Streptomyces

Streptomyces griseus, from the Actinomycetes Society of Japan.

Belongs within: Streptomycetaceae.

Streptomyces is a diverse genus of filamentous bacteria forming aerial mycelia.

Cutting up the excess
Published 15 November 2007
Generalised proteasome, from Answers.com.

Just before I start, I’d like to point out that I have very little understanding of the world of biochemistry. Most of it goes so far above my head that it causes detours on major airline routes. Therefore, I don’t expect you to believe a single word of what I’m about to say.

Three-domain phylogeny, from Palaeos.

Many of you who have passed through university biology will have seen some variation of the above diagram. It shows the Three Domain Hypothesis of Woese and colleagues, based on 18S rDNA analysis, dividing life into the eukaryotes, eubacteria and archaebacteria. In many cases, you’ll have seen it with a root on the branch dividing the eubacteria from the other two domains. A few years back, I wrote an essay for Palaeos on the phylogenetic relationships of bacteria. It wasn’t a bad piece of work, and I’m still reasonably happy with it. Toby White wrote a much longer essay last year that largely superseded what I’d written, though I’m not sure I’d agree with everything in it (for instance, we seem to have a tacit agreement to disagree on the usefulness of sequence data). Anyway, the upshot of these two pieces was that, while said division of life has been widely reiterated in textbooks and introductory courses, the actual evidence for it is not as strong as might be hoped. The vast majority of researchers agree that there is some sort of phylogenetic reality to the three domains, but whether a given domain represents a monophyletic clade or paraphyletic grade is a hotly debated topic. There is an even more significant amount of disagreement about what are the correct interrelationships within the eubacteria, and which members of the eubacteria are the most basal or closest to Neomura (archaebacteria + eukaryotes), whether eubacteria are monophyletic or paraphyletic. Both these points are something you may want to keep in mind as you read this post.

Proteasomes are enzymes that function to break down polypeptide strands within the cell. The average proteasome is roughly shaped like an open-ended barrel—the image at the top of this post shows an example looking down the barrel. Pieces of defective protein are taken in one end and chopped up into small fragments that are ejected at the other—doubtless to be recycled to make newer, better proteins. Proteasomes are a notable shared feature of eukaryotes and archaebacteria but were believed to be absent from eubacteria until they were identified from the actinobacterium Rhodococcus. Actinobacteria are a collection of Gram-positive bacteria with high G + C content that are generally accepted as forming a distinct clade. Since then, proteasomes have been identified from a range of other bacteria, including Mycobacterium and Streptomyces, and are probably found throughout the clade.

Simplified diagram of a 20S proteasome in sideview, showing the terminal α-subunits and the medial β-subunits (from Answers.com).

This is of some significance to the question of bacterial phylogeny because 18S rDNA trees place Actinobacteria as one of the closer groups of eubacteria to the other two domains. Whether proteasomes are a shared synapomorphy of Actinobacteria, archaea and eukaryotes, or a feature of the common ancestor of all living organisms that was subsequently lost in non-actinobacterial eubacteria, largely depends on where the root of the tree of living organisms turns out to be. Other eubacteria possess a different but closely related enzyme known as HslV, that authors such has Cavalier-Smith (2006) have suggested to be ancestral to proteasomes.

The possibility exists, I suppose, that proteasomes were transferred between Actinobacteria and Neomura by that bugbear of bacterial phylogenetics, horizontal or lateral gene transfer. In this case, I don’t think that’s likely. In general, the proteasome system replaces the HslV system or vice versa, which is highly suggestive of a single origin of one from the other. Only a single case appears to be known of co-existence of HslV and proteasomes in the one organism, with both being present in the Kinetoplastida, a clade of parasitic protozoans including the causative organisms of such spectacularly unpleasant diseases as sleeping sickness and leishmaniasis (Couvreur et al. 2002). Phylogenetic analysis of the kinetoplastid HslV sequence places it among the eubacterial HslV sequences rather than with the neomuran proteasomes, but is unable to resolve relationship to any particular eubacterial clade (ibid.) The available phylogenetic analyses indicate that kinetoplastids are relatively derived within the Neomura, and it seems more likely that the kinetoplastid HslV is the result of lateral gene transfer. It is worth noting that, confusingly, early diverging Actinobacteria such as Bifidobacterium seem to lack proteasomes (Cavalier-Smith 2006), but as they also seem to lack HslV they must be regarded as uninformative in this matter.

Structurally, the actinobacterial proteasome is comparable to the archaeal proteasome. In eukaryotes, seven each of different α-type and β-type subunuits go into forming the 20S proteasome, which forms the core, with the addition of several accessory proteins, of the final 26S proteasome. The archaeal and actinobacterial proteasomes, in contrast, are far more minimalist structures, with only a single α- and β-type subunit in archaebacteria, Mycobacterium and Streptomyces. Rhodococcus has a slightly intermediate state, with two different α- and β-type subunits (Nagy et al. 1998).

One particular finding of Nagy et al. deserves highlighting. Activity measures of the Streptomyces proteasome found that it was inhibited by lactacystin, a result found with proteasomes from other organisms. The interesting thing is that at least one Streptomyces strain produces lactacystin itself. Could the production of lactacystin function to regulate the activity of proteasomes in the living organism?

Systematics of Streptomyces

Characters (from Korn-Wendisch & Kutzner 1992): Aerial mycelium forming long chains of spores; verticils or pseudoverticils present; spore surface smooth, hairy, warty or spiny. LL-diaminopimelic acid present; glycine present in interpeptide bridge; peptidoglycan type A3γ; phospholipid type phosphatidylethanolamine only; mycolic acids absent; menaquinones 9(H6)/(H8); G+C content 69–78%.

Streptomyces Waksman & Henrici 1943 (see below for synonymy)K-WK92
|--+--S. thermolineatusNC18
| `--+--S. tyrosinilyticusNC18
| `--+--S. capparidisNC18
| `--S. vitaminophilus (Shomura et al. 1983) Goodfellow et al. 1986NC18
`--+--+--+--S. yeochonensis Kim et al. 2004NC18
| | `--+--S. guanduensisNC18
| | `--+--S. yanglinensis Xu et al. 2006NC18
| | `--+--S. bryophytorumNC18
| | `--S. cocklensisNC18
| `--+--S. rubidus Xu et al. 2006NC18
| `--+--S. paucisporeus Xu et al. 2006NC18
| `--+--+--S. abietisNC18
| | `--S. alni Liu et al. 2009NC18
| `--+--S. baliensisNC18
| `--+--S. erringtoniiNC18
| `--S. glaucinigerNC18
`--+--+--S. tateyamensisNC18
| `--S. xinghaiensisNC18
`--+--+--+--S. ferralitisNC18
| | `--S. rubrisoliNC18
| `--+--+--+--S. armeniacusNC18 [=Actinoplanes armeniacusK-WK92]
| | | `--S. artemisiaeNC18
| | `--+--S. pharmamarensisNC18
| | `--S. phytohabitansNC18
| `--+--+--S. qinglanensis Hu et al. 2012NC18
| | `--+--S. panacagriNC18
| | `--S. rubrusNC18
| `--+--+--S. flocculusNC18
| | `--+--*S. albus (Rossi Doria 1891) Waksman & Henrici 1943NC18
| | `--+--S. gibsoniiNC18
| | `--S. rangoonensisNC18
| `--+--+--+--S. aminophilusNC18
| | | `--S. cacaoiNC18
| | `--+--+--S. oryzaeNC18
| | | `--S. smyrnaeusNC18
| | `--+--+--S. halophytocolaNC18
| | | `--S. sulphureus (Gasperini 1894) Waksman 1953NC18
| | `--+--+--S. macrosporusNC18
| | | `--S. megasporus (ex Krassilnikov et al. 1968) Agre 1986NC18
| | `--+--+--S. mangroviNC18
| | | `--+--S. pini Madhaiyan et al. 2016NC18
| | | `--+--S. barkulensisNC18
| | | `--S. glaucosporusNC18
| | `--+--+--S. atacamensisNC18
| | | `--S. radiopugnansNC18
| | `--+--S. nanhaiensisNC18
| | `--+--S. chitinivoransNC18
| | |--S. fenghuangensisNC18
| | `--S. verrucosisporusNC18
| `--+--+--S. haliclonaeNC18
| | `--+--+--S. daqingensisNC18
| | | `--S. ovatisporusNC18
| | `--+--S. marinusNC18
| | `--+--+--S. glycovoransNC18
| | | `--S. karpasiensisNC18
| | `--+--S. xishensisNC18
| | `--+--S. abyssalisNC18
| | `--S. nanshensisNC18
| `--+--S. alkalithermotoleransNC18
| |--+--S. alkaliphilusNC18
| | `--S. calidiresistensNC18
| `--+--S. sodiidophilusNC18
| `--+--+--S. lopnurensisNC18
| | `--+--S. bohaiensisNC18
| | `--S. lonarensisNC18
| `--+--+--S. aidingensis Xia et al. 2013NC18
| | `--S. canalisNC18
| `--+--+--S. xiamenensisNC18
| | `--+--S. carpaticusNC18
| | `--+--S. cheonanensisNC18
| | `--S. harbinensisNC18
| `--+--+--S. hainanensisNC18
| | `--+--S. mayteniNC18
| | `--S. phaeoviridisNC18
| `--+--S. avicenniaeNC18
| `--+--+--S. sediNC18
| | `--S. zhaozhouensisNC18
| `--+--S. klenkiiNC18
| `--+--S. specialis Kämpfer et al. 2008NC18
| `--+--S. burgazadensisNC18
| `--S. hoynatensisNC18
`--+--+--+--+--S. palmaeNC18
| | | `--+--S. wellingtoniaeNC18
| | | `--+--+--S. cuspidosporusNC18
| | | | `--S. sparsogenesPG-V03
| | | `--+--S. aldersoniaeNC18
| | | `--+--S. milbemycinicusNC18
| | | `--S. sulfonofaciensNC18
| | `--+--+--S. aculeolatusNC18
| | | `--S. synnematoformansNC18
| | `--+--+--S. ascomycinicusNC18
| | | `--S. himastatinicusNC18
| | `--+--+--S. javensisNC18
| | | `--+--S. albiflavinigerNC18
| | | |--S. violaceusnigerPG-V03
| | | |--S. yogyakartensisNC18
| | | `--+--+--S. iranensisNC18
| | | | `--S. rapamycinicusNC18
| | | `--+--S. demainiiNC18
| | | `--+--S. hygroscopicusPG-V03 [incl. S. endusZL03]
| | | | |--S. h. ssp. hygroscopicusK-WK92
| | | | `--S. h. var. aureolacrimosusK-WK92
| | | `--S. sporocinereus (ex Krassilnikov 1970) Preobrazhenskaya 1986NC18
| | `--+--S. asiaticusNC18
| | |--S. cangkringensisNC18
| | |--S. griseinigerNC18
| | |--S. indosiensisNC18
| | |--S. rhizosphaericusNC18
| | `--+--+--S. malaysiensisPG-V03
| | | `--S. samsunensisNC18
| | `--+--S. melanosporofaciens Arcamone et al. 1959NC18
| | `--+--S. antimycoticusNC18
| | |--S. geldanamycininusNC18
| | |--S. sporoclivatusNC18
| | |--S. yatensisPG-V03
| | `--+--S. castelarensisNC18
| | `--S. mordarskiiNC18
| `--+--+--+--+--S. purpurogeneiscleroticusNC18
| | | | `--+--S. ochraceiscleroticus Pridham 1970NC18
| | | | `--S. violensNC18
| | | `--+--+--S. niger (Thirumalachar 1955) Goodfellow et al. 1986NC18
| | | | `--S. olivaceiscleroticusNC18
| | | `--+--S. sclerotialus Pridham 1970NC18
| | | `--+--S. ramulosusNC18
| | | `--+--+--S. albiaxialisNC18
| | | | `--S. chumphonensisNC18
| | | `--+--S. sparsusNC18
| | | `--+--S. daliensisNC18
| | | `--+--S. axinellaeNC18
| | | `--S. iconiensisNC18
| | `--+--+--S. monomycini Gause & Terekhova 1986NC18
| | | `--S. rimosus Sobin et al. 1953NC18
| | `--+--+--S. catenulae Davisson & Finlay 1961NC18
| | | `--S. misakiensisNC18
| | `--+--+--S. albospinusNC18
| | | `--+--S. celluloflavus Nishimura et al. 1953NC18
| | | `--+--S. kasugaensisPG-V03
| | | `--‘Streptoverticillium’ verticilliumNC18
| | | |--S. v. ssp. verticilliumNC18
| | | `--S. v. ssp. quintumNC18
| | `--+--+--S. staurosporininusNC18
| | | `--+--S. lunalinharesiiNC18
| | | `--+--S. yunnanensis Zhang, Li et al. 2003NC18, ZL03
| | | `--+--S. albulusNC18
| | | `--S. noursei Brown et al. 1953NC18
| | `--+--+--S. kronopolitisNC18
| | | `--S. lydicus De Boer et al. 1956NC18
| | |--+--S. albofaciensNC18
| | | |--S. chrestomyceticusNC18
| | | `--S. ‘rimosus ssp. paromomycinusNC18
| | `--+--S. caniferusNC18
| | |--S. ‘hygroscopicus ssp.’ rufusNC18
| | |--S. libaniNC18
| | | |--S. l. ssp. libaniNC18
| | | `--S. l. ssp. rufusNC18
| | `--S. platensis Tresner & Backus 1956NC18
| `--+--S. griseocarneusNC18
| |--+--S. ladakanumNC18
| | `--S. orinociNC18
| `--+--+--S. varsoviensis Kurylowicz & Woznick 1967NC18
| | `--+--S. griseoverticillatusNC18
| | `--S. sapporonensisNC18
| `--+--+--‘S. fervens ssp.’ melrosporusNC18
| | `--+--+--S. abikoensisNC18
| | | |--S. angustmyceticusNC18
| | | |--S. ehimensisNC18
| | | `--S. luteoverticillatusNC18
| | `--+--+--S. luteireticuliNC18
| | | `--+--S. youssoufiensisNC18
| | | `--S. zagrosensisNC18
| | `--+--S. mobaraensis (Nagatsu & Suzuki 1963) Witt & Stackebrandt 1991NC18
| | `--+--S. kishiwadensisNC18
| | `--S. mashuensisNC18
| `--+--S. kashmirensisNC18
| |--S. lilacinusNC18
| `--+--S. septatusNC18
| `--+--S. salmonisNC18
| |--+--S. roseoverticillatusNC18
| | `--+--S. luteosporeusNC18
| | |--‘Streptoverticillium’ cinnamoneumNC18
| | | |--S. c. ssp. cinnamoneumNC18
| | | |--S. c. ssp. lanosumNC18
| | | `--S. c. ssp. sparsumNC18
| | `--+--S. hiroshimensisNC18
| | |--+--+--S. aureoversilisNC18
| | | | `--S. rectiverticillatusNC18
| | | `--+--S. cinnamoneusNC18
| | | `--S. pseudoechinosporeusNC18
| | `--+--S. parvisporogenesNC18
| | `--+--S. ‘cinnamoneus ssp.’ albosporusNC18
| | |--S. ardusNC18
| | `--S. blastmyceticusNC18
| `--+--+--S. thioluteusNC18
| | `--+--S. morookaenseNC18
| | `--+--S. lacticiproducensNC18
| | |--S. lavenduligriseusNC18
| | `--‘Streptoverticillium’ reticulumNC18
| | |--S. r. ssp. reticulumNC18
| | `--S. r. ssp. protomycicumNC18
| `--+--+--S. olivoverticillatusNC18
| | `--‘Streptoverticillium’ viridoflavumNC18
| `--+--S. hachijoensisNC18
| `--+--S. distallicusNC18
| |--S. flavopersicusNC18
| |--S. kentuckensisNC18
| |--S. netropsisNC18
| |--Streptoverticillium Baldacci 1958NC18, K-WK92
| | |--S. mobaraenseK-WK92
| | |--S. netropsisK-WK92
| | |--S. rimofaciensBV16
| | |--S. salmonisPHK96
| | |--S. septatumK-WK92
| | `--S. syringiumNC18
| `--+--S. stramineusNC18
| `--+--S. eurocidicusNC18
| `--+--S. albireticuliNC18
| |--S. zhihengiiNC18
| `--+--S. baldaccii (Farina & Locci 1966) Witt & Stackebrandt 1991NC18
| |--S. biverticillatusNC18
| |--S. spitsbergensisNC18
| |--S. werraensisNC18
| `--+--S. fervensNC18
| `--+--S. ‘globisporus ssp. caucasicusNC18
| `--S. olivoreticuli (see below for synonymy)NC18
| |--S. o. ssp. olivoreticuliNC18
| `--‘Streptoverticillium’ o. ssp. cellulophilumNC18
`--+--+--+--S. lavendulae (Waksman & Curtis 1916) Waksman & Henrici 1948NC18
| | `--+--+--S. goshikiensisNC18
| | | |--S. sporoverrucosusNC18
| | | `--+--S. colombiensisNC18
| | | `--S. ‘lavendulae ssp.’ grasseriusNC18
| | `--+--S. nojiriensisNC18
| | |--S. spororaveusNC18
| | |--S. xanthophaeus Lindenbein 1952NC18
| | |--+--S. avidiniiNC18
| | | `--S. subrutilusNC18
| | |--+--S. cirratusNC18
| | | `--S. vinaceusNC18
| | `--+--S. cinnamonensisNC18
| | |--S. virginiae Grundy et al. 1952NC18
| | `--+--S. amritsarensis Sharma et al. 2014NC18
| | |--S. flavotriciniNC18
| | |--S. yangpuensisNC18
| | |--+--S. globosusNC18
| | | `--S. toxytriciniNC18
| | `--+--S. katraeNC18
| | |--S. polychromogenesNC18
| | `--S. racemochromogenesNC18
| `--+--+--+--S. coeruleoprunusNC18
| | | `--+--S. somaliensis (Brumpt 1906) Waksman & Henrici 1948NC18
| | | `--+--S. fradiaeNC18
| | | `--S. roseoflavusNC18
| | `--+--Kitasatoa kauaiensis Matsumae et al. 1968NC18
| | `--+--+--S. nashvillensisNC18
| | | `--+--S. gulbargensisNC18
| | | `--+--S. roseolusNC18
| | | |--+--S. filamentosusNC18
| | | | `--S. roseosporusNC18
| | | `--+--S. roseofulvusNC18
| | | `--+--S. laurentii Trejo et al. 1979NC18
| | | `--S. termitumNC18
| | `--+--+--S. vietnamensis Zhu et al. 2007NC18
| | | `--+--+--S. exfoliatus (Waksman & Curtis 1916) Waksman & Henrici 1948NC18
| | | | `--S. venezuelae Ehrlich et al. 1948NC18
| | | `--+--S. omiyaensisNC18
| | | `--S. wedmorensis (ex Milard & Burr 1926) Preobrazhenskaya 1986NC18
| | `--+--S. bikiniensis Johnstone & Waksman 1947NC18
| | `--+--S. hirsutusNC18
| | |--S. violaceorectusNC18
| | `--+--S. cinereoruberNC18
| | `--+--S. roseoviridisNC18
| | `--+--S. showdoensisNC18
| | `--S. viridobrunneusNC18
| `--+--+--+--S. spongiicolaNC18
| | | `--S. wuyuanensisNC18
| | `--+--S. clavuligerus Higgens & Kastner 1971NC18
| | `--+--S. tsukubensisNC18
| | `--+--S. amakusaensisNC18
| | `--S. inusitatusNC18
| `--+--+--S. cremeusNC18
| | `--S. spiroverticillatusNC18
| |--+--S. candidusNC18
| | `--+--‘Pilimelia columellifera ssp. pallidaNC18
| | `--+--S. beijiangensisNC18
| | `--S. fildesensisNC18
| |--+--S. graminofaciensNC18
| | |--S. kurssanoviiNC18
| | `--+--S. peuceticusNC18
| | | |--S. p. var. peuceticusK-WK92
| | | `--S. p. var. caesiusK-WK92
| | `--S. xantholiticusNC18
| `--+--+--+--+--S. pulveraceusNC18
| | | | `--+--S. gelaticusNC18
| | | | `--+--S. atratusNC18
| | | | |--S. sanglieriNC18
| | | | `--S. yaniiNC18
| | | `--+--+--S. drozdowicziiNC18
| | | | `--+--S.brevisporaNC18
| | | | `--S. laculatisporaNC18
| | | `--+--S. camponoticapitisNC18
| | | `--+--S. laceyiNC18
| | | `--+--S. niveusNC18
| | | `--S. spheroides Wallick et al. 1956NC18
| | `--+--S. sannanensisNC18
| | |--+--S. polyantibioticusNC18
| | | `--+--S. formicaeNC18
| | | `--+--S. lushanensisNC18
| | | `--S. scopuliridisNC18
| | `--+--+--S. mauvecolorNC18
| | | `--+--S. hundungensisNC18
| | | `--+--S. michiganensisNC18
| | | `--S. xanthochromogenesNC18
| | `--+--S. hypolithicusNC18
| | `--+--S. albidochromogenesNC18
| | |--S. chryseusNC18
| | |--S. enissocaesilisNC18
| | `--S. helvaticusNC18
| `--+--S. lunaelactisNC18
| `--+--S. albovinaceusNC18
| |--S. globisporusNC18
| |--S. griseinusNC18
| |--S. mediolaniNC18
| |--S. pluricolorescensNC18
| |--S. rubiginosohelvolusNC18
| |--S. tanashiensisNC18
| `--+--S. anulatusNC18
| |--S. griseoplanusNC18
| |--S. praecoxNC18
| |--S. pratensisNC18
| |--S. roseodiastaticusNC18
| |--+--+--S. finlayiNC18
| | | `--+--S. claviferNC18
| | | |--S. mutomyciniNC18
| | | `--+--S. atroolivaceus (Preobrazhenskaya et al. 1957) Pridham et al. 1958NC18
| | | `--S. olivoviridisNC18
| | `--+--S. halstedii (Waksman & Curtis 1916) Waksman & Henrici 1948 (see below for synonymy)NC18
| | `--+--S. flavogriseusNC18
| | |--S. flavovirensNC18
| | |--S. nigrifaciensNC18
| | `--S. nitrosporeusNC18
| `--+--S. cyaneofuscatus (Kudrina 1957) Pridham et al. 1958NC18
| `--+--+--S. citreofluorescens (Korenyako et al. 1960) Pridham 1970 (see below for synonymy)NC18
| | `--+--S. chrysomallusNC18
| | `--+--S. erumpensNC18
| | `--S. griseus (Krainsky 1914) Waksman & Henrici 1948NC18
| `--+--S. argenteolusNC18
| |--S. cinereorectusNC18
| `--+--S. alboviridisNC18
| |--S. ‘griseus ssp.’ alphaNC18
| |--S. californicus (Waksman & Curtis 1916) Waksman & Henrici 1948 (see below for synonymy)NC18
| |--S. ‘griseus ssp.’ cretosusNC18
| |--S. fulvissimusNC18
| |--S. fulvorobeusNC18
| |--S. lipmaniiNC18
| |--S. luridiscabiei Park, Kim et al. 2003NC18, PK03
| |--S. microflavusNC18
| `--S. willmoreiNC18
`--+--+--+--S. hebeiensisNC18
| | `--+--S. pathocidiniNC18
| | `--S. tremellaeNC18
| `--+--S. koyangensisNC18
| `--+--S. albidoflavus (Rossi Doria 1891) Waksman & Henrici 1948NC18
| `--S. champavatiiNC18
`--+--S. ‘albosporeus ssp.’ labilomyceticusNC18
`--+--+--S. indicus Luo et al. 2011NC18
| `--+--+--S. alfalfaeNC18
| | `--+--S. scopiformisNC18
| | `--+--S. silaceusNC18
| | `--+--S. alboflavus (Waksman & Curtis 1916) Waksman & Henrici 1948NC18
| | `--S. flavofunginiNC18
| `--+--+--S. aureocirculatus (Krassilnikov & Yuan 1965) Pridham 1970NC18
| | `--S. novaecaesareaeNC18
| `--+--S. vastusNC18
| `--+--S. cinereusNC18
| `--S. flaveusNC18
`--+--+--+--S. tendaePG-V03
| | `--+--S. rubrogriseusNC18
| | `--+--S. coelescensNC18
| | |--S. humiferusNC18
| | |--S. violaceolatusNC18
| | |--S. violaceoruberNC18
| | |--+--S. anthocyanicusNC18
| | | `--S. tricolorNC18
| | `--+--S. ‘diastaticus ssp.’ ardesiacusNC18
| | |--S. coelicoflavusNC18
| | `--+--S. flaveolusNC18
| | `--+--S. chilikensisNC18
| | `--S. fragilisNC18
| `--+--S. albogriseolusNC18
| |--S. chattanoogensisNC18
| |--S. lienomyciniNC18
| `--+--S. marokkonensisNC18
| |--S. thinghirensisNC18
| `--+--+--S. griseoaurantiacusNC18
| | `--S. jietaisiensisNC18
| `--+--+--S. variegatusNC18
| | `--+--S. spectabilisNC18
| | `--S. tacrolimicusNC18
| `--+--S. aureoverticillatusNC18
| `--+--S. intermediusNC18
| `--+--S. diastaticusNC18
| |--S. gougerotiiNC18
| `--S. rutgersensisNC18
`--+--+--+--S. kebangsaanensis Sarmin et al. 2013NC18
| | `--+--S. eurythermusNC18
| | `--+--S. nodosus Trejo 1961NC18
| | `--+--S. ‘achromogenes ssp.’ rubraridisNC18
| | `--+--S. glomeratusNC18
| | `--+--S. bangladeshensisNC18
| | `--S. cinerochromogenesNC18
| `--+--+--S. chartreusisNC18
| | `--+--S. similanensisNC18
| | `--+--S. seoulensis Chun et al. 1997NC18
| | `--+--S. griseoluteusNC18
| | `--S. recifensisNC18
| `--+--+--+--S. misionensis Cercos et al. 1962NC18
| | | `--S. phaeoluteichromatogenesNC18
| | `--+--S. costaricanusNC18
| | |--S. griseofuscus Sakamoto et al. 1962NC18
| | |--S. murinusNC18
| | `--S. phaeogriseichromatogenesNC18
| `--+--+--+--S. achromogenes Okami & Umezawa 1953NC18
| | | `--+--S. lucensisNC18
| | | `--S. niveoruberNC18
| | `--+--S. chibaensisNC18
| | `--S. corchorusii Ahmad & Bhuiyan 1958NC18
| `--+--+--S. bungoensis Eguchi et al. 1993NC18
| | `--S. capoamusNC18
| `--+--+--S. cellostaticus Hamada 1958NC18
| | `--+--S. yokosukanensis Nakamura 1961NC18
| | `--+--S. puniciscabiei Park, Kim et al. 2003NC18, PK03
| | |--+--S. durhamensis Gordon & Lapa 1966NC18
| | | `--S. filipinensisNC18
| | `--+--S. echinatusNC18
| | `--S. yaanensisNC18
| `--+--S. lannensisNC18
| `--+--S. chiangmaiensisNC18
| `--+--+--S. bulliiNC18
| | `--S. echinoruberNC18
| `--+--S. chromofuscusNC18
| |--+--S. canchipurensisNC18
| | |--S. cinereospinusNC18
| | `--S. coeruleofuscusNC18
| `--+--S. pluripotens Lee et al. 2014NC18
| `--+--S. mexicanus Petrosyan, García-Varela et al. 2003NC18, PG-V03
| `--+--S. thermocoprophilusPG-V03
| `--+--S. thermoviolaceusNC18
| | |--S. t. var. thermoviolaceusPG-V03
| | `--S. t. var. apingensPG-V03
| `--+--S. thermoalcalitoleransPG-V03
| `--+--S. thermogriseusPG-V03
| |--S. thermonitrificansNC18
| `--S. thermovulgarisPG-V03
`--+--+--S. ziwulingensisNC18
| `--+--+--+--S. malachitofuscusNC18
| | | `--+--S. leeuwenhoekiiNC18
| | | `--+--+--S. glaucusNC18
| | | | `--S. viridosporusNC18
| | | `--+--S. graminearusNC18
| | | |--S. griseomyciniNC18
| | | `--S. griseostramineusNC18
| | `--+--+--S. tritoleransNC18
| | | |--S. violaceorubidus Terekhova 1986NC18
| | | `--+--S. malachitospinusNC18
| | | |--+--S. hyderabadensisNC18
| | | | `--S. parvulusNC18
| | | `--+--S. olivaceus (Waksman 1923) Waksman & Henrici 1948NC18
| | | `--S. pactumNC18
| | `--+--+--S. luteus Luo et al. 2017NC18
| | | `--S. mutabilisNC18
| | `--+--S. viridochromogenesNC18
| | |--+--S. paradoxusNC18
| | | `--+--+--S. glaucescensNC18
| | | | `--S. pharetraeNC18
| | | `--+--+--S. albaduncusNC18
| | | | `--S. griseoloalbusNC18
| | | `--+--S. almquistiiNC18
| | | |--S. althioticusNC18
| | | `--S. matensisNC18
| | `--+--+--S. ambofaciens Pinnert-Sindico 1954NC18
| | | `--+--S. collinusNC18
| | | `--S. violaceochromogenesNC18
| | `--+--S. griseoflavusNC18
| | `--+--S. griseorubensNC18
| | `--+--S. erythrogriseusNC18
| | |--S. griseoincarnatusNC18
| | |--S. labedaeNC18
| | |--S. tunisiensisNC18
| | `--S. variabilisNC18
| `--+--+--S. geysiriensisNC18
| | |--S. ghanaensis Wallhäusser et al. 1966NC18
| | |--S. minutiscleroticusNC18
| | `--+--S. plicatusNC18
| | |--S. rocheiNC18
| | `--S. vinaceusdrappusNC18
| `--+--S. tuirusNC18
| `--+--S. djakartensisNC18
| `--+--+--+--S. calvusNC18
| | | |--S. virensNC18
| | | `--+--S. asterosporusNC18
| | | `--S. aureorectusNC18
| | `--+--S. aurantiogriseusNC18
| | `--+--S. andiiNC18
| | `--+--S. naganishiiNC18
| | `--+--S. nogalaterNC18
| | `--S. prasinosporusNC18
| `--+--+--S. flavoviridisNC18
| | `--S. pilosusNC18
| `--+--+--S. desertiNC18
| | |--S. fumigatiscleroticusNC18
| | `--+--S. poonensisNC18
| | `--+--S. roseiscleroticusNC18
| | `--S. ruberNC18
| `--+--+--S. arenaeNC18
| | `--S. violarusNC18
| |--+--S. griseoviridisNC18
| | `--S. zinciresistensNC18
| `--+--+--S. emeiensis Sun et al. 2007NC18
| | `--S. prasinopilosusNC18
| |--+--S. cyanoalbusNC18
| | `--+--S. bambergiensisNC18
| | `--S. prasinusNC18
| `--+--S. viridisNC18
| `--+--S. chlorusNC18
| `--+--S. pratensNC18
| `--+--S. herbaceusNC18
| `--S. incanusNC18
`--+--+--+--S. galbusNC18
| | `--S. longwoodensis Prosser & Palleroni 1981NC18
| `--+--+--S. griseoruber Yamaguchi & Saburi 1955NC18
| | `--+--S. acidiscabiesNC18
| | `--S. alanosinicusNC18
| `--+--S. phaeoluteigriseusNC18
| `--+--S. bobiliNC18
| `--S. galilaeusNC18
`--+--+--+--S. ‘cacaoi ssp.’ asoensisNC18
| | `--S. humidusNC18
| `--+--S. rishiriensisNC18
| `--+--+--S. phaeofaciensNC18
| | `--S. puniceusNC18
| `--+--+--S. ‘cinereoruber ssp.’ fructofermentansNC18
| | `--+--S. polymachusNC18
| | `--+--S. graminilatusNC18
| | `--+--S. lacrimifluminisNC18
| | `--+--S. reticuliscabieiNC18
| | `--S. turgidiscabiesNC18
| `--+--+--S. diastatochromogenesNC18
| | `--+--S. roseolilacinusNC18
| | `--+--S. indoligenesNC18
| | `--+--S. luridusNC18
| | `--+--S. gobitriciniNC18
| | `--S. lavendofoliaeNC18
| `--+--S. deccanensisNC18
| |--+--S. bottropensis Waksman 1961NC18
| | `--S. stelliscabieiNC18
| |--+--S. europaeiscabieiNC18
| | `--S. scabieiNC18 [=Oospora scabiesK-WK92]
| `--+--S. heilongjiangensisNC18
| `--+--S. neyagawaensisNC18
| `--+--S. decoyicusNC18
| |--S. ipomoeaeNC18
| |--S. ‘hygroscopicus ssp.’ ossamyceticusNC18
| `--S. torulosusNC18
`--+--+--+--S. graminifoliiNC18
| | `--+--S. hokutonensisNC18
| | `--S. prunicolor (Ryabova & Preobrazhenskaya 1957) Pridham et al. 1958NC18
| `--+--S. adustusNC18
| |--+--S. avermitilisNC18
| | `--+--S. atriruberNC18
| | `--+--S. kunmingensisNC18
| | `--+--S. endophyticusNC18
| | `--S. fractusNC18
| `--+--S. albonigerNC18
| |--S. spongiaeNC18
| `--+--+--S. olivochromogenes (Waksman 1923) Waksman & Henrici 1948NC18
| | `--+--S. kaempferiNC18
| | `--S. mirabilisNC18
| `--+--S. rhizosphaerihabitansNC18
| `--+--S. siamensisNC18
| `--+--S. albiflavescensNC18
| `--S. krungchingensisNC18
`--+--S. bicolorPG-V03
`--+--+--S. cinnabarinus (Ryabova & Preobrazhenskaya 1957) Pridham et al. 1958NC18
| `--+--S. phyllanthiNC18
| `--+--S. coacervatusNC18
| `--+--S. flavidovirensNC18
| `--S. longisporusNC18
`--+--+--S. antibioticus (Waksman & Woodruff 1941) Waksman & Henrici 1948NC18 (see below for synonymy)
| `--+--S. griseorubiginosus (Ryabova & Preobrazhenskaya 1957) Pridham et al. 1958 (see below for synonymy)NC18
| `--+--+--S. canus Heinemann et al. 1953NC18
| | `--S. ciscaucasicus Sveshnikova 1986NC18
| `--+--S. pseudovenezuelae (Kuchaeva et al. 1961) Pridham 1970NC18
| `--+--S. plumbiresistensNC18
| |--S. seymenliensisNC18
| `--+--+--S. phaeochromogenesNC18
| | `--+--S. ederensisNC18
| | `--S. umbrinusNC18
| `--+--S. tauricusNC18
| `--+--S. aurantiacusNC18
| `--S. glomeroaurantiacusNC18
`--+--+--S. curacoi Cataldi 1963NC18
| `--+--S. shaanxiensisNC18
| `--+--S. caeruleatusNC18
| `--S. osmaniensisNC18
`--+--+--+--S. cyaneusNC18
| | `--S. flavovariabilis (ex Korenkayo & Nikitina 1965) Sveshnikova 1986NC18
| `--+--S. viridodiastaticusNC18
| |--+--S. atrovirensNC18
| | `--S. heliomyciniNC18
| `--+--+--S. longispororuberNC18
| | `--S. speibonaeNC18
| `--+--S. iakyrusNC18
| `--+--+--S. coeruleorubidusNC18
| | `--+--S. bellusNC18
| | `--S. coerulescensNC18
| `--+--S. lomondensisNC18
| `--+--+--S. lusitanusNC18
| | `--S. thermocarboxydusNC18
| `--+--S. lavendulocolorNC18
| `--+--S. capillispiralisNC18
| |--S. cellulosaeNC18
| |--S. gancidicusNC18
| |--S. pseudogriseolusNC18
| |--S. rubiginosusNC18
| `--+--+--S. azureus Kelly et al. 1959NC18
| | `--S. caelestisNC18
| `--+--+--+--S. carpinensisNC18
| | | `--S. levisNC18
| | `--+--S. viridiviolaceusNC18
| | `--+--S. andamanensisNC18
| | `--S. fumanusNC18
| `--+--+--S. afghaniensisNC18
| | `--S. africanusNC18
| `--+--S. spiralisNC18
| `--+--S. brasiliensisNC18
| `--+--S. fimbriatusNC18
| `--+--S. thermodiastaticusNC18
| `--+--S. thermospinosisporusNC18
| `--+--S. bluensisNC18
| `--S. thermocarboxydovoransNC18
`--+--S. lincolnensis Mason et al. 1963NC18
`--+--+--S. rameusNC18
| `--S. regensisNC18
`--+--S. lanatusNC18
`--+--S. gilvifuscusNC18
|--+--+--S. actinomycinicusNC18
| | `--+--S. graminisoliNC18
| | |--S. hyaluromyciniNC18
| | `--+--S. jiujiangensisNC18
| | `--S. shenzhenensisNC18
| `--+--S. sasaeNC18
| `--+--S. panaciradicisNC18
| `--+--+--S. fuscichromogenesNC18
| | `--S. olivicoloratusNC18
| `--+--S. gramineusNC18
| `--S. rhizophilusNC18
`--+--S. niveiscabiei Park, Kim et al. 2003NC18, PK03
`--+--+--S. spinoverrucosusNC18
| `--+--S. muensisNC18
| `--+--S. fabaeNC18
| `--S. griseosporeusNC18
|--+--S. purpurascensNC18
| `--+--+--S. albosporeusNC18
| | |--S. janthinusNC18
| | |--S. roseoviolaceusNC18
| | `--S. violaceusNC18
| `--+--S. luteogriseusNC18
| `--+--S. hawaiiensisNC18
| `--+--S. indiaensisNC18
| `--S. massasporeusNC18
`--+--S. canescensNC18
|--S. coelicolorNC18
|--S. felleusNC18
|--S. griseochromogenesNC18
|--S. hydrogenansNC18
|--S. limosusNC18
|--S. resistomycificus Lindenbein 1952NC18
|--S. ‘griseus ssp.’ solvifaciensNC18
`--+--S. odoriferNC18
`--S. sampsoniiNC18

Streptomyces incertae sedis:
S. argillaceusPG-V03
S. auratusNC18
S. bambusaeNC18
S. caespitosusK-WK92
S. canariusNC18
S. cattleyBV16
S. cinnamomensisK-WK92
S. crystallinusNC18
S. cyslabdanicusNC18
S. erythrochromogenesK-WK92
S. espinosusPG-V03
S. fukangensisNC18
S. goraiensisPG-V03
S. heteromorphusPG-V03
S. indigocolorPG-V03
S. kathiraePG-V03
S. kitasoensisBV16
S. lateritiusK-WK92
S. lindensisBV16
S. litmocidiniK-WK92
S. litoralisNC18
S. maritimusPG-V03
S. mediterraneiBV16
S. melanogenesNC18
S. microstreptosporaPG-V03
S. narbonensisNC18
S. natalensisMN03
S. nigrescensK-WK92
S. noboritoensisNC18
S. olivaceoviridisNC18
S. orchidaceusBV16
S. padanusBV16
S. pallidusPG-V03
S. panayensisPG-V03
S. paraguayensisK-WK92
S. pelletieriKC01
S. peruviensisPG-V03
S. peucetiusBV16
S. polygonatiNC18
S. pristinaespiralisK-WK92
S. pulcherPHK96
S. purpureofuscusK-WK92
S. rectusK-WK92
S. ruaniiNC18
S. sanyensisNC18
S. sioyaensisNC18
S. steffisburgensisPG-V03
S. sudanensisBV16
S. tenjmariensisK-WK92
S. thermoflavusK-WK92
S. thermofuscusK-WK92
S. thermophilusK-WK92
S. thermosacchariPG-V03
S. thermotoleransPG-V03
S. tubercidicusNC18
S. vellosusPG-V03
S. verticillusMN03
S. yerevanensisNC18
S. zaomyceticusNC18
0--S. arcticusNC18
`--+--S. kanamyceticusNC18
`--+--S. aureusNC18
`--+--S. durmitorensisNC18
|--S. rectiviolaceusNC18
`--S. rosealbusNC18
0--S. acrimyciniNC18
|--S. baarnensisNC18
|--S. caviscabiesNC18
|--S. fimicariusNC18
|--S. flavofuscusNC18
|--S. setoniiNC18
`--+--S. badiusNC18
|--S. sindenensisNC18
|--+--S. parvusNC18
| `--S. sundarbansensisNC18
`--+--S. bacillarisNC18
`--+--S. albolongusNC18
|--S. araujoniaeNC18
|--S. cavourensisNC18
| |--S. c. ssp. cavourensisNC18
| `--S. c. ssp. washingtonensisNC18
`--S. griseobrunneusNC18

Streptomyces Waksman & Henrici 1943 [incl. Actinopycnidium, Actinosporangium, Chainia Thirumalachar 1955, Microellobospora]K-WK92

Streptomyces antibioticus (Waksman & Woodruff 1941) Waksman & Henrici 1948NC18 [=*Chainia antibioticaK-WK92]

Streptomyces californicus (Waksman & Curtis 1916) Waksman & Henrici 1948 [incl. S. floridae Bartz et al. 1951]NC18

Streptomyces citreofluorescens (Korenyako et al. 1960) Pridham 1970 [incl. S. fluorescens (Krassilnikov 1958) Pridham 1970]NC18

Streptomyces griseorubiginosus (Ryabova & Preobrazhenskaya 1957) Pridham et al. 1958 [incl. S. phaeopurpureus Shinobu 1957]NC18

Streptomyces halstedii (Waksman & Curtis 1916) Waksman & Henrici 1948 [incl. S. griseolus (Waksman 1923) Waksman & Henrici 1948]NC18

Streptomyces olivoreticuli [=Streptoverticillium olivoreticuli]NC18

*Type species of generic name indicated

References

[BV16] Barka, E. A., P. Vatsa, L. Sanchez, N. Gaveau-Vaillant, C. Jacquard, J. P. Meier-Kolthoff, H.-P. Klenk, C. Clément, Y. Ouhdouch & G. P. van Wezel. 2016. Taxonomy, physiology, and natural products of Actinobacteria. Microbiology and Molecular Biology Reviews 80 (1): 1–43.

Cavalier-Smith, T. 2006. Rooting the tree of life by transition analyses. Biology Direct 1: 19.

Couvreur, B., R. Wattiez, A. Bollen, P. Falmagne, D. Le Ray & J.-C. Dujardin. 2002. Eubacterial HslV and HslU subunits homologs in primordial eukaryotes. Molecular Biology and Evolution 19 (12): 2110–2117.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

[K-WK92] Korn-Wendisch, F., & H. J. Kutzner. 1992. The family Streptomycetaceae. In: Balows, A., H. G. Trüper, M. Dworkin, W. Harder & K.-H. Schleifer (eds) The Prokaryotes: A handbook on the biology of bacteria: Ecophysiology, isolation, identification, applications 2nd ed. vol. 1 pp. 921–995. Springer-Verlag: New York.

[MN03] Moffitt, M. C., & B. A. Neilan. 2003. Evolutionary affiliations within the superfamily of ketosynthases reflect complex pathway associations. Journal of Molecular Evolution 56: 446–457.

Nagy, I., T. Tamura, J. Vanderleyden, W. Baumeister & R. DeMot. 1998. The 20S proteasome of Streptomyces coelicolor. Journal of Bacteriology 180: 5448–5453.

[NC18] Nouioui, I., L. Carro, M. García-López, J. P. Meier-Kolthoff, T. Woyke, N. Kyrpides, C., R. Pukall, H.-P. Klenk, M. Goodfellow & M. Göker. 2018. Genome-based taxonomic classification of the phylum Actinobacteria. Frontiers in Microbiology 9: 2007.

[PK03] Park, D. H., J. S. Kim, S. W. Kwon, C. Wilson, Y. M. Yu, J. H. Hur & C. K. Lim. 2003. Streptomyces luridiscabiei sp. nov., Streptomyces puniciscabiei sp. nov. and Streptomyces niveiscabiei sp. nov., which cause potato common scab disease in Korea. International Journal of Systematic and Evolutionary Microbiology 53: 2049–2054.

[PG-V03] Petrosyan, P., M. García-Varela, A. Luz-Madrigal, C. Huitrón & M. E. Flores. 2003. Streptomyces mexicanus sp. nov., a xylanolytic micro-organism isolated from soil. International Journal of Systematic and Evolutionary Microbiology 53: 269–273.

[PHK96] Prescott, L. M., J. P. Harley & D. A. Klein. 1996. Microbiology 3rd ed. Wm. C. Brown Publishers: Dubuque (Iowa).

[ZL03] Zhang, Q., W.-J. Li, X.-L. Cui, M.-G. Li, L.-H. Xu & C.-L. Jiang. 2003. Streptomyces yunnanensis sp. nov., a mesophile from soils in Yunnan, China. International Journal of Systematic and Evolutionary Microbiology 53: 217–221.

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