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Cyanobacteria
From Basic Science to Applications

Cyanobacteria
From Basic Science to Applications
Edited by
A.K. Mishra, D.N. Tiwari
Department of Botany
Banaras Hindu University
Varanasi, India
A.N. Rai
Department of Biochemistry
North-Eastern Hill University
Shillong, India

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v
v
In Memory of
Professor Ram Nagina Singh, our revered teacher and
an internationally recognized authority on
phycology,
served the Banaras Hindu University as a faculty mem-
ber and superannuated as Professor and Head of the
Department of Botany in August 1976. He has been
credited with several original research contributions in
green algae and cyanobacteria (blue-green algae), most
of which have been published in reputed international
journals. Initially his research activities focused on tax-
onomy, life history, and cytology of certain green algae
(e.g., Fritschiella tuberosa and Draparnaldiopsis indica).
Subsequently, he focused his work on the role of nitrogen-fixing cyanobacteria in
nitrogen economy of Indian agriculture, particularly the cultivation of rice. He also
examined the application of diazotrophic cyanobacteria in reclamation of alkaline
“Usar” soils as elaborated in his book, “Role of Blue-Green Algae in Nitrogen Econ-
omy of Indian agriculture” (1961). During the 1960s and 1970s, he and his research
team researched and published extensively on physiology, cellular differentiation,
radiation biology, and genetics of cyanobacteria as well as cyanophages. During
this period, he and his associates produced several mutant strains of cyanobacteria
which have interesting implications for understanding of their morphogenesis. He
also worked in several famous labs abroad including University College London
(with F.E. Fritsch), University of Wisconsin (with R.H. Burris and Hans Ris), and
Michigan State University (with C.P. Wolk), and published original research
papers
on nitrogen fixation in cell free extracts of Mastigocladus species and confirmation
of prokaryotic cellular organization of cyanobacteria with the help of electron mi-
croscopic studies. Besides being an active researcher, Professor Singh was also an
outstanding teacher of microbiology, plant physiology, and phycology. It was a great
loss to botany, particularly phycology, when he suddenly expired just after 7 months
of his superannuation. We and others being his students always remember him with
great affection and gratitude.
Prof. Ram Nagina Singh
(1915–77)

xv
Contributors
Numbers in parentheses indicate the pages on which the authors’
contributions begin.
Gerard Abraham (347), Division of Microbiology, ICAR-
Indian Agriculture Research Institute, New Delhi, India
Haseen Ahmed (145), Laboratory of Photobiology and
Molecular Microbiology, Centre of Advanced Study in
Botany, Institute of Science, Banaras Hindu University,
Varanasi, India
N. Anand (43), Centre for Advanced Studies in Botany,
University of Madras, Chennai, India
Francisco Barona-Gómez (29), Evolution of Metabolic
Diversity Laboratory, Langebio, Cinvestav-IPN,
Guanajuato, Mexico
Edder D. Bustos-Díaz (29), Evolution of Metabolic
Diversity Laboratory; Ecological and Evolutionary
Genomics Laboratory, Langebio, Cinvestav-IPN,
Guanajuato, Mexico
Liliana Cepoi (217), Institute of Microbiology and
Biotechnology, Chisinau, Moldova
Jan Červený (193), Department of Adaptive
Biotechnologies, Global Change Research Institute,
Czech Academy of Sciences, Brno, Czech Republic
Sindhunath Chakraborty (383), Centre of Advanced
Study in Botany, Institute of Science, Banaras Hindu
University, Varanasi, India
Angélica Cibrián-Jaramillo (29), Ecological and
Evolutionary Genomics Laboratory, Langebio,
Cinvestav-IPN, Guanajuato, Mexico
Samuel Cirés (399), Department of Biology, Universidad
Autónoma de Madrid, Madrid, Spain
P.K. Dadheech (43), Department of Microbiology, School
of Life Sciences, Central University of Rajasthan,
Bandarsindri, Kishangarh, India
Josef Elster (277), Faculty of Sciences, University of
South Bohemia, České Budějovice; Institute of Botany,
Academy of Sciences of the Czech Republic, Třeboň,
Czech Republic
Francisca Fernández-Piñas (399), Department of Biology,
Universidad Autónoma de Madrid, Madrid, Spain
Lira A. Gaysina (1), Department of Bioecology and
Biological Education, M. Akmullah Bashkir State
Pedagogical University, Ufa; All-Russian Research
Institute of Phytopathology, Moscow, Russian
Federation
Cátia F. Gonçalves (359), i3S—Instituto de Investigação
e Inovação em Saúde; IBMC—Institute for Molecular
and Cell Biology, University of Porto, Porto, Portugal
Miguel González-Pleiter (399), Department of Biology,
Govindjee (79), Departments of Biochemistry and Plant
Biology, and Centre of Biophysics and Quantitative
Biology, University of Illinois at Urbana-Champaign,
Urbana, IL, United States
Donat-P. Häder (145), Emeritus from Friedrich-Alexander
University, Department of Biology, Möhrendorf,
Germany
Jara Hurtado-Gallego (399), Department of Biology,
Vinod K. Kannaujiya (207), Department of Botany, MMV,
Banaras Hindu University, Varanasi, India
Chandra Kant (347), Department of Botany, Dharam
Samaj College, Aligarh, Uttar Pradesh, India
Manish Singh Kaushik (173, 245), Laboratory of
Microbial Genetics, Centre of Advance Study
in Botany, Institute of Sciences, Banaras Hindu
Surbhi Kharwar (303), Laboratory of Microbial Genetics,
Centre ofAdvance Study in Botany, Institute of Sciences,
Jiří Komárek (277), Institute of Botany, Academy of
Sciences of the Czech Republic, Třeboň, Czech
Republic
O. Konur (419), Materials Engineering, Ankara Yildirim
Beyazit University, Ankara, Turkey

xvi Contributors
Vinod Kumar (131), Centre of Advanced Study in Botany,
Institute of Science, Banaras Hindu University,Varanasi,
India
Deepak Kumar (207), Laboratory of Photobiology and
Botany, Banaras Hindu University, Varanasi, India
Jay Kumar (327), School of Biotechnology, Institute of
Science, Banaras Hindu University, Varanasi, India
Ashok Kumar (327), School of Biotechnology, Institute of
Jana Kvíderová (277), Faculty of Sciences, University of
South Bohemia, České Budějovice, Czech Republic
Dušan Lazár (79), Department of Biophysics, Centre of
the Region Haná for Biotechnological and Agricultural
Research, Faculty of Science, Palacký University,
Olomouc, Czech Republic
Francisco Leganés (399), Department of Biology,
Steeve Lima (359), i3S—Instituto de Investigação e
Inovação em Saúde; IBMC—Institute for Molecular
and Cell Biology; ICBAS—Abel Salazar Institute
of Biomedical Sciences, University of Porto, Porto,
Portugal
Iris Maldener (65), Interfaculty Institute of Microbiology
and Infection Medicine Tübingen, Organismic
Interactions,UniversityofTübingen,Tübingen,Germany
Pankaj K. Maurya (131), Centre of Advanced Study in
Varanasi, India
Shashank Kumar Maurya (477), Biochemistry and
Molecular Biology Laboratory, Department of Zoology,
India
Arun Kumar Mishra (173, 245), Laboratory of Microbial
Genetics, Centre of Advance Study in Botany, Institute
of Sciences, Banaras Hindu University, Varanasi, India
Rajnikant Mishra (477), Biochemistry and Molecular
Biology Laboratory, Department of Zoology, Institute
of Science, Banaras Hindu University, Varanasi, India
Soumila Mondal (131), Centre of Advanced Study in
Varanasi, India
František Muzika (193), Department of Chemical
Engineering, University of Chemistry and Technology,
Prague, Czech Republic
Niveshika (477), Laboratory of Microbial Genetics, Centre
of Advance Study in Botany, Institute of Science,
Paulo Oliveira (359), i3S—Instituto de Investigação e
and Cell Biology, University of Porto, Porto, Portugal
George C. Papageorgiou (79), Institute of Biosciences and
Applications, National Centre for Scientific Research
Demokritos, Attikis, Greece
Parul Parihar (261), Ranjan Plant Physiology and
Biochemistry Laboratory, Department of Botany,
University of Allahabad, Allahabad, India
Anuradha Patel (261), Ranjan Plant Physiology and
Jainendra Pathak (145, 207), Laboratory of Photobiology
and Molecular Microbiology, Centre of Advanced
Sheo Mohan Prasad (261), Ranjan Plant Physiology
and Biochemistry Laboratory, Department of Botany,
A.N. Rai (459), Biochemistry Department, North-Eastern
Hill University, Shillong, India
Yattapu Prasad Reddy (347), Division of Microbiology,
ICAR-Indian Agriculture Research Institute, New
Delhi, India
Jacqueline Rücker (65), Department of Freshwater
Conservation of the Brandenburg University of
Technology, Cottbus-Senftenberg, Germany
David Šafránek (193), Systems Biology Laboratory,
Masaryk University, Brno, Czech Republic
Jakub Šalagovič (193), Systems Biology Laboratory,
Masaryk University, Brno, Czech Republic
Aniket Saraf (1), Department of Botany, Ramniranjan
Jhunjhunwala College, Mumbai, India
IgorSchreiber(193),DepartmentofChemicalEngineering,
University of Chemistry and Technology, Prague, Czech
Republic
Prashant Singh (1), Department of Botany, Institute of
Shailendra P. Singh (131, 145), Centre of Advanced
Prashant R. Singh (145), Laboratory of Photobiology and
Varanasi, India
Rachana Singh (261), Ranjan Plant Physiology and

Contributors xvii
Divya Singh (327), School of Biotechnology, Institute of
Savita Singh (347), Department of Botany, Babu Shivnath
Agrawal College, Mathura, India
Satya Shila Singh (383), Centre of Advanced Study in
Varanasi, India
A.K. Singh (459), Biochemistry Department, North-
Eastern Hill University, Shillong, India
Rajeshwar P. Sinha (131, 145, 207), Laboratory of
Photobiology and Molecular Microbiology, Centre of
Advanced Study in Botany, Institute of Science, Banaras
Hindu University, Varanasi, India
Meenakshi Srivastava (245), Laboratory of Microbial
Genetics, Centre of Advance Study in Botany, Institute
of Sciences, Banaras Hindu University, Varanasi, India
Alexandrina Stirbet (79), Newport News, VA, United States
Assaf Sukenik (65), TheYigalAllon Kinneret Limnological
Laboratory, Israel Oceanographic and Limnological
Research, Migdal, Israel
M.B. Syiem (459), Biochemistry Department, North-
Eastern Hill University, Shillong, India
Paula Tamagnini (359), i3S—Instituto de Investigação e
and Cell Biology; Faculty of Sciences, Department of
Biology, University of Porto, Porto, Portugal
N. Thajuddin (43), National Repository for Microalgae
and Cyanobacteria-Fresh Water, Department of
Microbiology, School of Life Sciences, Bharathidasan
University, Tiruchirappalli, India
D.N. Tiwari (173, 303), Laboratory of Microbial Genetics,
Centre ofAdvance Study in Botany, Institute of Sciences,
Sanjesh Tiwari (261), Ranjan Plant Physiology and
Balkrishna Tiwari (303), Laboratory of Microbial
Genetics, Centre of Advance Study in Botany,
Institute of Sciences, Banaras Hindu University,
Varanasi, India
Madhu B.Tyagi (327), Botany Department, MMV, Banaras
Hindu University, Varanasi, India
David Velázquez (399), Department of Biology,
Ekta Verma (383), Centre of Advanced Study in Botany,
India
Ravindra Kumar Yadav (347), Department of
Environmental Studies, University of Delhi, New Delhi,
India
Joao SarkisYunes (443), Federal University of Rio Grande,
Rio Grande, Brazil

xix
Preface
Cyanobacteria are a unique group of gram-negative prokaryotes having oxygenic photosynthesis and have the distinc-
tion of being the oldest known fossils, more than 3.5 billion years old. Many of them also carry out N2 fixation. They are
considered as the Earth’s architect because their oxygen evolving property has led to the formation of oxygen atmosphere
on this planet. They are thought to be the ancestors of plant chloroplast with the help of which plants make food for them-
selves. Despite being prokaryotic in nature, cyanobacteria exhibit a number of similarities with eukaryotic algae, hence are
often called “blue-green algae.” These organisms have been tremendously important in shaping the course of evolution and
ecological change throughout the Earth’s history. They occur as free-living forms as well as in a wide range of symbioses
with plants ranging from algae to angiosperms. The role of N2-fixing cyanobacteria as natural biofertilizers, particularly in
rice fields, as well as their application in reclamation of alkaline usar soils are well documented. Cyanobacteria are able to
coordinate and synchronize different bioenergetic and metabolic processes using temporal and spatial separations as a strat-
egy within and between their extremely small sized cells. Cyanobacteria also have been found to be an important source
of various bioactive compounds having pharmacological, industrial, and biotechnological applications. Cyanobacteria are
frequently used as bioremediation agents for the removal of toxic pollutants from the environment because they inhabit a
number of polluted sites and exhibit increased tolerance against those toxicants.
Considering these interesting aspects of their biology and their spectacular contributions to the environment as well as
human welfare, we have compiled this book which covers various important facts about cyanobacteria starting from diver-
sity, taxonomy, and distribution to their metabolic regulation and applications. This book logically explores different top-
ics related to cyanobacteria in well documented and profusely illustrated 24 individual chapters contributed by nationally
and internationally recognized authors having excellence in teaching and research in the field of cyanobacteria and plant
science. The book comprises details of cyanobacterial diversity, molecular taxonomy, the molecular basis underlying the
development and function of specialized structures such as heterocytes and akinetes, biochemical and physiological back-
ground of nitrogen fixation and hydrogen production, occurrence of cyanobacteria as symbionts and their ecophysiology
in extreme polar habitats. Moreover, the book is also enriched by skillful description about cyanobacterial photosynthesis,
photomorphogenesis, and photoprotection mechanisms along with siderophore dependent and independent strategies of
iron homeostasis. Besides, a significant portion of the book is dedicated to elucidate the cellular mechanisms of cyanobac-
teria to cope up with various environmental stresses (such as metal and pesticide toxicity), their plant growth promoting
properties, their secretion systems, commercial importance of their exopolysaccharides and phycobiliproteins, their poten-
tial as bioenergy and biofuel resources, and their biotechnological applications. Monitoring ecotoxicity of antibiotics in
fresh water using cyanobacteria and the inside story of cyanobacterial toxins are also catalogued in this book.
We hope that this edited volume will serve as a valuable resource to the undergraduate and postgraduate students as well
as researchers working on cyanobacteria. Using this book, teaching institutions should find it helpful in framing up to date
syllabi on cyanobacteria.
We affirm our indebtedness to all the contributors for their cooperation and valuable contributions without which this
book would have not been possible. We sincerely thank Dr. P.K. Singh, FNA, Prof. S.P. Singh, Prof. B.R. Chaudhary, Dr.
S.S. Singh (Banaras Hindu University), and Dr. Piyush Chaturvedi (Cambridge University) for their valuable suggestions.
We owe our deep felt gratitude and affection to our families whose blessings and supports inspired us to complete this
task. We also thank Head, Department of Botany, BHU, for providing necessary facilities.
A.K. Mishra
D.N. Tiwari
A.N. Rai

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1
Cyanobacteria. https://doi.org/10.1016/B978-0-12-814667-5.00001-5
© 2019 Elsevier Inc. All rights reserved.
Cyanobacteria in Diverse Habitats
Lira A. Gaysina⁎,†
, Aniket Saraf‡
, Prashant Singh§
⁎
Department of Bioecology and Biological Education, M. Akmullah Bashkir State Pedagogical University, Ufa, Russian Federation, †
All-Russian
Research Institute of Phytopathology, Moscow, Russian Federation, ‡
Department of Botany, Ramniranjan Jhunjhunwala College, Mumbai, India,
§
Department of Botany, Institute of Science, Banaras Hindu University, Varanasi, India
1. INTRODUCTION
Cyanobacteria are oxygen evolving, gram-negative, photoautotrophic bacteria that can be found in virtually all imaginable
habitats across the world. They can be found in habitats extending from the extremely cold deserts of the Arctic and
Antarctic Zones (Mataloni and Komárek, 2004) to the very extreme hot springs of the Yellowstone National Park, thus
representing an interesting form of life in a wide range of aquatic and terrestrial environments (Whitton and Potts, 2000a,b).
This extraordinary expanse of cyanobacteria is actually a result of an important character that the cyanobacteria can boast of
which we refer to as “adaptability.” Their amazing success in adapting to such diverse and varying habitats can be attributed
to a very long-standing evolutionary history that cyanobacteria enjoy. Some of the modern physiological and anatomical
features of cyanobacteria that have helped in supporting their long history on Earth include an extremely high tolerance
level of low oxygen and free sulfide, the ability to utilize H2S as a photoreductant in place of H2O, and the strong tolerance
capacity for ultraviolet B and C radiations (Whitton and Potts, 2000a,b). Thus, these are some of the major adaptive
features that have allowed the cyanobacteria to grow, flourish, and dominate in habitats where many other forms of life are
absolutely unknown or may struggle to even survive.
Interestingly, the origin of cyanobacteria and their basic anatomical features are typical bacterial type but their
ecological, biological, and morphological features are quiet specific and diverse too (Flores et al., 2006; Kalaitzis et al.,
2009; Flores and Herrero, 2010). The almost essential possession of some interesting features like the ability of buoyancy,
performing oxygenic photosynthesis, and fixation of atmospheric nitrogen (Walsby, 1994; Castenholz, 2001; Burja et al.,
2001; Berman-Frank et al., 2003) make the cyanobacteria more special and capable to adapt to varied conditions in both
normal and stressed habitats. As a consequence of such strong survival capabilities, they also act as prominent phototrophic
primary producers in natural ecosystems (Field et al., 1998; Bryant, 2003).
Cyanobacteria have played a major role in the evolution of different forms of life on Earth. The anaerobic earth was
converted to the aerobic one by the cyanobacteria about 2 billion years ago, thus finally supplying oxygen to the atmosphere
through oxygenic photosynthesis for approximately 1.5 billion years (Berman-Frank et al., 2003; Schopf et al., 1983). This
event of oxygenation of the Earth’s atmosphere allowed for an enormous degree of greater diversification of many forms of
life on the Earth. Apart from the oxygenation, cyanobacteria also influenced the advancement of life on Earth by shaping up
themselves as the progenitors of chloroplasts (Giovannoni et al., 1988; Sergeev et al., 2002), ultimately leading to a broader
diversification of algae and land plants.
In the current times, the cyanobacteria still continue to affect the life on Earth, as the major oxygen producers on this
planet. Apart from being the major oxygen producers, many of them are also efficient fixers of atmospheric nitrogen thus
making them extremely important components of nutrient-deprived ecosystems and extreme environments all throughout
the Earth. As a result of their high adaptive capacity, cyanobacteria are also well known to form a wide range of nitrogen-
fixing symbiotic relationships with representatives from almost all plant groups such as algae (Hemiaulus hauckii-Richelia
intracellularis) fungi (Geosiphon pyriforme-Nostoc), bryophytes (Anthoceros-Nostoc), pteridophytes (Azolla-Anabaena),
gymnosperms (Cycas-Nostoc), and angiosperms (Gunnera-Nostoc), thus supplying the host organisms with a lot of fixed
nitrogen and in return, acquiring carbohydrates from the host (Morot-Gaudry and Touraine, 2001), thus allowing the host
organisms to grow in regions where they may otherwise not survive at all.
Cyanobacterial diversity owing to their adaptive capacities occupies incredibly large canvas of the planet Earth.
The expanse of their diversification spreads in almost all the habitable ecosystems on the Earth. Along with having an
Chapter 1

2 Cyanobacteria

omnipresent occurrence with large ecological benefits, cyanobacteria are also one of the most important contributors to the
global nitrogen fixation and carbon fixation chains (Karl et al., 2002). Their physiological plasticity and level of flexibility
are of scales that are enormous in magnitude, thus enabling them to be active representatives of virtually all geographical
sections on the planet Earth (Kol, 1968; Castenholz, 1973; Whitton, 1973; Van Landingham, 1982; Reed et al., 1984; Kann,
1988; Dor and Danin, 1996; Laamanen, 1996; Skulberg, 1994; Weber et al., 1996). Many of them are luxuriantly growing
freshwater forms, but many of the cyanobacterial taxa thrive equally well in marine environments. Terrestrial habitat and
biological crusts are indeed common for the majority of the strains of the cyanobacteria along with a strong presence in
the benthic habitats. Due to their extraordinary physiologically adaptive capacity, cyanobacteria are also equally proficient
in forming frequent and useful endophytic and symbiotic associations (Rai, 1990; Adams, 2000; Bergman et al., 2007;
Thajuddin et al., 2010). They are known to form biofilms and microbial mats on shores, surface of the stones, plants,
and artificial objects especially in extreme habitats (Stal, 2000). The frequent and heavy occurrence of cyanobacteria in
freshwater bodies (Fig. 1A–C), seepages, rocky crevices, the bark of trees, rice fields, within limestone, salt subjugated
lands, marine ecosystems, deserts, polar environments (Fig. 1D–F), biological soil crusts (BSCs) (Fig. 1H and I), soil
surface (Fig. 1J), and in symbiotic associations, thus emphasizing their ability to survive and flourish in the above niches
with ease and competence (Whitton and Potts, 2000a,b; Elster et al., 1999).
However, in spite of their importance in many ecosystems, cyanobacteria have been neglected in many biodiversity
studies (Rejmánková et al., 2004). The hidden biodiversity of cyanobacteria is very high, especially in understudied and
less explored geographic regions (Nabout et al., 2013; Gaysina et al., 2018). As morphological features have traditionally
been the main criterion for classifying and identifying cyanobacteria, most of the studies carried out to date relied almost
exclusively on less discriminative morphology-based methods (Alvarenga et al., 2015).
2. CYANOBACTERIA IN AQUATIC HABITATS
2.1 Marine Cyanobacteria
Oceans and seas are the largest ecosystems on the planet. The oceans cover 71% of the Earth’s surface, with 50% of the
depth being below 3000m and a mean depth of about 3800m (Ramirez-Llodra et al., 2010). Cyanobacteria inhabit a wide
range of niches in marine ecosystems, where they occupy maritime coasts and the open ocean. Aquatic cyanobacteria
are divided into two large ecological groups: planktonic cyanobacteria that float freely in the water column and benthic
cyanobacteria that adhere to submerged solid surfaces (sediments, rocks, stones, algae, and aquatic plants) (Fogg et al.,
1973). The ecological conditions in these two habitats are different and that is why the biodiversity of the benthic and
planktonic species should be considered separately (Hoffmann, 1999).
2.1.1 Marine Plankton
Planktic cyanobacteria are adapted to vertical migrations using the phenomena of buoyancy regulation. Another adaptive
mechanism is the production of aerotopes, containing gas vesicles (Reynolds et al., 1987; Brookes and Ganf, 2001).
The marine cyanobacteria Prochlorococcus and Synechococcus are dominant in many ocean regions (Flombaum et al.,
2013). The cyanobacterial genus Prochlorococcus was isolated about 20 years ago. Probably, it is the most abundant
photosynthetic organism on the planet (Scanlan et al., 2009). Prochlorococcus is dominant in phytoplankton communities
in the central oceanic parts up to a depth of about 150m 40°N and 40°S. It has been hypothesized that its population size is
limited by latitudes, and hence Prochlorococcus is understandably absent at temperatures below 15°C (Moore et al., 1995;
Partensky et al., 1999; Zubkov et al., 2003; Johnson et al., 2006). This taxon also exhibits significant genetic diversity,
which has resulted in differences in light and temperature sensitivities and nitrogen adaption abilities (Johnson et al., 2006;
Moore et al., 2002, 2008; Moore and Chisholm, 1999; Rocap et al., 2003).
Synechococcus distribution is not as deep as Prochlorococcus, but it has a wider geographical distribution in both
polar and high-nutrient waters (Zubkov et al., 2003; Moore et al., 1995; Partensky et al., 1999). The genera Cyanobium
and Synechocystis are also widely distributed in marine planktonic communities (Costa et al., 2014). Some species of the
filamentous genus Romeria have also been found in marine planktons (Komárek, 2001).
In favorable conditions, cyanobacteria grow rapidly and form blooms (Sellner, 1997; De Figueiredo et al., 2006; Sciuto
and Moro, 2015). They exist in highly productive nutrient-rich waters by having gas vesicles supporting the migration
between the surface and bottom waters (Paerl et al., 2001).
Trichodesmium is an example of a colonial filamentous cyanobacteria. It is one of the most abundant bloom-forming
species in the marine pelagic zone, characterized by a panglobal distribution in oligotrophic environments in tropical and
subtropical oceans (Capone et al., 1997; LaRoche and Breitbarth, 2005). Trichodesmium has the ability to even grow above
20°C and thus may expand its distribution in the conditions of global warming (Hutchins et al., 2007; Stal and Zehr, 2008).

Cyanobacteria in Diverse Habitats Chapter | 1 3
FIG. 1 Cyanobacteria in diverse habitats: (A) freshwater dwelling cyanobacteria in small rivulets, India; (B) freshwater dwelling cyanobacteria in
shallow water bodies, India; (C) cyanobacteria in stagnant freshwater bodies, India; (D) freshwater dwelling cyanobacteria in shallow Arctic water body;
(E) freshwater dwelling cyanobacteria attached on stones in shallow Arctic water body; (F) almost dried cyanobacterial mat on decomposed soil in the
Arctic; (G) cyanobacteria-dominated pediceled algal crust in Grand Staircase-Escalante National Monument, Utah, United States; (H) cyanobacterial crust
in South Ural region, Russia; (I) polygonal lichen-dominated crust in steppe in South Ural region, Russia; (J) visible growth of algae and cyanobacteria
on wet soil in South Ural region, Russia.

4 Cyanobacteria
The nitrogen-fixing coccoid genus Crocosphaera has also been detected in tropical waters (Zehr et al., 2007).
Crocosphaera watsonii is well known as a significant contributor to oceanic nitrogen fixation (Zehr et al., 2001; Montoya
et al., 2004; Moisander et al., 2010). Crocosphaera is also able to reduce its iron-metalloenzyme inventory. It allows
Crocosphaera to inhabit regions low in iron (Saito et al., 2011).
In estuarine environments cyanobacterial blooms have been detected in all basins of the central Baltic Sea (Karjalainen
et al., 2007). Intensive growth of blooms has been reported in many areas, where these communities were rare before
(Kahru et al., 1994; Mazur-Marzec et al., 2006).
The genera Nodularia, Aphanizomenon, and Anabaena belong to the most important bloom-forming cyanobacteria
in the Baltic Sea (O’Neil et al., 2012). In this ecosystem, cyanobacteria Nodularia spumigena forms toxic blooms each
summer (Teikari et al., 2018). It is necessary to note that among Nodularia both planktonic and benthic species exist:
Nodularia baltica, N. litorea, and N. spumigena are planktonic species, while N. harveyana and N. spaerocapsa are benthic
species (Komárek et al., 1993).
2.1.2 Marine Benthos
Benthic filamentous cyanobacteria usually possess unique gliding motility (Hoiczyk, 2000). This group of cyanobacteria
can form mats—dense multilayered communities of many species, which grow on sediments of different environments
(Stal, 1995).
Filamentous cyanobacteria of the genus Lyngbya can be found in benthic communities worldwide, where it forms
dominant cover and blooms in tropical and subtropical coral reef and seagrass habitats (Paul et al., 2005; Jones et al., 2011;
O’Neil et al., 2012). Lyngbya is an important contributor to coral reef ecosystems, occasionally forming the dominant cover
and impacting the health of many other co-occurring organisms. Lyngbya majuscule belongs to the benthic cyanobacteria
and is widely distributed in tropics in reef and lagoons, forming dense mats (Hoffmann, 1994; Whitton and Potts, 1982,
2000a,b; Thacker and Paul, 2004). In Guam and the Republic of Palau, several Lyngbya species, including Lyngbya
majuscule, have been detected (Thacker and Paul, 2004). During the study of benthic bloom forming Lyngbya in Broward
County, Florida using polyphasic approach, it was found that this bloom contains a new species of Lyngbya. The second
most common Lyngbya in this habitat was similar to Lyngbya polychroa (Paul et al., 2005). The filamentous cyanobacterial
genus Moorea belongs to a cosmopolitan pantropical ecological group, abundant in the marine benthos. Members of this
genus are photosynthetic, nondiazotrophic, and often create mats or blooms in shallow water (Engene et al., 2012). The
genera Coleofasciculus, Hydrocoleum, and Lyngbya are dominant in all the sediment types in cyanobacterial populations in
intertidal flats of the German Wadden Sea (Vogt et al., 2018). Cyanobacteria Microcoleus ethnoplasts and representatives of
the genera Spirulina and Oscillatoria sp. belong to common species in marine intertidal and littoral habitats (Kulasooriya,
2011). The filamentous genus Nodosilinea, Leptolyngbya, Pseudanabaena, and Romeria constitute a large group of the
marine cyanobacterial strains, usually isolated from the Portugal coast (Costa et al., 2014). In all, 26 species of cyanobacteria
were included in the checklist of benthic marine algae and cyanobacteria of Northern Portugal with 15 species belonging to
the order Oscillatoriales (Sousa-Pinto, 2009). Lyngbya was the richest in species genus (Araujo et al., 2009).
Recent studies based on phylogenetic inference of small-subunit (SSU) rRNA genes have revealed a large extent of
novel biodiversity of benthic cyanobacteria from tropical and subtropical marine environments (Engene et al., 2013).
Among the 10 phylogenetic clades of natural product-producing strains from benthic environments, only two clades were
related to reference or type strains of cyanobacteria. Thus, the biodiversity in tropical marine forms of cyanobacteria are
unclear and definitely need further investigations.
2.1.3 Mangroves
Mangrove forest is an example of transitional ecosystems between the coast and the mainland (Rigonato et al., 2013). A
number of surveys of the cyanobacterial diversity of mangroves of several tropical and subtropical regions of the world
have been published (Branco et al., 1996, 1997; Silambarasan et al., 2012; Silva et al., 2014; Alvarenga et al., 2015).
Genera Aphanocapsa, Calothrix, Chroococcus, Coleofasciculus, Lyngbya, Oscillatoria, and Schizothrix comprise the most
widespread cyanobacterial mangrove dwellers across the world (Alvarenga et al., 2015).
It is necessary to note that using culture-independent approaches the estimates of the hidden biodiversity of cyanobacteria
can be evaluated up to a certain extent. Interesting data was published about the biodiversity of cyanobacteria of mangrove
soils in Southeast Brazil (Rigonato et al., 2013). Molecular-genetic analysis of 16S rRNA gene sequences showed high
cyanobacterial diversity. In all, 15 different cyanobacteria genera were detected in the mangrove soils, represented
by sequences of the genera Aphanocapsa, Cyanobium, Cyanothece, Gloeothece, Synechococcus, Prochlorococcus,
Leptolyngbya, Nostoc/Anabaena, Arthronema, Limnothrix, Oscillatoria, Phormidium, Tolypothrix, Tychonema, and

Xenococcus. In addition, sequences matching uncultured cyanobacterial 16S rRNA genes were abundant in all sites
sampled, indicating the existence of many potentially novel cyanobacteria. On sites close to seashore, mainly the genera
Prochlorococcus and Synechococcus were found while other mangrove niches were mostly inhabited by uncultured
cyanobacteria.
2.2 Freshwater Cyanobacteria
Cyanobacteria are known to flourish in freshwater habitats. They also inhabit in water bodies that maybe salty, brackish,
or fresh, in cold and hot springs, and in environments where almost no other microalgae can even exist (Mur et al., 1999).
There are several classifications of ecological groups of freshwater cyanobacteria. Mur et al. (1999) described several
ecostrategies of survival of freshwater cyanobacteria: scum-forming ecostrategists, stratifying ecostrategists, nitrogen-
fixing cyanobacteria, and small colonial and benthic cyanobacteria. We think that scum-forming ecostrategists, stratifying
ecostrategists, and small colonial cyanobacteria belong to the planktonic species, nitrogen-fixing cyanobacteria includes
both planktonic and benthic representatives. Below, we will focus on the biodiversity of these ecological groups.
2.2.1 Freshwater Plankton
The autotrophic picoplanktons represent the major primary producers in the ocean and large transparent lakes (Callieri
and Stockner, 2002; Ting et al., 2002). Single cells and microcolonies of picocyanobacteria (Pcy) are widely distributed in
lakes around the world. The single-celled Pcy populations are predominant in large, deep oligo-mesotrophic lakes while the
microcolonies prefer warmer, shallower, and more nutrient-rich lakes (Callieri, 2010). Picoplanktons consist predominantly
of cyanobacteria <3μm in size (Ivanikova et al., 2007). Representatives of these ecological groups very rarely create stable
scums through buoyancy regulation (Mur et al., 1999).
Autotrophic picoplanktons in oligotrophic lakes are dominated by the phycoerythrin-rich freshwater
Synechococcus (Fig. 2A) (Fahnenstiel and Carrick, 1992; Ting et al., 2002). Analysis of 16S ribosomal DNA of
freshwater Synechococcus species revealed that it is a polyphyletic genus (Urbach et al., 1998; Robertson et al.,
2001). In freshwater ecosystems, the genera Cyanobium and Synechocystis are also very important (Stockner, 1988;
Albertano et al., 1997; Komárek, 2003). The genus Aphanothece (Fig. 2B) forms large populations in freshwater
bodies (Mur et al., 1999). The cyanobacterial genera Chroococcus, Cyanodictyon, Rhabdoderma, Merismopedia
(Fig. 2C), Coelosphaerium, Coelomoron, Snowella, and Gomphosphaeria are also common in freshwater ecosystems
(Komárek and Anagnostidis, 1999; Komárek, 2003).
Scum-forming ecostrategists form large colonies of coccoid cells or filaments. The genera Microcystis, Anabaena,
and Aphanizomenon belong to this ecological group. Microcystis is one of the most distributed taxa, which forms blooms
in springs of the temperate zone and eutrophic lakes (Reynolds et al., 1981; Kurmayer et al., 2002). The high diversity
and wide global distribution of common ITS gene types together with the lack of phylogeographic structure confirms the
existence of intercontinental dispersal of Microcystis aeruginosa ITS types. This finding indicates that this species might
be a cosmopolitan cyanobacterium (van Gremberghe et al., 2011).
Filamentous species such as Planktothrix agardhii and Limnothrix redekei belong to the homogeneously dispersed
ecostrategists inhabiting eutrophic and hypertrophic shallow lakes with depth less than 3m (Mur et al., 1999). In the
eutrophic and hypertrophic shallow lakes with depth less than 3m, the cyanobacterial diversity comprises Planktothrix
agardhii and Limnothrix redekei (Mur et al., 1999). These cyanobacteria are extremely light sensitive and do not form
colonies at high light intensities (Reynolds, 1997). Stratifying ecostrategists, like Planktothrix rubescens, usually dominate
the intermediate zone of thermally stratified water bodies (Mur et al., 1999).
It is necessary to note that many of the cyanobacteria, including freshwater species, produce toxins (De Figueiredo
et al., 2004). Microcystis belongs to the major microcystin-producer taxa in Ugandan freshwater habitats in Africa, where
a significant amount of toxin has been detected in freshwater resources (Okello et al., 2010). Some species of the genera
Anabaena, Planktothrix, and Nostoc can also produce microcystins (Neilan et al., 1999; De Figueiredo et al., 2004).
2.2.2 Freshwater Benthos
Benthic cyanobacteria inhabiting the bottom sediments of the water bodies usually form coherent mats (Mur et al., 1999).
Benthic mats, consisting of the genera Aphanothece, Oscillatoria, and Phormidium, usually develop on the sediments
of lakes and ponds. These mats later become floating mats in standing water bodies (Komárek, 2003). Leptolyngbya,
Nostoc, Chroococcus, Aphanocapsa, and Aphanothece are the most distributed genera among epilithic cyanobacteria from
freshwater streams of Kakoijana Reserve Forest of Assam, India (Saha et al., 2007).

6 Cyanobacteria
FIG. 2 Freshwater and terrestrial cyanobacteria: (A) Synechococcus aeruginosus; (B) Aphanothece stagnina; (C) Merismopedia tenuissima; (D)
Lyngbya sp.; (E) Nostoc punensis; (F) Hormoscilla pringsheimii; (G) Phormidium uncinatum; (H) cf. Trichocoleus hospitus; (I) Cylindrospermum majus;
(J) Leptolyngbya voronichiniana; (K) Leptolyngbya foveolarum; (L) Microcoleus vaginatus; (M) Nostoc cf. punctiforme; (N) Phormidium breve; (O)
Phormidium dimorphum; (P) Trichormus variabilis. Scale bars=10μm, for (B–L, N–P) the bar is placed in (A).

Among benthic cyanobacteria, toxin-producing species also exist. In comparison with planktonic species, toxin
production and species composition of toxic benthic cyanobacterial communities have been studied insufficiently
(Gugger et al., 2005; Wood et al., 2007; Quiblier et al., 2013). Together with the genera Oscillatoria and Phormidium,
other oscillatorian cyanobacteria Lyngbya (Fig. 2D), Leptolyngbya, Microcoleus, Tychonema, and Schizothrix are usually
detected (Steppe et al., 1996; Mez et al., 1997, 1998; Hitzfeld et al., 2000; Aboal et al., 2005; Gugger et al., 2005). Together
with these benthic taxa, a toxin-producing Planktothrix has been detected from a biofilm in New Zealand (Wood et al.,
2010). In toxic freshwater benthic biofilms cyanobacteria, Aphanothece and Synechococcus from the order Chroococcales
together with nitrogen-fixing cyanobacteria Anabaena and Scytonema from order Nostocales are usually found (Krienitz
et al., 2003; Dasey et al., 2005; Mohamed et al., 2006; Mohamed, 2008; Smith et al., 2011).
Сyanobacteria using nitrogen-fixing strategy are usually dominant in deep, shallow ecosystems with low levels of
inorganic dissolved nitrogen. This group includes the genera Anabaena, Aphanizomenon, Cylindrospermopsis, Nodularia,
and Nostoc (Mur et al., 1999). Nostoc caeruleum, N. commune, N. microscopicum, N. parmelioides, N. pruniforme,
N. verrucosum, and N. zetterstedtii forming macroscopic colonies have been reported from inland aquatic ecosystems
(Mollenhauer et al., 1999). Another new species of Nostoc from freshwater habitats has been described recently from
Panchvati locality of Pune, India and has been named Nostoc punensis (Fig. 2E) (Singh et al., 2015).
3. SYMBIOTIC CYANOBACTERIA
Cyanobacteria are unique among the entire prokaryotes in their efficiency to form symbiotic relationships with a broad
and highly varying range of hosts. This ability is partly a result of their highly efficient photosynthetic apparatus which
allows them to continue this activity even at very low levels of light, which is common in the case of most of the symbiotic
relationships (Usher, 2008). They play decisively the role of “chloroplasts” in symbioses with an array of nonphotosynthetic
partners such as the marine invertebrates, where cyanobionts provide the host partners with carbon. Also, cyanobionts
usually perform the process of nitrogen fixation, especially in photosynthetic partners such as diatoms, thus supplying
the host with nitrogen. Interestingly, the symbiotic associations between cyanobacteria and marine organisms are one of
the most important interactions in the oligotrophic waters of the global ocean, where cyanobionts make it possible for an
extensive variety of marine organisms to survive in sometimes highly nutrient-deficient waters. The most common marine
organisms sheltering cyanobacteria include the corals, diatoms, dinoflagellates, seagrass, and sponges.
Marine symbiotic cyanobacteria and their interactions with potential hosts, as opposed to the terrestrial symbioses, have
largely remained neglected. In fact, there are just about a dozen contemporary scientific investigations of these associations
and a better understanding of cyanobionts of the marine environments is indeed needed. Recent findings, anyway, open
a plethora of interesting questions about the evolution of such symbiosis specificity, metabolic interactions, and cellular
adaptations, all of which are areas of great interest in symbiotic research, especially because they may eventually explain
the ecological impacts of cyanobionts living in the marine environment.
The symbiotic relationships are important constituents of the ecology and general life cycle pattern of many
cyanobacterial lineages. These include interactions with plants (Rai et al., 2000), fungi (Rai, 1990), animals (Wilkinson,
1992), and eukaryotic algae (Janson, 2002; Murakami et al., 2004). It has been anticipated that tightly integrated ecological
associations if maintained faithfully over evolutionary timescales can lead to coevolutionary patterns such as asymmetric
evolutionary rates (Law and Lewis, 1983), gene-for-gene interactions (Flor, 1955), or even cospeciation (Brooks, 1979).
Because the coevolution involves reciprocal evolutionary changes, it requires that each partner must have a significant
fitness effect on the other (Thompson, 1994). Thus, it has been suggested that if a species somehow restricts the number
of partners with which it interacts (specialization), this, in turn, may actually increase the response of that species to the
selection imposed by those partners, facilitating coevolution (Whitlock, 1996; Kawecki, 1998).
It has been well recognized that the colonies of the Caribean coral Montastraea cavernosa harbor endosymbiotic
cyanobacteria that express nitrogenase and, thus, fix nitrogen for the host (Lesser et al., 2007). Stable isotope results
have also showed that the zooxanthellae, which have an important mutualistic symbiotic relationship with corals,
actually use the products of this nitrogen fixation. As low oxygen concentrations are required to support cyanobacterial
respiration and nitrogen fixation, the nitrogen fixation is confined to those times of the day when physiological
hyperoxia or anoxia does not inhibit nitrogen fixation, thus maintaining an intricate balance between nitrogen fixation
and photosynthesis at the same time. Through this carefully controlled process, the endosymbiotic cyanobacteria help
the zooxanthellae corals to survive in oligotrophic waters by providing them with an important supplementary source
of a limiting element.
Heterocytous cyanobacteria Richelia intracellularis and Calothrix rhizosoleniae are examples of some interesting taxa
that live in a symbiotic relationship with many frequently occurring diatoms like Hemiaulus, Rhizosolenia, and Chaetoceros

8 Cyanobacteria
(Foster et al., 2011). These cyanobionts, in this case, are important because they provide fixed nitrogen to their nitrogen-
limited hosts. This impacts both the host and the ecosystem in a positive way.
Blooms of the diatom Hemiaulus hauckii and its cyanobacterial symbiont Richelia intracellularis can reach extremely
high densities and have important ecological impacts on the ecosystem. These blooms dominate the carbon deposition in
the offshore tropical river plumes and thus subsidize substantially to the new N inputs (Yeung et al., 2012). The blooms
of these diatom-diazotroph assemblages increase significantly the net primary productivity as well as the ecosystem
carbon export ratio, which ultimately leads to the biological uptake of dissolved inorganic carbon and silicates too. Also,
Hemiaulus and its nitrogen-fixing endosymbionts have been implicated as important donors to the formation of some
Mediterranean sapropels (Carpenter et al., 1999). As drivers of nitrogen fixation and carbon drawdown in river plumes
and oligotrophic waters, diatom-diazotroph interactions have had a global impact that is clearly an indicator of climatic
and anthropogenic influences. Diatoms belonging to the family Epithemiaceae possess a unicellular nitrogen-fixing
cyanobacterial endosymbiont (DeYoe et al., 1992). In case of limitations of external nitrogen, the number of endosymbionts
per diatom rises, at the same time, the mean endosymbiont surface area:volume also increases. This ultimately leads to
an increase in the contribution by the endosymbiont nitrogen fixation to the diatom nitrogen budget under N-limiting
conditions, thus helping the diatom partner to ward of nutrient stress. This flexibility of the endosymbiont load enables the
diatom to be more energy competent and contest more proficiently with other neighboring symbiotic diatoms in nitrogen-
stressed habitats. Also, the diatoms that host nitrogen-fixing cyanobacterial symbionts are high in protein, which benefits
the organisms that graze on these diatoms (Kupferberg, 1997). Thus, the diatom-cyanobacterial symbioses are quite
widespread and also important for oceanic primary productivity.
Cyanobionts are almost omnipresent in most of the marine sponges and their microenvironments and their presence can
cause noteworthy alterations in the morphology of the sponges. These cyanobacterial symbionts are generally intercellular
but sometimes may occur in specialized vacuoles which are termed as “cyanocytes” (Usher, 2008). In this symbiosis, the
photosynthates in the form of glycerol and organic phosphates are transferred from cyanobacterial partners to the sponge
hosts supplying up to 50% of the sponge’s energy budget and around 80% of its carbon budget. Unicellular Synechococcus
sp. is one of the most common cyanobacteria found in sponges, while nonheterocytous filamentous Oscillatoria spongeliae
has also been reported over a wide geographic range (Usher, 2008). Dysidea herbacea, a commonly occurring shallow-
water sponge present throughout the Indo-Pacific, harbors large numbers (up to 20% of the symbiotic association’s volume)
of the cyanobacterial symbiont Oscillatoria spongeliae. By using cellular localization of some selected compounds, it
has been well established that the polychlorinated metabolites attributed to Dysidea herbacea are actually localized in the
symbiont Oscillatoria spongeliae. Truly, this was one of the first demonstrations that marine natural products ascribed
to a sponge are actually localized in the symbiotic cells. Interestingly, these polychlorinated metabolites produced by
the symbiont prevent fish feeding and thus confer augmented fitness to the sponge-cyanobacterial symbiosis (Unson and
Faulkner, 1993).
Cyanobacteria-dinoflagellate symbioses are one of the foremost reasons for the widespread distribution of heterotrophic
dinoflagellates in the oligotrophic subtropical and tropical oceans (Farnelid et al., 2010). Histioneis sp. and Ornithocercus sp.
contain epi- or endobiotic cyanobacteria as partners. Immunolabeling-TEM using nitrogenase antisera have discovered that
many cyanobionts may have a nitrogen-fixing capacity and thus probably temporally segregate their physiological processes
to protect the nitrogenase enzyme (Foster et al., 2006). The mixed assemblage of a few nitrogen-fixing cyanobacteria
partners residing among a larger population of nonnitrogenase localizing cyanobionts as in Histioneis depressa confers a
benefit over other hosts that only contain non-N2-fixing cyanobionts partners. Thus, the dinoflagellate’s requirement for
fixed nitrogen as well as carbon is the driving force for this symbiotic relationship.
The dominance of cyanobacteria in epiphytic microbial communities has been well established by using light microscopy,
16S rRNA, and nifH gene analysis methods (Hamisi et al., 2013). The nifH gene expression patterns have also demonstrated
that the heterocytous phylotypes may be the dominant diazotrophs during the day while the nonheterocytous types may
establish their dominance at night. Cyanobionts usually exist associated with the leaves of the seagrass Cymodocea
rotundata as small attached patches of pigmented microbial aggregates of thin biofilms. Seagrass is benefitted from the
diazotrophic nitrogen-fixing ability of the cyanobacteria, while the diazotrophic activity of the cyanobacteria is stimulated
as indicated by the high nitrogenase levels recorded in the expression studies.
The filamentous heterocytous genus Nostoc is well known as a prolific symbiotic partner with many different
eukaryotic model systems like protists, animals, fungi, and plants (Rai et al., 2002). In many phylogenetic studies, using
an assortment of molecular markers (16S rRNA, rpoB, rbcLX, and nifD or nifH), the closest relatives to the genus Nostoc
were shown to be the genera Anabaena, Aphanizomenon, and Trichormus, also members of the family Nostocaceae
(Rajaniemi et al., 2005; Svenning et al., 2005; Henson et al., 2002, 2004; Lohtander et al., 2003; Singh et al., 2013, 2015).
The genus Nostoc is characterized by a complex life cycle that includes differentiation into heterocytes (nitrogen-fixing

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had to lie at their feet until one of them passed on, only a
few days later. Shortly afterwards the other ones died and
two of the Thirty-ninth had what was left which, while it
did not keep out the rain, did keep off the direct sun, a no
small comfort in that terribly hot place. We had two half
pieces of blankets, nearly used up and almost covered
with what Robert Burns called "crawlin' ferlies," the fearful
pests of our lives. I undertook one day to wash my shirt,
trying first one corner of it which went to pieces, so I
dried the garment carefully and without further effort at
washing wore it almost nine months.
First and last many tunnels were dug, in several of which I
bore a hand; I don't know how many succeeded in getting
out but there must have been several hundred;
bloodhounds were put on their track and those who were
brought back were put in the chain-gang. Among so many
men there must be some bad ones, a few very bad; they
even resorted to murder in their efforts to secure what
some of the prisoners possessed. To rid themselves of this
terrible set of evil men a vigilance committee of the well
disposed was organized and by sheer force of numbers,
overpowered and sentenced to death six of them. The
rebels, to their credit, furnished material for the gibbet
and the execution took place, much to the relief of those
who had to continue there.
Our drinking water came from holes in the ground four or
five feet deep; while it was pretty clear, there were many
dead maggots in the bottom, though we did not mind
them, thinking the water so much better than that in the
brook. One day in August a stream of water broke out just
inside of the inner stockade; it ran all of the time, but the
dead line was between us and the water; we procured
boards and made a trough and then got permission to put
it up, so that we had fine water all the rest of the time we
were there. To this day it is known as the Providence

Spring. Aside from scurvy, severe enough to loosen my
teeth, I was not sick a day while in the prison. Our rations
for the most part were a pint of boiled rice without any
salt for twenty-four hours and oftener it would be forty-
eight, for every time Captain Wirz discovered a new tunnel
he would punish all of us by skipping our rations.
Occasionally we would get some small black beans, such
as the planters raised for their hogs; these we would try
to cook with green pitch-pine with results that can be
imagined. I have blown myself black in the face many a
time trying to cook them and then had to eat them raw.
There was a sick call every day and when a man
answered the same, all he got for his pains was a dose of
sumach berries. No matter what the complaint might be
the remedy was always the same, for it was all they had
to give. Sometimes a man could be seen buried up to his
chin; he had the rheumatism and if he could endure the
antidote two or three days, he would come out cured. One
boy, to get some extra food, told the captain one day
where a new tunnel was in progress, and after the officer
had gone out, the men shaved one side of his head and
on his breast and back put big placards, bearing in big
black letters the word "Traitor." He was then marched all
over the camp and tormented almost to death; the enemy
finally took him outside, which was just what he wanted.
After Stoneman's raid, the rebels thinking Andersonville no
longer safe began to distribute us elsewhere and I
sampled the bull-pen of Savannah, Ga.; the stockade of
Millen, also in Georgia; and then was sent back to
Savannah where I was paroled and sent down the River,
to go on board a Union steamer; the sight of the Stars
and Stripes brought tears to every eye. On board, our
heads were shaved, we were bathed, clad anew and were
judiciously fed; our old clothes went overboard. After
reaching Annapolis I tipped the scales at seventy-five

pounds, less than half my weight when I enlisted. After a
brief stay in Parole Camp, I was paid off and sent home
on a thirty days' furlough, where I was sick all of the time,
but I returned to the camp at the end of the time to be
furloughed again, this time for sixty days. On getting back
to camp the second time, and wearying of it, I put my
name down among those to be returned to their
regiments and I reached mine the day after Lee
surrendered.

REGIMENTAL VETERAN
ASSOCIATION
The beginnings of the Association seem to have been lost in the
interval between 1867 and the present; it is agreed, however, that
the first four meetings were held in Boston hotels and that they
were not very largely attended. The time was too near the date of
getting home and the pleasures of that supreme event far
outweighed any rehearsal of common dangers in war-experiences.
Of the 5th gathering, the first basket picnic of the veterans of the
Thirty-ninth and their lady friends, there is in substance the following
account:
Downer's Landing, Hingham, was the place and Thursday, August
17th, the date; the party, numbering 300, left Litchfield's Wharf at
9.15 a. m. on steamers "Wm. Harrison" and "Emeline," arriving
about 11 o'clock; a half mile walk brought all parties to the "Melville
Gardens" which had been hired for the day. Noon saw the tables
spread with the many good things brought by the members. Before
repairing to the hall for dancing, Colonel C. H. Porter, President of
the Association, introduced as speakers, Lieut. Colonel Hutchins,
Major Graham, Captain Brigham, Lieuts. Mulligan and Mills,
Sergeants Eames and Gardner. At 3 p. m. came a dress parade with
one hundred and fifty men in line, Colonels Porter and Hutchins
dividing the honors of commanding. A letter was read from General
G. K. Warren, regretting his inability to be present, a disappointment
to the veterans, as they had expected to meet their former
commander once more; everyone of the Regiment and, for that
matter, every regiment in the Fifth Corps, holding the officer in the
highest esteem. On the formation of the line and led by Edmunds
Band, the company marched back to the landing, reaching Boston at

5.15 o'clock, all happy and conscious that the presence of the ladies
had added no little to the enjoyment of the day.
The reunions of 1872, '73 and '74 were held in Boston Hotels; in
1875, Oct. 6th in Woburn was held the most notable of the
Association's gatherings thus far, General Warren being the
distinguished guest and Company K, along with citizens of Woburn,
the hosts. There were present 166 men with General Peirson at their
head; drum corps and brass bands furnished music and everyone
joined heartily in the reception to the eminent soldier. At the rooms
of the selectmen, the public had a chance to meet General Warren.
At the armory the meeting was called to order by Captain Hutchins
and the chief feature was the presentation to General Warren of a
magnificent Maltese Cross in Gold, the badge of the Fifth Corps, the
cost $100.00 having been met by the veterans. In the afternoon a
banquet was served in Lyceum Hall to more than four hundred
guests, the good people of the town having vied with each other in
making the occasion memorable. At the post prandial exercises,
remarks were made by Captains Hutchins and Tidd; there was an
extended address by J. A. Harvey, Co. C, followed by the
introduction of Sergt. Abijah Thompson, Co. K, as Toastmaster, who
read an original poem after which, and the playing by the band of
"Hail to the Chief," General Warren spoke briefly to the following
effect, "I rise to acknowledge the kind attentions I have received to-
day. Those who have spoken have referred in such kind terms to
me, and your marks of approbation have been so many that I do not
feel prepared to speak for the Fifth Army Corps, as I could wish. I
hope you will excuse me. I shall carry from this place a sense of
having been honored more than I deserve. (Cries of no, no.) The
feelings which this day has inspired will always remain, and you have
laid on me a debt of gratitude I never can repay." Col. C. H. Porter
responded for General Peirson, letters were read from General J. C.
Robinson, and Colonels Farnham and Tilden of the Sixteenth Maine;
further responses to toasts were given by Maj. Ambrose Bancroft of
the Thirty-second Regiment, Capt. J. P. Crane of the Twenty-second,
Capt. C. S. Converse of the Fifth, Lieut. John L. Parker of the

Eleventh and others, the exercises terminating in an evening's levee
which lasted till midnight.
The Centennial year, 1876, found the veterans 150 strong in Natick,
the guests of Co. I.; 1877, August 28th, Co. D of Quincy did the
hospitable act with 220 comrades present. In 1878, Co. E of
Somerville, on the 6th of Sept, helped celebrate the 16th anniversary
of the departure of the Regiment with 225 veterans in attendance,
the event gaining unwonted interest through the presence of
General John C. Robinson who had been the Division Commander of
the Regiment at the Wilderness and at Spottsylvania, losing there a
leg; there was a spirited address by Mayor Bruce of Somerville, an
extended historical paper by Col. C. H. Porter with speeches of
greater or less length by Gov. A. H. Rice, Gen'l N. P. Banks, Collector
Beard, Secretary Pierce, Speaker Long, General Peirson and others,
the celebration continuing with music and dancing till after midnight.
A pleasant feature of the afternoon was the presentation of an
elegant punch bowl and ladle to General Robinson by Lieut. C. K.
Conn to whose words the General responded so happily that all
recognized him as a good talker as well as fighter.

Newport Sept. 1. 1878
Capt. Chas H Porter
No. 27 State Street
Boston Mass
My dear friend

I am very sorry that the necessity for being in New York City on the
6th prevents my being with you at Somerville.
The 39th Massachusetts have been exceptionally kind to me, and I
am grateful for it. I have got hard work to do to set right the record
of my army career, and the recollection of this friendly feeling helps
to keep up my nerves for going through with the undertaking.
I hope you will all have a fine day and a happy one.
Yours truly
G. K. Warren
August 27th, 1879, found the survivors of the Thirty-ninth in
Taunton, guests of Co. F with Capt. J. J. Cooper president of the day
and 125 veterans on hand; Adjutant O. A. Barker welcomed the old
soldiers to the city and after a short business meeting, line was
formed for the Agricultural Fair Grounds where Hiram Maxfield of
Silver Springs fame served one of his imitable clambakes. In 1880,
Sept. 15th, Medford was the entertaining place with Co. C at the
front, Jas. A. Harvey being President. Oct. 5th, '81, the "old boys"
came back to Woburn again, the reception being in the hands of the
following named men of Company K., C. K. Conn., Geo. E. Fowle,
Capt. L. R. Tidd, A. L. Richardson, J. F. Ramsdell, A. P. Barrett, J.
Fred Leslie, A. Thompson and A. R. Linscott. Again Woburn has the
honor of entertaining General G. K. Warren and he is accompanied
by General J. C. Robinson, the valiant Division Commander, along
with General Peirson, the ever popular regimental commander. At
the dinner which was served in Lyceum Hall, remarks were made by
those named above, Col. Porter and others. Before another reunion,
General Warren will have passed away.
The regimental line formed again in Natick, Oct. 10, 1882; again Co.
I plays the role of entertainers with fully 150 survivors to honor the
occasion. Dinner was served in Concert Hall; Col. C. H. Porter spoke
at length in praise of General Warren who had died the 8th day of
the preceding August; resolutions of sympathy and respect were

passed by the veterans and a contribution was made to a Fifth Corps
fund to honor the General's memory. Remarks followed by Comrades
Barrett, "K"; Beck, "C"; Locke, "E"; Eames, "C"; Oliver, "E," and
others. 1883 brought the veterans to Quincy again with Co. D. Point
Shirley in Winthrop was the place of meeting, August 26, 1884, with
Co. H as entertainers. Roxbury, the home of Co. B, entertained next,
Sept. 23, 1885. Company G came to the front Sept. 16, 1886, at
Nantasket. The ladies of Somerville, in behalf of Co. E, furnish the
dinner for the reunion of 1887, Sept. 6th. Through the selection of
the Executive Committee, Bass Point was the place of meeting in
1888. Sept. 11, 1889, brought the clans to Medford, once more, with
Co. C.
Sept. 24, 1890, the beginning of another decade, brings the
veterans to Woburn, the home of Co. K, for the third time. As usual,
great preparations were made for the reception, the principal guests,
aside from the veterans themselves, being the widow and daughter
of General Warren, and General Peirson and wife who with the wife
of Mayor Johnson and the wife of the Hon. John Cummings formed
the receiving line in the hall of the Y. M. C. A. The formalities of the
occasion were conducted by Colonel C. H. Porter and Sergeant
Abijah Thompson, "K," and 166 survivors pressed forward, glad of
the opportunity of grasping the hands of their friends. Dinner was
served in Lyceum Hall. The after-dinner exercises were presided over
most happily by Sergt. Thompson who introduced Mayor Johnson,
General Peirson, the Hon. John Cummings, Colonel Porter and
others. Company A was the host Sept. 7, 1891, at the old Lynnfield
camping-ground, and the occasion was rendered notable by the
following paper, prepared for the day by Lieut. Elbridge Bradshaw of
Co. H:
A VACATION IDYL
Some thirty years ago, leading a sedentary life and
gradually sinking into a semi-bituminated condition, my
medical adviser, alarmed at my symptoms, ordered travel
and change of scenery. Having learned that Virginia

contained more travel and scenery to the square mile than
any other spot on the globe, I determined on visiting that
State. Being of a timid nature and fond of Company, I
joined myself to about a thousand other invalids, similarly
afflicted, and seeking the same remedy, forming ourselves
into a methodical organization. For convenience we
divided ourselves up into groups of one hundred men
each, using for purposes of distinction the first eleven
letters of the alphabet, omitting the letter J. For menial
service, i. e. to look after our physical wants, each group
hired for such purpose, six servants, viz. a captain, two
lieutenants, with a cook, a drummer and a bugler or fifer,
the latter two being hired to wake the excursionists in the
morning. To keep these captains and lieutenants in order,
we placed over them a colonel, a lieut. colonel and a
major, at the same time they being our head servants or
butlers. These people added to themselves an adjutant to
run errands, a chaplain, a doctor and a pill-driver. To
insure a faithful discharge of duty, from each group were
chosen a dozen fellows called sergeants and corporals
who were set over the others.
Virginia at this time being in a tumultuous condition, and
the U. S. Government having heard of our organization's
plan of travel and objective points, invited us, through its
Chief Magistrate, to walk over Virginia as peace officers,
punching the heads of belligerants and arguing with the
discontented, an invitation which we accepted. When
President Lincoln secured our services he loaded us with
benefits, first massing us at Lynnfield, giving us canvass
houses to protect us from the dew and damp, sweet straw
to nestle in, a pretty blue uniform, a belt to keep us from
bursting, an iron toothpick, a tube of iron with a wooden
handle, a little black bureau, in which to keep our collars,
cuffs and bric-a-brac, a black cotton pantry for provisions
and plates, with a round tin vessel for whiskey. Uncle Sam

also gave our servants (the shoulder-strapped ones)
toasting forks to stick pigs with and red sashes with which
to gird their persons when running and chasing the pigs
down. Rendered proud and arrogant by their good clothes
and shoulder straps, our servants rose on us and captured
our organization, styling themselves our superior officers,
and our entire body the Thirty-ninth Regiment of
Massachusetts Volunteers. To give the usurpation a flavor
of legality, they procured from Governor Andrew
commissions indicating officially their rank and authority.
On the whole they exercised their powers with great
moderation and kindness.
Though to the last, we suffered them to think themselves
our superiors, yet in reality, they still continued to be our
servants, caring for our food, clothing and morals,
furnishing us clean, airy lodgings having adequate fire-
escapes, so that in fact we had nothing on our minds
worth mentioning and all we had to do was to travel and
fight; in a word, take our pleasure. They also taught us
many pretty and amusing tricks; how to stand up straight
in rows to be shot at; to abstain from whiskey (with
quinine in it); to use the pickaxe and spade with the least
expenditure of muscular energy and, in mud and night
marches, to say our prayers without even stopping. As
soon as we could march without scalping each other's
heels, we left Lynnfield for active service and mighty
active it proved on the start, our first engagement being a
footrace against time through Boston. We left Lynnfield
with cooked rations, meaning saltpork and hardtack.

How dear to this heart is the old army hardtack,
As Lynnfield's reunion presents them to view;
When eaten with raw pork or fried into doughnuts,
The rations that beat him, are scat'ring and few.
How oft in our marches, he's braced up our courage,
As with gnawing and growling we've hobbled along;
Oh! well may the hardtack, the old army hardtack
Prove a classical theme for an old veteran's song.
That dear army hardtack was a limber old codger,
In the hands of a Thirty-ninth's amateur cook;
In his grip, the old hardtack took metamorphosis
Not mentioned by Ovid, nor in Parloa's cook-book,
As a pudding or pie in a cob-house as a dumpling,
As a fry or a toast, or a raw on the shell;
That old army hardtack, that blessed old hardtack!
For every recipe turned out equally well.
I have eaten high banquets at Young's and at Parker's,
I have tasted their beef, roast turkey and lamb;
But all of these dishes are flat and insipid,
Beside the old hardtack of dear Uncle Sam;
For the old army hardtack is seasoned with memories
Of battles and sieges when wearing the blue;
Of marchings and flankings and digging of trenches,
And loving communion with old comrades too.
The old army hardtack speaks, too, of dear comrades,
Whose faces are missing to-day in our line;
Their battles all fought, their warfare all ended,
But whose virtues still live in mem'ry's pure shrine,
Then cheer the old hardtack, the square army hardtack,
Who was flinty and wormy at times, I must own,
But when at Mine Run, he took a vacation,
His absence was greeted with many a moan.

Chorus.
The old flinty hardtack, the iron bound hardtack;
The moss-covered hardtack, we all knew him well.
Travelling the next three years through Virginia and its
environments, we were often obstructed by mud and
other earthern impediments, and the scenery was much
disfigured and frequently obliterated by sulphurous clouds
of smoke, hence excursioning for health and pleasure was,
on the whole, a failure. Speaking for myself, individually,
the climate didn't agree with me a bit. This I attribute
largely to the horizontal metallic showers with which that
region was infested and against which no ordinary cotton
umbrella was an adequate protection. Indeed the
atmosphere was so impregnated with little pellets of lead
and ragged chunks of cast iron, that my system must have
absorbed about fifty-five pounds of old junk and brought it
home with me for, on my return, I weighed 190 lbs.
against 135 when I left Lynnfield.
Natick with its Company I entertained for the third time, October 5,
1892; Quincy and Co. D did the hospitable act, also for the third
time, in 1893, August 30; Roxbury and Company B were the
entertainers in 1894 and Co. H of Dorchester received at the U. S.
Hotel, Boston, Sept. 25, 1895; for 1896, no record is found, but
Sept. 6, 1897, Co. E and Somerville appear again; it is Medford and
Co. C in 1898; Woburn and her K Company in 1899. The old century
ends, as far as our Regiment is concerned, October 10, 1900, with I
Company and Natick, while the new one begins Sept. 7, 1901, on
the old campground at Lynnfield; Sept. 22, 1902, finds the veterans
again in Quincy; Sept. 24, 1903, in Roxbury; August 19, 1904, with
Co. H. at Nantasket. Then with no special company distinctions the
reunions follow, directed by the Executive Committee, at Squantum
Inn, Sept. 21, 1905; Bass Point, Sept. 6, 1906; in a Dorchester hotel,
October 23, 1907; again at Bass Point, Sept. 29, 1908; at Revere
Beach for three successive years, viz. August 30, 1909, August 18,

1910, and August 18, 1911. Fifty years after the departure of the
Regiment from Massachusetts, nearly a hundred (92) veterans
assembled again in Somerville with Company E and a large number
of prominent citizens to celebrate the semi-centennial; the state
armory was the gathering place and General Peirson was the marked
figure on the occasion while wives, daughters and other lady friends
added to the pleasures of the hour; Sergt. Abijah Thompson of Co. K
was the oldest man present, he having seen fully 90 years. After the
dinner, over which Comrade the Rev. John F. Locke said grace and at
which Lieut. J. H. Dusseault, Co. E, presided, there was speaking by
Ex-Mayors Edward Glines, C. A. Grimmons and John M. Woods, the
latter a veteran of the war, and the first named a brother of Fred
Glines of Co. E who died in Salisbury. Mayor Burns of Somerville
extended the courtesies of the city to the veterans and welcomed
them all most heartily. General Peirson was received with
accustomed enthusiasm and was heard with rapt attention. The half
century event was a great success. The 51st anniversary was
observed in Medford, with the survivors of Co. C, Sept. 6, 1913; the
day, the place, the guests, quite one hundred in numbers, made the
event notable; the forenoon's meeting was in the hall of the S. C.
Lawrence Post, G. A. R., while the dinner was served in the drill-
room of the magnificent armory, presented to Medford and the State
by General S. C. Lawrence.

REGIMENTAL ROSTER
Nothing in the story of a regiment is of greater importance than its
Roster, for therein appears the record of the individual whether the
same be good or bad. One man alone makes a small appearance,
yet a thousand men make a regiment and every volunteer, whether
commissioned or enlisted, is entitled to the best that can be said of
him. If, in addition to his military service, his career in civil life may
be given in outline so much the better, for in America every able
bodied man is potentially a soldier. The foundation for the following
Roster is found upon the muster rolls, carefully preserved in the
State House, Boston, and additions have been made thereto through
the information afforded by members of the Veteran Association.
The careful reader will observe in scanning the data afforded by the
Roster that the ages of the soldiers almost entirely range between
those of eighteen and forty-five years, these being the respective
limits of legal enlistment; at the same time everyone is well aware
that a large part of the army was made up of boys in their early
teens; also we know full well that many a man went in long after
reaching the maximum age for military service. As a fact, then, very
many men lied their ages up or down; so far as the grand average,
however, is concerned the "over" age compensated for or offset
those who were "under." Since the muster-in rolls or enlistment
papers are sources of all data concerning the age of volunteer and,
it being well known that very many of them were and are incorrect,
the wonder rises as to the source of statements that have gone the
rounds of the public press in late years, wherein the ages
represented by the soldiers are carefully tabulated. However, from
whatever source obtained, as worthy of presentation here the
following alleged facts are given:

Discussion has elicited an official statement that about
2,800,000 Union men enlisted; there were about
5,000,000 men called out on both sides. Of these nearly
4,500,000 were under twenty-one; there were about
332,000 who were under sixteen and there were 1,500 in
the Union Army who were not fifteen years old. Less
attention has been given to the men who were over age,
but every regiment can give its cases of men fifty, sixty
and even seventy years of age whose great excess would
average up many a juvenile volunteer. When, however, the
rolls afford no such statements, where is the statistician
acquiring his alleged facts?
For the sake of brevity and economy of space the following
abbreviations are used:
A. A. G. = Assistant Adjutant General; b. = born; bur. = buried; bvt.
= brevet; batt. = battalion; Capt. = Captain; Co. = Company; Col. =
Colonel; com. = commission or committee; Corp. = Corporal; cr. =
credited; d. = died or dead; des. = deserted; det. Serv. = detached
Service; dis. = discharged; disa. = disability; en. = enlisted; ex. of s.
= expiration of service; F. & S. = Field and Staff; G. O. = General
Order; H. Arty. = Heavy Artillery; Infty. = Infantry; k. = killed; lat.
add. = latest address; Lt. or Lieut. = Lieutenant; M. = married; M. I.
= Mustered-in; M. O. = Mustered-out; mos. = months; mus. =
musician; M. V. M. = Mass. Vol. Militia; N. F. R. = no further record;
N. G. = National Guard; O. W. D. = Order, War Department; Pris. =
Prisoner; prom. = promoted; re-en. = re-enlisted; rep. = reported;
res. = resigned; S. = single; S. H. = Soldiers' Home; S. S. =
sharpshooters; S. O. = Special Order; Sergt. = Sergeant; trans. =
transferred; U. S. C. T. = U. S. Colored Troops; V. R. C. = Veteran
Reserve Corps; w. = widower; wd. = wounded; W. D. = War
Department.
In reciting facts pertaining to each name, the same order obtains
throughout the Roster; first comes the family name of the soldier,
next his Christian appellation; in some instances time and place of

birth are given; as a rule, age, whether married or single, occupation
and place of residence follow in order; next, date of enlistment or
muster-in; incidents of army life are next in place, and then the time
and manner of leaving the army; finally are given incidents of civil
life and latest address if the same be known. The application of
abbreviations and the order are seen in the following supposed case:
Jones, John, 20, S.; shoemaker, Natick; Aug. 22, '62; wd.
May 5, '64, Wilderness; dis. disa., Aug. 20, '64; Selectman,
Natick, 1880, '81; 1913, Natick.
Printed in full the foregoing would be as follows:
Jones, John, at the age of twenty years, single, a
shoemaker living in Natick, enlisted August 22, 1862, or
was mustered in on that date; he was wounded in the
battle of the Wilderness and, on account of wounds or
disability therefrom, was discharged August 20, 1864; he
was a Selectman in Natick in 1880 and '81 and in 1913 is
still residing there.

FIELD AND STAFF
COLONELS
P. Stearns Davis, 44, M.; stationer, Cambridge; August 29,
1862; Phineas Stearns Davis was born in Brookline, June
23, 1818, his Christian names coming to him from an
ancestor who bore a part in the Boston Tea Party; his
earlier education, received in the Brookline public schools,
was supplemented by a journey around the world; in the
publishing of schoolbooks he was long associated with his
brother, Robert, on Washington Street, Boston; deeply
interested in Free Masonry, Colonel Davis had been Master
of Putnam Lodge, Cambridge, was a member of St. Paul
Chapter, Royal Arch, and was a charter member of St.
Bernard's Commandery, Knights Templar of Boston;
entering the Militia at a very early age, the beginning of
the War found him Division Inspector on the Staff of
General Samuel Andrews of the First Division; later
promoted to the rank of Brigadier General, he was serving
in 1862 on a Board of Examination, thereby rendering
signal aid to Governor Andrew; he passed thence to the
Thirty-ninth Regiment. On leaving his home, he said to his
mother who had expressed wonder, if not regret, at his
going, "Mother, if I should live to see the end of this war
without going and doing my whole duty to my country, I
should never rest," and he went away with her blessing.
Perhaps no man throughout the strife entered the service
with higher motives than those which prompted Colonel
Davis. Possessing as high an ideal of discipline and drill as
he had of morality and patriotism, he proceeded to
enforce them with the result that few if any organizations

in the volunteer service excelled the Thirty-ninth in true
soldierly qualities. Early called to the command of a
brigade, it was truly said of him that he never was
assigned to any position which he did not fill. The
particulars of his death, July 11, 1864, have appeared in
the body of this book; his funeral, held with Masonic
honors in the Unitarian Church of Cambridge, was on July
18, the entire city being in mourning, with all places of
business closed; flags were at half-mast and in the
audience assembled to honor his memory were the City
Council of Cambridge, Governor Andrew and Staff,
Adjutant General Schouler, Mayor Lincoln of Boston and a
wide range of other civil and military officers; Free
Masonry in which he was so prominent was represented
by Putnam Lodge to which he belonged, officers of the
Grand Lodge, St. Bernard's Encampment of Boston, and
the National Lancers also were present. Speakers at the
services were the Rev. Chandler Robbins, who had
officiated at his marriage, and Chaplain E. B. French who
had accompanied the remains of his commander home.
With the long escort, the body of Colonel Davis was borne
to Mt. Auburn Cemetery, having as bearers General
Samuel C. Lawrence, Colonels C. L. Holbrook and L. B.
Marsh, Postmaster Leighton and Deputy Sheriff L. L.
Parker, the burial being with Masonic rites.
Charles L. Peirson, from Lieut. Colonel July 13, 1864;
owing to the stress of the "Battle Summer" campaign, his
severe wound at the Weldon R. R. August 18, '64, and
subsequent absence from the Regiment, not to mention
the red tape that ever did hedge military matters about, it
was not till the 23d of November, 1864, that Colonel
Peirson was mustered in to his rank: the Records of the
War Department, Washington, D. C., state:
Peirson is now held and considered by this
Department, under the provisions of the Act of

Congress, approved February 24, 1897, to have
been mustered into the service of the United
States in the grade of Colonel, Thirty-ninth
Massachusetts Infantry, to take effect from July
13, 1864, and to have held that rank until the
date of his discharge from service.
Upon the recommendation of Major General G. K.
Warren, Peirson was commissioned Colonel of
Volunteers by brevet, to date from March 13,
1865, for meritorious conduct in the battles of the
Wilderness and Spottsylvania in May, 1864, and
as Brigadier General of Volunteers, by brevet, to
date from March 13, 1865, for gallant and
meritorious conduct in the battle of the Weldon
Railroad in August, 1864.
After months of prostration, incident to his
wound, and on the clear evidence of his inability
to return to the Regiment, Colonel Peirson
resigned and was mustered out of the service
January 11, 1865. Subsequent to the war,
General Peirson was long in the iron business,
Boston; on his retirement therefrom, he found
occupation for his well earned leisure in historical
studies, particularly with reference to the Civil
War, being a member of the Loyal Legion, which
he commanded, 1895, and the Massachusetts
Military Historical Society. His city residence is at
191 Commonwealth Avenue; his summer abode
is at Pride's Crossing, city of Beverly.

Colonel Charles L. Pierson
B'v't Brigadier-General

LIEUTENANT COLONELS
Charles L. Peirson, 28, S.; civil engineer, Salem; wd. May 8
and 10, '64, Spottsylvania; prom. Colonel; Charles
Lawrence Peirson was born in Salem; was graduated from
Lawrence Scientific School, Harvard, 1853; was a Corporal
in the Fourth Battalion, under Major T. G. Stevenson,
which in the spring of 1861 did gratuitous service in Fort
Warren, Boston Harbor; later commissioned First Lieut.
and Adjutant in the Twentieth Massachusetts, he was
taken prisoner at the Battle of Ball's Bluff and suffered
three months' confinement in Libby Prison, Richmond; on
his return to his regiment he was detailed for special
service on the staff of General N. J. T. Dana and also later
upon that of General John Sedgwick, thus passing through
the Peninsula campaign; it was while on sick leave from
such service that he was notified of his appointment to his
new position in the Thirty-ninth Regiment.
Henry M. Tremlett, from Major July 13, 1864; absent at
the time on detached service in Boston Harbor he did not
rejoin the Regiment until October following; wd. March 31,
'65, at Gravelly Run, he died of wounds at his home in
Boston, June 6th following, the very day of the return of
the Thirty-ninth. The six weeks immediately following the
battle were spent in the hospital at City Point; thence he
returned to Boston, getting there May 9th, apparently on
the road to recovery, but the setting in of intermittent
fever proved to be too great a trial of his strength; his
body was buried in Forest Hills Cemetery. Of him a writer
in a Boston paper wrote at the time:—

His standard of manliness was one of noble
action rather than of puling pretension, and his
whole life showed him to be a loving son, a dear
brother, a kind and generous companion, a
devoted friend and a truly loyal man, willing to
sacrifice his life for the noble cause for which he
contended.

MAJORS
Henry M. Tremlett, b. Dorchester, July 15, 1833; 29, S.;
merchant, Boston; Aug. 28, 1862; educated at Chauncy
Hall School, Boston, he succeeded his father in mercantile
life on Foster's Wharf; when Governor Andrew called for
volunteers to serve in Fort Warren in the spring of 1862,
he was one of those who filled the ranks of the Fourth
Battalion, serving therein as First Sergeant. On the
organization of the Twentieth Regiment, he was
commissioned Captain and in that capacity bore his part in
the fatal day at Ball's Bluff and was with the Army of the
Potomac through the Seven Days' Fight. With the Thirty-
ninth he participated in all of its experiences till, in the fall
of '63, he was ordered to Boston where for quite a year,
as Provost Marshal, he had charge of the draft rendezvous
till after the death of Colonel Davis and the severe
wounding of Colonel Peirson his return was necessary,
serving thereafter as Lieut. Colonel.
Frederick R. Kinsley, July 13, '64, from Captain, Co. E; not
mustered; captured, Aug. 19, '64, at the Weldon R. R.,
was held until the following March; came home in
command of the Regiment; M. O. as Capt., June 2, 1865;
soon after the war, with two brothers, he bought and
worked a large farm in Dorchester, N. H.; represented the
town in the Legislature; in 1911 he removed to Lowell
where, in 1913, he makes his home.

ADJUTANTS
Henry W. Moulton, 21, M.; currier, So. Danvers; Aug. 18,
'62; was first commissioned in the Thirty-fifth, Aug. 12,
'62, and was trans. as above. Owing to the detailing of
Adjutant Washburn, Lieutenant Moulton took his place;
wd. May 10, 1864, Laurel Hill; absent, sick, until his
discharge; dis. disa., Dec. 5, 1864.
Orville A. Barker, from Co. C, Dec. 5, '64; prom. Captain,
April 3, '65; not mustered; M. O. June 2, 1864; a druggist
for many years in Taunton, Captain Barker found time to
serve as Treasurer of Morton Hospital and for thirty-five
years was Clerk of the Baptist Church; he died Feb. 21,
1912.

QUARTERMASTER
Edward E. White, 34, —; —, Cambridge; August 25, 1862;
prom. Captain, April 3, '65; not mustered; brevet Capt.
and Major, U. S. Volunteers, March 13, 1865; M. O. as 1st
Lieut., June 2, 1865.

SURGEONS
Calvin G. Page, 33, —; physician, Boston; August 22,
1862; dis. as Major, disa., Nov. 16, '63; an A. B., Harvard,
1852, he took his M. D. there in 1854; d. March 29, 1869.
William Thorndike, 29, M.; surgeon, Beverly; Nov. 17,
1863; an A. B. from Harvard, 1854, he also gained there
his M. D., 1857; had seen service as Ass't Surgeon, Thirty-
fourth Massachusetts Volunteers, whence he came to the
Thirty-ninth; his efficiency in the Regiment was thoroughly
appreciated by the men, and General Peirson affirms that
recovery from the wound received at the Weldon R. R.
was the result of the care and attention of his surgeon;
the son of the latter, William, Jr., also Harvard, 1892, and
M. D., 1896, is a Boston practitioner, whose wife is a
daughter of the late General William Tecumseh Sherman;
Surgeon Thorndike died in 1887.

ASSISTANT SURGEONS
James L. Chipman, 31, —; physician, Milford; August 25,
1862; dis. as 1st Lieut. disa., May 23, '64; later, June 26,
'65, 1st Lieut. and Ass't Surg. Forty-third U. S. C. T.; M. O.
Oct. 20, '65.
Henry H. Mitchell, 23, —; physician, East Bridgewater;
August 25, 1862; res. Nov. 3, '64, as 1st Lieut. for prom.
as Major and Surgeon, Thirty-sixth U. S. C. T.; res. June
15, 1864.
John F. Butler, —, —; physician, Chesterfield, N. H.; 1st
Lieut. May 27, 1863; an M. D. from Harvard, 1854, a
classmate of Surgeon Thorndike, he was M. O. June 2,
1865.

CHAPLAIN
Edward Beecher French, 29, M.; clergyman, Chatham;
August 18, 1862; a graduate of Harvard's Divinity School,
1859, Chaplain French enlisted as a private from his
pastorate, and was commissioned from the ranks; of him
Thomas E. Small remarks, "At the battle of the Wilderness
the Chaplain was right up at the front with the boys and
when Daniel Burnham of our Company was shot and
about to die, the Chaplain took his last message and
whatever he had to send to his wife and family and
comforted him in his last few moments of life; he
accompanied the remains of Colonel Davis from
Petersburg to Cambridge and spoke at the funeral; M. O.
June 2, 1865; he was born in Lowell, Nov. 20, 1832; his
earlier years were spent in Holliston; his first pastorate
was in Chatham, whence he was the first man to enlist in
the Thirty-ninth; after the war he served pastorates in
Babylon, L. I., and Perth Amboy, N. J., but his health,
enfeebled by exposures at the front, broke and recovery
was sought in Texas and Wisconsin, but without avail. He
died July 14, 1907, in Harwich with relatives of his wife,
who had preceded him to the other world, and his body
was laid by the side of hers in the Harwich burial ground."

NON-COMMISSIONED STAFF
Sergeant Majors
Charles Henry Chapman, 21, S.; student, Cambridge;
prom. 2d Lieut. Co. G, Nov. 11, 1862; Brown University,
Class of 1861.
T. Cordis Clarke from Co. B; Dec. 8, '62; prom. 2d Lieut.
Nov. 13, '62; vide Co. E.
Charles W. Hanson, from Co. A; Dec. 6, '62; prom. 2d
Lieut. Jan. 25, '63; vide Co. H.
Joseph A. Merrifield, from Co. A, Feb. 20, '63; prom. 2d
Lieut. Sept. 20, '63; vide Co. F.
Edwin Mills, from Co. E; Sept., '63; prom. 2d Lieut. Jan. 8,
'64; vide Co. A.
Charles K. Conn, from Co. K, April 28, '64; wd. and pris.
May 8, '64; prom. 2d Lieut. Feb. 1, '65; vide Co. H.
George H. Dennett, from Co. K, Feb. 1, '65; prom. 2d
Lieut.; not mustered; M. O. June 2, 1865, as Sergt. Major;
d. Malden.
Quartermaster Sergeant
Henry B. Leighton, 25, —; —, Cambridge; Sept. 4, 1862;
prom. 2d Lieut. April 3, '65; not mustered; M. O. as Q. M.
Sergt. June 2, 1865.
Commissary Sergeant
Lucius W. Hilton, 21, —; —, —; Sept. 4, 1862; M. O. June
2, 1865.
Hospital Steward

Frederick Harvey, 27, M.; apothecary, Dorchester; dis.
Sept. 7, 1863, S. O. W. D.
Orville A. Barker, from Co. F, Oct. 13, '62; prom. 2d Lieut.
Nov. 8, '63; vide Co. C.
George A. Stuart, 22, S.; chemist, Boston; March 9, 1864;
trans. June 2, 1865, to Thirty-second Infantry.
Principal Musician
Matthew Woodward, from Co. F, Nov. 1, '63; M. O. June 2,
1865.
(To avoid needless repetition of dates in regard to
transfers to and from the Regiment the following facts are
stated here:—June 25, '64, on the M. O. of the Twelfth
Massachusetts Infantry, the men whose enlistments had
not expired were trans. to the Thirty-ninth, and on the
13th, of July, '64, under similar circumstances, men were
received from the Thirteenth Massachusetts. When, June
2, '65, the Thirty-ninth was preparing to go home all
members whose terms were not expiring were trans. to
the Thirty-second Massachusetts and were M. O. with that
organization June 29, 1865).
In battle-names, Spottsylvania may include both Alsop's
Farm and Laurel Hill.

COMPANY A
From South Danvers, after the War to become the town of
Peabody.
Captains
George S. Nelson, 27, M.; tanner, South Danvers; August
18, '62; res. March 2, 1865; had been commissioned
Captain in the Thirty-fifth, August 12, '62, and was trans.
as above; at last account, Capt. Nelson's address was 880
Seminary Avenue, Chicago.
As Acting Captain, 1st Lieut. Henry F. Felch of Company E
commanded the Company on its return to Boston.
First Lieutenants
Emory Washburn, Jr., 24, —; lawyer, Cambridge; Aug. 25,
'62; the son of Ex-Governor Emory Washburn, he was
born in Worcester, Oct. 1, 1837; graduating from Harvard
College in 1860, he had just taken his degree of LL. B. in
1862 when he was commissioned in the new regiment
then forming; evidently his direct service, if any, in the
Thirty-ninth was brief, for on the first Monthly Report he
appears as detached and a member of the staff of General
Charles Devens, also a Worcester man; in this capacity he
did excellent work, as appears in the report of General
Devens, after the battle of Fredericksburg, written Dec.
17, '62, wherein he says, "I am under especial obligations,
for their zeal and fidelity, to my staff," including with two
others, "my aide, Lieut. E. Washburn, Jr." It would appear
that Adjutant Washburn returned to the Regiment for one
week at Poolesville, Md., resigning, January 24, 1864; he
died in 1885.

Charles H. Porter, from 2d Lieut. Co. D, Jan. 25, '63; prom.
Captain, Sept. 8, '64; not mustered; M. O. as 1st Lieut.
June 2, 1865; as a member of the Loyal Legion, Captain
Porter was conspicuous in promoting its interests; was
Junior Vice-Commander, 1897; Registrar, 1903-5;
Recorder, 1906-11; no veteran of the Regiment took more
interest in its annual reunions than did Captain Porter, and
for years he was practically its motive power. His papers
on the campaigns in which he bore a part were valuable
contributions to the Massachusetts Military Historical
Society. Born in Weymouth, 1843, he was only six weeks
old when the family removed to Quincy; his early
education was had in the Quincy High School; his business
life was that of insurance; he was almost constantly in
public life, twelve years on the School Board, three years
Selectman, the First Mayor of Quincy, 1888, he was re-
elected; several years on the State Board of Health; first
Commander, Paul Revere Post, G. A. R.; commissioned as
Lieut. Colonel in Seventh M. V. M. by Governor Andrew, he
was widely known as Colonel Porter; he was seven years
Trustee of the Chelsea Soldiers' Home and was ever
prominent in local business organizations and in Masonic
Circles; d. Aug. 10, 1911.
Second Lieutenants
George H. Wiley, 23, M.; shoemaker, So. Danvers; Aug.
18, '62; res. Jan. 7, '64; had been commissioned in the
Thirty-fifth Aug. 12, '62, and was trans. as above; was 3d
Lieut. Co. H, Fifth M. V. M., three mos. service; d. May 19,
1910, Boston.
L. F. Wyman, Feb. 23, '64, from Co. K; returned to "K,"
Mar. 2, '64.
Edwin Mills from Sergeant Major, Jan. 8, '64; dis. on
account of wds. rec'd May 10, '64, Oct. 19, '64.

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