View source for Trypanosomatida

{{short description|Group of single-celled parasitic organisms}}
{{automatic taxobox
| name = Trypanosomes
| fossil_range = [[Albian]] to [[Holocene|recent]] {{fossilrange|100|0}}
| image = Trypanosoma_cruzi_crithidia.jpeg
| image_caption = ''[[Trypanosoma cruzi]]''
| display_parents = 2
| parent_authority = Kent 1880
| taxon = Trypanosomatidae
| authority = Doflein 1901
| subdivision_ranks = Subfamily
| subdivision =
* [[Blechomonadinae]]
* [[Leishmaniinae]]
** Clade Crithidiatae
** Clade Leishmaniatae
* [[Paratrypanosomatinae]]
* [[Phytomonadinae]]
* [[Strigomonadinae]]
* [[Trypanosomatinae]]
}}

'''Trypanosomatida''' is a group of [[kinetoplastid]] [[unicellular organism]]s distinguished by having only a single [[flagellum]]. The name is derived from the [[Greek language|Greek]] ''trypano'' (borer) and ''soma'' (body) because of the corkscrew-like motion of some trypanosomatid species. All members are exclusively [[parasite|parasitic]], found primarily in [[insect]]s.<ref>{{cite journal |author=Podlipaev S |title=The more insect trypanosomatids under study-the more diverse Trypanosomatidae appears |journal=[[International Journal for Parasitology]] |volume=31 |issue=5–6 |pages=648–52 |date=May 2001 |doi=10.1016/S0020-7519(01)00139-4|pmid=11334958}}</ref> A few genera have life-cycles involving a secondary host, which may be a [[vertebrate]], [[invertebrate]] or [[plant]]. These include several species that cause major diseases in humans.<ref>{{cite journal |vauthors=Simpson AG, Stevens JR, Lukes J |title=The evolution and diversity of kinetoplastid flagellates |journal=[[Trends in Parasitology]] |volume=22 |issue=4 |pages=168–74 |date=April 2006 |pmid=16504583 |doi=10.1016/j.pt.2006.02.006 }}</ref> Some trypanosomatida are [[intracellular parasite]]s, with the important exception of ''[[Trypanosoma brucei]]''.

== Medical importance ==

The three major [[human]] diseases caused by trypanosomatids are; African [[trypanosomiasis]] ([[African trypanosomiasis|sleeping sickness]], caused by ''[[Trypanosoma brucei]]'' and transmitted by [[Tsetse fly|tsetse flies]]<ref>{{cite web |title=Trypanosomiasis, human African (sleeping sickness) |url=https://www.who.int/en/news-room/fact-sheets/detail/trypanosomiasis-human-african-(sleeping-sickness) |website=www.who.int |access-date=14 May 2020 |language=en |archive-date=20 April 2018 |archive-url=https://web.archive.org/web/20180420223033/http://www.who.int/mediacentre/factsheets/fs259/en/ |url-status=live }}</ref>), South American trypanosomiasis ([[Chagas disease]], caused by ''[[Trypanosoma cruzi|T. cruzi]]'' and transmitted by [[Triatominae|triatomine]] bugs), and [[leishmaniasis]] (a set of trypanosomal diseases caused by various species of ''[[Leishmania]]'' transmitted by [[Phlebotominae|sandflies]]<ref name="Measure vectors' attraction to auxiliary hosts">
:{{Cite journal|s2cid=233743387|pmid=33950136|doi=10.1590/0001-37652021XXXX|title=''Lutzomyia longipalpis'': an update on this sand fly vector|last1=Rêgo|first1=Felipe D.|last2=Soares|first2=Rodrigo Pedro|journal=Anais da Academia Brasileira de Ciências|volume=93|issue=3|id=e20200254|year=2021|pages=e20200254 |doi-access=free|hdl=11336/166702|hdl-access=free}}
:
:This review cites this research.
:
:{{Cite journal|last1=Abbasi|first1=Ibrahim|last2=Trancoso Lopo de Queiroz|first2=Artur|last3=Kirstein|first3=Oscar David|last4=Nasereddin|first4=Abdelmajeed|last5=Horwitz|first5=Ben Zion|last6=Hailu|first6=Asrat|last7=Salah|first7=Ikram|last8=Mota|first8=Tiago Feitosa|last9=Fraga|first9=Deborah Bittencourt Mothé|date=2018-11-13|title=Plant-feeding phlebotomine sand flies, vectors of leishmaniasis, prefer ''Cannabis sativa''|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=115|issue=46|s2cid=53112660|pages=11790–11795|doi=10.1073/pnas.1810435115|pmc=6243281|pmid=30373823|bibcode=2018PNAS..11511790A |doi-access=free}}
</ref>).

== Evolution ==

The family is known from fossils of the extinct genus ''[[Paleoleishmania]]'' preserved in Burmese [[amber]] dating to the [[Albian]] (100 [[Mya (unit)|mya]]) and [[Dominican amber]] from the [[Burdigalian]] (20–15 mya) of [[Hispaniola]].<ref name="Poinar2008">{{cite journal |last1=Poinar |first1=G. |year=2008 |title=''Lutzomyia adiketis'' sp. n. (Diptera: Phlebotomidae), a vector of ''Paleoleishmania neotropicum'' sp. n. (Kinetoplastida: Trypanosomatidae) in Dominican amber |volume=1 |doi=10.1186/1756-3305-1-22 |pmid=18627624 |pmc=2491605 |journal=Parasites & Vectors |page=22 |issue=1 |doi-access=free }}</ref> The genus ''[[Trypanosoma]]'' is also represented in Dominican amber in the extinct species ''[[Trypanosoma antiquus|T. antiquus]]''.<ref name="Poinar2005">{{cite journal |last= Poinar |first=G. |title = ''Triatoma dominicana'' sp. n. (Hemiptera: Reduviidae: Triatominae), and ''Trypanosoma antiquus'' sp. n. (Stercoraria: Trypanosomatidae), the First Fossil Evidence of a Triatomine-Trypanosomatid Vector Association |journal= [[Vector-Borne and Zoonotic Diseases]] |volume= 5|issue= 1|pages= 72–81|year= 2005 |doi=10.1089/vbz.2005.5.72 |pmid=15815152}}</ref>

== Taxonomy ==

Three genera are [[heteroxenous|dixenous]] (two hosts in the life cycle) – ''[[Leishmania]]'', ''[[Phytomonas]]'' and ''[[Trypanosoma]]'', while The remainder are [[Monoxenous development|monoxenous]] (one host in the life cycle).{{citation needed|reason=Unsourced statements|date=April 2025}}  ''[[Paratrypanosoma]]'' appears to be the first evolving branch in this order. Fifteen genera are recognised in the ''Trypanosomatidae'' and there are three subfamilies – ''[[Blechomonadinae]]'', ''[[Leishmaniinae]]'' and ''[[Strigomonadinae]]''.{{clarification needed|reason=Contradicts list which has six subfamilie with many more genera|date=April 2025}}  The genera in the subfamily ''[[Strigomonadinae]]'' are characterised by the presence of obligatory intracellular bacteria of the [[Kinetoplastibacterium]] genus.<ref name="pmid23345457"/>
* Family '''Trypanosomatidae''' <small>Calkins 1926</small> [Trypanomorphidae <small>Woodcock 1906</small>; Trypanosomataceae <small>Senn 1911</small>]
** Genus ''[[Agamomonas]]'' <small>Grassé 1952</small>
** Genus ''[[Batracoleishmania]]'' <small>Dasgupta 2011</small>
** Genus ''[[Blastocrithidia]]'' <small>Laird 1959</small>
** Genus ''[[Cercoplasma]]'' <small>Roubaud 1911</small>
** Genus ''[[Cystotrypanosoma]]'' <small>Roubaud 1911</small>
** Genus ''[[Jaenimonas]]'' <small>Votypka & Hamilton 2015</small>
** Genus ''[[Lamellasoma]]'' <small>Davis 1947</small>
** Genus ''[[Leptowallaceina]]'' <small>Podlipaev & Frolov 2000</small>
** Genus ''[[Lewisonella]]'' <small>Chalmers 1918 nomen dubium</small>
** Genus ''[[Malacozoomonas]]'' <small>Nicoli, Penaud & Timon-David 1972</small>
** Genus ''[[Nematodomonas]]'' <small>Nicoli, Penaud & Timon-David 1972</small>
** Genus †''[[Paleoleishmania]]'' <small>Poinar & Poinar, 2004</small>
** Genus †''[[Paleotrypanosoma]]'' <small>Poinar 2008</small>
** Genus ''[[Paramecioides]]'' <small>Grassé 1882</small>
** Genus ''[[Sauroleishmania]]'' <small>Ranque 1973</small>
** Genus ''[[Sergeia]]'' <small>Svobodová et al. 2007 non Stimpson 1860 non Nasonov 1923 non Sergio Manning & Lemaitre 1994</small>
** Genus ''[[Trypanomonas]]'' <small>Danilewsky 1885</small>
** Genus ''[[Trypanomorpha]]'' <small>Woodcock 1906</small>
** Genus ''[[Undulina]]'' <small>Lankester 187</small>
** Genus ''[[Wallaceina]]'' <small>Bulat, Mokrousov & Podlipaev 1999</small> [''[[Proteomonas]]'' <small>Podlipaev, Frolov & Kolesnikov 1990 non Hill & Wetherbee 1986</small>]
** Genus ''[[Wallacemonas]]'' <small>Kostygov & Yurchenko 2014</small>
** '''Subfamily [[Paratrypanosomatinae]]''' <small>Votýpka & Lukeš 2013</small>
*** Genus ''[[Paratrypanosoma]]'' <small>Votypka & Lukes 2013</small>
** '''Subfamily [[Trypanosomatinae]]'''
[[File:Trypanosoma (248 09) Trypanosoma equiperdum.jpg|thumb|right|''[[Trypanosoma equiperdum]]'']]
::* Genus ''[[Trypanosoma]]'' <small>Gruby 1843</small>
:* '''Subfamily [[Blechomonadinae]]''' <small>Votypka & Suková 2013</small>
::* Genus ''[[Blechomonas]]'' <small>Votypka & Suková 2013</small>
[[File:Leishmania donovani 01.png|thumb|right|''[[Leishmania donovani]]'']]
:* '''Subfamily [[Leishmaniinae]]''' <small>sensu Maslov & Lukeš 2012</small>
::* '''Clade [[Crithidiatae]]''' <small>Maslov & Lukeš 2012</small>
[[File:CRITHIDIA 2.jpg|thumb|right|''[[Crithidia luciliae]]'']]
:::* Genus ''[[Crithidia]]'' <small>Léger 1902</small>
:::* Genus ''[[Leptomonas]]'' <small>Kent 1880</small>
:::* Genus ''[[Lotmaria]]'' <small>Schwarz 2015</small>
::* '''Clade [[Leishmaniatae]]''' <small>Maslov & Lukeš 2012</small>
:::* Genus ''[[Borovskyia]]'' <small>Kostygov & Yurchenko 2017</small>
:::* Genus ''[[Endotrypanum]]'' <small>Mesnil & Brimont 1908</small>
:::* Genus ''[[Leishmania]]'' <small>Ross 1903</small>
:::* Genus ''[[Novymonas]]'' <small>Votýpka et al. 2015</small>
:::* Genus ''[[Paraleishmania]]'' <small>Cupolillo et al. 2000</small>
:::* Genus ''[[Zelonia]]'' <small>Shaw, Camargo et Teixeira 2016</small>
:* '''Subfamily [[Phytomonadinae]]''' <small>Kostygov & Yurchenko 2015</small>
::* Genus ''[[Herpetomonas]]'' <small>Kent 1880 non Donovan 1909</small>
::* Genus ''[[Lafontella]]'' <small>Kostygov & Yurchenko 2015</small>
[[File:Epifluorescence microscopy of Phytomonas serpens.png|thumb|right|''[[Phytomonas serpens]]'']]
::* Genus ''[[Phytomonas]]'' <small>Donovan 1909</small>
:* '''Subfamily [[Strigomonadinae]]''' <small>Votypka et al. 2014</small>
[[File:Angomonas deanei structure.TIF|thumb|right|''[[Angomonas deanei]]'']]
::* Genus ''[[Angomonas]]'' <small>Souza & Corte-Real 1991</small>
::* Genus ''[[Kentomonas]]'' <small>Votypka et al. 2014</small>
::* Genus ''[[Strigomonas]]'' <small>Lwoff & Lwoff 1931</small>

== Life cycle ==

Some trypanosomatids only occupy a single [[Host (biology)|host]], while many others are [[heteroxenous]]: they live in more than one host species over their life cycle. This heteroxenous life cycle typically includes the [[intestine]] of a bloodsucking [[insect]] and the blood and/or tissues of a [[vertebrate]]. Rarer hosts include other bloodsucking invertebrates, such as [[leech]]es,<ref>{{cite journal | url=https://www.sciencedirect.com/science/article/abs/pii/S0020751905000020 | doi=10.1016/j.ijpara.2004.12.005 | title=A new lineage of trypanosomes from Australian vertebrates and terrestrial bloodsucking leeches (Haemadipsidae) | journal=International Journal for Parasitology | date=April 2005 | volume=35 | issue=4 | pages=431–443 | last1=Hamilton | first1=P. B. | last2=Stevens | first2=J. R. | last3=Gidley | first3=J. | last4=Holz | first4=P. | last5=Gibson | first5=W. C. | pmid=15777919 | url-access=subscription }}</ref> and other organisms such as [[plant]]s. Different species go through a range of different morphologies at different stages of the life cycle, with most having at least two different morphologies. Typically the promastigote and epimastigote forms are found in insect hosts, trypomastigote forms in the mammalian [[bloodstream]] and amastigotes in [[intracellular]] environments. {{cn|date=March 2023}}

Among commonly studied examples, ''[[Trypanosoma brucei|T. brucei]]'', ''[[Trypanosoma congolense|T. congolense]]'', and ''[[Trypanosoma vivax|T. vivax]]'' are extracellular, while ''[[Trypanosoma cruzi|T. cruzi]]'' and ''[[Leishmania]]'' spp. are intracellular.<ref name = "Tissue-Tropism-in-Parasitic-Diseases" /> Trypanosomatids with intracellular stages express {{ Visible anchor |δ-amastin}} proteins on their surfaces.<ref name = "Tissue-Tropism-in-Parasitic-Diseases" /> de Paiva ''et al.'', 2015 illuminates δ-amastins' roles in intracellular success.<ref name = "Tissue-Tropism-in-Parasitic-Diseases" >
{{ Cite journal
 | doi=10.1098/rsob.190036
 | pmid=31088251
 | pmc=6544988
| title=Tissue tropism in parasitic diseases
 | date=2019
 | last1=Silva Pereira
 | first1=Sara
 | last2=Trindade
 | first2=Sandra
 | last3=De Niz
 | first3=Mariana
 | last4=Figueiredo
 | first4=Luisa M.
 | journal=Open Biology
 | volume=9
 | issue=5
 | article-number=190036
 }}</ref>

===Sexual reproduction===

Trypanosomatids that cause globally known diseases such [[leishmaniasis]] (''[[Leishmania]]'' species), [[African trypanosomiasis]] referred to as sleeping sickness (''[[Trypanosoma brucei]]''), and [[Chagas disease]] (''[[Trypanosoma cruzi]]'') were found to be capable of [[meiosis]] and [[genetic recombination|genetic exchange]].<ref name = Silva2022>{{cite journal | doi=10.1590/1678-4685-GMB-2022-0065 | title=Sex in protists: A new perspective on the reproduction mechanisms of trypanosomatids | date=2022 | last1=Silva | first1=Verônica Santana da | last2=Machado | first2=Carlos Renato | journal=Genetics and Molecular Biology | volume=45 | issue=3 | pages=e20220065 | pmid=36218381 | pmc=9552303 }}</ref> These findings indicate the capability for [[sexual reproduction]] in the Trypanosomatida.<ref name = Silva2022/>

== Morphologies ==
[[File:TrypanosomatidMorphologies PlainSVG.svg|thumb|right|Six main [[morphology (biology)|morphologies]]]]
A variety of different morphological forms appear in the life cycles of trypanosomatids, distinguished mainly by the position, length and the cell body attachment of the [[flagellum]]. The [[kinetoplast]] is found closely associated with the [[basal body]] at the base of the flagellum and all species of trypanosomatid have a single nucleus. Most of these morphologies can be found as a life cycle stage in all trypanosomatid genera however certain morphologies are particularly common in a particular genus. The various morphologies were originally named from the genus where the morphology was commonly found, although this terminology is now rarely used because of potential confusion between morphologies and genus. Modern terminology is derived from the Greek; "mastig", meaning whip (referring to the flagellum), and a prefix which indicates the location of the flagellum on the cell. For example, the amastigote (prefix "a-", meaning no flagellum) form is also known as the leishmanial form as all ''[[Leishmania]]'' have an amastigote life cycle stage.{{cn|date=March 2023}}
* '''[[Amastigote]]''' (leishmanial).<ref name="Hoare1966">{{cite journal | title = Developmental Stages of Trypanosomatid Flagellates: a New Terminology | journal = [[Nature (journal)|Nature]] | year = 1966 | last1 = Hoare | volume = 212 | pages = 1385–6 | doi=10.1038/2121385a0 | issue=5068 | first1 = Cecil A. | last2 = Wallace | first2 = Franklin G.| bibcode = 1966Natur.212.1385H | s2cid = 4164112}}</ref> Amastigotes are a common morphology during an intracellular lifecycle stage in a mammalian host. All ''[[Leishmania]]'' have an amastigote stage of the lifecycle. ''Leishmania'' amastigotes are particularly small and are among the smallest eukaryotic cells. The flagellum is very short, projecting only slightly beyond the flagellar pocket.
* '''{{ Visible anchor |Promastigote}}''' (leptomonad).<ref name="Hoare1966" /> The promastigote form is a common morphology in the insect host. The flagellum is found anterior of nucleus emerging directly from the anterior cell body. The kinetoplast is located in front of the nucleus, near the anterior end of the body.
* '''{{ Visible anchor |Epimastigote}}''' (crithidial).<ref name="Hoare1966" /> Epimastigotes are a common form in the insect host and ''[[Crithidia]]'' and ''[[Blastocrithidia]]'', both parasites of insects, exhibit this form during their life cycles. The flagellum exits the cell anterior of nucleus and is connected to the cell body for part of its length by an undulating membrane. The kinetoplast is located between the nucleus and the anterior end.
* '''{{ Visible anchor |Trypomastigote}}''' (trypanosomal).<ref name="Hoare1966" /> This stage is characteristic of the genus ''[[Trypanosoma]]'' in the mammalian host bloodstream as well as infective metacyclic stages in the fly vector. In trypomastigotes the [[kinetoplast]] is near the posterior end of the body, and the flagellum lies attached to the cell body for most of its length by an undulating membrane.
* '''{{ Visible anchor |Opisthomastigote}}''' (herpetomonad).<ref name="Hoare1966" /> A rarer morphology where the flagellum posterior of nucleus, passing through a long groove in the cell.
* '''{{ Visible anchor |Endomastigote}}'''.<ref>{{Cite journal |last1=Merzlyak |first1=Ekaterina |last2=Yurchenko |first2=Vyacheslav |last3=Kolesnikov |first3=Alexander A. |last4=Alexandrov |first4=Kirill |last5=Podlipaev |first5=Sergei A. |last6=Maslov |first6=Dmitri A. |date=2001-03-01 |title=Diversity and Phylogeny of Insect Trypanosomatids Based on Small Subunit rRNA Genes: Polyphyly of ''Leptomonas'' and ''Blastocrithidia'' |journal=The Journal of Eukaryotic Microbiology |language=en |volume=48 |issue=2 |pages=161–169 |doi=10.1111/j.1550-7408.2001.tb00298.x |pmid=12095103 |s2cid=13880469 }}</ref> A morphotype where the flagellum does not extend beyond the deep flagellar pocket.

<gallery mode="packed" widths="360px" heights="220">
Image:LeishmaniaMexicana Amastigote SEM.jpg|'''Amastigote''': False colour [[Scanning electron microscope|SEM]] micrograph of [[amastigote]] form ''[[Leishmania mexicana]]''. The cell body is shown in orange and the flagellum is in red. 219 pixels/μm.
Image:LeishmaniaMexicana Promastigote SEM.jpg|'''Promastigote''': False colour [[Scanning electron microscope|SEM]] micrograph of [[promastigote]] form ''[[Leishmania mexicana]]''. The cell body is shown in orange and the flagellum is in red. 119 pixels/μm.
Image:TrypanosomaBrucei_ProcyclicTrypomastigote_SEM.jpg|'''Trypomastigote''': False colour [[Scanning electron microscope|SEM]] micrograph of procyclic form ''[[Trypanosoma brucei]]''. The cell body is shown in orange and the flagellum is in red. 84 pixels/μm.
</gallery>

== Other features ==
Notable characteristics of trypanosomatids are the ability to perform [[trans-splicing]] of RNA and possession of [[glycosome]]s, where much of their [[glycolysis]] is confined to. The [[acidocalcisome]], another [[organelle]], was first identified in trypanosomes.<ref>{{cite journal |vauthors=Docampo R, de Souza W, Miranda K, Rohloff P, Moreno SN |title=Acidocalcisomes — conserved from bacteria to man |journal=[[Nature Reviews Microbiology]] |volume=3 |issue=3 |pages=251–61 |date=March 2005 |pmid=15738951 |doi=10.1038/nrmicro1097 |s2cid=31935658 }}</ref>

=== {{Anchor|Kinetoplastibacterium}}Bacterial endosymbiont ===
{{Automatic taxobox|name=''Kinetoplastibacterium''|taxon=Kinetoplastibacterium|authority=Du et al., 1994}}
Six species of trypanosomatids are known to carry an additional proteobacterial [[endosymbiont]], termed TPE (trypanosomatid proteobacterial endosymbionts). These trypanosomatids (''[[Strigomonas oncopelti]]'', ''[[Strigomonas culicis|S. culicis]]'', ''[[Strigomonas galati|S. galati]]'', ''[[Angomonas desouzai]]'', and ''[[Angomonas deanei|A. deanei]]'') are in turn known as SHTs, for symbiont-harboring trypanosomatids. All such symbionts have a shared evolutionary origin and are classified in the [[Candidatus]] genus "'''Kinetoplastibacterium'''".<ref name="pmid23345457">{{cite journal | last1=Alves |first1=JM |last2=Serrano |first2=MG |last3=Maia da Silva |first3=F |last4=Voegtly |first4=LJ |last5=Matveyev |first5=AV |last6=Teixeira |first6=MM |last7=Camargo |first7=EP |last8=Buck |first8=GA | journal = Genome Biology and Evolution |date=2013 |volume=5 |issue=2 |pages=338–50 |doi=10.1093/gbe/evt012 |doi-access=free |pmid=23345457|pmc=3590767 |title=Genome evolution and phylogenomic analysis of ''Candidatus'' Kinetoplastibacterium, the betaproteobacterial endosymbionts of ''Strigomonas'' and ''Angomonas'' }}</ref>

As with many symbionts, the bacteria have a much reduced genome compared to their free-living relatives of genera ''[[Taylorella]]'' and ''[[Achromobacter]]''. ([[GTDB]] finds the genus sister to ''[[Proftella]]'', a symbiont of ''[[Diaphorina citri]]''.)<ref>{{cite web |title=GTDB - Tree at g__Kinetoplastibacterium |url=https://gtdb.ecogenomic.org/tree?r=g__Kinetoplastibacterium |website=gtdb.ecogenomic.org |access-date=2022-12-20 |archive-date=2022-12-20 |archive-url=https://web.archive.org/web/20221220182712/https://gtdb.ecogenomic.org/tree?r=g__Kinetoplastibacterium |url-status=live }}</ref> Reflecting their inability to live alone, they have lost genes dedicated to essential biological functions, relying on the host instead. They have modified their division to become synchronized with the host. In ''S. culicis'' at least, the TPE helps the host by synthesizing [[heme]]<ref name="pmid23345457"/> and producing essential enzymes, staying tethered to the [[kinetoplast]].<ref name=":4">{{Cite journal|last1=de Souza|first1=W.|last2=Motta|first2=M. C.|date=1999|title=Endosymbiosis in protozoa of the Trypanosomatidae family|journal=FEMS Microbiology Letters|volume=173|issue=1|pages=1–8|doi=10.1111/j.1574-6968.1999.tb13477.x|pmid=10220875|doi-access=free}}</ref>
{{clear}}

== References ==
{{Reflist}}

{{refbegin}}
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* {{cite journal |author=Matthews KR |title=The developmental cell biology of ''Trypanosoma brucei'' |journal=[[Journal of Cell Science]] |volume=118 |issue=Pt 2 |pages=283–90 |date=January 2005 |pmid=15654017 |pmc=2686837 |doi=10.1242/jcs.01649 }}
* {{cite journal |author=Matthews KR, Gull K |title=Evidence for an interplay between cell cycle progression and the initiation of differentiation between life cycle forms of African trypanosomes |journal=[[Journal of Cell Biology]] |volume=125 |issue=5 |pages=1147–56 |date=June 1994 |pmid=8195296 | doi = 10.1083/jcb.125.5.1147 |pmc=2120053 |last2=Gull }}
* {{cite journal |vauthors=Morrison LJ, Marcello L, McCulloch R |title=Antigenic variation in the African trypanosome: molecular mechanisms and phenotypic complexity |journal=[[Cellular Microbiology]] |volume=11 |issue=12 |pages=1724–34 |date=December 2009 |pmid=19751359 |doi=10.1111/j.1462-5822.2009.01383.x |s2cid=26552797 |url=http://eprints.gla.ac.uk/8339/1/8339.pdf |access-date=2019-07-10 |archive-date=2021-10-20 |archive-url=https://web.archive.org/web/20211020115950/http://eprints.gla.ac.uk/8339/1/8339.pdf |url-status=live }}
* {{cite journal |author=Seed JR, Wenck MA |title=Role of the long slender to short stumpy transition in the life cycle of the African trypanosomes |journal=[[Kinetoplastid Biology and Disease]] |volume=2 |issue=1 |page=3 |date=June 2003 |pmid=12844365 |pmc=165594 |doi=10.1186/1475-9292-2-3 |last2=Wenck |doi-access=free }}
* {{cite journal |author=Shadan S |title=Microbiology: Signals for change |journal=Nature |volume=459 |issue=7244 |page=175 |date=May 2009 |pmid=19444199 |doi=10.1038/459175a |bibcode=2009Natur.459..175S |doi-access=free }}
* {{cite journal |author=Sherwin T, Gull K |title=The cell division cycle of ''Trypanosoma brucei brucei'': timing of event markers and cytoskeletal modulations |journal=[[Philosophical Transactions of the Royal Society B]] |volume=323 |issue=1218 |pages=573–88 |date=June 1989 |pmid=2568647 |doi=10.1098/rstb.1989.0037|last2=Gull |bibcode=1989RSPTB.323..573S |doi-access= }}
* {{cite web |title=African trypanosomiasis |date=August 2006 |publisher=[[World Health Organization]] |url=https://www.who.int/mediacentre/factsheets/fs259/en/ |access-date=2020-10-05 |archive-date=2016-12-04 |archive-url=https://web.archive.org/web/20161204153318/http://www.who.int/mediacentre/factsheets/fs259/en/ |url-status=live }}
* {{Cite EB1911|wstitle=Trypanosomes|volume=27|pages=340–347|first=Harold Mellor|last=Woodcock}} ([https://archive.org/details/encyclopaediabri27chisrich/page/340/mode/2up online]). A comprehensive survey of the organisms' natural history.
{{refend}}

== External links ==
{{Commons category|Trypanosomatida}}
* [https://web.archive.org/web/20090414211811/http://knoesis.wright.edu/internal/wiki/index.php/Trykipedia Trykipedia], Trypanosomatid specific ontologies
* [http://tolweb.org/Trypanosomatida/98015 Tree of Life: Trypanosomatida]
* [http://www.boldsystems.org/index.php/Taxbrowser_Taxonpage?taxid=494532 Taxonomy at BOLD Systems]
* [http://taxonomicon.taxonomy.nl/TaxonName.aspx?id=199492&tree=0.1&syn=1 Taxonomy at Taxonomicon]
* [https://tree.opentreeoflife.org/taxonomy/browse?id=5257404 Open Tree Taxonomy]
* [http://zipcodezoo.com/index.php/Trypanosomatida ZipcodeZoo]

{{Discoba}}
{{Taxonbar|from=Q132954}}

[[Category:Trypanosomatida| ]]
[[Category:Parasitic excavates]]
[[Category:Extant Albian first appearances]]
[[Category:Euglenozoa orders]]