Spirochaete

Spirochaetes
	Treponema pallidum, a spirochaete which causes syphilis
Scientific classification
Domain:	Bacteria
Phylum:	Spirochaetota; Garrity and Holt 2021[1]
Class:	Spirochaetia; Paster 2020[2][3]
Orders
	Brachyspirales; Brevinematales; Leptospirales; Spirochaetales; "Treponematales";
Synonyms
	Spirochaetota: "Protozoobacteriales" Prévot 1958; "Spirochaetae" Cavalier-Smith 2002; "Spirochaetaeota" Oren et al. 2015; "Spirochaetes" Garrity and Holt 2001; "Spirochaetota" Whitman et al. 2018; ; Spirochaetia: Spirochaetes Cavalier-Smith 2002; "Spirochaetia" Cavalier-Smith 2020; "Leptospiria" Cavalier-Smith 2020; ;

A spirochaete (/ˈspaɪroʊˌkiːt/)[4] or spirochete is a member of the phylum Spirochaetota (/-ˈkiːtiːz/), (synonym Spirochaetes) [5] which contains distinctive diderm (double-membrane) gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or spiraled, hence the name) cells.[6] Spirochaetes are chemoheterotrophic in nature, with lengths between 3 and 500 μm and diameters around 0.09 to at least 3 μm.[7]

Cross section of a spirochaete cell

Endoflagella Components. Legend: Fig. 1 A cross-section of a typical spirochete cell showing endoflagella located in the periplasm between the inner cytoplasmic membrane and the outer membrane. Periplasm, consisting of a gel-like matrix, provides a semi-stable medium to secure endoflagella during rotation. The axial filament, indicated in red, is composed of bundles of endoflagella. Fig. 2 A side-view of a spirochete cell which shows two axial filaments in opposing motion. One axial filament rotates in a clockwise orientation; an adjacent axial filament rotates in a counter-clockwise orientation. Rotation of the endoflagella creates torsion and drives the corkscrew rotation of the cell. Fig. 3 An expanded view of the cellular membranes that surround endoflagellum. Both the inner and outer membrane contain a phospholipid bi-layer, with non-polar fatty acid chains in-ward of polar phosphorus heads. Peptidoglycan, the cell wall, provides structure in bacterial microorganisms. Axial filaments are superior to the peptidoglycan.

Spirochaetes are distinguished from other bacterial phyla by the location of their flagella, called endoflagella which are sometimes called axial filaments.[8] Endoflagella are anchored at each end (pole) of the bacterium within the periplasmic space (between the inner and outer membranes) where they project backwards to extend the length of the cell.[9] These cause a twisting motion which allows the spirochaete to move about. When reproducing, a spirochaete will undergo asexual transverse binary fission. Most spirochaetes are free-living and anaerobic, but there are numerous exceptions. Spirochaete bacteria are diverse in their pathogenic capacity and the ecological niches that they inhabit, as well as molecular characteristics including guanine-cytosine content and genome size.[10][11]

Pathogenicity

Many organisms within the Spirochaetota phylum cause prevalent diseases. Pathogenic members of this phylum include the following:

Leptospira species, which causes leptospirosis[12]
Borrelia burgdorferi, B. mayonii, B. bissettiae, B. garinii, B. afzelii, B. spielmanii, B. lusitaniae, which cause Lyme disease.[13]
Borrelia recurrentis, which causes relapsing fever[14]
Treponema pallidum subspecies which cause treponematoses such as syphilis and yaws.
Brachyspira pilosicoli and Brachyspira aalborgi, which cause intestinal spirochaetosis[15]

Salvarsan, the first partially organic synthetic antimicrobial drug in medical history, was effective against spirochaetes and primarily used to cure syphilis. Additionally, oral spirochaetes are known to play a significant role in the pathogenesis of human periodontal disease.[16]

Taxonomy and molecular signatures

The class currently consists of 14 validly named genera across 4 orders and 5 families.[17][18][19] The orders Brachyspirales, Brevinematales and Leptospirales each contain a single family, Brachyspiraceae, Brevinemataceae and Leptospiraceae, respectively. The Spirochaetales order harbours two families, Spirochaetaceae and Borreliaceae. Molecular markers in the form of conserved signature indels (CSIs) and CSPs have been found specific for each of the orders, with the exception of Brevinimetales, that provide a reliable means to demarcate these clades from one another within the diverse phylum.[18] Additional CSIs have been found exclusively shared by each family within the Spirochaetales. These molecular markers are in agreement with the observed phylogenetic tree branching of two monophyletic clades within the Spirochaetales order.[18] CSIs have also been found that further differentiate taxonomic groups within the Borreliaceae family that further delineate evolutionary relationships that are in accordance with physical characteristics such as pathogenicity (viz. Borrelia emend. Borreliella gen. nov.).[20] However, this study has been criticized, and other studies using different approaches do not support the proposed split.[21] The new naming system for the Lyme and relapsing fever Borrelia has not been adopted by the scientific literature.[21]

A CSI has also been found exclusively shared by all Spirochaetota species.[18] This CSI is a 3 amino acid insert in the flagellar basal body rod protein FlgC which is an important part of the unique endoflagellar structure shared by Spirochaetota species.[22] Given that the CSI is exclusively shared by members within this phylum, it has been postulated that it may be related to the characteristic flagellar properties observed among Spirochaetota species.[18][22]

Historically, all families belonging to the Spirochaetota phylum were assigned to a single order, the Spirochaetales.[10][11] However, the current taxonomic view is more connotative of accurate evolutionary relationships. The distribution of a CSI is indicative of shared ancestry within the clade for which it is specific. It thus functions as a synapomorphic characteristic, so that the distributions of different CSIs provide the means to identify different orders and families within the phylum and so justify the phylogenetic divisions.[18]

Phylogeny

16S rRNA based LTP_12_2021[23][24][25]

120 single copy marker proteins based GTDB 08-RS214[26][27][28]

Spirochaetia

Leptospirales

Leptospiraceae

Turneriella

	Leptonema

	Leptospira

Brevinematales

Brevinemataceae	Brevinema

Brachyspirales

Brachyspiraceae	Brachyspira

Spirochaetales

Exilispira

Rectinemataceae	"Rectinema"

Treponemataceae	Treponema

Sphaerochaetaceae

Spirochaeta thermophila

Sediminispirochaeta

Marispirochaeta

Pleomorphochaeta

	Bullifex

	Sphaerochaeta

Spirochaetaceae

	Spirochaeta lutea

	Salinispira

Spirochaeta

	Spirochaeta halophila

	Alkalispirochaeta

Spirochaeta species-group 2

Oceanispirochaeta

	Spirochaeta cellobiosiphila

	Thiospirochaeta

Spirochaeta aurantia

Borreliaceae

	Borrelia

	Borreliella

"Leptospirae"

"Leptospiria"

"Turneriellales"

"Turneriellaceae"	Turneriella

Leptospirales

"Leptonemataceae"	Leptonema

Leptospiraceae	Leptospira

"Euspirochaetae"

"Brevinematia"

Brevinematales

Brevinemataceae	Brevinema

"Brachyspirae"

Brachyspirales

Brachyspiraceae	Brachyspira

"Exilispiria"

"Exilispirales"

"Exilispiraceae"	Exilispira

Spirochaetia

"Borreliales"

Borreliaceae

	Borrelia

	Borreliella

WRBN01

WRBN01	"Entomospira"

DSM‑17781

DSM‑17781	Spirochaeta cellobiosiphila

Spirochaetales_E

DSM‑19205

Thiospirochaeta

Spirochaetaceae_B

	Spirochaeta species-group 2

	Oceanispirochaeta

"Sphaerochaetales"

Sphaerochaetaceae

Parasphaerochaeta

Sphaerochaeta

	Bullifex

	"Ca. Ornithospirochaeta"

JC444

"Marispirochaetaceae"	Marispirochaeta

DSM‑16054

"Sediminispirochaetaceae"	Sediminispirochaeta

Spirochaetales_A

Spirochaetaceae_A	Spirochaeta thermophila

DSM‑27196

"Salinispiraceae"

	Spirochaeta lutea

	Salinispira

DSM‑8902	Spirochaeta africana

"Alkalispirochaetaceae"	Alkalispirochaeta

"Treponematales"

"Rectinemataceae"	Rectinema

Breznakiellaceae

Gracilinema

Breznakiella

	Leadbettera

	Treponema primitia

Treponemataceae

"Ca. Gallitreponema"

Treponema species-group 2

	"Ca. Avitreponema"

	Brucepastera

Treponema

Taxonomy

The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN)[29] and National Center for Biotechnology Information (NCBI).[30]

Phylum Spirochaetota Garrity and Holt 2021
- Class Spirochaetia Paster 2020
  - Genus ?"Spirosymplokos" Guerrero et al. 1993
  - Order Leptospirales Gupta et al. 2014
    - Family Leptospiraceae Hovind-Hougen 1979
      - Genus Leptonema Hovind-Hougen 1983
      - Genus Leptospira Noguchi 1917
      - Genus Turneriella Levett et al. 2005
  - Order Brachyspirales corrig. Gupta et al. 2014
    - Family Brachyspiraceae Paster 2012
      - Genus Brachyspira Hovind-Hougen et al. 1982
      - Genus ?"Ca. Maribrachyspira" Matsuyama et al. 2017
  - Order Brevinematales Gupta et al. 2014
    - Family Brevinemataceae Paster 2012
      - Genus Brevinema Defosse et al. 1995
    - Family "Longinemaceae" Karnachuk et al. 2021
      - Genus ?"Longinema" Karnachuk et al. 2021
    - Family "Thermospiraceae" Ben Ali Gam et al. 2023
      - Genus ?Thermospira Ben Ali Gam et al. 2023
  - Order "Exilispirales" Pallen, Rodriguez-R & Alikhan 2022
    - Family "Exilispiraceae" Pallen, Rodriguez-R & Alikhan 2022
      - Genus Exilispira Imachi et al. 2008
  - Order Spirochaetales Buchanan 1917 ["Entomospirales" Pallen, Rodriguez-R & Alikhan 2022; "Marispirochaetales" Pallen, Rodriguez-R & Alikhan 2022; "Salinispirales" Pallen, Rodriguez-R & Alikhan 2022; "Sediminispirochaetales" Pallen, Rodriguez-R & Alikhan 2022]
    - Family "Pillotinaceae" Margulis & Hinkle 1992
      - Genus ?Pillotina Hollande and Gharagozlou 1967 ex Bermudes et al. 1988
    - Family Borreliaceae Gupta et al. 2014
      - Genus Borrelia Swellengrebel 1907 (relapsing fever Borrelia, Lreptile-associated Borrelia and Echidna-associated Borrelia)
      - Genus Borreliella Adeolu & Gupta 2015 (yme borreliosis Borrelia)
      - Genus ?Cristispira Gross 1910
    - Family Sphaerochaetaceae Hördt et al. 2020 ["Marispirochaetaceae" Pallen, Rodriguez-R & Alikhan 2022; "Sediminispirochaetaceae" Pallen, Rodriguez-R & Alikhan 2022]
      - Genus Bullifex Wylensek et al. 2021
      - Genus Marispirochaeta Shivani et al. 2017
      - Genus "Ca. Ornithospirochaeta" Gilroy et al. 2021
      - Genus Parasphaerochaeta Bidzhieva et al. 2020
      - Genus ?Pleomorphochaeta Arroua et al. 2016
      - Genus Sediminispirochaeta Shivani et al. 2016
      - Genus Sphaerochaeta Ritalahti et al. 2012
    - Family Spirochaetaceae Swellengrebel 1907 ["Entomospiraceae" Pallen, Rodriguez-R & Alikhan 2022; "Salinispiraceae" Pallen, Rodriguez-R & Alikhan 2022; "Thiospirochaetaceae" Pallen, Rodriguez-R & Alikhan 2022]
      - Genus ?"Canaleparolina" Wier, Ashen & Margulis 2000
      - Genus ?Clevelandina Bermudes et al. 1988
      - Genus ?Diplocalyx Gharagozlou 1968 ex Bermudes et al. 1988 non Richard 1850 non Presl 1845
      - Genus ?Hollandina To et al. 1978 ex Bermudes et al. 1988 non Haynes 1956
      - Genus ?"Mobilifilum" Margulis et al. 1990
      - Genus Alkalispirochaeta Sravanthi et al. 2016
      - Genus "Entomospira" Grana-Miraglia et al. 2020 non Enderlein 1917
      - Genus Oceanispirochaeta Subhash & Lee 2017b
      - Genus Salinispira Ben Hania et al. 2015
      - Genus Spirochaeta Ehrenberg 1835 em. Pikuta et al. 2009 non Turczaninow 1851
      - Genus Thiospirochaeta Dubinina et al. 2020
  - Order "Treponematales" Song et al. 2021
    - Family Breznakiellaceae Brune et al. 2022
      - Genus Breznakiella Song et al. 2022
      - Genus Gracilinema Brune et al. 2022
      - Genus ?Helmutkoenigia Brune et al. 2022
      - Genus Leadbettera Brune et al. 2022
      - Genus ?Zuelzera Brune et al. 2022
    - Family "Rectinemataceae" Song et al. 2021
      - Genus Rectinema Koelschbach et al. 2017
    - Family Treponemataceae Robinson 1948
      - Genus "Ca. Avitreponema" Gilroy et al. 2021
      - Genus "Brucepastera" Song et al. 2022
      - Genus "Ca. Gallitreponema" Gilroy et al. 2021
      - Genus ?"Teretinema" Song et al. 2022
      - Genus Treponema Schaudinn 1905 em. Abt et al. 2013

References

Oren A, Garrity GM (2021). "Valid publication of the names of forty-two phyla of prokaryotes". Int J Syst Evol Microbiol. 71 (10): 5056. doi:10.1099/ijsem.0.005056. PMID 34694987. S2CID 239887308.
Paster BJ (2010). "Class I. Spirochaetia class. nov.". In Krieg NR, Staley JT, Brown DR, Hedlund BP, Paster BJ (eds.). Bergey's Manual of Systematic Bacteriology. Vol. 4—The Bacteroidetes, Spirochaetes, Tenericutes (Mollicutes), Acidobacteria, Fibrobacteres, Fusobacteria, Dictyoglomi, Gemmatimonadetes, Lentisphaerae, Verrucomicrobia, Chlamydiae, and Planctomycetes (2nd ed.). New York, NY: Springer. doi:10.1007/978-0-387-68572-4. ISBN 978-0-387-95042-6.
Oren A, Garrity GM (2020). "Validation list no. 195. List of new names and new combinations previously effectively, but not validly, published". Int J Syst Evol Microbiol. 70 (9): 4844–4847. doi:10.1099/ijsem.0.004366. PMID 32993851. S2CID 222147003.
"SPIROCHAETE | Meaning & Definition for UK English | Lexico.com". Lexico Dictionaries | English. Archived from the original on 27 January 2021.
Dorland's Illustrated Medical Dictionary. Elsevier.
Ryan KJ, Ray CG, eds. (2004). Sherris Medical Microbiology (4th ed.). McGraw Hill. ISBN 978-0-8385-8529-0.
Margulis L, Ashen JB, Solé M, Guerrero R (August 1993). "Composite, large spirochetes from microbial mats: spirochete structure review". Proceedings of the National Academy of Sciences of the United States of America. 90 (15): 6966–6970. Bibcode:1993PNAS...90.6966M. doi:10.1073/pnas.90.15.6966. PMC 47056. PMID 8346204.
Carroll KC, Hobden JA, Miller S (2019). "Spirochetes and Other Spiral Microorganisms". Jawetz, Melnick, & Adelberg’s Medical Microbiology. McGraw-Hill Education. Retrieved 9 May 2021.
Madigan MT (2019). Brock biology of microorganisms (Fifteenth, Global ed.). NY, NY: Pearson. p. 519. ISBN 9781292235103.
Paster BJ (2011). "Phylum XV. Spirochaetes Garrity and Holt.". In Brenner DJ, Krieg NR, Garrity GM, Staley JT (eds.). Bergey's Manual of Systematic Bacteriology. New York: Springer. p. 471.
Paster BJ (2011). "Family I. Sprochaetes Swellengrebel 1907, 581AL.". In Brenner DJ, Krieg NR, Garrity GM, Staley JT (eds.). Bergey's Manual of Systematic Bacteriology. New York: Springer. pp. 473–531.
McBride A, Athanazio D, Reis M, Ko A (2005). "Leptospirosis". Current Opinion in Infectious Diseases. 18 (5): 376–86. doi:10.1097/01.qco.0000178824.05715.2c. PMID 16148523. S2CID 220576544.
Wolcott KA, Margos G, Fingerle V, Becker NS (September 2021). "Host association of Borrelia burgdorferi sensu lato: A review". Ticks and Tick-Borne Diseases. 12 (5): 101766. doi:10.1016/j.ttbdis.2021.101766. PMID 34161868.
Schwan TG (June 1996). "Ticks and Borrelia: model systems for investigating pathogen-arthropod interactions". Infectious Agents and Disease. 5 (3): 167–181. PMID 8805079.
Amat Villegas I, Borobio Aguilar E, Beloqui Perez R, de Llano Varela P, Oquiñena Legaz S, Martínez-Peñuela Virseda JM (January 2004). "[Colonic spirochetes: an infrequent cause of adult diarrhea]". Gastroenterol Hepatol (in Spanish). 27 (1): 21–3. doi:10.1016/s0210-5705(03)70440-3. PMID 14718105.
Yousefi L, Leylabadlo HE, Pourlak T, Eslami H, Taghizadeh S, Ganbarov K, et al. (July 2020). "Oral spirochetes: Pathogenic mechanisms in periodontal disease". Microbial Pathogenesis. 144: 104193. doi:10.1016/j.micpath.2020.104193. PMID 32304795. S2CID 215818931.
Schoch CL, Ciufo S, Domrachev M, Hotton CL, Kannan S, Khovanskaya R, et al. "TherSpirochaetia". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 25 October 2016.
Gupta RS, Mahmood S, Adeolu M (2013). "A phylogenomic and molecular signature based approach for characterization of the phylum Spirochaetes and its major clades: proposal for a taxonomic revision of the phylum". Front Microbiol. 4 (217): 217. doi:10.3389/fmicb.2013.00217. PMC 3726837. PMID 23908650.
Oren A, Garrity GM (2014). "List of new names and new combinations previously effectively, but not validly, published" (PDF). Int. J. Syst. Evol. Microbiol. 64 (3): 693–696. doi:10.1099/ijs.0.062521-0.
Adeolu M, Gupta RS (2014). "A phylogenomic and molecular marker based proposal for the division of the genus Borrelia into two genera: the emended genus Borrelia containing only the members of the relapsing fever Borrelia, and the genus Borreliella gen. nov. containing the members of the Lyme disease Borrelia (Borrelia burgdorferi sensu lato complex)". Antonie van Leeuwenhoek. 105 (6): 1049–1072. doi:10.1007/s10482-014-0164-x. PMID 24744012.
Winslow C, Coburn J (2019). "Recent discoveries and advancements in research on the Lyme disease spirochete Borrelia burgdorferi". F1000Research. 8: 763. doi:10.12688/f1000research.18379.1. PMC 6545822. PMID 31214329.
Macnab RM (2003). "How bacteria assemble flagella". Annu Rev Microbiol. 57: 77–100. doi:10.1146/annurev.micro.57.030502.090832. PMID 12730325.
"The LTP". Retrieved 23 February 2021.
"LTP_all tree in newick format". Retrieved 23 February 2021.
"LTP_12_2021 Release Notes" (PDF). Retrieved 23 February 2021.
"GTDB release 08-RS214". Genome Taxonomy Database. Retrieved 10 May 2023.
"bac120_r214.sp_label". Genome Taxonomy Database. Retrieved 10 May 2023.
"Taxon History". Genome Taxonomy Database. Retrieved 10 May 2023.
Euzéby JP. "Spirochaetes". List of Prokaryotic names with Standing in Nomenclature (LPSN). Retrieved 20 July 2018.
Schoch CL, Ciufo S, Domrachev M, Hotton CL, Kannan S, Khovanskaya R, et al. "Spirochaetes". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 20 July 2018.

External links

Introduction to the Spirochetes University of California Museum of Paleontology (UCMP)

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[1] Oren A, Garrity GM (2021). "Valid publication of the names of forty-two phyla of prokaryotes". Int J Syst Evol Microbiol. 71 (10): 5056. doi:10.1099/ijsem.0.005056. PMID 34694987. S2CID 239887308.

[Paster_2010-2] Paster BJ (2010). "Class I. Spirochaetia class. nov.". In Krieg NR, Staley JT, Brown DR, Hedlund BP, Paster BJ (eds.). Bergey's Manual of Systematic Bacteriology. Vol. 4—The Bacteroidetes, Spirochaetes, Tenericutes (Mollicutes), Acidobacteria, Fibrobacteres, Fusobacteria, Dictyoglomi, Gemmatimonadetes, Lentisphaerae, Verrucomicrobia, Chlamydiae, and Planctomycetes (2nd ed.). New York, NY: Springer. doi:10.1007/978-0-387-68572-4. ISBN 978-0-387-95042-6.

[3] Oren A, Garrity GM (2020). "Validation list no. 195. List of new names and new combinations previously effectively, but not validly, published". Int J Syst Evol Microbiol. 70 (9): 4844–4847. doi:10.1099/ijsem.0.004366. PMID 32993851. S2CID 222147003.

[4] "SPIROCHAETE | Meaning & Definition for UK English | Lexico.com". Lexico Dictionaries | English. Archived from the original on 27 January 2021.

[Dorlands-5] Dorland's Illustrated Medical Dictionary. Elsevier.

[Sherris-6] Ryan KJ, Ray CG, eds. (2004). Sherris Medical Microbiology (4th ed.). McGraw Hill. ISBN 978-0-8385-8529-0.

[7] Margulis L, Ashen JB, Solé M, Guerrero R (August 1993). "Composite, large spirochetes from microbial mats: spirochete structure review". Proceedings of the National Academy of Sciences of the United States of America. 90 (15): 6966–6970. Bibcode:1993PNAS...90.6966M. doi:10.1073/pnas.90.15.6966. PMC 47056. PMID 8346204.

[McGraw-8] Carroll KC, Hobden JA, Miller S (2019). "Spirochetes and Other Spiral Microorganisms". Jawetz, Melnick, & Adelberg’s Medical Microbiology. McGraw-Hill Education. Retrieved 9 May 2021.

[Brock-9] Madigan MT (2019). Brock biology of microorganisms (Fifteenth, Global ed.). NY, NY: Pearson. p. 519. ISBN 9781292235103.

[Paster_2011-10] Paster BJ (2011). "Phylum XV. Spirochaetes Garrity and Holt.". In Brenner DJ, Krieg NR, Garrity GM, Staley JT (eds.). Bergey's Manual of Systematic Bacteriology. New York: Springer. p. 471.

[Paster_2011b-11] Paster BJ (2011). "Family I. Sprochaetes Swellengrebel 1907, 581AL.". In Brenner DJ, Krieg NR, Garrity GM, Staley JT (eds.). Bergey's Manual of Systematic Bacteriology. New York: Springer. pp. 473–531.

[McBride_2005-12] McBride A, Athanazio D, Reis M, Ko A (2005). "Leptospirosis". Current Opinion in Infectious Diseases. 18 (5): 376–86. doi:10.1097/01.qco.0000178824.05715.2c. PMID 16148523. S2CID 220576544.

[13] Wolcott KA, Margos G, Fingerle V, Becker NS (September 2021). "Host association of Borrelia burgdorferi sensu lato: A review". Ticks and Tick-Borne Diseases. 12 (5): 101766. doi:10.1016/j.ttbdis.2021.101766. PMID 34161868.

[Schwan_1996-14] Schwan TG (June 1996). "Ticks and Borrelia: model systems for investigating pathogen-arthropod interactions". Infectious Agents and Disease. 5 (3): 167–181. PMID 8805079.

[pmid14718105-15] Amat Villegas I, Borobio Aguilar E, Beloqui Perez R, de Llano Varela P, Oquiñena Legaz S, Martínez-Peñuela Virseda JM (January 2004). "[Colonic spirochetes: an infrequent cause of adult diarrhea]". Gastroenterol Hepatol (in Spanish). 27 (1): 21–3. doi:10.1016/s0210-5705(03)70440-3. PMID 14718105.

[16] Yousefi L, Leylabadlo HE, Pourlak T, Eslami H, Taghizadeh S, Ganbarov K, et al. (July 2020). "Oral spirochetes: Pathogenic mechanisms in periodontal disease". Microbial Pathogenesis. 144: 104193. doi:10.1016/j.micpath.2020.104193. PMID 32304795. S2CID 215818931.

[NCBI-17] Schoch CL, Ciufo S, Domrachev M, Hotton CL, Kannan S, Khovanskaya R, et al. "TherSpirochaetia". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 25 October 2016.

[pmid23908650-18] Gupta RS, Mahmood S, Adeolu M (2013). "A phylogenomic and molecular signature based approach for characterization of the phylum Spirochaetes and its major clades: proposal for a taxonomic revision of the phylum". Front Microbiol. 4 (217): 217. doi:10.3389/fmicb.2013.00217. PMC 3726837. PMID 23908650.

[Oren_&_Garrity-19] Oren A, Garrity GM (2014). "List of new names and new combinations previously effectively, but not validly, published" (PDF). Int. J. Syst. Evol. Microbiol. 64 (3): 693–696. doi:10.1099/ijs.0.062521-0.

[pmid24744012-20] Adeolu M, Gupta RS (2014). "A phylogenomic and molecular marker based proposal for the division of the genus Borrelia into two genera: the emended genus Borrelia containing only the members of the relapsing fever Borrelia, and the genus Borreliella gen. nov. containing the members of the Lyme disease Borrelia (Borrelia burgdorferi sensu lato complex)". Antonie van Leeuwenhoek. 105 (6): 1049–1072. doi:10.1007/s10482-014-0164-x. PMID 24744012.

[Winslow2019-21] Winslow C, Coburn J (2019). "Recent discoveries and advancements in research on the Lyme disease spirochete Borrelia burgdorferi". F1000Research. 8: 763. doi:10.12688/f1000research.18379.1. PMC 6545822. PMID 31214329.

[Macnab03-22] Macnab RM (2003). "How bacteria assemble flagella". Annu Rev Microbiol. 57: 77–100. doi:10.1146/annurev.micro.57.030502.090832. PMID 12730325.

[23] "The LTP". Retrieved 23 February 2021.

[24] "LTP_all tree in newick format". Retrieved 23 February 2021.

[25] "LTP_12_2021 Release Notes" (PDF). Retrieved 23 February 2021.

[about-26] "GTDB release 08-RS214". Genome Taxonomy Database. Retrieved 10 May 2023.

[tree-27] "bac120_r214.sp_label". Genome Taxonomy Database. Retrieved 10 May 2023.

[taxon_history-28] "Taxon History". Genome Taxonomy Database. Retrieved 10 May 2023.

[29] Euzéby JP. "Spirochaetes". List of Prokaryotic names with Standing in Nomenclature (LPSN). Retrieved 20 July 2018.

[30] Schoch CL, Ciufo S, Domrachev M, Hotton CL, Kannan S, Khovanskaya R, et al. "Spirochaetes". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 20 July 2018.