692 FAMILY I.
ENTEROBACTERIACEAE
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(Sakazaki et al., 1989a). Apart from ornithine decarboxylase test,
these strains are biochemically undistinguishable from K. plan-
ticola, and are identical to K. planticola by DNA hybridization
(Farmer et al., 1985a; D.J. Brenner, personal communication).
Farmer et al. (1985a) questioned the validity of K. ornithinolytica
as a species distinct from K. planticola. Some ornithine decar-
boxylase positive strains have been shown to belong in K. pneu-
moniae by DNA relatedness criteria (Lindh and Frederiksen,
1990).
Finally, Calymmatobacterium granulomatis, which has not been
cultured on bacteriological media, has been transferred to the
genus Klebsiella as Klebsiella granulomatis, mostly based on rrs se-
quences (Carter et al., 1999). Partial rpoB sequencing suggested
that K. granulomatis is very close to K. pneumoniae (Drancourt et
al., 2001) and phoE sequencing showed K. granulomatis close to
K. pneumoniae subsp. rhinoscleromatis (Carter et al., 1999). Thus,
there is presently no proof that K. granulomatis is a species distinct
DIFFERENTIATION OF THE SPECIES OF THE GENUS
Table BXII.c.229 presents the characteristics differentiating the
species and subspecies of Klebsiella.
List of species of the genus Klebsiella
1. Klebsiella pneumoniae (Schroeter 1886) Trevisan 1887,
94AL (Hyalococcus pneumoniae Schroeter 1886, 1952.)
pneu.mo⬘ni.ae. Gr. n. pneumonia pneumonia, inflammation
of the lungs; M.L. gen. n. pneumoniae of pneumonia.
The characteristics are as described for the genus and
as listed in Table BXII.c.229. K. pneumoniae can be divided
into three sequence clusters (Brisse and Verhoef, 2001) un-
related to the present subdivision in subspecies. K. pneu-
moniae is normally found in the intestinal tract of humans
and animals. It may be isolated in association with several
pathological processes in humans, e.g., community-ac-
quired pneumonia or nosocomial urinary tract infection.
Serotype K2 is the most common K-type isolated from pa-
tients with urinary tract infection, pneumonia, or bacte-
remia. In animals, K. pneumoniae may be isolated from me-
tritis in mares, bovine mastitis, or generalized infections in
captive monkeys. Environmental strains generally utilize
more carbon sources than clinical strains, but often fail to
utilize d-adonitol. Intraspecies DNA relative reassociation
values among strains is 80–90% (Brenner et al., 1972c)
or 73–100% (Woodward et al., 1979).
The mol% G Ⳮ C of the DNA is: 56–58 (Tm) (Seidler et
al., 1975).
Type strain: ATCC 13883, CIP 82.9, DSM 30104, JCM
1662.
GenBank accession number (16S rRNA): X87276, Y17656,
AB004753, AF130981.
a. Klebsiella pneumoniae subsp. pneumoniae (Schroeter
1886) Trevisan 1887, 94AL (Hyalococcus pneumoniae
Schroeter 1886, 1952.)
law.
Distinguished from the subspecies subsp. ozaenae and
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Proteobacteria, Part B: The Gammaproteobacteria
Account: s5037934
from K. pneumoniae, and instead there are arguments to make it
part of K. pneumoniae.
Sequence data are increasingly used to suggest or support
nomenclatural changes and some of these changes can be an-
ticipated. The taxonomic significance of the three sequence clus-
ters observed in K. pneumoniae or the two sequence clusters in
K. oxytoca (Brisse and Verhoef, 2001) is still unknown, and DNA–
DNA hybridization data are needed before a proposal can be
made to design new species or subspecies. Since the phylogenetic
tree obtained by rrs gene (encoding 16S rRNA) sequence com-
parison showed the genus Klebsiella to be split in at least two
groups, a new genus, Raoultella, was proposed to contain R. plan-
ticola (type species), R. ornithinolytica, and R. terrigena (Drancourt
et al., 2001). More sequencing work needs to be done to evaluate
this proposal since partial rpoB sequence failed to group Raoultella
species in a single cluster and these sequences were only 512
nucleotides long.
KLEBSIELLA
subsp. rhinoscleromatis by the characteristics listed in Ta-
ble BXII.c.229. Clinical strains may produce an ex-
tended-spectrum b-lactamase.
The mol% G Ⳮ C of the DNA is: 56–58 (Tm) (Seidler
et al., 1975).
Type strain: ATCC 13883, CIP 82.9, DSM 30104, JCM
1662.
GenBank accession number (16S rRNA): X87276, Y17656,
AB004753, AF130981.
b. Klebsiella pneumoniae subsp. ozaenae (Abel 1893) Ør-
skov 1984c, 355VP (Effective publication: Ørskov 1984b,
463) (Klebsiella ozaenae (Abel 1893) Bergey, Harrison,
Breed, Hammer and Huntoon 1925, 266; Bacillus mu-
cosus ozaenae Abel 1893, 167; Bacillus ozaenae (Abel 1893)
Lehmann and Neumann 1896, 204.)
o.zae⬘nae. L. fem. n. ozaena ozena; L. gen. n. ozaenae of
ozena.
Distinguished from the subspecies subsp. pneumoniae
and subsp. ozaenae by the characteristics listed in Table
BXII.c.229. Occurs in ozena and other chronic diseases
of the respiratory tract.
The mol% G Ⳮ C of the DNA is: not available.
Type strain: ATCC 11296, CIP52.211, JCM 1663, LMG
3113.
GenBank accession number (16S rRNA): Y17654,
AF130982.
c. Klebsiella pneumoniae subsp. rhinoscleromatis (Trevisan
1887) Ørskov 1984c, 355VP (Effective publication: Ør-
skov 1984b, 464) (Klebsiella rhinoscleromatis Trevisan 1887,
95; Bacterium rhinoscleromatis (Trevisan 1887) Migula
1900, 352.)