS broth [180].
The type species P. selangorensis was isolated from a Malaysian
Isolated from sauerkraut, fermented plant material, fermented
food ingredient, chili bo [177] and later transferred to the
seafood, cold smoked salmon, fermented or refrigerated meat
genus Lactobacillus, proposing Lactobacillus selangorensis
products, spontaneous sourdoughs, and prepacked finished
comb. nov [178]. Given the data presented in the present
study we adopt the proposal of Leisner et al. [177]. dough [181]. It is used commercially as starter culture for
fermented meats [58, 59].
Two subspecies are recognised: Latilactobacillus sakei subsp.
Description of Paralactobacillus carnosus and Latilactobacillus sakei subsp. sakei.
selangorensis comb. nov.
Paralactobacillus selangoresis (se.lan.gor.en’sis. N.L. masc.
adj. selangorensis, belonging to the province of Selangor, Description of Latilactobacillus sakei
Malaysia); Leisner et al. 2000, Haakensen et al. 2011, 2982VP subsp. carnosus comb. nov
Cells are able to grow on acetate agar and can lower the pH to Latilactobacillus sakei subsp. carnosus (car.no’sus. L. masc. adj.
below 4.15 in LA broth [179]. No growth occurs with 6.5 % carnosus pertaining to meat).
NaCl [177]. The genome size of the type strain is 2.09 Mbp; Basonym: Lactobacillus sakei subsp. carnosus Torriani et al.
the mol% G+C content of DNA is 46. 1996, 1162VP
The species was isolated from a Malaysian food ingredient L. sakei subsp. carnosus was formerly also referred to as Lacto-
called chili bo. bacillus curvatus subsp. melibiosus [182]. Characteristics of
The type strain is ATCC BAA-66T = CCUG 43347T = CIP the species are described in [183]. The genome size of the type
106482T=DSM 13344T = LMG 17710T. strain is 1.99 Mbp. The mol% G+C content of DNA is 41.0.
Genome sequence accession number: JQAZ00000000. Isolated from fermented meat products, vacuum-packaged
meat, sauerkraut, and other fermented plant material.
The type strain is R 14b/aT=LMG 17302T = DSM
15831T=CCUG 31331T = CIP 105422T=JCM 11031T.
Description of Latilactobacillus gen. Genome sequence accession number: AZFG00000000.
nov. 16S rRNA gene accession number: AY204892.
Latilactobacillus (La.ti.lac.to.ba.cil'lus. L. masc. adj. latus wide,
broad; N.L. masc. n. Lactobacillus a bacterial genus name;
N.L. masc. n. Latilactobacillus a widespread lactobacillus).
Description of Latilactobacillus sakei
Species of Latilactobacillus are homofermentative, their mol% subsp. sakei comb. nov.
G+C content is between 40 and 42 and the genome size ranges Latilactobacillus sakei subsp. sakei (sa’ke.i. N.L. gen. n. sakei
from 1.82 to 2.12 Mbp; they produce both d(−)- and l-(+)- of sake).
lactic acid with the exception of L. fuchuensis, which produces
only the l(+)- isomer. Strains in the species lead a free-living Basonym: Lactobacillus sakei subsp. sakei corrig. Katagiri,
lifestyle and are mesophilic; many strains are psychrotrophic Kitahara and Fukami 1934, 157 (Approved Lists); emend.
and grow below 8 °C. L. sakei and L. curvatus have commer- Klein et al., 1996
cial importance as meat starter cultures [58, 59].
L. sakei subsp. sakei strains produce ammonia from arginine
A phylogenetic tree on the basis of 16S rRNA genes of all and acetoin from glucose and they grow in the presence of 10
species in the genus Latilactobacillus is provided in Fig. S6G. % NaCl. The genome size of the type strain is 1.91 Mbp. The
The type species is Latilactobacillus sakei comb. nov.; Latilac-
tobacillus was previously referred to as L. sakei group. Isolated from sake starter, fermented meat products, vacuum-
packaged meat, sauerkraut and other fermented plant mate-
rial, and human faeces [183].
Description of Latilactobacillus sakei The type strain is T.S [K. Kitahara 37]T = ATCC 15521T= LMG
comb. nov. 9468T = DSM 20017T=CCUG 30501T = CIP 103139T=IFO
Latilactobacillus sakei (sa’ke.i. N.L. gen. n. sakei of sake). (now NBRC) 15893T = JCM 1157T.
2815
Genome sequence accession number: AZDN00000000. Basonym: Lactobacillus graminis Beck et al. 1989, 93VP (Effec-
tive publication: Beck et al. 1988, 282)
16S rRNA gene accession number: AM113784.
Strains of this species show a flocculant sediment after 3
days of growth in MRS broth [188]. The genome size of the
Description of Latilactobacillus type strain is 1.84 Mbp. The mol% G+C content of DNA
curvatus comb. nov. is 40.3.
Latilactobacillus curvatus (cur.va′tus. L. masc. adj. curvatus
Isolated from grass silage, meat products, sourdough, gut of
curved).
snail Cornum aspersum and grapes.
Basonym: Lactobacillus curvatus (Bacterium curvatum Troili-
The type strain is G90(1)T = LMG 9825T = DSM 20719T=ATCC
Petersson 1903, 137) Abo-Elnaga and Kandler 1965; Troili-
Petersson 1980 (Approved Lists); emend. Klein et al. 1996 51150T= CCUG 32238T = CIP 105164T=JCM 9503T = NRRL
B-14857T.
Some L. curvatus strains are motile [184]; they occur in pairs,
short chains, and frequently in horseshoe forms. Characteris- Genome sequence accession number: AYZB00000000.
tics of the species are provided by [170, 180, 185]. The genome 16S rRNA gene accession number: AM113778.
of DNA is 42.0.
Isolated from cow dung, fermented and vacuum-packaged Description of Loigolactobacillus gen.
refrigerated meat and fish products, dairy products such nov.
as milk and cheese, fermented plant products like sauer- Loigolactobacillus (Loi.go.lac.to.ba.cil’lus. Gr. masc. n. loigos
kraut, sourdough (including prepacked finished dough destruction, ruin, havoc; N.L. masc. n Lactobacillus a bacte-
and pressed yeast), radish, pickles and kimchi, other plant- rial genus; Loigolactobacillus, a lactobacillus with spoilage
derived materials like honey and from the environmental potential).
fermentation process of corn or grass silage [186].
Cells are are non- motile, non- spore-
forming, Gram-
The type strain is 1T = LMG 9198T = DSM 20019T=LMG
positive, catalase negative rods, found singly and in pairs.
13553T = ATCC 25601T= CCUG 30669T = CIP 102992T=IFO
They are homofermentative and produce both d-(−)- and
(now NBRC) 15884T = JCM 1096T=NRRL B-4562T.
l-(+)-lactic acid isomers. Pentose fermentation is species
Genome sequence accession number: AZDL00000000. specific; most of species produce acid from d-mannose and
16S rRNA gene accession number: AM113777. d-mannitol. The mol% G+C content is between 40.6 and
44.3.
Description of Latilactobacillus species in the genus Loigolactobacillus is provided in Figure
fuchuensis comb. nov. S6H.
Latilactobacillus fuchuensis (fu.chu.en’sis. N.L. masc. adj. fuch-
uensis of Fuchu, the city where this bacterium was originally The type species of the genus is L. coryniformis comb. nov.;
isolated). Loigolactobacillus was previously referred to as L. coryniformis
group.
Basonym: Lactobacillus fuchuensis Sakala et al. 2002, 1153VP
Original characteristics of L. fuchuensis strains are described
in [187]. The genome size of the type strain is 2.12 Mbp. The Description of Loigolactobacillus
mol% G+C content of DNA is 41.8. coryniformis comb. nov.
Isolated from vacuum-packaged refrigerated beef, common Loigolactobacillus coryniformis (co.ry.ni.for′mis. Gr. fem. n.
carp intestine and other seafood products. coryne a club; L. fem. n. forma shape; N.L. masc. adj. coryni-
formis club-shaped).
The type strain is B5M10T = DSM 14340T = CCUG
47133T=JCM 11249T. Basonym: Lactobacillus coryniformis Abo-Elnaga and Kandler
1965, 18 (Approved Lists)
Genome sequence accession number: AZEX00000000.
L. coryniformis cells are short, coccoid rods, frequently
pear-shaped. Pantothenic acid, niacin, riboflavin, biotin and
p-aminobenzoic acid are essential for the growth of all or the
Description of Latilactobacillus majority of the strains tested [170].
graminis comb. nov. Two subspecies are recognized: Loigolactobacillus coryni-
Latilactobacillus graminis (gra’mi.nis. L. gen. n. graminis of formis subsp. coryniformis and Loigolactobacillus coryniformis
grass). subsp. torquens.
2816
Description of Loigolactobacillus aminopeptidase, cystine aminopeptidase, acid phosphatase,

coryniformis subsp. coryniformis naphthol-AS-BI-phosphohydrolase, β-galactosidase,
β-glucosidase and N-acetyl-β-glucosaminidase activities
comb. nov. [189]. The genome size of the type strain is 2.78 Mbp. The
Loigolactobacillus coryniformis subsp. coryniformis (co.ry′ni. mol% G+C content of DNA is 40.7.
for′mis. Gr. n. coryne a club; L. adj. formis shaped; N.L. adj.
coryniformis club-shaped). Isolated from a spoiled lager beer.
Basonym: Lactobacillus coryniformis subsp. coryniformis Abo- The type strain is L-1062=JCM 18665T = LMG 23555T=DSM
Elnaga and Kandler 1965, 18 (Approved Lists) 18080T = L1062T.
L. coryniformis subsp. coryniformis strains produce l-(+) Genome sequence accession number: ASM166367v1.
isomer of the lactic acid at amounts of 15–20 % of total lactic For bioinformatics analysis, the closed genome of strain
acid [170]. The genome size of the type strain is 2.71 Mbp. TMW1.1989 was used: CP014873 (chromosome), CP014874
The mol% G+C content of DNA is 42.9. (plasmid) and CP014875 (second plasmid).
Isolated from silage, cow dung, dairy barn air and sewage, and 16S rRNA gene accession number: AB779648.
from table olives, wheat, pickled vegetable, cheese and ting, a
fermented sorghum porridge.
Description of Loigolactobacillus
The type strain is ATCC 25602T = CIP 103133T=DSM 20001T
= CCUG 30666T=JCM 1164T = LMG 9196T=NRRL B-4391T. bifermentans comb. nov.
Loigolactobacillus bifermentans (bi.fer.men′tans. L. pref.
Genome sequence accession number: AZCN00000000. bi twice; L. pres. part. fermentans leavening; N.L. part. adj.
16S rRNA gene accession number: M58813. bifermentans doubly fermenting).
Basonym: Lactobacillus bifermentans (ex Pette and van
Beynum 1943) Kandler et al. 1983, 896VP
coryniformis subsp. torquens comb. L. bifermentans strains are non-motile irregular rods with
rounded or often tapered ends; clumps are often observed.
nov. Lactic acid is fermented to acetic acid, ethanol, traces of
Loigolactobacillus coryniformis subsp. torquens (tor’quens. L. propionic acid, carbon dioxide and free H2 [190]. The genome
part. adj. torquens, twisting). size of the type strain is 3.14 Mbp. The mol% G+C content
Basonym: Lactobacillus coryniformis subsp. torquens Abo- of DNA is 44.3.
Elnaga and Kandler 1965, 18 (Approved Lists) Isolated from spoiled Edam and Gouda cheeses where it
L. coryniformis subsp. torquens strains exclusively produce forms undesired small cracks [191], from fermented masau
d(−)-lactic acid [170]. The genome size of the type strain is fruits and from Himalayan fermented milk products.
2.78 Mbp. The mol% G+C content of DNA is 42.9. The type strain is N2T = ATCC 35409T = CCUG 32234T=CIP
Isolated from cheese, yaks’ milk cheese, silage and tomato 102811T = DSM 20003T=JCM 1094T = LMG 9845T.
pomace silage. Genome sequence accession number: AZDA00000000.
The type strain is CECT 4129=ATCC 25600T = CCUG 16S rRNA gene accession number: JN175330.
30667T=CIP 103134T = DSM 20004T=JCM 1166T = LMG
9197T=NRRL B-4390T.
Genome sequence accession number: AZDC00000000. Description of Loigolactobacillus
16S rRNA gene accession number: AJ575741. iwatensis comb. nov.
Loigolactobacillus iwatensis (i.wa.ten’sis. N.L. masc. adj. iwat-
ensis of or belonging to Mount Iwate, where the first strains
Description of Loigolactobacillus were isolated).
backii comb. nov. Basonym: Lactobacillus iwatensis Tohno et al. 2013, 3856VP
Loigolactobacillus backii (back′i.i. N.L. gen. n. backii, named L. iwatensis strains are facultatively anaerobic, non-spore-
in recognition of Werner Back, a German microbiologist who
forming and non-motile rods. They are homofermentative;
contributed to the microbiological and technological develop-
they are positive for C4 esterase, leucine aminopeptidase,
ment of brewing).
valine aminopeptidase and acid phosphatase [189]. The
Basonym: Lactobacillus backii Tohno et al. 2013, 3858VP genome size of the type strain is 2.62 Mbp. The mol% G+C
L. backii strains are rod-shaped and occur singly, in pairs
and in chains; they show leucin aminopeptidase, valine Isolated from orchardgrass silage.
2817
The type strain is IWT246T = JCM 18838T = DSM 26942T. Isolated from fermented Chinese cabbage.
Genome sequence accession number: RHNP00000000. The type strain is 187–3T = NCIMB 15172T=CCM 8910T.
16S rRNA gene accession number: AB773428. Genome sequence accession number: RHOE00000000.
jiayinensis comb. nov.
Description of Dellaglioa gen. nov.
Loigolactobacillus jiayinensis (jia.yin.en’sis. N.L. masc. adj.
jiayinensis, pertaining to Jiayin, a county in the Heilongjiang Dellaglioa (Del.la.gli.o’a. N.L. fem. n. Dellaglioa, named after
province of China). Franco Dellaglio, an Italian microbiologist, former chairman
of the Subcommittee on taxonomy of Bifidobacterium, Lacto-
Basonym: Lactobacillus jiayinensis Long and Gu 2019, 2348VP bacillus and related organisms, known for his significant
Cells are non-motile, rod-shaped, singly or in pairs [158]. The research contributions to the taxonomy of the lactic acid
genome size of the type strain is 3.04 Mbp. The mol% G+C bacteria).
content of DNA is 42.6. Strains of Dellaglioa are facultatively anaerobic, psychrophilic,
Isolated from a fermentation Chinese cabbage. nonmotile, homofermentative rod-shaped bacteria. Strains of
the only species included in the genus are psychrotrophic and
The type strain is 257–1T = NCIMB 15166T=CCM 8904T = occur as meat spoilage organism.
LMG 31065T.
The type species is Dellaglioa algida.
Genome sequence accession number: RHOF00000000.
Description of Dellaglioa algida comb.
nov.
Description of Loigolactobacillus Dellaglioa algida (al’gi.da. L. fem. adj. algida cold, referring to
rennini comb. nov. the ability to grow at low temperature).
Loigolactobacillus rennini (ren.ni’ni. N.L. gen. n. rennini, of
Basonym: Lactobacillus algidus Kato et al. 2000, 1148VP
rennet).
Characteristics are described in [193]. The genome size of
Basonym: Lactobacillus rennini Chenoll et al. 2006, 451VP
the type strain is 1.59 Mbp, the mol % G+C content of DNA
L. rennini strains are non-motile rods, found singly and in is 36.
pairs. They grow at pH 3.7, 4.5 and 8.0, and in medium with
Isolated as spoilage organisms from refrigerated beef and
5 and 10 % (w/v) NaCl [192]. The genome size of the type
pork meat.
The type strain is M6A9T= JCM 10491T= LMG 19872T=DSM
Isolated from rennet and are associated with cheese spoilage.
15638T= IP 106688T.
The type strain is 1–7T = CECT 5922T=DSM 20253T = JCM
14279T. Genome sequence accession number: AZDI00000000.
Genome sequence accession number: AYYI00000000. 16S rRNA gene accession number: AB033209.
16S rRNA gene accession number: LC258150.

Description of Liquorilactobacillus
Description of Loigolactobacillus gen. nov.
zhaoyuanensis comb. nov. Liquorilactobacillus (Li.quo.ri.lac.to.ba.cil’lus L. masc. n.
liquor, liquid; N.L. masc. n. Lactobacillus a bacterial genus;
Loigolactobacillus zhaoyuanensis (zhao.yuan.en’sis. N.L. masc. N.L. masc. n. Liquorilactobacillus a lactobacillus from
adj. zhaoyuanensis, pertaining to Zhaoyuan, a county in the liquids, referring to the isolation of most species from liquids
Heilongjiang province of China). including water, plant sap, and alcoholic beverages).
Basonym: Lactobacillus zhaoyuanensis Long and Gu, 2019, Homofermentative, their mol% G+C content is between
2348VP 33.9 and 40.0; most of the species are motile with the excep-
Cells are non-motile rods, usually singly or in pairs. They tion of L. cacaonum, L. hordei, L. mali. They contain meso-
grow at 15–33°C and pH 5–10. Both isomers of lactic acid are diaminopimelic acid. Liquorilactobacillus species were mostly
produced (93 % d(−)-lactate and 7 % l(+)-lactate) [158]. The isolated from fermented plant materials including alcoholic
genome size of the type strain is 2.70 Mbp. The mol% G+C fermentations and water kefir. Many strains of Liquorilacto-
content of DNA is 42.7. bacillus produce dextran from sucrose [194].
2818
A phylogenetic tree on the basis of 16S rRNA genes of all Description of Liquorilactobacillus
species in the genus Liquorilactobacillus is provided in Fig. cacaonum comb. nov.
S6I.
Liquorilactobacillus cacaonum (ca.ca.o′num. N.L. gen. pl. n.
The type species is Liquorilactobacillus mali comb. nov.; cacaonum, of cacao beans).
Liquorilactobacillus species were previously considered part
Basonym: Lactobacillus cacaonum De Bruyne et al. 2009, 11VP
of the L. salivarius group.
L. cacaonum cells are non-motile small rods, they grow at
37C in MRS broth at pH 3.9; no growth is observed in MRS
Description of Liquorilactobacillus supplemented with NaCl [198]. The genome size of the type
mali comb. nov. strain is 1.92 Mbp. The mol% G+C content of DNA is 33.9.
Liquorilactobacillus mali (ma’li. L. gen. n. mali, of an apple). Isolated from cocoa fermentation.
Basonym: Lactobacillus mali Carr and Davies 1970, 769 The type strain is R-34119T = DSM 21116T = LMG 24285T.
(Approved Lists); emend. Kaneuchi et al. 1988, 272 Genome sequence accession number: AYZE00000000.
L. mali cells are non-motile or weakly motile with a few peri- 16S rRNA gene accession number: AM905389.
trichous flagella. Liquid cultures are turbid after few days,
with subsequent clearing and sediment. They produce acetoin
and dextran; malic acid is decomposed to lactic acid and CO2. Description of Liquorilactobacillus
Most strains have a pseudocatalase activity when they grow capillatus comb. nov.
on MRS agar with 0.1 % (w/v) glucose [195, 196]. The genome
Liquorilactobacillus capillatus (ca.pil.la’tus. L. masc. adj. capil-
size of the type strain is 2.59. The mol% G+C content of DNA latus, hairy, referring to the characteristic of having long,
is 36.1. hairy flagella).
Isolated from wine must, fermenting cider, fermented Basonym: Lactobacillus capillatus Chao et al. 2008, 2557VP
molasses, water kefirs, cocoa bean fermentations and table
olives. L. capillatus cells are motile by means of peritrichous flagella;
they grow at pH 4.0 and pH 8.0 but they do not grow at 8 %
The type strain is ATCC 27053T = CCUG 30141T=CCUG NaCl [199]. The genome size of the type strain is 2.24 Mbp.
32228T = CIP 103142T=DSM 20444T = JCM 1116T=LMG The mol% G+C content of DNA is 37.6.
6899T = NBRC 102159T=NCIB (now NCIMB) 10560T =
NRRL B-4563T = VKM B-1600T. Isolated from fermented brine used for stinky tofu production.
Genome sequence accession number: AYYH00000000. The type strain is YIT 11306T = BCRC 17811T=DSM 19910T
= JCM 15044T.
Genome sequence accession number: AZEF00000000.
aquaticus comb. nov.
Liquorilactobacillus aquaticus (a.qua′
ti.
cus. L. masc. adj.
aquaticus, living, growing or found in water, or an aquatic
ghanensis comb. nov.
environment). Liquorilactobacillus ghanensis (gha.nen′sis. N.L. masc. adj.
ghanensis, pertaining to Ghana, where the species was first
Basonym: Lactobacillus aquaticus Mañes-Lázaro et al. 2009, isolated).
2217VP
Basonym: Lactobacillus ghanensis Nielsen et al. 2007, 1471VP
L. aquaticus cells are motile rods, which grow at pH 4.5–8.0
but not at pH 3.3. There is no growth in 5 or 10 % NaCl. L. ghanensis cells are motile with peritrichous flagella, and
colonies have slightly uneven edges after 3–4 days of anaerobic
The species is described in [197]. The genome size of the growth. They do not grow at pH 8.0, nor at 6.5 % NaCl [200].
type strain is 2.41 Mbp. The mol% G+C content of DNA The genome size of the type strain is 2.61 Mbp. The mol%
is 37.4. G+C content of DNA is 37.1.
Isolated from a eutrophic freshwater pond. Isolated from cocoa fermentations.
The type strain is IMCC1736 = CECT 7355 = DSM
T T
The type strain is L489T = CCUG 53453T = DSM 18630T=JCM
21051T=JCM 16869T. 15611T.
Genome sequence accession number: AYZD00000000. Genome sequence accession number: AZGB00000000.
16S rRNA gene accession number: DQ664203. 16S rRNA gene accession number: DQ523489.
2819
Description of Liquorilactobacillus The type strain is 59bT=CECT 7334T=DSM 19972T=JCM

hordei comb. nov. 18036T.
Liquorilactobacillus hordei (hor’de.i. L. gen. n. hordei, from Genome sequence accession number: AZEH00000000.
barley). 16S rRNA gene accession number: AY681127.
Basonym: Lactobacillus hordei Rouse et al. 2008, 2016VP
L. hordei cells are non-motile rods, they grow at pH 4 and 8 Description of Liquorilactobacillus
but not at pH 3 and 9. They produce bacteriocins [201]. The satsumensis comb. nov.
content of DNA is 34.8. Liquorilactobacillus satsumensis (sat.su.men′sis. N.L. masc.
adj. satsumensis, pertaining to Satsuma, old name for the
Isolated from malted barley, water kefirs and in Turkish tradi- southern part of Kyushu in Japan, from where the type strain
tional fermented gilaburu fruit juice. was isolated).
The type strain is UCC128T = DSM 19519T = JCM Basonym: Lactobacillus satsumensis Endo and Okada 2005,
16179T=LMG 24241T. 85VP
Genome sequence accession number: AZDX00000000. L. satsumensis cells are motile rods with peritrichous flagella.
Growth is observed in MRS broth at pH 3.5 containing 5 %
16S rRNA gene accession number: EU074850.
(w/v) NaCl but not with 10 % (v/v) ethanol. Dextran is formed
from sucrose [204]. The genome size of the type strain is 2.65
nagelii comb. nov. Isolated from mashes of shochu, a traditional Japanese
distilled spirit made from fermented rice and other starchy
Lacitilactobacillus nagelii (na.gel’i.i. N.L. gen. n. nagelii, of materials.
Nagel, after Charles W. Nagel, Washington State University,
USA, for his contributions to the science of wines). The type strain is DSM 16230T = JCM 12392T=NRIC 0604T.
Basonym: Lactobacillus nagelii Edwards et al. 2000, 700VP Genome sequence accession number: AZFQ00000000.
L. nagelii cells are motile rods, they grow in MRS broth with 16S rRNA gene accession number: AB154519.
5%(w/v) NaCl (pH 4.5) at 25 °C; both citrate and malate are
utilized in the presence of glucose and dextran is formed from
sucrose [202]. The genome size of the type strain is 2.50 Mbp. Description of Liquorilactobacillus
The mol% G+C content of DNA is 36.7. sicerae comb. nov.
Isolated from partially fermented wine, spontaneous cocoa Liquorilactobacillus sicerae (si’ce.rae. L. gen. n. sicerae of cider).
bean fermentations, water kefirs, fermented cassava food and
Basonym: Lactobacillus sicerae Puertas et al. 2014, 2954VP
silage fermentation of fruit residues.
L. sicerae cells are motile rods with polar flagella, they produce
The type strain is LuE10T = ATCC 700692T = CCUG
exopolysaccharides from sucrose and both malic and citric
43575T=DSM 13675T = JCM 12492T.
acids are utilized in the presence of glucose [205]. The genome
Genome sequence accession number: AZEV00000000. size of the type strain is 2.49 Mbp. The mol% G+C content
of DNA is 37.5.
16S rRNA gene accession number: Y17500.
Isolated from spoiled apple cider.
The type strain is CUPV261T = CECT 8227T = KCTC 21012T.
oeni comb. nov. Genome sequence accession number: PRJEB5073.
Liquorilactobacillus oeni (oe′ni. Gr. masc. n. oinos, wine; N.L. 16S rRNA gene accession number: HG794492.
gen. n. oeni, of wine).
Basonym: Lactobacillus oeni Mañes-Lázaro et al. 2009, 2013VP Description of Liquorilactobacillus
L. oeni cells are mostly motile and they grow at pH 4.5–8.0 sucicola comb. nov.
but not at pH 3.3 and with 10 % ethanol. l-Malic acid is trans- Liquorilactobacillus sucicola (su.ci.co’la. L. n. sucus, juice, sap;
formed into l(+)-lactic acid. They produce exopolysaccharide L. suff. -cola (from L. masc. or fem. n. incola), inhabitant,
from sucrose [203]. The genome size of the type strain is 2.12 dweller; N.L. masc. n. sucicola, a sap-dweller).
Basonym: Lactobacillus sucicola Irisawa and Okada 2009,
Isolated from Bobal wine. 2664VP
2820
L. sucicola cells are motile rods by means of peritrichous Description of Ligilactobacillus gen.
flagella; they do not grow in GYP broth containing 5 % (w/v) nov.
NaCl [206]. The genome size of the type strain is 2.46 Mbp.
The mol% G+C content of DNA is 38.5. Ligilactobacillus (Li.gi.lac.to.ba.cil'lus. L. v. ligare to tie, unite;
N.L. masc. n. Lactobacillus a bacterial genus name; N.L. masc.
Isolated from the sap of an oak (Quercus sp.). n. Ligilactobacillus a lactobacillus with a host-associated life
The type strain is NRIC 0736T = DSM 21376T=JCM 15457T. style).
Genome sequence accession number: AYZF00000000. Species of Ligilactobacillus are homofermentative, their mol%
G+C content is between 32.5 and 43.3. Several Ligilactobacillus
16S rRNA gene accession number: AB433982. species include strains that are motile. Most Ligilactobacillus
species have been isolated from animals and humans and are
adapted to vertebrate hosts. Several strains of Ligilactobacillus
Description of Liquorilactobacillus express urease, this enzyme is the most powerful bacterial tool
uvarum comb. nov. to withstand gastric acidity; in lactobacilli, urease activity is
Liquorilactobacillus uvarum (u.va′rum. L. gen. pl. n. uvarum associated with a vertebrate host-adapted lifestyle [210, 211].
of grapes). Several Ligilactobacillus species also commonly occur in
fermented foods and are used commercially as starter cultures
Basonym: Lactobacillus uvarum Mañes-Lázaro et al. 2008, or probiotic cultures.
2129VL
L. uvarum cells are motile rods; they grow at pH 4.5 and 8 but species in the genus Ligilactobacillus is provided in Figure S6J.
not at pH 3.3. They produce exopolysaccharide from sucrose
[207]. The genome size of the type strain is 2.69 Mbp. The The type species is Ligilactobacillus salivarius comb. nov.;
mol% G+C content of DNA is 36.9. Ligilactobacillus was previously referred to as L. salivarius
group.
Isolated from from Bobal grape musts.
The type strain is 8T = Lb8T= CECT 7335T = DSM 19971T=JCM
16870T. Description of Ligilactobacillus
Genome sequence accession number: AZEG00000000. salivarius comb. nov.
Ligilactobacillus salivarius (sa.li.va’ri.us. L. masc. adj. salivarius
salivary).
Basonym: Lactobacillus salivarius Rogosa et al. 1953, 691
Description of Liquorilactobacillus (Approved Lists); emend. Li et al. 2006
vini comb. nov.
L. salivarius strains require pteroylglutamic acid and niacin
Liquorilactobacillus vini (vi’ni. L. gen. n. vini of wine).
for good growth. Riboflavin is also markedly stimulatory.
Basonym: Lactobacillus vini Rodas et al. 2006, 516VP Good growth does not take place in media unless polyoxy-
The cells are motile rods, they do not produce exopolysac- ethylene sorbitan monooleate (Tween 80) is added [61, 212].
charide from sucrose, but they utilize citric and malic acids The genome size of the type strains is 1.98 Mbp. The mol%
[208]. L. vini was the first organism for which metabolism of G+C content of DNA is 37.5.
pentoses via the pentose phosphate pathway to lactate as sole Isolated from the mouth and intestinal tract of humans, cats,
end product was described [18]. The metabolism for homo- hamsters, chickens and swine, and from dairy products; the
fermentative metabolism of pentoses was initially described species shows no adaptation to specific hosts [213].
for strains later classified as L. vini [18, 209]. The genome
size of the type strain is 2.24 Mbp. The mol% G+C content The type strain is H066T = ATCC 11741T= CCUG 31453T
of DNA is 37.5. = CIP 103140T=DSM 20555T = JCM 1231T=LMG 9477T =
NRRL B-1949T.
Isolated from fermenting Spanish grape must and from
bioethanol industrial processes in different distilleries of Genome accession number: AYYT00000000.
Brazil. 16S rRNA gene accession number: AF089108.
The type strain is Mont 4 = CECT 5924 = DSM 20605 =JCM
T T T
14280T.
Description of Ligilactobacillus
Genome sequence accession number: AYYX00000000.
acidipiscis comb. nov.
16S rRNA gene accession number: AJ576009. Ligilactobacillus acidipiscis (a.ci.di.pis’cis. L. masc. adj. acidus
sour; L. n. piscis fish; N.L. gen. n. acidipiscis of a sour fish, an
isolation source of strains of this species).
2821
Basonym: Lactobacillus acidipiscis Tanasupawat et al. 2000, Isolated from dental plaques and intestines of animals.
1481VP. The species includes strains previously designated as
The type strain is 535T = ATCC 35046T=CCUG 33906T =
L. cypricasei [214]
CIP 103152T=DSM 20602T = IFO (now NBRC) 15882T =
L. acidipiscis strains grow in 10 % NaCl; some strains grow in JCM 5670T=LMG 9843T = NCIMB 13278T (formerly NCDO
the presence of 12 % NaCl. They show no reaction in litmus 2425T)=NRRL B-14176T.
milk and do not form slime from sucrose. Niacin and calcium
pantothenate are required for growth [215]. The genome size Genome sequence accession number: AYYW00000000.
of the type strain is 2.33 Mbp. The mol% G+C content of 16S rRNA gene accession number: AB326350.
DNA is 39.1.
Isolated from fermented fish (pla-ra and pla-chom) in Thai-
land but also found in dairy products, soy sauce mash, table Description of Ligilactobacillus
olives, sake starter, tropical grasses, forage crops, bee pollen, apodemi comb. nov.
and Chinese DaQu, a saccharification starter for production Ligilactobacillus apodemi (a.po.de′mi. N.L. gen. n. apodemi, of
of vinegar and liquor from cereals. Apodemus speciosus, the field mouse from which the organism
The type strain is FS60-1T = CCUG 46556T = CIP was first isolated).
106750T=DSM 15836T = HSCC 1411T=JCM 10692T = NBRC Basonym: Lactobacillus apodemi Osawa et al. 2006, 1695VP
102163T=NRIC 0300T = PCU 207T=TISTR 1386T.
L. apodemi strains are non-motile, they are tannase-positive
Genome sequence accession number: AZFI00000000. and they produce gallic acid from tannic acid but they do
16S rRNA gene accession number: AB023836. not convert gallic acid to pyrogallol [217]. The genome size
of the type strain is 2.10 Mbp. The mol% G+C content of
DNA is 38.6.
Description of Ligilactobacillus agilis
Isolated from the faeces of a wild mouse.
comb. nov.
Ligilactobacillus agilis (a’gi.lis. L. masc. adj. agilis agile, motile). The type strain is ASB1T = CIP 108913T = DSM 16634T=JCM
16172T.
Basonym: Lactobacillus agilis Weiss et al. 1982, 266VP (Effec-
tive publication: Weiss et al. 1981, 252) Genome sequence accession number: AZFT00000000.
L. agilis strains are motile with peritrichous flagella; motility 16S rRNA gene accession number: AJ871178.
is easily demonstrated in MRS broth [175]. The genome size
of the type strain is 2.06 Mbp. The mol% G+C content of
DNA is 41.7. Description of Ligilactobacillus
Isolated from municipal sewage; L. agilis is the dominant araffinosus sp. nov.
species in the pigeon crops and it is also found in the gut Ligilactobacillus araffinosus (a.raf.fi.no’sus. Gr. pref. a not; N.L.
and cecum of birds, human gut and vagina, porcine intestinal masc. adj. raffinosus of raffinose; N.L. masc. adj. araffinosus,
mucin and fermented food products such as masau fruits, not fermenting raffinose).
Nigerian ogi, and cheese. Basonym: Lactobacillus aviarius subsp. araffinosus Fujisawa
The type strain is CIP 101264T = CCUG 31450T=DSM 20509T et al. 1985, 223VP; Fujisawa et al. 1986, (Effective publication:
= JCM 1187T=LMG 9186T = NRRL B-14856T. Fujisawa et al., 1984, 419).
Genome sequence accession number: AYYP00000000. Placement of L. araffinosus in a separate species is supported
by ANI values and digital DNA–DNA hybridization [10]. The
availability of the genome sequences of the type strains of
both subspecies revealed that they are only distantly related
Description of Ligilactobacillus with an ANI value of 88.98%, substantially below the 95–96
% threshold recognised for the species level. Furthermore,
animalis comb. nov.
their core nucleotide identity (CNI) [10] is 91.1, which is
Ligilactobacillus animalis (a.ni.ma′lis. L. gen. n. animalis, of below the 94 % threshold at which two strains are allocated
an animal). to the same species. Finally, their in silico DDH value is 37.80,
Basonym: Lactobacillus animalis Dent and Williams 1983, which additionally supports that these two subspecies should
439VP; Effective publication: Dent and Williams 1982, 384 be recognised as different species [10].
L. animalis strains grow at 45 °C, cells in exponential growth Cells are Gram- positive, non-motile, non-spore-forming
phase occur singly or in pairs [216]. The genome size of the rods with rounded ends, usually occurring singly or in short
type strain is 1.89 Mbp. The mol% G+C content of DNA is chains. L. araffinosus strains ferment trehalose and cellobiose,
41.1 they do not produce acid from galactose, lactose, melibiose
2822
and raffinose [51]. The genome size of the type strain is 1.48 Strains of this species are non-motile rods and some of them
Mbp. The mol% G+C content of DNA is 38.1. contain filamentous cells [219]. The genome size of the type
Isolated from the intestine and faeces of birds.
Isolated from faeces of horses.
The type strain is ML2T = ATCC 43235T= DSM 20653T =
CCUG 32231T=CIP 103145T = JCM 5667T The type strain is YIT 0455T = ATCC BAA-261T = DSM
15833T = CCUG 47129T=JCM 10991T.
Genome sequence accession number: AYYZ00000000.
Genome accession number: AZFH00000000.
Description of Ligilactobacillus faecis
aviarius comb. nov.
comb. nov.
Ligilactobacillus aviarius (a.vi.a’ri.us. L. masc. adj. aviarius,
pertaining to birds). Ligilactobacillus faecis (fae’cis. L. gen. n. faecis of faeces).
Basonym: Lactobacillus aviarius Fujisawa et al. 1985, 223VP; Basonym: Lactobacillus faecis Endo et al. 2013, 4505VP.
Fujisawa et al. 1986; Effective publication: Fujisawa et al., Original characteristics of the species described in [220]. The
1984, 419 mol% G+C content of DNA is 41.1.
The cells are non-motile rods with rounded ends, occurring Isolated from faeces of a jackal (Canis mesomelas) and
singly or in short chains. The final pH of glucose broth is raccoons (Procyron lotor).
3.9–4.0. This species is strictly anaerobic [51]. The genome
size of the type strain is 1.68 Mbp. The mol% G+C content The type strain is AFL13-2T = JCM 17300T = DSM 23956T.
of DNA is 40.1. Genome accession number: not available at time of publication
Isolated from the intestine and faeces of birds. 16S rRNA gene accession number: AB812750.
The type strain is 75 = ATCC 43234 =DSM 20655 = CCUG
T T T
32230T=CIP 103144T = JCM 5666T=LMG 10753T = NBRC

102162T. Description of Ligilactobacillus
Genome sequence accession number: AYZA00000000.
hayakitensis comb. nov.
Ligilactobacillus hayakitensis (ha.ya.ki.ten′sis. N.L. masc. adj.
16S rRNA gene accession number: M58808. hayakitensis, of Hayakita, the name of the area where the
bacterium was originally isolated).
Basonym: Lactobacillus hayakitensis Morita et al. 2007, 2838VP
Description of Ligilactobacillus ceti
Original characteristics of the species are described in [221].
comb. nov.
Ligilactobacillus ceti (ce′ti. L. gen. n. ceti, of a whale). G+C content of DNA is 34.1.
Basonym: Lactobacillus ceti Vela et al. 2008, 893VP Isolated from the faeces of a thoroughbred as predominant
Original characteristics of the species are described in [218]. species in the intestinal microbiota.
The genome size of the type strain is 1.40 Mbp. The mol% The type strain is KBL13T = DSM 18933T = JCM 14209T.
Genome accession number: AZGD00000000.
Isolated from the lungs of a beaked whale.
The type strain is 142–2T = CCUG 53626T=DSM 22408T =
CECT 7185T=JCM 15609T.
Genome accession number: JQBZ00000000. Description of Ligilactobacillus
16S rRNA gene accession number: AM292799. murinus comb. nov.
Ligilactobacillus murinus (mu.ri’nus. L. adj. murinus of mice).
Basonym: Lactobacillus murinus Hemme et al., 1982, 384VP.
Description of Ligilactobacillus equi
comb. nov. L. murinus strains are non-motile rods which slowly ferment
ribose and arabinose. l-LDH is activated by FDP and Mn2+.
Ligilactobacillus equi (e′qui. L. gen. n. equi, of the horse).
They do not hydrolyse urea and hippurate; they decarboxy-
Basonym: Lactobacillus equi Morotomi et al. 2002, 214VP late malate. Riboflavin is a required growth factor [222]. The
2823
genome size of the type strain is 2.20 Mbp. The mol% G+C 16S rRNA gene accession number: AB326354.
Isolated from the intestinal tract of mice and rats and from Description of Ligilactobacillus
sourdough.
saerimneri comb. nov.
The type strain is 313T = ATCC 35020T=CCUG 33904T = Ligilactobacillus saerimneri (sae.rim’ne.ri. N.L. gen. masc. n.
CIP 104818T=CNRZ 220T = DSM 20452T=IFO (now NBRC) saerimneri of Saerimner, a pig occurring in Nordic mythology,
14221T = JCM 1717T = LMG 14189T. because the organism was isolated from pigs).
Genome accession number: AYYN00000000. Basonym: Lactobacillus saerimneri Pedersen and Roos 2004,
16S rRNA gene accession number: AJ621554. 1367VP
Strains of this species grows aerobically in MRS agar but
at a lower rate compared to anaerobic growth. They do not
Description of Ligilactobacillus hydrolize aesculin [226]. The genome size of the type strain is
pobuzihii comb. nov. 1.69 Mbp. The mol% G+C content of DNA is 42.6.
Ligilactobacillus pobuzihii (po.bu.zi’hi.i. N.L. gen. n. pobuzihii Isolated from pig faeces. The habitat of L. saerimneri is the
referring to the isolation of the type strain from pobuzihi, intestines of pigs, the human gut and vagina and the cecum
fermented cummingcordia). of chicken.
Basonym: Lactobacillus pobuzihii Chen et al. 2010, 1916VP The type strain is GDA154T = CCUG 48462T = DSM
16049T=JCM 15955T = LMG 22087T.
Original characteristics of the species are described in [223].
The genome size of the type strain is 2.35 Mbp. The mol% Genome accession number: AZFP00000000.
Isolated from pobuzihi, fermented cummincordia, fermented
fish, and traditional vinegar.
The type strain is E100301T = RIFY 6501T = JCM 18084T=KCTC salitolerans comb nov.
13174T = NBRC 103219T.
Ligilactobacillus salitolerans (sa.li.to’le.rans. L. masc. n. sal salt;
Genome accession number: JQCN00000000. L. pres. part. tolerans tolerating; N.L. part. adj. salitolerans
16S rRNA gene accession number: AB326358. salt tolerating).
Basonym: Lactobacillus salitolerans Tohno et al. 2019, 967VP
Characteristics of the species [227] are based on the descrip-
Description of Ligilactobacillus tion of one strain. The genome size of the type strain is 2.30
ruminis comb. nov. Mbp. The mol% G+C content of DNA is 41.7.
Ligilactobacillus ruminis (ru’mi.nis. N.L. gen. n. ruminis of
Isolated from spent mushrooms substrates.
rumen).
The type strain is YK43T = JCM 31331T= DSM 103433T.
Basonym: Lactobacillus ruminis Sharpe et al. 1973, 47
(Approved Lists) Genome accession number: BFFP01000000.
Strains of L. ruminis are motile by peritrichous flagella, anaer- 16S rRNA gene accession number: LC127508.
obic, and grow on surface only under reduced oxygen pres-
sure; growth in liquid media is supported with the addition
of cysteine–HCl. Strains isolated from sewage are nonmotile Description of Lactiplantibacillus
and do not grow at 45 °C [224]. The genome size of the type gen. nov.
strain is 2.01 Mbp. The mol% G+C content of DNA is 43.4. Lactiplantibacillus (Lac.ti.plan.ti.ba.cil'lus. L. neut. n. lactis
Isolated from rumen of cow and from sewage; also commonly milk; L. fem. n. planta plant, referring to the plantarum-group
found in the gut of humans, horses and pigs and bovine lactobacilli; L. masc. n. bacillus a rod; N.L. masc. n. Lacti-
uterus. Genomic analyses indicated a differentiation of plantibacillus a milk derived rodlet from the (Lactobacillus)
specific phylogenetic lineages of the species to specific verte- plantarum group.
brate hosts [225]. Lactiplantibacillus species are Gram-positive, non-spore-
forming, homofermentative and non-motile rods. Lactiplan-
The type strain is RFI =ATCC 27780 = CCUG 39465 =CIP
T T T
tibacillus species ferment a wide range of carbohydrates; most
103153T = DSM 20403T=JCM 1152T = LMG 10756T=NBRC
species metabolise phenolic acids by esterase, decarboxylase
102161T = NRRL B-14853T.
and reductase activities. Lactiplantibacillus plantarum is atyp-
Genome accession number: AYYL00000000. ical for its pseudocatalase activity and reduction of nitrate. For
2824
discrimination between the two subspecies of L. plantarum, Basonym: Lactobacillus plantarum subsp. plantarum Bringel
sequencing of the recA and cpn60 genes or AFLP profiling is et al. 2005, 1633VP
necessary [228, 229]. The mol % G+C content of DNA ranges
The description is that of the species [2, 231]. The genome
between 42.9 and 48.7.
Lactiplantibacillus species are isolated from many different of DNA is 44.2.
fermented foods including fermented vegetables, meats, dairy
Isolated from dairy products and dairy environments, silage,
products, and fermented cereals [58, 59], but they are also
sauerkraut, pickled vegetables, sourdough, cow dung, the
found in insect-associated habitats or as temporary residents
human mouth, intestinal tract and stools, and from sewage.
of vertebrate intestinal microbiota and are characterized by
a nomadic behaviour [230]. L. plantarum has been widely The type strain is ATCC 14917T = CCUG 30503T=CIP
used as a model species for metabolic, ecological, and genetic 103151T = DSM 20174T=IFO (now NBRC) 15891T = JCM
studies in lactobacilli. L. plantarum is of commercial impor- 1149T=LMG 6907T = NCIMB 11974T=NRRL B-4496T.
tance as starter culture for multiple food fermentations, and
Genome sequence accession number: AZEJ00000000.
is applied as probiotic culture.
16S rRNA gene accession number: AJ965482.
species in the genus Lactiplantibacillus is provided in Figure
S6K.
Description of Lactiplantibacillus
The type species is Lactiplantibacillus plantarum comb. nov.; plantarum subsp. argentoratensis
Lactiplantibacillus was previously referred to as L. plantarum
group. comb. nov
Lactiplantibacillus plantarum subsp. argentoratensis (ar.gen.
to.ra.ten′sis. N.L. masc. adj. argentoratensis, of or pertaining
Description of Lactiplantibacillus to Argentoratus, the Roman name of the City of Strasbourg
plantarum comb. nov. in Alsace, France).
Lactiplantibacillus plantarum (plan.ta’rum. L. gen. pl. n. plan- Basonym: Lactobacillus plantarum subsp. argentoratensis
tarum of plants). Bringel et al. 2005, 1633VP
Basonym: Lactobacillus plantarum Bergey et al. 1923, 250 Strains of this species differ from L. plantarum subsp. plan-
(Approved Lists) tarum strains due to the absence of melezitose fermentation
Previously designated as Streptobacterium plantarum [2]. [229]. The genome size of the type strain is 3.20 Mbp. The
Cells are non-motile rods occurring singly, in pairs, or in mol% G+C content of DNA is 45.
short chains. Some strains reduce nitrate in limited glucose Isolated from starchy food, fermenting food of plant origin,
concentration and pH 6.0 or higher. Some strains exhibit timothy, orchardgrass and elephant grass silage, fermented
pseudocatalase activity, or true catalase when haem is Uttapam batter, fermented idli batter.
present. They require calcium pantothenate and niacin to
grow [2, 231]. The type strain is DKO 22T = CCUG 50787T=CIP 108320T =
DSM 16365T=JCM 16169T.
The species has a nomadic lifestyle; it is a dominant member
of the microbiota in spontaneous vegetable and olive fermen- Genome sequence accession number: AZFR00000000.
tations and also occurs in sourdough, dairy fermentations, 16S rRNA gene accession number: AJ640078.
and fermented meats [58, 59]. L. plantarum contributes to
spoilage of beer and wine. L. plantarum also is part of the
microbiota of insects [232], and is isolated from the human Description of Lactiplantibacillus
intestinal tract, particularly the oral cavity.
daoliensis comb. nov.
Two subspecies are recognized: Lactiplantibacillus plantarum Lactiplantibacillus daoliensis (dao.li.en’sis. N.L. masc. adj.
subsp. plantarum and Lactiplantibacillus plantarum subsp. daoliensis, pertaining to Daoli, a district in Harbin city,
argentoratensis. China).
Basonym: Lactobacillus daoliensis Liu and Gu 2019, 3258VP
Characteristics are similar to L. pingfangensis but the strain
plantarum subsp. plantarum comb. does not grow at 37 °C [233]. The genome size of the type
nov. strain is 2.63 Mbp. The mol% G+C content of DNA is 43.7.
Lactiplantibacillus plantarum subsp. plantarum (plan.ta’rum.
L. gen. pl. n. plantarum of plants).
The type strain is 116-
1AT=LMG 31171T = NCIMB
15181T=CCM 8934T.
2825
Genome sequence accession number: BJDH00000000. Isolated from cocoa bean heap fermentation, fermented
grapes, and fermented cereals.
The type strain is R-34115T = DSM 21115T = LMG 24284T.
Genome sequence accession number: AYGX00000000.
daowaiensis comb. nov. 16S rRNA gene accession number: AM905388.
Lactiplantibacillus daowaiensis (dao.wai.en’sis. N.L. masc. adj.
daowaiensis, pertaining to Daowai, a district in Harbin city, Description of Lactiplantibacillus
China). herbarum comb. nov.
Basonym: Lactobacillus daowaiensis Liu and Gu 2019, 3258VP Lactiplantibacillus herbarum (
her.
ba’rum. L. gen. pl. n.
herbarum, of herbs).
Characteristics are similar to L. pingfangensis but the strain
does not grow at 37 °C [233]. The genome size of the type Basonym: Lactobacillus herbarum Mao et al. 2015, 4685VP
strain is 2.85 Mbp. The mol% G+C content of DNA is 44.0. Cells are non-motile rods, usually singly, in pairs or in short
Isolated from fermented Chinese cabbage. chains. The cell wall contains meso-diaminopimelic acid,
alanine, glutamic acid, galactose and an unidentified sugar.
The type strain is 203–3T =LMG 31172T=NCIMB 15183T = They produce acetoin from pyruvate. L. herbarum can be
CCM 8933T. distinguished from related species on the basis of sucrose
Genome sequence accession number: BJDJ00000000. fermentation (it does not ferment sucrose) and growth
temperature (it cannot grow at 37 °C) [234]. The genome
16S rRNA gene accession number: LC438517. size of the type strain is 2.90 Mbp. The mol% G+C content
of DNA is 43.5.
Description of Lactiplantibacillus Isolated from fermented radish.

dongliensis comb. nov. The type strain is TCF032-E4T = CCTCC AB2015090T = DSM
Lactiplantibacillus dongliensis (dong.li.en’sis. N.L. masc. adj. 100358T.
dongliensis pertaining to a district in Harbin, China). Genome sequence accession number: LFEE00000000.
Basonym: Lactobacillus dongliensis Liu and Gu 2019, 3259 VP
16S rRNA gene accession number: NR_145899.
Characteristics are similar to L. pingfangensis but the type
strain of L. dongliensis also f.erments ribose [233]. The Description of Lactiplantibacillus
genome size of the type strain is 3.11 Mbp. The mol% G+C modestisalitolerans comb. nov.
content of DNA is 44.7. Lactiplantibacillus modestisalitolerans (mo.des.ti.sa.li.to’le.
rans. L. masc. adj. modestus moderate; L. masc. n. sal salis
salt; L. pres. part. tolerans tolerating; N.L. part. adj. modestis-
The type strain is 218–3T = LMG 31173T=NCIMB 15184T = alitolerans moderately salt tolerating).
CCM 8932T.
Basonym: Lactobacillus modestisalitolerans Miyashita et al.
Genome sequence accession number: BJDK00000000 2015, 2489VP
16S rRNA gene accession number: LC438518. Cells are non- motile, non- spore-forming, facultatively
anaerobic rods. They grow at 15–40 °C, pH 3.5–7.5, and 0–9 %
NaCl. They produce both lactic acid isomers. They hydrolyse
aesculin, are tellurite and bile-aesculin tolerant and convert
Description of Lactiplantibacillus arginine to ornithine [235]. The genome size of the type strain
fabifermentans comb. nov. is 2.75 Mbp. The mol% G+C content of DNA is 48.6.
Lactiplantibacillus fabifermentans (fa.bi.fer.men’tans. L. fem. Isolated from Pla-som (fermented fish).
n. faba a bean; L. pres. part. fermentans fermenting; N.L. part.
adj. fabifermentans fermenting beans). The type strain is NB446T = NBRC 107235T = BCC 38191T.
Basonym: Lactobacillus fabifermentans De Bruyne et al. 2009, Genome sequence accession number: BJEA00000000.
10VP 16S rRNA gene accession number: AB907192.
Cells are non-motile long rods, usually singly, in pairs or in
short chains. Colonies are circular with a convex elevation
and an entire margin. They grow at 37 °C, pH 3.9 and with 6
% NaCl [198]. The genome size of the type strain is 3.28 Mbp.
2826
Description of Lactiplantibacillus faecal microbiota of healthy dogs, traditional fura processing,

mudanjiangensis comb. nov. wine and sow milk.
Lactiplantibacillus mudanjiangensis (mu.dan.ji.ang.en’sis. N.L. The type strain is CST 10961T = ATCC 700211T=CCUG
masc. adj. mudanjiangensis pertaining to the Mudanjiang 35983T = CIP 104668T = CNRZ 1885T = CST 10961T = DSM
River, a river flowing through the Heilongjiang province of 10667T=JCM 12533T = LMG 16673T=NRRL B-23115T.
China where the bacterium was isolated).
Genome sequence accession number: AZEO00000000.
Basonym: Lactobacillus mudanjiangensis Gu et al. 2013,
4703VP 16S rRNA gene accession number: AJ306297.
Cells are facultatively anaerobic rods occurring singly. Growth

is observed at 30 and 37°C and pH 4, they are resistant to 6 % Description of Lactiplantibacillus
(w/v) NaCl [176]. The genome size of the type strain is 3.58
pentosus comb. nov.
Lactiplantibacillus pentosus (pen.to′sus. N.L. masc. adj.
Isolated from fermented Chinese cabbage [176] and fermented pentosus, of pentose, pertaining to pentoses).
carrot juice [236] .
Basonym: Lactobacillus pentosus (ex Fred et al. 1921) Zanoni
The type strain is 11050T = LMG 27194T=CCUG 62991T et al. 1987, 339VP
Genome sequence accession number: BJDY00000000. The cells are non-motile straight rods and produce acid and
16S rRNA gene accession number: HF679037. clot in litmus milk [238]. The genome size of the type strain
is 3.65 Mbp. The mol% G+C content of DNA is 46.3.
Description of Lactiplantibacillus nangangensis Isolated from diverse sources including corn silage, fermenting
comb. nov. olives, sewage, fermented mulberry leaf powders, fermented
Lactiplantibacillus nangangensis (nan.gang.en’sis. N.L. masc. teas, glutinous rice dough, corn noodles, chili sauce, mustard
adj. nangangensis, pertaining to Nangang, a district in Harbin pickles, stinky tofu, dairy products, mustard pickle, fermented
city, China). idli batter, tempoyak, human vagina, human stools, and
Basonym: Lactobacillus nangangensis Liu and Gu 2019, 3258VP sourdoughs.
Characteristics are similar to L. pingfangensis but the strain The type strain is 124–2T = ATCC 8041T=CCUG 33455T =
does not grow at 37 °C [233]. The genome size of the type CIP 103156T=DSM 20314T = JCM 1558T=LMG 10755T =
strain is 2.90 Mbp. The mol% G+C content of DNA is 44.3. NCAIM B.01727T = NCCB 32014T = NCIMB 8026T (formerly
NCDO 363)=NRRL B-227T = NRRL B-473T.
Genome sequence accession number: AZCU00000000.
The type strain is 381–7T = NCIMB 15186T=CCM 8930T.
16S rRNA gene accession number: D79211.
Genome sequence accession number: BJDI00000000.
16S rRNA gene accession number: LC438520
pingfangensis comb. nov.
Description of Lactiplantibacillus Lactiplantibacillus pingfangensis (ping.fang.en’sis. N.L. masc.
paraplantarum comb. nov. adj. pingfangensis, pertaining to a district in Harbin city,
Lactiplantibacillus paraplantarum (pa.ra.plan.tar’um. Gr. prep. China).
para resembling; N.L. gen. n. plantarum, a species epithet;
N.L. gen. pl. n. paraplantarum, resembling L. plantarum). Basonym: Lactobacillus pingfangensis Liu and Gu 2019, 7VP
Basonym: Lactobacillus paraplantarum Curk et al. 1996, 598VP Growth is observed at 30 and 37 but not at 45 °C; dl-lactic
acid is produced from a wide range of hexoses and disac-
The cells are non-motile rods, usually singly, in pairs and charides but not from pentoses [233]. The genome size of
sometimes in short chains. They grow at pH 5 and 7 and they the type strain is 2.90 Mbp; the mol% G+C content of DNA
tolerate NaCl up to a concentration of 8 % [237]. The genome is 44.2.
of DNA is 43.7. Isolated from fermented Chinese cabbage.
Isolated as spoilage organisms from beer and from human The type strain is 382–1T = LMG 31176T=NCIMB 15187T =
faeces. They are also found in grape marmalade, dairy prod- CCM 8935T.
ucts, jangajji (a Korean fermented food), fermented vegeta-
Genome sequence accession number: BJDG00000000.
bles, fermented fruits, fermented dates, rice bran pickles,
silage, cocoa beans, fermented sourdough, fermented slurry, 16S rRNA gene accession number: LC438521.
2827
Description of Lactiplantibacillus 16S rRNA gene accession number: HM443954.

plajomi comb. nov.
Lactiplantibacillus plajomi (pla.jom’i. N.L. gen. n. plajomi of Heterofermentative Lactobacillaceae
Pla-jom, referring to the isolation of the type strain from a
traditional fermented fish product in Thailand).
Description of Furfurilactobacillus
Basonym: Lactobacillus plajomi Miyashita et al. 2015, 2488VP
gen. nov.
Cells are non-motile, facultatively anaerobic rods. They grow Furfurilactobacillus (Fur.fu.ri.lac.to.ba.cil’lus. L. masc. n.
at 15–37 °C, at pH 4.0–7.5 and with 0–8 % NaCl. They are furfur bran, relating to the origin of furfurilactobacilli from
positive for Voges–Proskauer test, tellurite tolerance, bile- cereal fermentations; N.L. masc. n. Lactobacillus a bacterial
aesculin tolerance test and deamination of arginine [235]. genus name; N.L. masc. n. Furfurilactobacillus a lactobacillus
The genome size of the type strain is 2.76 Mbp. The mol% from bran).
G+C content of DNA is 48.7
Heterofermentative and aerotolerant. Growth is observed
Isolated from Pla-jom (fermented fish) collected in Yasothon, at 15 and 37°C but not at 45 °C. The two species in the
Thailand. genus with genome sequences available have a genome size
The type strain is NB53T = NBRC 107333T = BCC 38054T. of 2.9–3.0 Mbp and a mol% G+C content of DNA of 43–44
%. Species in the genus were isolated from sourdough or
Genome sequence accession number: BJDZ00000000. spoiled beer and have an exceptional capacity to metabolize
16S rRNA gene accession number: AB907190. phenolic compounds [240, 241]. The ecology of the genus
remains largely unexplored but appears to be similar to the
nomadic lifestyle of L. plantarum.
Description of Lactiplantibacillus A phylogenetic tree on the basis of 16S rRNA genes of all
songbeiensis comb. nov. species in the genus Lactiplantibacillus is provided in Figure
Lactiplantibacillus songbeiensis (song.bei.en’sis. N.L. masc. S6L.
adj. songbeiensis pertaining to Songbei, a district in Harbin The type species of the genus is Furfurilactobacillus rossiae
city, China). comb. nov.; Furfurilactobacillus was previously referred to as
Basonym: Lactobacillus songbeiensis Liu and Gu 2019, 3259VP L. rossiae group.
Characteristics are similar to L. pingfangensis but the type
strain of L. songbeiensis also ferments ribose [233]. The
Description of Furfurilactobacillus
content of DNA is 44.4. rossiae comb. nov.
Furfurilactobacillus rossiae (ros’si.ae. N.L. gen. n. rossiae of
Rossi, named in recognition of Jone Rossi, a microbiologist
The type strain is 398–2T = LMG 31174T=NCIMB 15189T = working at the University of Perugia).
CCM 8931T.
Basonym: Lactobacillus rossiae Corsetti et al. 2005, 39VP. The
Genome sequence accession number: BJDL00000000. species was initially named ‘L. rossii’
16S rRNA gene accession number: LC438523. F. rossiae grows at 15 °C but not at 45 °C [242]. The genome
of DNA is 43.3.
Description of Lactiplantibacillus Isolated from wheat sourdough and from related cereal
xiangfangensis comb. nov. fermentations, beer, fruit, and fecal samples of children and
Lactiplantibacillus xiangfangensis (xi.ang.fang.en′sis. N.L. swine. Owing to its capacity for metabolism of phenolic
masc. adj. xiangfangensis, pertaining to Xiangfang, a district compounds and flavonoids, it was used experimentally as
of Harbin city in China). starter culture for cactus pear fermentation [243].
Basonym: Lactobacillus xiangfangensis Gu et al. 2012, 860VP The type strain is CS1T = ATCC BAA-822T = DSM 15814T =
JCM 16176T.
Cells are non-spore-forming, facultatively anaerobic rods.
They grow at 30 and 37 °C [239]. The genome size of the type Genome sequence accession number: AZFF00000000.
Isolated from pickle and sourdough.
The type strain is 3.1.1T = LMG 26013T=NCIMB 14687T.
Genome sequence accession number: JQCL00000000.
2828
Description of Furfurilactobacillus G+C content of DNA ranges from 35.6 to 43.5; and the
curtus comb. nov. genome size ranges from 1.75 to 2.57 Mbp.
Furfurilactobacillus curtus (cur’tus. L. masc. adj. curtus short, A phylogenetic tree on the basis of 16S rRNA genes of all
referring to the short cell morphology of the type strain). species in the genus Paucilactobacillus is provided in Figure
S6M.
Basonym: Lactobacillus curtus Asakawa et al. 2017, 3905VP
The type species of the genus is Paucilactobacillus vaccinos-
It grows over a wide pH range (pH 3.5–8.5) and at 15 °C but tercus comb. nov.; Paucilactobacillus was previously referred
not at 45 °C [244]. The mol% G+C content of DNA is 43. to as L. vaccinostercus group.
Isolated from spoiled beer.
The type strain is JCM 1149T = ATCC 14917T= DSM 20174 Description of Paucilactobacillus
T
= LMG 6907T = NCIMB 11974T=NRRL B-4496T. vaccinostercus comb. nov.
Genome sequence accession number: not available at the time Paucilactobacillus vaccinostercus (vac.ci.no.ster’cus; L. masc.
of publication. adj. vaccinus, from cows; L. neut. n. stercus, manure; N.L.
masc. adj. vaccinostercus, from cow manure).
Basonym: Lactobacillus vaccinostercus Okada et al. 1979,
439VL. Some strains of the species were previously designated
Description of Furfurilactobacillus as Lactobacillus durianis [246]
siliginis comb. nov. Growth occurs between 15 and 40 °C, pH 4.4 and 7.5, and
Furfurilactobacillus siliginis (si.li’gi.nis. L. gen. n. siliginis, of with pentoses, glucose and maltose as carbon source [247].
wheat flour, referring to the origin of the type strain in wheat The genome size of the type strain is 2.57 Mbp. The mol%
sourdough). G+C content of DNA is 43.9.
Basonym: Lactobacillus siliginis Aslam et al. 2006, 2212VP Isolated from cow dung and from fermented tea leaves and
fermented cereals.
This species has a narrow temperature range of growth (20–37
°C) but a wide pH range (pH 4.0–8.0) [245]. The genome The type strain is X-94T = TUA 055BT=ATCC 33310T = DSM
size of the type strain is 2.07 Mbp. The mol% G+C content 20634T= LMG 9215T.
of DNA is 44.1. Genome sequence accession number: AYYY00000000.
Isolated from a wheat sourdough. 16S rRNA gene accession number: AJ621556.
The type strain is M1-212T = JCM 16155T = DSM
22696T=KCTC 3985T = NBRC 101315T. Description of Paucilactobacillus
Genome sequence accession number: JQCB00000000. hokkaidonensis comb. nov.
16S rRNA gene accession number: AB370882. Paucilactobacillus hokkaidonensis (hok.kai.do.nen’sis. N.L.
masc. adj. hokkaidonensis of Hokkaido in northern Japan
from where the type strain was isolated).
Description of Paucilactobacillus gen.
Basonym: Lactobacillus hokkaidonensis Tohno et al. 2013,
nov. 2529VP
Paucilactobacillus (Pau.ci.lac.to.ba.cil’lus. L. masc. adj. paucus
Growth is observed in the range of 4 to 37°C and in the pH
few; N.L. masc. n. Lactobacillus a bacterial genus name; N.L.
range of 4.0–7.0; strains ferment pentoses, maltose and, vari-
masc. n. Paucilactobacillus a lactobacillus that ferments few
ably, melibiose [248]. The genome size of the type strain is
carbohydrates).
2.33 Mbp. The mol% G+C content of DNA is 38.1.
Gram positive, rod-shaped, catalase negative, heterofermen-
Isolated from grass silage.
tative, and aerotolerant. Growth is observed at between 20
and 37 °C. Several species are psychrotrophic but Pauci- The type strain is LOOC260T = JCM 18461T = DSM 26202T.
lactobacillus suebicus grows at 45 °C. Strains in this genus
Genome sequence accession number: JQCH00000000.
were predominantly isolated from fermented plant material
including silage, pickles, and fruit mashes. The adaptation to 16S rRNA gene accession number: AB721549.
hexose depleted habitats is indicated by the lack of mannitol
dehydrogenase in many strains of the genus; among hetero-
fermentative lactobacilli, this property is shared only with
Secundilactobacillus that is isolated from related habitats.
Paucilactobacillus species preferentially metabolise pentoses
and many strains do not ferment disaccharides. The mol%
2829
Description of Paucilactobacillus 16S rRNA gene accession number: AY733084.

kaifaensis comb. nov.
Paucilactobacillus kaifaensis (kai.fa.en’sis. N.L. masc. adj. Description of Paucilactobacillus
kaifaensis, pertaining to Kaifa, a district in Harbin city, China). suebicus comb. nov.
Basonym: Lactobacillus kaifaensis Liu and Gu 2019, 3259VP Paucilactobacillus suebicus (su.e’bi.cus. L. masc. adj. suebicus,
Growth is observed at 30 and 37 but not at 45 °C. dl-Lactic from Swabia, a region in the South West of Germany where
acid is produced only from ribose, xylose, maltose, arabitol the type strain was isolated).
and gluconate [233]. The genome size of the type strain is 1.75 Basonym: Lactobacillus suebicus Kleynmans et al. 1989, 495VL
Mbp; the mol% G+C content of DNA is 38.0.
The type strain exhibits high tolerance to acid and ethanol;
Isolated from fermented Chinese cabbage. growth is observed at pH 2.8 or at pH 3.3 and 14% ethanol.
The type strain is 778–3T=LMG 31177T=NCIMB The type strain ferments pentoses, glucose, and maltose;
15191T=CCM 8929T. growth is observed at 10 °C and up to 45 °C for some strains
Genome sequence accession number: BJDM00000000. mol% G+C content of DNA is 39.0.
16S rRNA gene accession number: LC438525. Isolated from fermented cherry mashes, from cider and silage.
The type strain is I, WC-t4-15T = ATCC 49375T= DSM 5007T
= JCM 9504T=KCTC 3549T = LMG 11408T.
Description of Paucilactobacillus
nenjiangensis comb. nov. Genome sequence accession number: AZGF00000000.
Paucilactobacillus nenjiangensis (nen.ji.ang.en’sis. N.L. masc. 16S rRNA gene accession number: AJ575744.
adj. nenjiangensis, pertaining to the Nenjiang River in the
Chinese Heilongjiang province where the type strain was
isolated). Description of Paucilactobacillus
Basonym Lactobacillus nenjiangensis Gu et al. 2013, 4704VP
wasatchensis comb. nov.
Paucilactobacillus wasatchensis (wa.satch.en’sis. N.L. masc.
The type strain ferments several pentoses, hexoses, maltose, adj. wasatchensis of the Wasatch mountains in Utah, USA,
and sucrose. Growth is not observed at 45 °C or below pH the origin of the type strain).
4.0 [176]. The genome size of the type strain is 1.99 Mbp. The
mol% G+C content of DNA is 38.7. Basonym: Lactobacillus wasatchensis Oberg et al. 2016, 163VP
Isolated from pickle. Growth is observed at 30 and 37°C but not at 45 °C. In MRS,
only ribose and galactose are fermented [251]. The genome
The type strain is 11 102T=LMG 27192T=NCIMB 14833T. size of the type strain is 1.90 Mbp. The mol% G+C content
Genome sequence accession number: BJEB00000000. of DNA is 39.8.
16S rRNA gene accession number: HF679039. Isolated from spoiled cheddar cheese and from silage.
The type strain is WDC04T = DSM 29958T = LMG 28678T.
Description of Paucilactobacillus Genome sequence accession number: AWTT00000000.

oligofermentans comb. nov. 16S rRNA gene accession number: NR_147709.
Paucilactobacillus oligofermentans (o.li.go.fer.men’tans, Gr.
masc. adj. oligos few; L. pres. part. fermentans, fermenting; N.L.
part. adj. oligofermentans, fermenting few [carbohydrates]).
Description of Limosilactobacillus
gen. nov.
Basonym: Lactobacillus oligofermentans Koort et al. 2005,
Limosilactobacillus (Li.mo.si.lac.to.ba.cil'lus. L. masc. adj.
2236VL
limosus, slimy, referring to the property of most strains in
Strains grow at 4 and 15°C but not at 37 °C [249]. The genome the genus to produce exopolysaccharides from sucrose; N.L.
size of the type strain is 1.83 Mbp. The mol% G+C content masc. n. Lactobacillus a bacterial genus name; N.L. masc. n.
of DNA is 35.6. Limosilactobacillus, a slimy lactobacillus.
Isolated from marinated poultry meat at the end of its shelf Gram-positive, rod- or coccoid-shaped, catalase-negative,
life, and from fermented olives. heterofermentative, and anaerobic or aerotolerant. Growth
is observed at 37 °C and, for most species, at 45 °C but not at
The type strain is AMKR18T = DSM 15707T = JCM
15 °C. Strains in the genus have very small genomes ranging
16175T=LMG 22743T.
from 1.6 Mbp for L. equigenerosi to 2.25 Mbp for L. mucosae;
Genome sequence accession number: AZFE00000000. the mol % G+C content as calculated from whole genome
2830
shotgun sequences ranges from 38.6.1 to 53.4. In comparison The species was effectively but not validly published as Lacto-
to other heterofermentative lactobacilli, Limosilactobacillus bacillus alvi Kim et al. 2011 [256]. Growth at 40 °C is faster
species ferment a relatively broad spectrum of carbohydrates, than at 37 °C [256]. The mol% G+C content of DNA is 42.7.
however, several species do not ferment glucose. Acid resist-
Isolated from the gizzard of hens.
ance is typically mediated by expression of urease, glutami-
nase, glutamate decarboxylase and / or arginine deiminase The type strain is R54T = KCCM 90099T = LMG 31569T=JCM
activities [252]. With the exception of L. fermentum and L. 17644T.
secaliphilus, strains in the genus were isolated from intestinal Genome sequence accession number: Not available at time
habitats, or were shown experimentally to have adapted to of publication.
the intestine of vertebrate animals. Limosilactobacillus reuteri
and other strains in the genus produce exopolysaccharides 16S rRNA gene accession number: NR_118032.
from sucrose to support biofilm formation on non-secretory
epithelia in the upper intestinal tract [14, 253]. In their
natural habitat, Limosilactobacillus species generally form
stable associations with Lactobacillus spp.; the same asso- antri comb. nov
ciation is observed in food fermentations with Limosilacto- Limosilactobacillus antri (an’tri. L. gen. n. antri of a cave, refer-
bacillus. Limosilactobacillus species., particularly L. reuteri, ring to the antrum region of the stomach).
are produced commercially for use as starter culture and as Basonym: Lactobacillus antri Roos et al 2005, 81VP
probiotic culture.
A phylogenetic tree on the basis of 16S rRNA genes of all G+C content of DNA is 51.1.
species in the genus Limosilactobacillus is provided in Figure
S6N. Isolated from a biopsy of a healthy human gastric mucosa;
strains of this species were also isolated from the intestine of
The type species of the genus is Limosilactobacillus fermentum other vertebrate animals [98].
comb. nov.; Limosilactobacillus was previously referred to as
the Lactobacillus reuteri group. The type strain is Kx146A4T = LMG 22111T = DSM 16041T=
CCUG 48456T.
Genome sequence accession number: AZDK00000000.
fermentum comb. nov.
Limosilactobacillus fermentum (fer.men′tum. L. neut. n.
fermentum which causes fermentation, leaven, ferment). Description of Limosilactobacillus
Basonym: Lactobacillus fermentum Beijerinck 1901, 233
caviae comb. nov.
(Approved Lists). The species includes strains previously Limosilactobacillus caviae (ca’vi.ae. N.L. gen. n. caviae of a
classified as Lactobacillus cellobiosus [254] cavia, of a guinea pig).
L. fermentum is the only species in Limosilactobacillus that is Basonym: Lactobacillus caviae Killer et al. 2017, 2908VP
not adapted to the intestine of vertebrates [17]. The genome Description provided by [257]. The mol% G+C content of
size of the type strain is 1.90 Mbp. The mol% G+C content DNA is 37.7.
of DNA is 52.4.
Isolated from the oral cavity of a guinea pig.
Occurs widely in spontaneously fermented cereals and
other fermenting plant materials [58, 255], also in dairy The type strain is MOZM2T = CCM 8609T = DSM
100239T=LMG 28780T.
products, manure and sewage, and the faeces and vagina
of humans. Genome sequence accession number: Not available at time
of publication.
The type strain is F (352)T = ATCC 14931T = DSM 20052T=CIP
102980T = JCM 1173T=LMG 6902T. 16S rRNA gene accession number: KT343143.
Genome sequence accession number: JQAU00000000.
16S rRNA gene accession number: JN175331. Description of Limosilactobacillus
coleohominis comb. nov.
Limosilactobacillus coleohominis (co.le.o.ho’mi.nis. Gr. masc.
Description of Limosilactobacillus n. koleos vagina; L. gen. n. hominis of humans; N.L. gen. n.
alvi sp. nov. coleohominis of the vagina of humans).
Limosilactobacillus alvi (al’vi. L. gen. n. alvi, of the stomach, Basonym: Lactobacillus coleohominis Nikolaitchouk et al.
of the digestive organs). 2001, 2084VP.
2831
The type strain was reported not to produce gas from glucose Acid tolerant, anaerobic and thermophilic organisms [98].
[258], however, the genome encodes for enzymes of the phos- The genome size of the type strain is 1.85 Mbp. The mol%
phoketolase pathway but lacks phosphofructokinase, the key G+C content of DNA is 41.6.
enzyme of the Embden-Meyerhoff pathway [14]. The genome
Isolated from a biopsy of a human stomach, and from human
milk.
of DNA is 41.1.
The type strain is Kx156A7T = LMG 22113T = DSM
Isolated from the human vagina; in addition, in human intes-
16045T=CCUG 48454T.
tinal microbiota and from swine.
Genome sequence accession number: AZFN00000000.
The type strain is DSM 14060T = CCUG 44007T=CIP 106820T.
Genome sequence accession number: AZEW00000000.
gorillae comb. nov.
Description of Limosilactobacillus Limosilactobacillus gorillae gorillae ( go.
ril’lae. L. gen. n.
equigenerosi comb. nov. gorillae of the western lowland gorilla).
Limosilactobacillus equigenerosi (e.qui.ge.ne.ro’si. L. masc. n. Basonym: Lactobacillus gorillae Tsuchida et al. 2014, 4005VP
equus, horse; L. masc. adj. generosus, of noble birth, well-bred;
N.L. gen. n. equigenerosi, from a thoroughbred horse). Thermophilic organism [261]. The genome size of the type
Basonym: Lactobacillus equigenerosi Endo et al. 2008, 917VP
Isolated from the faeces of a captive gorillas and from wild
Acid tolerant and thermophilic organism; cells have a coccoid western lowland gorillas.
morphology [259]. The genome size of the type strain is 1.60
Mbp. The mol% G+C content of DNA is 42.7. The type strain is KZ01T = JCM 19575T = DSM 28356T.
Isolated from the intestinal tract of a thoroughbred horse. Genome sequence accession number: BCAH00000000.
The type strain is NRIC 0697T = JCM 14505T=DSM 18793T. 16S rRNA gene accession number: AB904716.
Genome sequence accession number: AZGC00000000.

ingluviei comb. nov.
Limosilactobacillus ingluviei (in.glu’vi.ei. L. gen. n. ingluviei
Description of Limosilactobacillus of a crop sac).
frumenti comb. nov. Basonym Lactobacillus ingluviei Baele et al. 2003, 135VP. The
Limosilactobacillus frumenti (fru.men’ti. L. gen. n. frumenti species includes strains previously named as L. thermotolerans
from cereal). [262].
Basonym: Lactobacillus frumenti Müller et al. 2000, 2132VP Thermophilic species with optimal growth at 42 °C and poor
growth at 30 °C or below [263]. The genome size of the type
Description provided in [260]. The genome size of the type strain is 2.16 Mbp. The mol% G+C content of DNA is 49.9.
Isolated from the crop of a pigeon [263], birds (turkeys,
Isolated from an industrial rye bran fermentation but also chickens, geese) but also from cattle, carnivore faeces, and
identified in the intestine of poultry and swine, and in must Korean rice wine (makgeolii).
and wine.
The type strain is KR3T = DSM 15946T = LMG 20380T=CCUG
The type strain is TMW 1.666T = DSM 13145T=LMG 19473T. 45722T.
Genome sequence accession number: AZER00000000. Genome sequence accession number: AZFK00000000.
16S rRNA gene accession number: AJ250074. 16S rRNA gene accession number: AF333975.
Description of Limosilactobacillus Description of Limosilactobacillus

gastricus comb. nov. mucosae comb. nov.
Limosilactobacillus gastricus (gas’ tri.
cus. N.L. masc. adj. Limosilactobacillus mucosae (mu.co’sae. N.L. gen. n. mucosae
gastricus from Gr. adj. gastrikos of the stomach). of mucosa).
Basonym: Lactobacillus gastricus Roos et al. 2005, 80VP Basonym: Lactobacillus mucosae Roos et al. 2000, 256VP
2832
Many strains of the species carry a mucus binding protein, a Not all strains of L. pontis ferment glucose [267]; growth is
putative colonization factor of L. mucosae. The genome size observed at 15 and at 45 °C. The genome size of the type strain
of the type strain is 2.25 Mbp. The mol% G+C content of is 1.67 Mbp. The mol% G+C content of DNA is 43.5.
DNA is 46.4.
Isolated from type I and type II sourdough but also as typical
Isolated from the intestine of a pig [264] but also found in representatives of the intestinal microbiota of swine [76].
the intestine of other vertebrates including humans, type II Other sources of isolation include silage, dairy products,
sourdough, and related cereal fermentations. mezcal fermentation and wet wheat distillers' grain.
The type strain is S32T = CCUG 43179T = CIP 106485T=DSM The type strain is LTH 2587T = DSM 8475T=LMG 14187T.
13345T = JCM 12515T.
Genome sequence accession number: AZGO00000000.
Genome sequence accession number: AZEQ00000000.
16S rRNA gene accession number: X76329.
reuteri comb. nov.
oris comb. nov.
Limosilactobacillus reuteri (reu′te.ri. N.L. gen. n. reuteri, of
Limosilactobacillus oris (or'is. L. gen. n. oris, of the mouth). Reuter; named for G. Reuter, a German bacteriologist).
Basonym: Lactobacillus oris Farrow et al. 1988, 116VP Basonym: Lactobacillus reuteri Kandler et al. 1982, 266VL
The genome size of the type strain is 2.03 Mbp. The mol% Prior to 1980, strains of the species were classified as L.
G+C content of DNA is 50.0. fermentum Biotype II [268]. The species has been studied
Isolated from the human saliva [265], and, less frequently, as model species to determine host-adaptation of intestinal
from other human body sites including the vagina and lactobacilli, and co-adaptation of Limosilactobacillus and
mother’s milk, and from foods such as corn dough and bran. Lactobacillus species (e.g. L. taiwanensis) that coexist in
biofilms in intestinal ecosystems and preferentially different
The type strain is 5A1T = ATCC 49062T= CCUG 37396T = substrates [55, 56, 269]. L. reuteri is divided in host-adapted
CIP 103255T=CIP 105162T = DSM 4864T=JCM 7507T = JCM lineages that are equivalent to the taxonomic rank of sub-
11028T=LMG 9848T. species [269, 270] and differ in their ability to form biofilms
Genome sequence accession number: AZGE00000000. in the mouse forestomach [271]. Few strains of L. reuteri
produce the antimicrobial compound reutericyclin with a
16S rRNA gene accession number: X94229. polyketide synthase, which is a unique ability among lacto-
bacilli [272]. The genome size of the type strain is 1.94 Mbp.
Description of Limosilactobacillus The mol% G+C content of DNA is 38.6.
panis comb. nov. Isolated as dominant members of the intestinal microbiota
Limosilactobacillus panis (pa’nis. L. gen. n. panis, of bread). of rodents, birds, swine, and in other intestinal ecosystems;
also occurs in cereal fermentations, particularly type II sour-
Basonym: Lactobacillus panis Wiese et al. 1996, 452VP doughs [58]. Food isolates are of intestinal origin [273].
The genome size of the type strain is 2.01 Mbp. The mol% The type strain of the species is F 275T = ATCC 23272T=DSM
G+C content of DNA is 48.1. 20016T = JCM 1112T=LMG 9213T = LMG 13557T.
Isolated from a type II sourdough [266], other strains of Genome sequence accession number: AZDD00000000.
the species occur in fermenting plant material, and in the
intestine of birds. 16S rRNA gene accession number: AP007281.
The type strain is CCUG 37482T = DSM 6035T=JCM 11053T.

Genome sequence accession number: AZGM00000000. Description of Limosilactobacillus
16S rRNA gene accession number: X94230.
secaliphilus comb. nov.
Limosilactobacillus secaliphilus (se.ca.li.phi’lus. L. neut. n.
secale rye; Gr. masc. adj. philos loving; N.L. masc. adj. seca-
Description of Limosilactobacillus liphilus rye-loving).
pontis comb. nov. Basonym: Lactobacillus secaliphilus Ehrmann et al. 2007,
Limosilactobacillus pontis (pon’tis. L. gen. n. pontis, of a 748VP
bridge, referring to BRIDGE, which was the acronym of an
The type strain does not ferment glucose [274]. The genome
EU funded research project).
Basonym: Lactobacillus pontis Vogel et al. 1994, 228VP of DNA is 47.7.
2833
Isolated from a type II sourdough; although other lactobacilli a trait which differentiates Secundilactobacillus from all other
in type II sourdoughs are predominantly of intestinal origin heterofermentative lactobacilli except Paucilactobacillus.
[114], the habitat of L. secaliphilus remains unknown [274]. Many strains in the genus metabolize diols via diol-hydratase
and convert agmatine, a metabolite of arginine decarboxyla-
The type strain is TMW 1.1309T = DSM 17896T=CCUG
tion, via the agmatine deiminase pathway. Strains in the genus
53218T.
generally harbour genes coding for transaldolase/transketo-
Genome sequence accession number: JQBW00000000. lase which mediate metabolism of pentoses to pyruvate.
16S rRNA gene accession number: AM279150. A phylogenetic tree on the basis of 16S rRNA genes of all
species in the genus Secundilactobacillus is provided in Fig.
S6O.
The type species of the genus is Secundilactobacillus male-
vaginalis comb. nov. fermentans comb. nov.; Secundilactobacillus was previously
Limosilactobacillus vaginalis (va.gi.na’lis. L. gen. n. vaginalis, referred to as the Lactobacillus collinoides group.
of the vagina).
Basonym: Lactobacillus vaginalis Embley et al. 1989, 368VP
Description of Secundilactobacillus
Description provided in [275]. The genome size of the type
malefermentans comb. nov.
Secundilactobacillus malefermentans (ma.le.fer.men’tans. L.
Frequently isolated as member of the microbiota of the adv. male, bad; L. part. pres. fermentans fermenting; N.L. part.
human vagina. adj. malefermentans badly fermenting, referring to spoiled
The type strain of the species is Lac 19T = ATCC 49540T=CCUG beer).
31452T = CIP 105932T=DSM 5837T = JCM 9505T=LMG Basonym: Lactobacillus malefermentans (ex Russell and
12891T. Walker 1953) Farrow et al. 1989, 371VL. The species was
Genome sequence accession number: AZGL00000000. initially described in 1953 [277]; the species name was later
revived with a new type strain [278]
Strains of this species have a restricted carbohydrate fermen-
tation pattern. Growth is observed between 10 and 37°C and
Proposed species in the genus between pH 4.1 and 8.0. The genome size of the type strain is
Limosilactobacillus 2.05 Mbp. The mol% G+C content of DNA is 41.0.
The name ‘Lactobacillus timonensis’ was effectively published Isolated from beer.
for a species that clusters phylogenetically within the genus The type strain is D2 MF1T = ATCC 49373T= DSM 5705T =
Limosilactobacillus [276]; however, its species description JCM 12497T= LMG 11455T.
lacks essential information and a type strain was not deposited.
Genome sequence accession number: AZGJ00000000.
gen. nov.
Secundilactobacillus (Se.cun.di.lac.to.ba.cil’lus. L. adj. Description of Secundilactobacillus
secundus, second or next, following; N.L. masc. n. Lactoba- collinoides comb. nov.
cillus a bacterial genus name; N.L. masc. n. Secundilactoba-
Secundilactobacillus collinoides (col.lin.o.i’des. L. masc. adj.
cillus a lactobacillus that occurs in a secondary fermentation
collinus hilly; Gr. suff. eides resembling, similar; N.L. masc.
or as a spoilage organism after primary fermenters depleted
adj. collinoides hill-shaped, pertaining to colony form).
hexoses and disaccharides).
Basonym: Lactobacillus collinoides Carr and Davies 1972, 470
Gram-positive, rod-shaped, catalase negative, heterofermen-
(Approved Lists). Prior to the species description, strains of
tative. Most strains grow at 15 °C and some also grow at 45
S. collinoides were referred to as L. brevis var. quinicus or L.
°C, the pH range of growth is highly variable. The genome size
pasteurianus var. quinicus
ranges from 1.85 Mbp for S. oryzae to 3.62 Mbp for S. colli-
noides, the mol% G+C content of DNA ranges from 41.03–47 Characteristics as described [279]; the genome size of the type
%. Strains in the genus lead a free-living lifestyle and were strain is 3.62 Mbp. The mol% G+C content of DNA is 46.1.
isolated as secondary fermentation or spoilage organisms
Isolated from compost, as spoilage organisms in apple cider,
from hexose-depleted habitats including silage, beer, liquor
table olives, dairy products, fermented durian fruit, and wines.
mashes and apple cider. Metabolic properties of Secundilacto-
bacillus species match adaptation to hexose-depleted habitats. The type strain is Cl3aT=ATCC 27612T = DSM 20515T=JCM
Many strains of the genus do not reduce fructose to mannitol, 1123T = LMG 9194T.
2834
Genome sequence accession number: AYYR00000000. The type strain is YIT 11304T = BCRC 17810T=DSM 19909T
= JCM 15043T.
Genome sequence accession number: AZEE00000000.
kimchicus comb. nov.
Secundilactobacillus kimchicus (kim.chi’cus. N.L. neut. n. Description of Secundilactobacillus
kimchium, kimchi; L. masc. suff. -icus pertaining to; N.L. oryzae comb. nov.
masc. adj. kimchicus pertaining to or isolated from kimchi, Secundilactobacillus oryzae (o.ry’zae. L. gen. n. oryzae of rice).
a type of sauerkraut produced in Korea).
Basonym: Lactobacillus oryzae Tohno et al. 2013, 2961VP
Basonym: Lactobacillus kimchicus Liang et al. 2011, 896 VP
Growth is observed between 4 and 45°C and between pH 5.0
Growth is observed at 15 and at 45 °C and between pH 5.0 and 8.0 [283]. The genome size of the type strain is 1.85 Mbp.
and 9.0 [280]. The genome size of the type strain is 2.59 Mbp. The mol% G+C content of DNA is 42.8.
Isolated from fermented rice grains in Tochigi, Japan.
The species was isolated from kimchi.
The type strain is SG293T = JCM18671T=DSM 26518T.
The type strain is DCY51 = JCM 15530 = KCTC 12976 .
T T T
Genome sequence accession number: BBJM00000000.
Genome sequence accession number: AZCX00000000.
16S rRNA gene accession number: EU678893.
Description of Secundilactobacillus paracollinoides comb. nov.
mixtipabuli comb. nov. Secundilactobacillus paracollinoides (pa.ra.col.li.no’i.des. Gr.
Secundilactobacillus mixtipabuli (mi.xti.pa’bu.li. L. masc. pref. para beside; N.L. masc. adj. collinoides, a species epithet;
N.L. masc. adj. paracollinoides beside S. collinoides, referring
adj. mixtus mixed; L. neut. n. pabulum fodder; N.L. gen. n.
to the close relationship with S. collinoides).
mixtipabuli of mixed fodder, referring to the isolation of the
type strain from silage). Basonym: Lactobacillus paracollinoides Suzuki et al. 2004,
116VP. Strains of this species have been referred to as L. pasto-
Basonym: Lactobacillus mixtipabuli Tohno et al. 2015, 1983VP
rianus prior to 2005 [284]
Characteristics as described by [281]. The genome size of the
Growth is observed at 15 but not at 45 °C [285]. The genome
type strain is 2.52 Mbp. The mol% G+C content of DNA is
43.7.
of DNA is 46.8.
Isolated from silage.
Isolated as beer or cider spoilage organisms, and from
The type strain is IWT30T = JCM 19805T = DSM 28580T. fermented olives.
Genome sequence accession number: BCMF00000000. The type strain is LA2T = DSM 15502T = JCM 11969T.
16S rRNA gene accession number: AB894863. Genome sequence accession number: AZFD00000000.
odoratitofui comb. nov. Description of Secundilactobacillus
Secundilactobacillus odoratitofui (o.do.ra.ti.to’fu.i. L. part adj. pentosiphilus comb. nov.
odoratus smelly; N.L.N.L. gen. n. tofui of tofu; N.L. gen. n. Secundilactobacillus pentosiphilus (pen.to.si’phi.lus. N.L. neut.
odoratitofui, related to stinky tofy from which the type strain n. pentosum pentose; Gr. masc. adj. philos, friend, loving; N.L.
was isolated). masc. adj. pentosiphilus, preferring pentoses).
Basonym: Lactobacillus odoratitofui Chao et al. 2010, 2905VP Basonym: Lactobacillus pentosiphilus Tohno et al. 2017, 3643VP
Characteristics of the species are as described by [282]. The Strains of this species do not produce acid or gas from hexoses
genome size of the type strain is 2.76 Mbp. The mol% G+C including glucose but ferment pentoses; growth is observed
content of DNA is 44.3. between 30 and 37 °C [286]. The genome size of the type
Isolated from fermented brine used for stinky tofu production
in Taipei County, Taiwan. Isolated from silage.
2835
The type strain is IWT25T = JCM 31145T = DSM 102974T. 16S rRNA gene accession number: AB282889.
Genome sequence accession number: BCMI00000000.
16S rRNA gene accession number: LC085284. Description of Levilactobacillus gen.
nov.
Description of Secundilactobacillus Levilactobacillus (Le.vi.lac.to.ba.cil'lus. L. v. levare to lift up,
release, relieve; N.L. masc. n. Lactobacillus a bacterial genus
silagei comb. nov. name; N.L. masc. n. Levilactobacillus a lactobacillus with
Secundilactobacillus silagei (si.la’ge.i. N.L. gen. n. silagei of leavening potential, referring to the occurrence of multiple
silage, from which the type strain was isolated). species in the genus in type I sourdoughs that are used as sole
Basonym: Lactobacillus silagei Tohno et al. 2013, 4616VP leavening agent).
Characteristics as described by [287]. The genome size of the Gram-positive, rod-shaped, catalase negative, heterofer-
type strain is 2.66 Mbp. The mol% G+C content of DNA is mentative. Most strains grow at 15 °C but not at 45 °C, are
44.9. acid tolerant, growing generally in the pH range of 4.0–7.0
and produce dl-lactic acid. The genome size ranges from
1.97 Mbp for Levilactobacillus bambusae to 3.38 Mbp for
The type strain is IWT126T = JCM 19001T = DSM 27022T. Levilactobacillus cerevisiae, the mol% G+C content ranges
from 46.3–55.9 %. Strains in the genus lead a free-living
Genome sequence accession number: BCMG00000000.
lifestyle and were isolated from sourdough or fermented
16S rRNA gene accession number: AB786910. vegetable products but also occur as spoilage organisms in
alcoholic beverages. Strains in the genus generally harbour
genes coding for transaldolase/transketolase which mediate
Description of Secundilactobacillus metabolism of pentoses to pyruvate. Levilactobacillus brevis
silagincola comb. nov. is used commercially as starter culture in food and feed
Secundilactobacillus silagincola (si.lag.in’co.la. N.L. neut. n. applications.
silagum, silage; L. masc. n. incola, inhabitant; N.L. masc. n. A phylogenetic tree on the basis of 16S rRNA genes of all
silagincola, living in or isolated from silage). species in the genus Levilactobacillus is provided in Figure
Basonym: Lactobacillus silangincola Tohno et al. 2017, 3642VP S6P.
Growth is observed between 10 and 37°C and between pH 4.0 The type species of the genus is Levilactobacillus brevis comb.
and 7.5 [286]. The genome size of the type strain is 2.62 Mbp. nov.; Levilactobacillus was previously referred to as L. brevis
The mol% G+C content of DNA is 43.1. group.
The type strain is IWT5T=JCM 31144T=DSM 102973T. Description of Levilactobacillus
Genome sequence accession number: BCMJ00000000. brevis comb. nov.
16S rRNA gene accession number: LC085283. Levilactobacillus brevis (bre′vis. L. adj. brevis short).
Basonym: Lactobacillus brevis (Betabacterium breve Orla-
Jensen 1919, 175) Bergey et al. 1934, 312 (Approved Lists)
The species was described in [2, 289]. Strains of L. brevis
similis comb. nov. widely occur in vegetable and cereal fermentations and as
Secundilactobacillus similis (si’mi.lis. L. masc. adj. similis, beer-spoilage organisms and were also isolated from insects.
similar, as the type strain is similar to closely related species). Niche adaptation is mediated by acquisition of plasmids
Basonym: Lactobacillus similis Kitahara et al. 2010, 189VP [290]. The genome size of the type strain is 2.47 Mbp. The
S. similis grows at 15 but not at 45 °C [288]. The genome size
of the type strain is 3.49 Mbp. The mol% G+C content of Isolated from milk, cheese, sauerkraut and rrelated vegetable
DNA is 47.0. fermentations, sourdough, silage, cow manure, faeces, and the
mouth and intestinal tract of humans and rats.
Isolated from fermented cane molasses at alcohol plants in
Thailand and from rice wine (makgeolii). The type strain is 14, Bb14T=DSM 20054T=ATCC 14869T=JCM
1059T=LMG 6906T=LMG 7944T= NRRL B-4527T.
The type strain is M36T = DSM 23365T = JCM 2765T=LMG
23904T. Genome sequence accession number: AZCP00000000.
Genome sequence accession number: AYZM00000000. 16S rRNA gene accession number: M58810.
2836
Description of Levilactobacillus Description of Levilactobacillus

acidifarinae comb. nov. fujinensis comb. nov.
Levilactobacillus acidifarinae (a.ci.di.fa.ri’nae. L. masc. adj. Levilactobacillus fujinensis (fu.jin.
en’sis. N.L. masc. adj.
acidus sour; L. fem. n. farina; N.L. gen. n. acidifarinae of fujinensis, pertaining to Fujin, a county in the Heilongjiang
sourdough). province of China).
Basonym: Lactobacillus acidifarinae Vancanneyt et al. 2005, Basonym: Lactobacillus fujinensis Long and Gu 2019, 2351VP
619VP
Characteristics of the species are as described [158]. The
Characteristics of the species are as described [291]. The genome size of the type strain is 3 Mbp. The mol% G+C
genome size of the type strain is 2.92 Mbp. The mol% G+C content of DNA is 47.4.
Isolated from type I wheat sourdough and fermented rice
The type strain is 218–6T = CCM 8908T=KCTC 21134T =
bran.
LMG 31067T.
The type strain is R-19065T = DSM 19394T = CCM
Genome sequence accession number: RHNX00000000.
7240T=CCUG 50162T = JCM 15949T=LMG 2220T.
Genome sequence accession number: AZDV00000000.
Description of Levilactobacillus
fuyuanensis comb. nov.
Description of Levilactobacillus Levilactobacillus fuyuanensis (fu.yuan.en’sis. N.L. masc.
bambusae comb. nov. adj. fuyuanensis, pertaining to Fuyuan, a county in the
Levilactobacillus bambusae (bam.bu’sae. N.L. gen. n. bambusae Heilongjiang province of China).
of the plant genus Bambusa from which the type strain was
Basonym: Lactobacillus fuyuanensis Long and Gu 2019,
isolated).
2351VP
Basonym: Lactobacillus bambusae Guu et al. 2018, 2428VP
Characteristics of the species are as described [292]; L. genome size of the type strain is 2.72 Mbp. The mol% G+C
bambusae is more distantly related to other species in Levilac- content of DNA is 48.6.
tobacillus than any other species in the genus as documented
by AAI, G+C content and genome size. The genome size of
the type strain is 1.97 Mbp. The mol% G+C content of DNA The type strain is 244–4T = CCM 8906T=KCTC 21137T=LMG
is 46.3. 31052T.
Isolated from traditional fermented bamboo shoots. Genome sequence accession number: RHNZ00000000.
The type strain is BS-W1 = BCRC 80970 = NBRC 112377 .
T T T
Genome sequence accession number: QCXQ00000000.
16S rRNA gene accession number: KX400838. Description of Levilactobacillus
hammesii comb. nov.
Levilactobacillus hammesii (ham.me’si.i. N.L. gen. n. hammesii
Description of Levilactobacillus of Hammes, in recognition of the scientist Walter P. Hammes,
cerevisiae comb. nov. who contributed significantly to the taxonomy of lactic acid
Levilactobacillus cerevisiae (ce.re.vi’si.ae. L. gen. n. cerevisiae, bacteria as well as the technology and microbial ecology of
of beer). fermented foods).
Basonym: Lactobacillus cerevisiae Koob et al. 2017, 3456VP Basonym: Lactobacillus hammesii Valcheva et al. 2005, 766VP
Characteristics as described [293]. The genome size of the Characteristics of the species are as described [294]. Strains
type strain is 3.38 Mbp. The mol% G+C content of DNA is produce antifungal fatty acids from linoleic acid [295]. The
49.6. genome size of the type strain is 2.82 Mbp. The mol% G+C
Isolated from spoiled beer.
Isolated from wheat and rye sourdoughs, ryegrass silages, and
The type strain is 2301T = DSM 100836T=LMG 29073T.
a municipal biogas plant.
Genome sequence accession number: RHNN00000000.
The type strain is LP38T=TMW 1.1236T= DSM 16381T=CCUG
16S rRNA gene accession number: KT445896. 51325T=CIP 108387T=JCM 16170T.
2837
Genome sequence accession number: AZFS00000000. Genome sequence accession number: RHOD00000000.
16S rRNA gene accession number: AJ632219. 16S rRNA gene accession number: MK110856.

huananensis comb. nov. mulengensis comb. nov.
Levilactobacillus huananensis (hua.nan.en’sis. N.L. masc. Levilactobacillus mulengensis (mu.leng.en’sis. N.L. masc.
adj. huananensis, pertaining to Huanan, a county in the adj. mulengensis, pertaining to Muleng, a county in the
Heilongjiang province of China). Heilongjiang province of China).
Basonym: Lactobacillus huananensis Long and Gu 2019, Basonym: Lactobacillus mulengensis Long and Gu 2019,
2350VP 2352VP
Characteristics are described in [158]. The genome size of Characteristics of the species are as described [158]. The
the type strain is 2.29 Mbp. The mol% G+C content of DNA genome size of the type strain is 3.11 Mbp. The mol% G+C
is 47.5. content of DNA is 50.3.
Isolated from fermented Chinese cabbage. Isolated from fermented Chinese cabbage.
The type strain is 151-2BT=CCM 8913T= KCTC 21129T=LMG The type strain is 112–3T=CCM 8909T=KCTC 21123T= LMG
31063T. 31049T.
Genome sequence accession number: RHOC00000000. Genome sequence accession number: RHNW00000000.
16S rRNA gene accession number: MK110857. 16S rRNA gene accession number: MK110866.
namurensis comb. nov.
koreensis comb. nov.
Levilactobacillus namurensis (na.mur.en’sis. N.L. masc. adj.
Levilactobacillus koreensis (ko.re.en’sis. N.L. masc. adj. kore- namurensis, of the province of Namur, Belgium, the source
ensis of Korea, where the type strain was isolated). of isolation of the type strain).
Basonym: Lactobacillus koreensis Naam Bui et al. 2011, 774VP Basonym: Lactobacillus namurensis Scheirlinck et al. 2007,
Characteristics of the species are as described [296]. The 226VP
genome size of the type strain is 2.47 Mbp. The mol% G+C Characteristics of the species are as described [297]. The
content of DNA is 49.6. genome size of the type strain is 2.48 Mbp. The mol% G+C
Isolated from cabbage kimchi and from sourdough. content of DNA is 52.0.
The type strain is DCY50T= JCM 16448T=KCTC 13530T. Isolated from wheat sourdough and from vegetable
fermentations.
Genome sequence accession number: AZDP00000000.
The type strain is DSM 19117T= CCUG 52843T=JCM
16S rRNA gene accession number: FJ904277. 15612T=LMG 23583T.
Genome sequence accession number: AZDT00000000.
Description of Levilactobacillus 16S rRNA gene accession number: AM259119.
lindianensis comb. nov.
Levilactobacillus lindianensis (lin.dian.en’sis. N.L. masc.
adj. lindianensis, pertaining to Lindian, a county in the
Heilongjiang province of China). parabrevis comb. nov.
Levilactobacillus parabrevis (pa.ra.bre’vis. Gr. pref. para
Basonym: Lactobacillus lindianensis Long and Gu, 2019,
beside; L. masc. adj. brevis, a species epithet; N.L. masc. adj.
2349VP
parabrevis beside L. brevis, referring to the close relationship
Characteristics are described in [158]. The genome size of with L. brevis).
the type strain is 2.33 Mbp. The mol% G+C content of DNA
Basonym: Lactobacillus parabrevis Vancanneyt et al. 2006,
is 49.2.
1556VP
The type strain is 220–4T= NCIMB 15163T=CCM genome size of the type strain is 2.61 Mbp. The mol% G+C
8902T=KCTC 21136T. content of DNA is 49.1.
2838
The species is isolated from farmhouse red Cheshire cheese, Characteristics as described [301]. The genome size of the
wheat sourdough, fermented vegetables, and a municipal type strain is 2.75 Mbp. The mol% G+C content of DNA is
biogas plant. 55.9.
The type strain is RODS-
DWT=SLB MAST=ATCC Isolated from wheat and rice sourdoughs, from fermented
53295T=LMG 11984T. vegetables and a municipal biogas plant.
Genome sequence accession number: AZCZ00000000. The type strain is LTH 5753T=DSM 15429T=JCM
15956T=LMG 21871T.
Genome sequence accession number: AZFC00000000.
Description of Levilactobacillus 16S rRNA gene accession number: AJ534844.

paucivorans comb. nov.
Levilactobacillus paucivorans (pau.ci.vo’rans. L. masc. adj. Description of Levilactobacillus
paucus few, low in number; L. part. pres. vorans devouring; suantsaii comb. nov.
N.L. part. adj. paucivorans, eating few [carbohydrates], refer- Levilactobacillus suantsaii (suan.tsai’i. N.L. gen. n. suantsaii
ring to the few sugars that are fermented by the type strain). of suan-tsai, a fermented mustard product, from which the
Basonym: Lactobacillus paucivorans Ehrmann et al. 2010, type strain was isolated).
2356VP Basonym: Lactobacillus suantsaii Liou et al. 2019, 1489VP
Characteristics of the species are as described [299]. The Characteristics of the species are as described [302]. The
genome size of the type strain is 2.37 Mbp. The mol% G+C genome size of the type strain is 2.43 Mbp. The mol% G+C
content of DNA is 49.1. content of DNA is 51.1.
Isolated from a storage tank of a brewery. Isolated from suan-tsai, a traditional Taiwanese fermented
The type strain is TMW 1.1424 =DSM 22467 =JCM
T T mustard green.
18045T=LMG 25291T. The type strain is L88T=BCRC 12945T=NBRC 113535T.
Genome sequence accession number: JQCA00000000. Genome sequence accession number: QXIL00000000.
16S rRNA gene accession number: FN185731. 16S rRNA gene accession number: MH730159.

senmaizukei comb. nov. suantsaiihabitans comb. nov.
Levilactobacillus senmaizukei (sen.mai.zu’ke.i. N.L. gen. n. Levilactobacillus suantsaiihabitans (suan.tsai.i.ha'bi.tans.
senmaizukei of senmaizuke, a fermented pickle). N.L. neut. n. suantsaium suan-tsai, the name of a traditional
Basonym: Lactobacillus senmaizukei Hiraga et al. 2008, 1627VP fermented mustard green product of Taiwan; L. pres. part.
habitans, inhabiting; N.L. part. adj. suantsaiihabitans inhab-
Characteristics of the species are as described [300]. The iting suan-tsai).
content of DNA is 48.6. Basonym: Lactobacillus suantsaiihabitans Lin et al. 2019, 8VP
Isolated from senmaizuke, a fermented turnip product. Growth is observed in the range of 20–37°C but not at 15 °C
The type strain is L13T= DSM 21775T=NBRC 103853T=TISTR mol% G+C content of DNA is 51.8.
1847T.
Isolated from a fermented mustard green product.
Genome sequence accession number: AYZH00000000.
The type strain is R19T=BCRC 81129T=NBRC 113532T.
Genome sequence accession number: RKLX00000000.
16S rRNA gene accession number: NH810313.
spicheri comb. nov.
Levilactobacillus spicheri (spi’.cher.i. N.L. gen. n. spicheri of Description of Levilactobacillus
Spicher. The name honours Gottfried Spicher, a German tangyuanensis comb. nov.
scientist who pioneered the microbiological and biochemical
Levilactobacillus tangyuanensis (tang.yuan.en’sis. N.L. masc.
characterization of sourdough fermentation).
adj. tangyuanensis, pertaining to Tangyuan, a county in the
Basonym: Lactobacillus spicheri Meroth et al. 2004, 631VL Heilongjiang province of China).
2839
Basonym: Lactobacillus tangyuanensis Long and Gu 2019, Description of Levilactobacillus

2350VP zymae comb. nov.
Characteristics of the species are as described [158]. The Levilactobacillus zymae (zy’mae. Gr. fem. n. zyme leaven,
genome size of the type strain is 2.19 Mbp. The mol% G+C sourdough; N.L. gen. n. zymae of sourdough).
Basonym: Lactobacillus zymae Vancanneyt et al. 2005, 619VP
The type strain is 137–3T=CCM 8907T=KCTC 21125T=LMG genome size of the type strain is 2.71 Mbp. The mol% G+C
31053T. content of DNA is 53.6.
Genome sequence accession number: RHOA00000000. Isolated from a type I wheat sourdough, forages and fermented
16S rRNA gene accession number: MK110859. onions.
The type strain is R-18615T=DSM 19395T=CCM 7241T=CCUG
50163T= JCM 15957T=LMG 22198T.
Description of Levilactobacillus Genome sequence accession number: AZDW00000000.
tongjiangensis comb. nov.
Levilactobacillus tongjiangensis (tong.ji.ang.en’sis. N.L. masc.
adj. tongjiangensis, pertaining to Tongjiang, a county in the
Basonym: Lactobacillus tongjianensis Long and Gu 2019, Description of Fructilactobacillus
2351VP gen. nov.
Characteristics of the species are as described [158]. The Fructilactobacillus (Fruc.ti.lac.to.ba.cil’lus. L. masc. n. fructus,
genome size of the type strain is 2.72 Mbp. The mol% G+C fruit; N.L. masc. n. Lactobacillus a bacterial genus name; N.L.
content of DNA is 47.7. masc. n. Fructilactobacillus, fruit-loving lactobacillus, refer-
ring to the preference for growth in presence of fructose as
Isolated from fermented Chinese cabbage. electron acceptor).
The type strain is 218–10T=CCM 8905T=KCTC 21135T=LMG Gram- positive, rod- shaped, catalase negative, heterofer-
31055T. mentative and aerotolerant. Growth is observed at 15 °C but
Genome sequence accession number: RHNY00000000. not at 37 °C. Strains in the genus have very small genomes
ranging from 1.23 Mbp for Fructilactobacillus sanfranciscensis
16S rRNA gene accession number: MK110863. to 1.44 Mbp for Fructilactobacillus lindneri; the mol% G+C
content of DNA as calculated from whole genome shotgun
sequences ranges from 34.1 to 38.9. Comparable to other
Description of Levilactobacillus insect-associated lactobacilli, strains in the genus ferment
yonginensis comb. nov. only few carbohydrates; some strains ferment only maltose
Levilactobacillus yonginensis (yong.in.en’sis. N.L. masc. adj. and sucrose. Most strains of the species are fructophilic and
yonginensis of Yongin, the city in Korea where the organism use fructose as electron acceptor rather than as carbon source.
was isolated). Several species in the genus are core member of insect intes-
tinal microbiota, and were isolated from insects, flowers, and
Basonym: Lactobacillus yonginensis Yi et al. 2013, 3278VP spoiled or fermented foods.
Characteristics of the species are as described [303]; ginse- A phylogenetic tree on the basis of 16S rRNA genes of all
noside Rb1 is hydrolysed to Rd by β-glucosidase activity. The species in the genus Fructilactobacillus is provided in Fig.
genome size of the type strain is 2.69 Mbp. The mol% G+C S6Q.
The type species of the genus is Fructilactobacillus fructivorans
Isolated from kimchi. comb. nov.; Fructilactobacillus was previously referred to as
The type strain is THK-
V8T = KACC 16236T = JCM L. fructivorans group.
18023 =ATCC 16236 .
T T
Genome sequence accession number: RHNO00000000. Description of Fructilactobacillus

16S rRNA gene accession number: JN128640. fructivorans comb. nov.
Fructilactobacillus fructivorans (fruc.ti.vo’rans. L. masc.
n. fructus, fruit; L. part. pres. vorans, eating, devouring;
N.L. part. adj. fructivorans, fruit-eating, intended to mean
fructose-devouring).
2840
Basonym: Lactobacillus fructivorans Charlton et al. 1934, 1 Isolated from a flower (West-Indian jasmine).
(Approved Lists)
The type strain is Ru20-1T=LMG 29008T=NBRC
The type species of Fructilactobacillus was described as 111239T=PCU 346T= TISTR 2381T.
Lactobacillus fructivorans in 1934 [304] and includes strains
Genome sequence accession number: not available at the time
that were previously described as Lactobacillus heterohi-
of publication.
ochii, Lactobacillus trichodes [305], and L. homohiochii
[306]. Lactobacillus homohiochii was isolated from Saké 16S rRNA gene accession number: LC094494.
mashes and described as homofermentative Lactobacillus
species [307]. The heterofermentative L. heterohiochii (later
classified as F. fructivorans comb. nov.) was isolated from Description of Fructilactobacillus
the same mash. The type strain of L. homohiochii, which lindneri comb. nov.
likely represents an organism related to Lactobacillus
Fructilactobacillus lindneri (lind'ne.ri. N.L. gen. n. lindneri
acetotolerans, has been lost [308]. F. fructivorans tolerates
of Lindner, recognizing the German bacteriologist Lindner).
ethanol concentrations of up to 15 %. The genome size of
the type strain is 1.37 Mbp. The mol% G+C content of DNA Basonym: Lactobacillus lindneri (Henneberg 1901); Back et al.
is 38.9. 1997, 601VL Earlier isolates were referred to as Bacillus lindneri
or Bacterium lindneri [312]
F. fructivorans is stable member of the intestinal microbiota
of fruit flies [309] as well as spoiled sake mashes [58]; it was The description is provided in [312]. The genome size of the
also isolated from spoiled mayonnaise, salad dressings, sour- type strain is 1.44 Mbp. The mol% G+C content of DNA is
dough, dessert wines and aperitifs. 34.1.
The type strain is IFO (now NBRC) 13954T=ATCC Isolated from spoiled beer and from wine.
8288T=CCUG 32260T=CIP 103042T=DSM 20203T=JCM The type strain is KPAT=CIP 102983T=DSM 20690T=JCM
1117T=LMG 9201T=NRRL B-1841T. 11027T=LMG 14528T.
Genome sequence accession number: AZDS00000000. Genome sequence accession number: JQBT00000000.
16S rRNA gene accession number: NR_036789. 16S rRNA gene accession number: X95421.
Description of Fructilactobacillus Description of Fructilactobacillus

florum comb. nov. sanfranciscensis comb. nov.
Fructilactobacillus florum (flo’rum. L. gen. pl. n. florum, of Fructilactobacillus sanfranciscensis (san.fran.cis.cen’sis. N.L.
flowers, from which the type strain was isolated). masc. adj. sanfranciscensis, from San Francisco).
Basonym: Lactobacillus florum Endo et al. 2010, 2481VP Basonym: Lactobacillus sanfranciscensis (ex Kline and Sugi-
The species was described as composed by three fructophilic hara 1971) Weiss and Schillinger 1984, 503VL. The species
strains [310]. The genome size of the type strain is 1.35 Mbp. was described as Lactobacillus sanfrancisco in 1971 [313] and
The mol% G+C content of DNA is 41.1. re-named to Lactobacillus sanfranciscensis [314]. Isolates of
F. sanfranciscensis described prior to 1978 were designated as
Iolated from peony and bietou flowers, and from grapes and Lactobacillus brevis subsp. lindneri [315]
wine.
Virtually all isolates of the species originate from traditional
The type strain is F9-1T=DSM 22689T=CM 16035T=NRIC sourdoughs that are used as leavening agent [114]; one
0771T. isolate was obtained from agave mash [151]. The extremely
Genome sequence accession number: AYZI00000000. small genome size of 1.2–1.3 Mbp, the restricted carbohy-
drate fermentation pattern, which for includes only maltose
16S rRNA gene accession number: AB498045. for some strains, and the narrow pH range of growth (pH
3.9–6.6) [316, 317] document adaptation to a very narrow
ecological niche. An insect origin of the species is suggested
Description of Fructilactobacillus by culture-independent analysis of the microbiota of grain
ixorae comb. nov. beetles [318]. The genome size of the type strain is 1.23
Fructilactobacillus ixorae (i.xo’rae. N.L. gen. n. ixorae of Ixora
coccinea, West Indian Jasmine as source of isolation). The type strain is L-12T=ATCC 27651T=DSM 20451T=LMG
16002T.
Basonym: Lactobacillus ixorae Techo et al. 2016, 5504VP
Genome sequence accession number: AYYM00000000.
The description is provided in [311]. The mol% G+C content
of DNA is 47.8. 16S rRNA gene accession number: X76327.
2841
Description of Fructilactobacillus 16S rRNA gene accession number: KT783533.

vespulae comb. nov.
Fructilactobacillus vespulae (ves’pu.lae. N.L. gen. n. vespulae
of the wasp Vespula). Description of Apilactobacillus gen.
Basonym: Lactobacillus vespulae Hoang et al. 2015, 3330 VP nov.
The description is provided in [319]. The mol% G+C content Apilactobacillus (A.pi.lac.to.ba.cil'lus. L. fem. n. apis bee; N.L.
of DNA is 35.5. masc. n. Lactobacillus a bacterial genus name; N.L. masc. n.
Apilactobacillus a lactobacillus from bees).
Isolated from a queen wasp.
Gram positive, rod-shaped, heterofermentative. Growth is
The type strain is DCY75T=KCTC 21023T=JCM 19742T. generally observed in the range of 15–37 °C; many strains
Genome sequence accession number: not available at the time grow at acidic conditions below pH 3.0. Strains in the genus
of publication. have small genomes ranging from 1.42 Mbp for Apilactoba-
cillus kosoi to 1.58 Mbp for Apilactobacillus quenuiae; the
mol% G+C content of DNA ranges from 30.5 to 36.4. All
strains in the genus convert fructose to mannitol; strains
Description of Acetilactobacillus gen. in the genus typically ferment very few carbohydrates
nov. including the pollen and bee-associated carbohydrates
fructose, glucose, and sucrose but not maltose or pentoses.
Acetilactobacillus, (A.ce.ti.lac.to.ba.cil’lus. L. neut. n. acetum Comparable to Fructilactobacillus, Apilactobacillus has an
vinegar, referring to the isolation of the sole representative of
insect-associated lifestyle and occurs on flowers, which
this genus from a grain vinegar mash; N.L. masc. n. Lactoba-
serve as hub for dispersal of lactobacilli, and insects
cillus a bacterial genus name; N.L. masc. n. Acetilactobacillus,
[320]. In contrast to Fructilactobacillus, Apilactobacillus
a lactobacillus from vinegar).
has adapted to bees including honeybees (A. apinorum,
Gram- positive, rod- shaped, catalase negative, heterofer- A. kunkeei) and wild bees (A. timberlakei, A. micheneri,
mentative. Mesophilic, with a narrow temperature range of A. quenuiae). In the bee and bumblebee gut, Apilactoba-
growth, and acid tolerant with an optimum pH of growth at cillus is associated with its homofermentative sister genus
4.0. The spectrum of carbohydrates that support acid forma- Bombilactobacillus.
tion is unusual as it includes sugar alcohols and disaccharides
but excludes most hexoses and all pentoses. The unusual A phylogenetic tree on the basis of 16S rRNA genes of all
substrate requirements and pH optimum for growth may species in the genus Apilactobacillus is provided in Figure
reflect an ecological association with acetic acid bacteria. S6R.
The sole representative of the genus was isolated from a grain
The type species of the genus is Apilactobacillus kunkeei comb.
vinegar mash (vinegar pei) and has a genome size of 2.33 Mbp
nov. Apilactobacilus was previously referred to as L. kunkeei
with a mol% G+C content of 41.7.
group.
The type species of the genus is Acetilactobacillus jinshanensis
sp. nov.
Description of Apilactobacillus
Description of Acetilactobacillus kunkeei comb. nov.
jinshanensis sp. nov. Apilactobacillus kunkeei (kun’kee.i; N.L. gen.n. kunkeei of
Kunkee, named in honor of the Californian microbiologist
Acetilactobacillus jinshanensis (jin.shan’en.sis. N.L. masc.
Ralph Kunkee).
adj. jinshanensis, referring to Jinshan, the site of isolation in
Zhenjiang province, China). Basonym: Lactobacillus kunkeei Edwards et al. 1998, 1083VL
The species was effectively but not validly published as A facultative anaerobic organism [321]; the genome size of
Lactobacillus jinshani Yu et al. 2020 [41]. Growth is observed the type strain is 1.52 Mbp. The mol% G+C content of DNA
between 20 and 40 °C with optimal growth at 35 °C [41]. The is 36.4.
optimum pH for growth is 4.0; growth occurs in the range of
3.0–5.0. The strain ferments several disaccharides and sugar Isolated form a sluggish grape wine fermentation but the
alcohols but no pentoses. The genome size of the type strain species is typically associated with honey bees and flowers.
is 2.33 Mbp. The mol% G+C content of DNA is 41.7. The type strain is YH-15T=ATCC 700308T= DSM 12361T=JCM
Isolated from a grain vinegar mash. 16173T.
The type strain is HSLZ-75T=CICC 6269T=JCM 33270T. Genome sequence accession number: AZCK00000000.
Genome sequence accession number: CP034726. 16S rRNA gene accession number: Y11374.
2842
Description of Apilactobacillus Description of Apilactobacillus

apinorum comb. nov. ozensis comb. nov.
Apilactobacillus apinorum ( a.
pi.
no’rum. N.L. gen. pl. n. Apilactobacillus ozensis (o.zen’sis. N.L. masc. adj. ozensis of
apinorum of the Apini, referring to the isolation of this species Oze, the Japanese National Park from where the type strain
from several species of honey bees). was isolated).
Basonym: Lactobacillus apinorum Olofsson et al. 2014, 3113VP Basonym: Lactobacillus ozensis Kawasaki et al. 2011, 2437VP
Growth is observed at anaerobic conditions in the range of A. ozensis grows anaerobically but not aerobically; growth is
15–50°C and pH 3.0–12.0 [89]. The genome size of the type inhibited by 1.5 % NaCl [325]. The genome size of the type
strain is 1.36 Mbp. The mol% G+C content of DNA is 34.9. strain is 1.48 Mbp. The mol% G+C content of DNA is 31.9.
Isolated from honey stomach of the honeybee. Isolated from a chrysanthemum flower.
The type strain is Fhon13NT=DSM 26257T = CCUG 63287T. The type strain is Mizu2-1T=DSM 23829T=JCM 17196T.
Genome sequence accession number: JXCT00000000. Genome sequence accession number: AYYQ00000000.
16S rRNA gene accession number: JX099541. 16S rRNA gene accession number: AB572588.
Description of Apilactobacillus kosoi Description of Apilactobacillus

comb. nov. quenuiae comb. nov.
Apilactobacillus kosoi (ko’so.i. N.L. gen. n. kosoi, of kôso, a Apilactobacillus quenuiae (que.nu’i.ae. N.L. gen. fem. n.
high sucrose fermented beverage in Japan, the origin of the quenuiae, named in recognition of Cécile Plateaux-Quénu’s
type strain). contributions to the biology of bees).
Basonym: Lactobacillus kosoi Chiou et al., 2018, 2707VL Basonym: Lactobacillus quenuiae Frederick et al. 2018, 1884VP
The species is described in [322, 323]. A. kosoi is most closely A description of the speices is provided in [324]. The genome
related to A. micheneri; major physiological properties are size of the type strain is 1.58 Mbp. The mol% G+C content
shared with A. micheneri, A. timberlakei and A. quenuiae. The of DNA is 30.3.
genome size of the type strain is 1.42 Mbp. The mol% G+C Isolated as part of the core microbiome of wild bees and from
content of DNA is 30.5. flowers.
Isolated from koso, a Japanese sugar-vegetable fermented The type strain is HV_6T=DSM 104127T=NRRL B-65474T.
beverage.
Genome sequence accession number: POSN00000000.
The type strain is 10HT=NBRC 113063T=BCRC 81100T.
16S rRNA gene accession number: KX656667.
Genome sequence accession number: BEXE01000000.
16S rRNA gene accession number: LC318484
Description of Apilactobacillus
timberlakei comb. nov.
Description of Apilactobacillus Apilactobacillus timberlakei (tim.ber.lake’i. N.L. gen. masc. n.
micheneri comb. nov. timberlakei names in honor of Philip Timberlake’s work on
Apilactobacillus micheneri (mi.che’ne.ri. N.L. gen. masc. n. bee taxonomy).
micheneri named in honor of Charles D. Michener‘s contribu- Basonym: Lactobacillus timberlakei McFrederick et al. 2018,
tions to the study of bees in natural habitats). 1884VP
Basonym: Lactobacillus micheneri McFrederick et al. 2018, Aerotolerant [324]; the genome size of the type strain is 1.54
1993VP Mbp. The mol% G+C content of DNA is 30.5.
Major physiological properties are shared with A. timberlakei Isolated as part of the core microbiome of wild bees, and from
and A. quenuiae [324]. The genome size of the type strain is flowers.
1.46 Mbp. The mol% G+C content of DNA is 30.3.
The type strain is HV_12T=DSM 104128T=NRRL B-65472T.
Isolated from bees and flowers.
Genome sequence accession number: POST00000000.
The type strain is Hlig3T=DSM 104126T=NRRL B-65473T.
16S rRNA gene accession number: KX656650.
Genome sequence accession number: POSO00000000.
16S rRNA gene accession number: KT833121.
2843
Description of Lentilactobacillus gen. Isolated from pressed yeast, milk, cheese, fermenting plant
nov. material and the human mouth. L. buchneri is used commer-
cially as silage inoculant.
Lentilactobacillus (Len.ti.lac.to.ba.cil’lus. L. masc. adj. lentus
slow, sluggish, referring to the slow growth of species in the The type strain is ATCC 4005T=CCUG 21532T=CIP
genus with lactate or propanediol as carbon course; N.L. 103023T=DSM 20057T=JCM 1115T=LMG 6892T=NCAIM
masc. n. Lactobacillus a bacterial genus name; N.L. masc. n. B.01145T=NRRL B-1837T=VKM B-1599T.
Lentilactobacillus a slow [growing] lactobacillus). Genome sequence accession number: AZDM00000000.
Gram-positive, rod-shaped, catalase negative, heterofer- 16S rRNA gene accession number: AB205055.
mentative. Most strains grow at 15 °C and some also grow at
45 °C, the pH range of growth is highly variable but optimal
growth is typically observed below pH 6.0. The genome
Description of Lentilactobacillus
size ranges from 1.57 Mbp for Lentilactobacillus senioris
to 4.91 Mbp for Lentilactobacillus parakefiri, the mol% curieae comb. nov.
G+C content of DNA ranges from 39.1–45.2 %. Strains in Lentilactobacillus curieae (cu.rie’ae. N.L. fem. gen. n. curieae of
the genus were isolated from silage, fermented vegetables, Curie, named after Marie Curie, to acknowledge a role model
particularly as secondary fermentation or spoilage micro- for women in science).
biota, (spoiled) wine and cereal mashes. Generally, strains Basonym: Lactobacillus curieae Lei et al. 2013, 2504VP
in the species lead a free-living lifestyle; individual species
of the genus, particularly L. senioris, appear to transition Description: Growth is observed at 15 and 45°C and up to 8
to a specialized, host adapted lifestyle. Lentilactobacillus % NaCl [327]. The genome size of the type strain is 2.10 Mbp.
species generally metabolize a broad spectrum of pentoses, The mol% G+C content of DNA is 39.8.
hexoses, and disaccharides. Most species metabolize lactate Isolated from a stinky tofu brine, cocoa bean fermentations
to 1,2-propanediol and / or 1,2-propanediol to propanol and from cheese curd powder.
and propionate; these pathways are virtually exclusive to
heterofermentative lactobacilli and, among these, most The type strain is CCTCC M 2011381T=S1L19T=JCM 18524T.
common and most frequent in Lentilactobacillus. Most Genome sequence accession number: CP018906.
Lentilactobacillus species also convert agmatine, a product
of bacterial decarboxylation of arginine, via the agmatine 16S rRNA gene accession number: JQ086550.
deiminase pathway [14, 326]; in lactobacilli, this pathway
is associated with a free-living lifestyle. Lentilactobacillus
species are used commercially as silage starter cultures, and Description of Lentilactobacillus
in few dairy and cereal applications. diolivorans comb. nov.
A phylogenetic tree on the basis of 16S rRNA genes of all Lentilactobacillus diolivorans (di.o.li.vo’rans. N.L. n. diol from
species in the genus Lentilactobacillus is provided in Figure 1,2-propanediol; L. pres. part. vorans eating; N.L. part. adj.
S6S. diolivorans eating diols).
The type species of the genus is Lentilactobacillus buchneri Basonym: Lactobacillus diolivorans Kroonemen et al. 2001,
comb. nov.; Lentilactobacillus was previously referred to as 645VP
L. buchneri group. Growth is observed between 20 and 40 °C, and between
pH 3.8 and 7. L. diolivorans was the first lactic acid bacte-
rium for which the metabolic pathway for propanediol
and glycerol metabolism to propanol and propionate or
Description of Lentilactobacillus
1,3 propanediol and 3-hydroxypropionate, respectively,
buchneri comb. nov. was thoroughly described [328]. This metabolic pathway
Lentilactobacillus buchneri (buch’ne.ri. N.L. gen. n. buch- is functional is multiple other lactobacilli [14, 18]. The
neri of Buchner, referring to the German microbiologist genome size of the type strain is 3.27 Mbp. The mol% G+C
Buchner). content of DNA is 40.0.
Basonym: Lactobacillus buchneri (Bacillus buchneri [sic] Isolated from maize silage and as a spoilage organism in
Henneberg 1903, 163) Bergey et al. 1923, 251 (Approved vegetable (cucumber) fermentations and fermented dairy
Lists) products.
Biochemical characteristics of L. buchneri were described The type strain is JKD6T=DSM 14421T=JCM 12183T=LMG
as being similar to L. brevis [11, 231] but L. buchneri differs 19667T.
with regards to the production of 1,2 propanediol from
Genome sequence accession number: AZEY00000000.
lactate. The genome size of the type strain is 2.45 Mbp. The
mol% G+C content of DNA is 44.4. 16S rRNA gene accession number: AF264701.
2844
Description of Lentilactobacillus Genome sequence accession number: AYYV00000000.

farraginis comb. nov. 16S rRNA gene accession number: AJ621553.
Lentilactobacillus farraginis (far.ra’gi.nis. L. gen. n. farraginis of
mash, referring to soshu mash as an ingredient of the compost
from which the type strain was isolated). Description of Lentilactobacillus
kisonensis comb. nov.
Basonym: Lactobacillus farraginis Endo and Okada 2007,
Lentilactobacillus kisonensis (ki.so.nen’sis. N.L. masc. adj.
711VP
kisonensis from Kiso, the county in Japan, from where the
Growth is observed at 15 and 45°C and up to pH 9.0 for some type strain was isolated).
strains [329]. The genome size of the type strain is 2.86 Mbp.
Basonym: Lactobacillus kisonensis Watanabe et al. 2009, 757VP
Description. Growth is observed at 10 but not at 45 °C and
Isolated from isolated from a compost of distilled shochu
between pH 4.5 to 8.0 [333]. The genome size of the type
residue.
The type strain is DSM 18382T= JCM 14108T=NRIC 0676T.
Isolated from a pickle brine.
Genome sequence accession number: AZFY00000000.
The type strain is YIT 11168T=DSM 19906T=JCM
16S rRNA gene accession number: AB262731. 15041T=NRIC 074T.
Genome sequence accession number: AZEB00000000.
Description of Lentilactobacillus 16S rRNA gene accession number: AB366388.
hilgardii comb. nov.
Lentilactobacillus hilgardii (hil.gar’di.i. N.L. gen. n. hilgardii
of Hilgard, named after E.W. Hilgard, a pioneer of enology Description of Lentilactobacillus
in the State of California, USA). otakiensis comb. nov.
Lentilactobacillus otakiensis (o.ta.ki.en’sis. N.L. masc. adj.
Basonym: Lactobacillus hilgardii Douglas and Cruess 1936,
otakiensis from Otaki, the village in Japan, from where the
115 (Approved Lists); the species was initially described in
type strain was isolated).
1936; the description was emended in 1949 after the original
type strain was lost [330, 331] Basonym: Lactobacillus otakiensis Watanabe et al. 2009, 758VP
Strains grow optimally between pH 4.5 and 5.5 and in the Physiological characteristics and origin are similar to L. kison-
range of 15 to 40 °C [330, 331]. The genome size of the type ensis [333]. The genome size of the type strain is 2.35 Mbp.
strain is 2.60 Mbp. The mol% G+C content of DNA is 39.6. The mol% G+C content of DNA is 42.4.
Isolated from spoiled wine, kefir grains, mezcal fermentations Isolated from sunki, a fermented turnip product and from
and silage. kefir.
The type strain is 9T=ATCC 8290T=CIP 103007T=DSM The type strain is YIT 11163T=DSM 19908T=JCM
20176T=JCM 1155T=LMG 6895T. 15040T=NRIC 0742T.
Genome sequence accession number: AZDF00000000. Genome sequence accession number: AZED00000000.
16S rRNA gene accession number: M58821. 16S rRNA gene accession number: AB366386.
Description of Lentilactobacillus Description of Lentilactobacillus

kefiri comb. nov. parabuchneri comb. nov.
Lentilactobacillus kefiri (ke.fi’ri. N.L. gen. n. kefiri, of kefir, Lentilactobacillus (Gr. pref. para beside; N.L. gen. n. buch-
referring to the source of isolation). neri a species epithet; N.L. gen. n. parabuchneri beside
L. buchneri, referring to the close relationship with this
Basonym: Lactobacillus kefir Kandler and Kunath 1983, 672VL
organism).
Characteristics are described in [332]. Growth is observed at
Basonym: Lactobacillus parabuchneri Farrow et al. 1988,
15 but not at 45 °C; the genome size of the type strain is 2.23
371VL; strains of the species were previously classified as L.
ferintoshensis [334] or L. frigidus [64, 335]
Isolated from kefir as part of the core microbiota.
Characteristics are described in [278]; growth is observed at
The type strain is A/KT=ATCC 35411T=CIP 103006T= DSM 10 and 40 °C. The genome size of the type strain is 2.61 Mbp.
20587T=JCM 5818T=LMG 9480T. The mol% G+C content of DNA is 43.5.
2845
Isolated from dairy products, saliva, silage, spoiled beer, and Description of Lentilactobacillus
some strains were shown to persist over month in whiskey raoultii sp. nov.
mashes in Scottish distilleries [334].
Lentilactobacillus raoultii (ra.oul’ti.i. N.L. gen. n. raoultii of
The type strain is 6ET=ATCC 49374T= DSM 5707T=JCM Raoult, named after the microbiologist Didier Raoult).
12493T=LMG 11457T.
The species was effectively but not validly published as Lacto-
Genome sequence accession number: AZGK00000000. bacillus raoultii Nicaise et al. 2019 [337]. Growth is observed
16S rRNA gene accession number: AB205056. between 25–45°C and between pH 5 and 7.5; the genome
of DNA is 41.4.
Description of Lentilactobacillus Isolated from the vagina of a healthy women.
parafarraginis comb. nov. The type strain is Strain Marseille P4006T = CSUR
Lentilactobacillus parafarraginis (Gr. pref. para beside; L. P4006T=LMG 31598T = CCUG 71848T.
gen. n. farraginis a species epithet; N.L. gen. n. parafarraginis Genome sequence accession number: OVSN00000000.
beside L. farraginis, referring to the close relationship with
this organism). 16S rRNA gene accession number: LT854294.
Basonym: Lactobacillus parafarraginis Endo and Okada 2007,
711VP Description of Lentilactobacillus rapi
Physiological characteristics are similar to L. farraginis but comb. nov.
growth is observed only between 20 and 37 °C [329]. The Lentilactobacillus rapi (ra’pi. L. gen. n. rapi of a turnip, the
genome size of the type strain is 3.08 Mbp. The mol% G+C substrate for fermented sunki products).
Basonym: Lactobacillus rapi Watanabe et al. 2009, 759VP
Isolated from a compost of distilled shochu residue, silage,
fermented vegetables, and kefir grains. Origin and physiological properties are similar to L. kison-
ensis [333]. The genome size of the type strain is 2.86 Mbp.
The type strain is NRIC 0676T=DSM 18390T=JCM The mol% G+C content of DNA is 43.0.
14109T=NRIC 0677T.
Isolated from sunki and other vegetable fermentations.
Genome sequence accession number: AZFZ00000000.
16S rRNA gene accession number: AB262734. 15042T=NRIC 0743T.
Genome sequence accession number: AZEI00000000.
Description of Lentilactobacillus 16S rRNA gene accession number: AB366389.
parakefiri comb. nov.
Lentilactobacillus parakefiri (pa.ra.ke'fir.i. Gr. pref. para
beside; N.L. gen. n. kefiri a species epithet; N.L. masc. adj. Description of Lentilactobacillus
parakefiri beside L. kefiri, referring to the close relationship senioris comb. nov.
with this organism). Lentilactobacillus senioris (se.ni.o’ris. L. gen. n. senioris of an
Basonym: Lactobacillus parakefiri Takizawa et al. 1994, elderly person, indicating the source of the type strain).
438VP. The classification of L. parakefiri as separate species Basonym: Lactobacillus senioris Oki et al., 2012, 606VP
was disputed as the first published genome sequence of the
type strain was virtually identical to L. kefiri [14, 16] but Growth is observed at 15 and 37 °C but not at 45 °C; the type
re-sequencing of the type strain supported classification of strain was isolated from human faeces [173]. L. senioris is the
L. parakefiri as separate species [336] most distantly related member of the genus, has the smallest
genome size (1.57 Mbp) and lowest G+C content (39.9)
Growth is observed at 15 but not at 45 °C [101]. The genome among species in the genus, and does not share the signature
size of the type strain is 4.91 Mbp. The mol% G+C content genes that are shared by other lentilactobacilli, which may
of DNA is 42.6. indicate that L. senioris transitions from a free-living to a
Isolated from kefir grains. host-adapted lifestyle.
The type strain is GCL 1731T=ATCC 51648T=CCUG 39468T Isolated from from the faeces of a 100-year-old female.
= CIP 104242T=DSM 10551T = LMG 15133T. The type strain is YIT 12364T=DSM 24302T=JCM 17472T.
Genome sequence accession number: AZEN00000000. Genome sequence accession number: AYZR00000000.
16S rRNA gene accession number: AY026750. 16S rRNA gene accession number: AB602570.
2846
Description of Lentilactobacillus 16S rRNA gene accession number: MK110813.

sunkii comb. nov.
Lentilactobacillus sunkii (sun’ki.i. N.L. gen. n. sunkii of sunki, Description of Lapidilactobacillus
non-salted Japanese traditional pickle). mulanensis comb. nov.
Basonym: Lactobacillus sunkii Watanabe et al. 2009, 759VP Lapidilactobacillus mulanensis (mu.lan.en´sis. N.L. masc. adj.
Physiological properties and origin are comparable to L. mulanensis, pertaining to Mulan, a county in Heilongjiang
kisonensis [333]. The genome size of the type strain is 2.69 Province, PR China).
Mbp. The mol% G+C content of DNA is 42.1. Basonym: Lactobacillus mulanensis Long et al. 2020, 14VP.
Isolated from sunki, a fermented turnip product, and from L. mulanensis grows between 15 and 32 °C; the strain produces
kefir. acid from hexoses and disaccharides but not from pentoses
mol% GC content of DNA is 42.5.
15039T=NRIC 0744T.
Isolated from a vegetable fermentation [338].
Genome sequence accession number: A
ZEA00000000.mol%
G+C content of DNA is 36.9. The type strain is 143-6T = NCIMB 15162T = CCM 8951T =
JCM 33274T = CCTCC AB 2018411T.
Genome sequence accession number: RHOW00000000.
Description of Lapidilactobacillus
achengensis comb. nov. 16S rRNA gene accession number: MK110808.
Lapidilactobacillus achengensis (a.cheng.en´sis. N.L. masc. adj.
achengensis, pertaining to Acheng, a county in Heilongjiang Description of Lapidilactobacillus
Province, P.R. China). wuchangensis comb. nov.
Basonym: Lactobacillus achengensis Long et al. 2020, 15VP. Lapidilactobacillus wuchangensis (wu.chang.en´sis. N.L.
L. achengensis grows between 10 and 37°C but not at 5 and masc. adj. wuchangensis, pertaining to Wuchang, a county in
45°C and metabolises several pentoses and a broad spectrum Heilongjiang Province, PR China).
of disaccharides [338]. The genome size of the type strain is Basonym: Lactobacillus wuchangensis Long et al. 2020, 15VP.
2.64 Mbp. The mol% GC content of DNA is 49.0.
L. mulanensis grows between 15 and 37 °C but not at 10
Isolated from a vegetable fermentation [338]. and 45°C; the strain produces L(+)-lactate from d-ribose,
d-glucose, N-acetylglucosamine and cellobiose [338]. The
The type strain is 247-4T = NCIMB 15155T = CCM 8897T =
genome size of the type strain is 2.53 Mbp. The mol% GC
LMG 31059T = CCTCC AB 2018410T.
Genome sequence accession number: RHOV00000000.
JCM 33271T = CCTCC AB 2018406T.
Description of Lapidilactobacillus Genome sequence accession number: RHOU00000000.
gannanensis comb. nov. 16S rRNA gene accession number: MK110811.
Lapidilactobacillus gannanensis (gan.nan.en´sis. N.L.
masc. adj. gannanensis, pertaining to Gannan, a county in
Heilongjiang Province, PR China). Description of Lacticaseibacillus
daqingensis comb. nov.
Basonym: Lactobacillus gannanensis Long et al. 2020, 16VP.
Lacticaseibacillus daqingensis (da.qing.en´sis. N.L. masc. adj.
L. gannanensis grows between 15 and 37 °C but not at 10 and daqingensis, pertaining to Daqing, a city in in Heilongjiang
45°C; the strain produces acid from D-glucose, N‑acetylglu- Province, PR China).
cosamine, salicin, cellobiose and gentiobiose but not from
Basonym: Lactobacillus daqingensis Long et al. 2020, 14VP.
pentoses [338]. The genome size of the type strain is 2.39 Mbp.
The mol% GC content of DNA is 40.8. L. daqingensis grows at 10 - 37°C but not at 5° and 45°C [338].
The genome size of the type strain is 2.74 Mbp. The mol% GC
CCTCC AB 2018409T.
Genome sequence accession number: RHOT00000000.
2847
The type strain is 143-4(a)T = NCIMB 15173T = CCM 8948T The type strain is 33-1T = NCIMB 15169T = CCM 8947T =
= JCM 33273T = CCTCC 2018390T. JCM 33272T = CCTCC 2018405T.
Genome sequence accession number: RHOH00000000. Genome sequence accession number: RHOG00000000.
16S rRNA gene accession number: MK110842. 16S rRNA gene accession number: MK110845.
Description of Lacticaseibacillus Description of Loigolactobacillus

hegangensis comb. nov. binensis comb. nov.
Lacticaseibacillus hegangensis (he.gang.en´sis. N.L. masc. Loigolactobacillus binensis (bin.en´sis. N.L. masc. adj. binensis,
adj. hegangensis, pertaining to Hegang, a city in Heilongjiang pertaining to Bin, a county in Heilongjiang Province, PR
Province, PR China). China).
Basonym: Lactobacillus hegangensis Long et al. 2020, 12VP. Basonym: Lactobacillus binensis Long et al. 2020, 16VP.
Properties of L. hegangensis are similar to L. daqingensis [338]. L. binensis grows between 10 and 37 °C but not at 5 and 45°C;
The genome size of the type strain is 2.50 Mbp. The mol% GC the strain produces acid from several hexoses, sugar alcohols
content of DNA is 55.5. and maltose, but not from pentoses [338]. The genome size
of the type strain is 2.85 Mbp. The mol% GC content of DNA
Isolated from a vegetable fermentation [338]. is 43.5.
The type strain is 73-4T = NCIMB 15177T = CCM 8912T = Isolated from a vegetable fermentation [338].
CCTCC AB 2018407T.
Genome sequence accession number: RHOL00000000. LMG 31186T.
16S rRNA gene accession number: MK110833. Genome sequence accession number: BJDN00000000.
suibinensis comb. nov. Description of Lactiplantibacillus
Lacticaseibacillus suibinensis (sui.bin.en´sis. N.L. masc. adj. garii comb. nov.
suibinensis, pertaining to Suibin, a county in Heilongjiang Lactiplantibacillus garii (ga’ri.i N.L. gen. n. garii of gari, the
Province, PR China). fermented cassava product from which the type strain was
Basonym: Lactobacillus suibinensis Long et al. 2020, 12VP. isolated).
Properties of L. suibiensis are similar to L. daqingensis but L. Basonym: Lactobacillus garii Diaz et al. 2020, 5VP.
suibiensis deaminates arginine [338]. The genome size of the L. garii grows in the temperature range of 6–42 °C and in the
type strain is 2.56 Mbp. The mol% GC content of DNA is 53.0. pH range of 4.0 to 8.8. The type strain produces D(-)-lactate
Isolated from a vegetable fermentation [338]. from a wide spectrum of pentoses, hexoses, and disaccharides
The type strain is 247-3T = NCIMB 15176T = JCM 33275T. mol% GC content of DNA is 48.3
Genome sequence accession number: RHOK00000000. Isolated from gari, a fermented cassava product [339].
16S rRNA gene accession number: MK110834. The type strain is FI11369T = NCIMB 15148T = DSM 108249T.
Genome sequence accession number: QWZQ00000000.
16S rRNA gene accession number: MN81791.
yichunensis comb. nov.
Lacticaseibacillus yichunensis (yi.chun.en´sis. N.L. masc.
adj. yichunensis, pertaining to Yichun, a city in Heilongjiang
Province, PR China). angrenensis comb. nov.
Levilactobacillus angrenensis (ang.ren.en´sis. N.L. masc. adj.
Basonym: Lactobacillus yichunensis Long et al. 2020, 14VP.
angrenensis, pertaining to Angren, a county in the Tibet
Properties of L. yichunensis are similar to L. daqingensis, but Autonomous Region, PR China, where the bacterium was
it does not grow at 37°C. [338]. The genome size of the type isolated).
strain is 2.56 Mbp. The mol% GC content of DNA is 55.4.
Basonym: Lactobacillus angrenensis Long et al. 2020, 16VP.
2848
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Funding information
Jinshui Zheng acknowledges support from the National Natural 17. Duar RM, Lin XB, Zheng J, Martino ME, Grenier T et al. Lifestyles
Science Foundation of China (NSFC) (31970003 and 31770003); Stijn in transition: evolution and natural history of the genus Lactoba-
Wittouck acknowledges stipend support from the Fonds voor Weten- cillus. FEMS Microbiol Rev 2017;41:S27–S48.
schappelijk Onderzoek - Vlaanderen. Michael Gänzle acknowledges 18. Gänzle MG. Lactic metabolism revisited: metabolism of lactic
financial support from the Canada Research Chairs Program. Sarah acid bacteria in food fermentations and food spoilage. Curr Opin
Lebeer acknowledges funding through from Flanders Innovation and Food Sci 2015;2:106–117.
Entrepreneurship (VLAIO) IWT-SBO project 150052. 19. Vandamme P, Pot B, Gillis M, de Vos P, Kersters K et al. Poly-
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We are indebted to Aharon Oren (Jerusalem, Israel) for nomenclatural
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Conflicts of interest
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The authors declare no conflict of interest. Bruno Pot is employed by
igal: prokaryotic gene recognition and translation initiation site
Yakult Europe but has no conflict of interest for this paper. identification. BMC Bioinformatics 2010;11:119.
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TAXONOMIC DESCRIPTION

Zheng et al., Int. J. Syst. Evol. Microbiol. 2020;70:2782–2858

A taxonomic note on the genus Lactobacillus: Description

Introduction anaerobic and non-­spore-­forming micro-­organisms. The

004107 © 2020 The Authors

Leuconostocaceae, including the genera Convivina, Fructoba-

only one genome, making it impossible to assess their

Phylogenomic evaluation of Lactobacillaceae and

Lactobacillus crispatus Lactobacillus gallinarum

Genome sequence accession number: AYZO00000000. 15019T=JCM 1120T=LMG 6413T=LMG 13555T=NRRL

Lactobacillus jensenii Lactobacillus kefiranofaciens

16S rRNA gene accession number: JX099549. Lactobacillus mulieris

16S rRNA gene accession number: AB107638.

Lactobacillus pasteurii The type strain is MYMRS/TLU1T = DSM 24759T=CCM

Description of Amylolactobacillus Description of Holzapfelia gen. nov.

Description of Bombilactobacillus Isolated from the digestive tract of laboratory-­

Description of Companilactobacillus Isolated from kimchi.

Effective publication Schuster et al., 2019, as Lactobacillus Description of Companilactobacillus

Description of Companilactobacillus Description of Companilactobacillus

Description of Companilactobacillus Description of Companilactobacillus

Description of Companilactobacillus Description of Companilactobacillus

Description of Lacticaseibacillus Description of Lacticaseibacillus

Description of Lacticaseibacillus Description of Lacticaseibacillus

Description of Loigolactobacillus aminopeptidase, cystine aminopeptidase, acid phosphatase,

16S rRNA gene accession number: LC258150.

Description of Liquorilactobacillus The type strain is 59bT=CECT 7334T=DSM 19972T=JCM

32230T=CIP 103144T = JCM 5666T=LMG 10753T = NBRC

Description of Lactiplantibacillus Isolated from fermented radish.

Description of Lactiplantibacillus faecal microbiota of healthy dogs, traditional fura processing,

Cells are facultatively anaerobic rods occurring singly. Growth

Description of Lactiplantibacillus 16S rRNA gene accession number: HM443954.

Description of Paucilactobacillus 16S rRNA gene accession number: AY733084.

Description of Paucilactobacillus Genome sequence accession number: AWTT00000000.

Genome sequence accession number: AZGC00000000.

Description of Limosilactobacillus Description of Limosilactobacillus

The type strain is CCUG 37482T = DSM 6035T=JCM 11053T.

Description of Levilactobacillus Description of Levilactobacillus

Description of Levilactobacillus Description of Levilactobacillus

Description of Levilactobacillus 16S rRNA gene accession number: AJ534844.

Description of Levilactobacillus Description of Levilactobacillus

Basonym: Lactobacillus tangyuanensis Long and Gu 2019, Description of Levilactobacillus

Genome sequence accession number: RHNO00000000. Description of Fructilactobacillus

Description of Fructilactobacillus Description of Fructilactobacillus

Description of Fructilactobacillus 16S rRNA gene accession number: KT783533.

Description of Apilactobacillus Description of Apilactobacillus

Description of Apilactobacillus kosoi Description of Apilactobacillus

Description of Lentilactobacillus Genome sequence accession number: AYYV00000000.

Description of Lentilactobacillus Description of Lentilactobacillus

Description of Lentilactobacillus 16S rRNA gene accession number: MK110813.

Description of Lacticaseibacillus Description of Loigolactobacillus

Find out more and submit your article at microbiologyresearch.org.

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Introduction anaerobic and non-spore-forming micro-organisms. The

Description of Bombilactobacillus Isolated from the digestive tract of laboratory-