Bergey’s Manual of Systematic Bacteriology

Bergey’s manual, which first appeared in 1923 and, at present, is in its 9th edition under the
title Bergey’s Manual of Systematic Bacteriology, is a major taxonomic treatment of bacteria
(prokaryotes). This manual has served the community of microbiologists since more than 80
years and is a compendium of information on all recognized species of bacteria (prokaryotes).

Any discussion dealing with bacterial (prokaryotic) classification cannot go without a

thorough acquaintance with this manual. Each chapter of this treatise, written by an expert,
contains tables, figures, and other systematic information useful for identification of bacteria.

Many schemes for identification of bacteria have been devised prior to 1923 but all were
usually fragmentary. There was need for a single scheme which could cover all the described
bacteria. David Hendricks Bergey, a professor of bacteriology at the university of
Pennsylvania (USA), proceeded in this direction and began preparing a complete review of
the enormous literature of bacterial taxonomy.

To aid the publication of this work, the Society of American Bacteriologists (now called the
American Society of Microbiologists) appointed an Editorial Board headed by Bergey. This
resulted in the publication of the first edition of Bergey’s Manual of Determinative
Bacteriology in 1923. The second edition of the manual was published in 1925 and the third
edition in 1930.

In 1934, the Society of American Bacteriologists transferred to Dr. Bergey all its rights, title,
and interests in the Manual in order to allow Bergey to create an independent, non-profit
trust, namely, The Bergey’s Manual Trust. Throughout the years, this trust continues to
prepare and publish successive editions of the manual and promotes research in the field of
bacterial taxonomy.

The first eight editions of this manual appeared under the title ‘Bergey’s Manual of
Determinative Bacteriology’. The 9th edition was retitled and was published as the 1st edition
under the title. ‘Bergey’s Manual of Systematic Bacteriology’, which consisted of four
volumes published in 1984, 1986, 1989, and 1991, respectively. This infect was the first
edition of the Manual under the changed title.

The change of the title of the manual from determinative bacteriology to systematic
bacteriology indicates that manual’s usefulness is no longer restricted to a determinative role,
i.e., mere identification, but it is now aimed at a systematic classification of bacteria.
However, Bergey’s Manual of Systematic Bacteriology has appeared in the form of its 2nd
edition consisting of five volumes; its first volume was published in 2001, second in 2005
and three additional volumes expected shortly.

Bergey’s Manual of Systematic Bacteriology (First Edition):

Phenetic Classification:
The first edition of Bergey’s Manual of Systematic Bacteriology is primarily phenetic, i.e.,
based on the natural similarity of phenotypic characteristics of microorganisms, and has
appeared in four volumes consisting of sections.

The four volumes, their year of publication, and the sections and the groups of bacteria
included in each of them are the following:
Vol. I: 1984 (sections 1-11): Gram-negative bacteria of general, medical, or industrial
importance.

Vol. II: 1986 (sections 12-17): Gram-positive bacteria other than actinomycetes.

Vol. Ill: 1989 (sections 18-25): Gram-negative bacteria with distinctive properties,
cyanobacteria, and archaea.

Vol. IV: 1991 (sections 26-23): Actinomycetes (Gram-positive filamentous bacteria).

All prokaryotes were retained in this edition in a single kingdom Prokaryotiae, divided into
four divisions called Gracilicutcs, Firmicutes, Tenericutes, and Mendosicutes.

Despite many other differences in characteristics among the divisions, the Gracilicutes (thin
skin) possess gram-negative cell wall; the Firmicutes (thick and strong skin) have gram-
positive cell wall; the tenericutes (soft or tender skin) lack cell wall and represented by
mycoplasmas; and the Mendosicutes (skin with faults) accommodating archaeobacteria that
lack conventional peptidoglycan.

Each of the 33 sections in the four volumes contain prokaryotes that share a few easily
determined characteristics and bears a title that either describes these properties or provides
the vernacular names of the prokaryotes included.

The characteristics used to define sections are normally features such as general shape and
morphology, Gram-staining properties, oxygen relationships, motility, the presence of
endospores the mode of energy production, and so forth.
Bergey’s Manual of Systematic Bacteriology (Second Edition):

Phylogenetic Classification:
Prokaryotic taxonomy enjoyed enormous progress after the publication of the first edition of
Bergey’s Manual of Systematic Bacteriology. It became possible with the aid of newer
molecular techniques such as the sequencing of ribosomal RNA (rRNA), DNA, and proteins.
These techniques has made phylogenetic analysis of prokaryotes practicable.

In the light of the availability of considerable knowledge of phylogenetic relationships

amongst prokaryotes, the second edition of Bergey’s Manual of Systematic Bacteriology is
largely phylogenetic rather than phenetic.

As a result, the second edition is quite different from the first edition in the characteristics
chosen as basis for classification. However, the second edition consists of five volumes. Its
first volume was published in 2001, second in 2005, and three additional volumes expected
shortly.

The details of the five volumes are the following:

Vol. I: The Archaea, and the Deeply Branching and Phototropic Bacteria

Vol. II: The Proteobacteria

Vol. III: The Low G + C Gram-positive Bacteria (Fermicutes)

Vol. IV: The High G + C Gram-positive Bacteria (Actinobacteria)

Vol. V: The Planctomycetes, Spirochaetes, Fibrobacteres, Bacteriodetes, and Fusobacteria

(Vol. V also contains a section accommodating the update descriptions and phylogenetic
arrangements that have been revised since publication of volume I)

A Concise Account of the Classification:

Volume I. The Archaea and the Deeply Branching and Phototrophic Bacteria:
Domain Archaea:

Phylum AI: Crenarchaeota:

The phylum Crenarchaeota contains thermophilic and hyperthcrmophilic sulfur-metabolizing
prokaryotes arranged in single class Thermoprotei divided into three orders and five families.
The important genera are Thermoproteus, Desulgurococcus, Pyrodictium, and Sulfolobus.
Phylum A II: Euryarchaeota:
The phylum Euryarchaeota contains primarily methanogens and halophiles; thermophilic,
sulfur-reducers also are included in this phylum. This phylum is divided into seven classes
(Methanobacteria, Methanococci. Halobacteria, Thermoplasmata, Thermococci, Archeoglobi,
and Methanopyri), nine orders, and sixteen families.

The representative genera of each class are Methanobacterium, Methanococcus,

Halobacterium, Thermophasma, Thermococcus, Archaeoglobus, and Methanopyrus,
respectively.

Domain: Bacteria:

Phylum BI. Aquificae:

The phylum Aquificae has one class (Aquificae) that contains autotrophic bacteria possessing
ability to use hydrogen for energy production. Aquifex and Hydrogenobacter are the
representative genera. Aquifex means “water maker” as this genus reduces oxygen by using
hydrogen and produces water.

Phylum BII: Thermotogae:

The phylum thermotogae consists of one class (Thermotogae). It contains anaerobic,
thermophilic, fermentative, gram-negative bacteria. The representative genera are
Thermotoga, Geotoga, Fervidobacterium, etc.

Phylum Bill: Thermodesulfobacteria:

This phylum contains one class (Thermodesulfobacteria) and only two genera,
Thermodesulfobacterium and Thermodesulfatator. The bacteria are anaerobic, thermophilic,
and sulfate-reducing.

Phylum BIV: Dienococcus-Thermus:

The phylum contains one class (Dienococci). The representative genera are Dienococcus,
Thermus, Marinithermus, etc. The bacteria are gram-positive and extraordinarily radiation
resistant.

Phylum BVI: Chloroflexi:

The phylum Chloroflexi consists of two classes (Chloroflexi and Anaerolieae). Many
members of this phylum are gram-negative and are called green nonsulfur bacteria. The
important genera are Chloroflexus, Oscillochloris, Herpetosiphon and Anaerolinea.
Phylum BVII: Thermomicrobia:
The phylum contains one class (Thermomicrobia) and is represented by a single genus,
Thermomicrobium. Thermomicrobium is an aerobic thermophilic chemotroph possessing
unusual lipids that contain 1, 2-dialcohols instead of glycerol, and have neither ester nor ether
linkages.

This is in contrast to the lipids of Bacteria and Eukarya that contain fatty acids esterified to
glucose. Also, the cells of Thermomicrobium differ from those of Bacteria in that they lack
peptidoglycan in their cell walls.

Phylum B VIII: Nitrospirae:

The phylum consists of one class (Nitrospira) and genera like Nitrospira and Leptospirillum.

Phylum BIX: Deferribacteres:

The phylum Deferribacteres contains one class (Deferribacteres) and genera such as
Deferribacter, Denitrovibrio, etc.

Phylum BX: Cyanobacteria:

The phylum Cyanobacteria consists of a single class (Cyanobacteria) divided into five
subsections. The taxonomic position of the cyanobacteria is left open in Bergey s Manual.
The cyanobacteria are oxygenic phototrophs and show a distant relationship to gram-positive
bacteria.

These organisms were the first oxygen-evolving phototrophs on Earth and were responsible
for the conversion of the anoxic atmosphere of our planet to oxic. Genera belonging to this
phylum are mentioned as “form genus” in the Manual because the latter refers to a group of
cyanobaceria with very characteristic morphology found worldwide but not all isolates of
such a type may actually fit into the same genus.

The important “form genera” are Microcytis, Cyanocytis, Lyngbya, Oscillatoria, Spirulina,
Anabaena, Nostoc, Scytonema, Calothrix, Tolypothrix, etc.

Phylum B XI. Chlorobi:

Only one class (Chlorobia) constitutes the phylum. The representative genera are
Chlorobium, Pelodictyon, Chlorobaculum, etc. This phylum accommodates anoxygenic
photosynthetic bacteria known as the green sulfur bacteria, which can assimilate CO2 through
the reductive (reverse) tricarboxylic acid cycle rather than calvin cycle and oxidize sulfide to
sulfur granules that accumulate outside the cell.
Volume II:

The Proteobacteria:

Phylum B XII: Proteobacteria:

The phylum Proteobacteria consists of five classes (Alfaproteobacteria, Betaproteobacteria,
Gammaproteobacteria, Deltaproteobacteria, and Epsilonproteobacteria).

Proteobacteria contains over 400 genera which are all gram-negative, show extreme
metabolic diversity, and represent the majority of bacteria of pharmaceutical, industrial,
agricultural, and biological research significance. It is believed by many that the
proteobacteria evolved from a photosynthetic ancestor and, presumably, many strains lost
photosynthetic activity during adaptation to new ecological niches.

Class Alfaproteobacteria
Alphaproteobacteria (∝-proteobacteria) consist of the most bacteria that are capable of
growing at low nutrient levels (i.e., oligotrophic forms); the representative genera of this class
are Rhodospirillum, Azospirillum, Rickettsia, Rhizobium, Agrobacterium, Nitrobacter,
Hyphomicrobium, Methylobacterium, etc.

Class Betaproteobacteria
Betaproteobacteria (β-proteobacteria) utilize the substances that diffuse from organic
decomposition in the anaerobic zone of their habitats. Some of its genera are pathogenic.

The important genera included in betaprotobacteria are Thermothrix, Bordetella, Leptothrix,

Neissaria, Aquaspirillum, Nitrosomonas, Gallionella, Spirillum, etc.

Class Gammaproteobacteria
Gammaproteobacteria (γ-proteobacteria) are chemoorganotrophic, facultative anaerobic, and
the representative genera are Chromatium, Xanthomonas, Beggiatoa, Pseudomonas, Vibrio,
Photobacterium, Escherichia, Klebsiella, Erwinia, etc.

Class Deltaproteobacteria
Deltaproteobacteria (δ-proteobacteria) can mainly be categorized as of three groups. Many of
the deltaproteobacteia are anaerobic and cause desulfurication (generate sulfide from sulfate
and sulfur); representative genera of such bacteria are Desulfovibrio, Deslfuromonas,
Desulfobaca, etc. Some deltaproteobacteria (e.g., Bdellovibrio, Bacteriovorax) are predators
on other bacteria.
Representatives such as Myxococcus, Polyangium and Chondromyces are the
deltaproteobacteria that form fruiting bodies and also prey on other bacteria.

Class Epsilonproteobacteria
Epsilonproteobacteria (Ɛ- proteobacteria) is a small class and its members like Helicobacter
and Campylobacter are important human pathogens causing intestinal infections.

Volume III:

Low G + C Gram-Positive Bacteria:

Phylum B XIII: Firmicutes:

Phylum Firmicutes consists of the gram-positive bacteria with low G + C content (mol%
value below 50%) in their DNA. Most of the bacteria of this phylum are heterotroptic.
Though mycoplasmas lack cell wall and stain gram-negative, they are classified under this
phylum because of their close relationship to low G + C gram-positive bacteria. The phylum
is classified into three classes, Clostridia, Mollicutes, and Bacilli.

Class Clostridia contains a very wide variety of gram-positive bacteria that vary in
morphology and size but tend to be anaerobic. Some produce endospores but others do not.
Important genera are Clostridium, Acetobacterium, Desulfotomaculum, Eubacterium,
Heliobacterium, Syntrophomonas, etc.

Class Mollicutes consists of the members that are wall-less and are generally called
mycoplasmas. These bacteria are pleomorphic in shape, normally nonmotile, stain gram-
negative due to absence of cell wall, and require sterols for growth. The important genera
include Mycoplasma, Phytoplasma, Spiroplasma. that contain several important animal and
plant pathogens.

Class Bacilli comprises a variety of bacteria which are aerobic or facultatively anaerobic
cocci and rods, and many of them are endospore-forming (e.g., Bacillus, Sporosarcina,
Paenibacillus). Many genera of this phylum are pharmaceutical and industrially significant
(e.g., Bacillus, Lactobacillus, Enterococcus, Leuconostoc, Streptococus, Lactococcus).

Volume IV:

High G + C Gram-Positive Bacteria:

Phylum B XIV: Actinobacteria:

The phylum Actinobacteria is classified into a single class Actinobacteria which contains
high G + C gram-positive bacteria with mole% value above 50 to 55%. These bacteria show
enormous morphological variations ranging from cocci to regular or irregular rods to
extensively branching hyphae. Endospore formation lacks but many genera do form a variety
of asexual spores.

The phylum consists of five subclasses, six orders, forteen suborders, forty four families, and
many a number of genera.

The important genera of actinobacteria are Actinomyces, Arthrobacter, Brevibacterium,

Cellulomonas, Clavibacter, Corynebacterium, Mycobacterium, Nocardia, Actinoplanes,
Propionibacterium, Streptomyces, Frankia, etc. The largest and the most complex genes is
Streptomyces containing over 500 species.

Volume V:

Plantomycetes, Spirochaetes, Fibrobacters, Bacteroidetes, and Fusobacteria:

Phylum B XV: Planctomycetes:

This phylum consists of a single class (Planctomycetacia) containing four genera named
Planctomyces, Gemmata, Pirellula and Isosphaera. Members lack peptidoglycan and their
cell walls are of an S-layer type consisting of protein. They divide by budding and may
produce non- prosthecate appendages called stalks.

Phylum B XVI: Chlamydias:

Phylum Chlamydiae contains a single class (Chlamydiae) and seven genera. The genus
Chlamydia is by far the most important genus; its species cause many human diseases.

Phylum B XVII: Spirochaetes:

Phylum Spirochaetes is classified into one class (Spirochaetes) containing thirteen genera, the
important ones are Spirochaeta, Borellia, Treponema, Cristispira, Leptospira, etc. They are
gram-negative, motile, helically-shaped bacteria characterized by a unique motility
behaviour.

Phylum B XVIII: Fibrobacteres:

This phylum is represented by a single class (Fibrobacteres) and a single genus (Fibrobacter).
Fibrobacter is a cellulolytic gram-negative bacterium occurring in the rumen of ruminant
animals where it breaks down cellulose.

Phylum B XIX: Acidobacteria:

The phylum contains only one class (Acidobacteria) represented by three genera named
Acidobacterium, Geothrix and Holophaga.
Phylum B XX: Bacteroidetes:
The phylum is divided into three classes (Bacteroides, Flavobacteria, and Sphingobacteria)
exemplified by genera like Bacteroides, Flavobacterium, Flexibacter, Cytophaga, Toxothrix,
etc. Flexibacter and Cytophaga are gliding bacteria and are ecologically significant.

Phylum B XXI: Fusobacteria:

The phylum contains one class (Fusobacteria) with genera like Fusobacterium, Leptotrichia,
and Cetobacterium. Fusobacterium is filamentous and occurs in the oral cavity of humans.

Phylum B XXII: Verrucomicrobia:

This phylum consists of a single class (Verrucomicrobiae) and genera such as
Verrucomicrobium, Prosthecobacter, and Xiphinematobacter. Members form cytoplasmic
appendages called prosthecae and divide symmetricaly.

Phylum B XXIII: Dictyoglomus:

The phylum is represented by a single class (Dictyoglomi) consisting of a single genes named
Dictyogomus.