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{{short description|Family of flowering plants in the Eudicot order Saxifragales}}
{{Short description|Family of flowering plants in the Eudicot order Saxifragales}}
{{automatic taxobox
{{Automatic taxobox
|image = Cleaned-Illustration Myriophyllum spicatum.jpg
|image = Cleaned-Illustration Myriophyllum spicatum.jpg
|image_caption = ''Myriophyllum spicatum''
|image_caption = ''Myriophyllum spicatum''
Line 7: Line 7:
|subdivision_ranks = Genera
|subdivision_ranks = Genera
|subdivision = See text
|subdivision = See text
}}
{{Infobox
|bodystyle = width:18em
|subheader=[[Floral formula]]
|data1={{resize|100%|✶<math>\mathrm{K}^{3-4}</math><math>\mathrm{C}^{3-4}</math> or <math>\mathrm{C}^0</math><math>\mathrm{A}^{2-8}</math><math>\mathrm{G}^{\overline{(2-4)}}</math>}}
}}
}}


'''Haloragaceae''' (the watermilfoil [[family (biology)|family]]) is a [[dicotyledon]] [[flowering plant]] family in the [[order (biology)|order]] [[Saxifragales]], based on the [[phylogenetic]] [[APG system]]. In the [[Cronquist system]], it was included in the order [[Haloragales]].
'''Haloragaceae''' (the watermilfoil [[family (biology)|family]]) is a [[eudicot]] [[flowering plant]] family in the [[order (biology)|order]] [[Saxifragales]], based on the [[phylogenetic]] [[APG system]]. In the [[Cronquist system]], it was included in the order [[Haloragales]].


== Description ==
== Description ==


The Haloragaceae are very diverse in habit, including both small trees and submerged aquatics. Most members of the Haloragaceae are herbaceous, and most of those in turn are [[Perennial plant|perennial]]s, though some species are [[annual plant|annual]]s. In contrast however, members of the genus ''Haloragodendron'' are woody. Most species of ''Myriophyllum'' are [[monoecious]] while most other taxa have hermaphrodite flowers. The flowers are usually small and inconspicuous, but some genera can have more "showy" conspicuous flowers (''Haloragodendron'', ''Glischrocaryon''). Flowers are usually [[radially symmetrical]], and unusual for [[core Eudicots]], [[merosity]] is (2-3)-4 parted. Petals are usually keeled or hooded when present. In ''Myriophyllum'' female flowers usually lack a perianth. They have (2-)4-8 stamens and an inferior ovary of (2-)4 carpels. In ''Myriophyllum'' the fruit is a schizocarp of 1-seeded 'nutlets' other genera can have nuts or drupes that can be winged or inflated.{{sfn|Orchard|1975}}{{sfn|Moody|Les|2007}}
The Haloragaceae ( alternate spelling Halorrhagidaceae) are very diverse in habit, including both small trees and submerged aquatics. Most members of the Haloragaceae are herbaceous, and most of those in turn are [[Perennial plant|perennial]]s, though some species are [[annual plant|annual]]s. In contrast however, members of the genus ''Haloragodendron'' are woody. Most species of ''Myriophyllum'' are [[monoecious]] while most other taxa have hermaphrodite flowers. The flowers are usually small and inconspicuous, but some genera can have more "showy" conspicuous flowers (''Haloragodendron'', ''Glischrocaryon''). Flowers are usually [[radially symmetrical]], and unusual for [[core eudicot]]s, [[merosity]] is (2-3)-4 parted. Petals are usually keeled or hooded when present. In ''Myriophyllum'' female flowers usually lack a perianth. They have (2-)4-8 stamens and an inferior ovary of (2-)4 carpels. In ''Myriophyllum'' the fruit is a schizocarp of 1-seeded 'nutlets' other genera can have nuts or drupes that can be winged or inflated.{{sfn|Orchard|1975}}{{sfn|Moody|Les|2007}} The genus [[Gunnera]], formerly included here are now the separate family [[Gunneraceae]].

[[Floral formula]]:
<math>\star \mathrm{K}^{3-4} \; \mathrm{C}^{3-4}</math> or <math>\mathrm{C}^0 \; \mathrm{A}^{2-8} \; \mathrm{G}^{\overline{(2-4)} }</math>


== Taxonomy ==
== Taxonomy ==


=== History ===
=== Phylogeny ===


[[Molecular phylogenetic]] studies, in particular, the [[APG system]], placed the Haloragaceae within the [[core eudicot]] [[Order (biology)|order]], [[Saxifragales]].{{sfn|APG IV|2016}} Earlier versions of the APG had allowed either the broader circumscription (Haloragaceae ''[[Sensu|s.l.]]'') or a narrower Haloragaceae ''s.s.''.{{sfn|Chen et al|2014}}
Historically, the Haloragaceae included many disparate genera, since [[Segregate (taxonomy)|segregated]]. A major [[Circumscription (taxonomy)|circumscription]] was carried out by [[Anton Karl Schindler |Schindler]] in 1905, dividing the "Halorrhagaceae" into two subfamilies (Halorrhagoideae and Gunneroideae) and the former into two tribes (Halorrhageae and Myriophylleae), with a total of seven genera. He removed some of the disparate genera and merged ''Gonocarpus'' and ''Meionectes'' into ''Haloragis''.{{sfn|Schindler|1905}} This classification long remained the standard till Shaw (1966) removed ''[[Gunnera]]'' (into its own family Gunneraceae), the sole genus in Gunneroideae, leaving six genera.{{sfn|Willis|Shaw|1966}} This situation remained untill the monograph of Orchard (1975). Orchard restored ''Gonocarpus'' and split ''Haloragodendron'' from ''Haloragis'', leaving 8 genera.{{sfn|Orchard|1975}}{{sfn|Moody |Les|2007}}
{{cladogram
|title= Cladogram of Saxifragales families{{sfn|Jian et al|2008}}{{sfn|Stevens|2019}}{{sfn|APG IV|2016}}
|caption=[[Cynomorium]] (Cynomoriaceae) remains unplaced within this tree
|align=left
|cladogram=
{{clade | style=font-size:100%;line-height:120%;width:700px;
|label1='''Saxifragales'''
|1={{clade
|1=[[Peridiscaceae]] (4)
|label2=&nbsp;97&nbsp;
|2={{clade
|label1=
|1={{clade
|1=''[[Peony|Paeonia]]'' (Paeoniaceae)
|label2=&nbsp;woody&nbsp;clade&nbsp;
|2={{clade
|1=''[[Liquidambar]]'' (Altingiaceae)
|label2=&nbsp;69&nbsp;
|2={{clade
|label1=&nbsp;98&nbsp;
|1=[[Hamamelidaceae]] (27)
|label2=&nbsp;95&nbsp;
|2={{clade
|1=''[[Cercidiphyllum]]'' (Cercidiphyllaceae)
|2=''[[Daphniphyllum]]'' (Daphniphyllaceae)
}}
}}
}}
}}
|label2=&nbsp;core&nbsp;Saxifragales&nbsp;
|2={{clade
|1={{clade
|1=[[Crassulaceae]] (34)
|label2=&nbsp;Haloragaceae&nbsp;''s.l.''
|2={{clade
|1=''[[Aphanopetalum]]'' (Aphanopetalaceae)
|2={{clade
|1=''[[Tetracarpaea]]'' (Tetracarpaeaceae)
|2={{clade
|1=''[[Penthorum]]'' (Penthoraceae)
|2='''Haloragaceae''' ''s.s.'' (8)
}}
}}
}}
}}
|label2=&nbsp;Saxifragaceae&nbsp;alliance&nbsp;
|2={{clade
|1={{clade
|1=[[Iteaceae]] (including [[Pterostemonaceae]]) (2)
}}
|2={{clade
|1=''[[Ribes]]'' (Grossulariaceae)
|2=[[Saxifragaceae]] (33)
}}
}}
}}
}}
}}
}}


}}
=== Molecular era ===

{{clear}}

=== Subdivision ===

==== History ====

Historically, the Haloragaceae included many disparate genera, since [[Segregate (taxonomy)|segregated]]. A major [[Circumscription (taxonomy)|circumscription]] was carried out by [[Anton Karl Schindler|Schindler]] in 1905, dividing the "Halorrhagaceae" into two subfamilies (Halorrhagoideae and Gunneroideae) and the former into two tribes (Halorrhageae and Myriophylleae), with a total of seven genera. He removed some of the disparate genera and merged ''Gonocarpus'' and ''Meionectes'' into ''Haloragis''.{{sfn|Schindler|1905}} This classification long remained the standard till Shaw (1966) removed ''[[Gunnera]]'' (into its own family Gunneraceae, within the order Gunnerales), the sole genus in Gunneroideae, leaving six genera.{{sfn|Willis|Shaw|1966}} This situation remained until the monograph of Orchard (1975). Orchard restored ''Gonocarpus'' and split ''Haloragodendron'' from ''Haloragis'', leaving 8 genera.{{sfn|Orchard|1975}}{{sfn|Moody |Les|2007}}<br />
'''List of genera, habitat, ''distribution'' (Number of species)'''
* Terrestrial
** ''[[Glischrocaryon]]'' <small>[[Endl.]]</small> ''Australia'' (4)
** ''[[Gonocarpus]]'' <small>[[Thunb.]]</small> ''Australia, New Zealand, S. E. Asia'' (36)
** ''[[Haloragis]]'' <small>[[J.R.Forst.]] & [[G.Forst.]]</small> ''Australia, New Zealand, S. Pacific'' (26)
** ''Haloragodendron'' <small>Orchard</small> ''Australia'' (5)
* Semiaquatic
** ''Laurembergia'' <small>[[P.J.Bergius]]</small> ''Pantropical'' (4)
* Aquatic
** ''[[Meziella]]'' <small>[[Schindl.]]</small> ''S. W. Australia'' (1)
** ''[[Myriophyllum]]'' <small>[[Carl Linnaeus|L.]]</small> ''Cosmopolitan'' (60)
** ''[[Proserpinaca]]'' <small>L.</small> ''New World'' (3)

==== Molecular era ====

A molecular study resolved the infrafamilial relationships among the genera, resulting in some taxonomic revision, including redistribution of species. In addition, ''Meionectes'' was reinstated, separating two species from ''Haloragis'' and creating a new monotypic genus, ''Trihaloragis'' by segregating ''Gonocarpus hexandrus''. Consequently the number of genera is increased to ten, with the addition of:{{sfn|Moody |Les|2007}}
* ''Meionectes'' <small>[[R.Br.]]</small> (2)
* ''Trihaloragis'' <small>Moody & Les</small> (1)

''Glischrocaryon''-''Haloragodendron'' is resolved as the [[basal node]], [[sister group|sister]] to the remaining family. While [[monophyly]] of this group is well supported, monophyly of the two separate genera is less well supported, and suggests some [[paraphyly]]. Thus the generic limits remain unresolved.{{sfn|Moody |Les|2007}}

A subsequent, more detailed study of ''Myriophyllum'' demonstrated that the [[monotypic]] genus ''Meziella'' was embedded within it, leading to its submersion within the former as ''Myriophyllum'' subgenus ''Meziella'', thereby reducing the number of genera within the family to 9.{{sfn|Moody|Les|2010}}


==== Species ====
The modern Haloragaceae includes about 7–11 genera, depending on [[Circumscription (taxonomy)|circumscription]], with about 145 known species:{{sfn|Christenhusz|Byng|2016}}{{sfn|Christenhusz et al|2017}}{{sfn|POWO|2019}}{{sfn|Byng|2014}}{{sfn|Moody |Les|2007}}
*''[[Glischrocaryon]]''
*''[[Gonocarpus]]''
*''[[Haloragis]]''
*''[[Haloragodendron]]'', with five species of bushes, endemic to Australia
*''[[Laurembergia]]''
*''[[Meionectes]]''
*''[[Myriophyllum]]'', with about 69 species of water plants
*''[[Proserpinaca]]''
*''[[Trihaloragis]]''


As of 2014, the family has 138 species, distributed among the nine genera as follows:{{sfn|Chen et al|2014}}
The taxa of the earlier family Cercodiaceae and Myriophyllaceae are now included in the family Haloragaceae. Earlier, the genus ''[[Gunnera]]'' was included in this family, but it is not related and now placed in [[Gunneraceae]], a family of the order [[Gunnerales]].{{citation needed|date=2019-11-14}}
{{div col|colwidth=20em}}
* ''Glischrocaryon'' (4)
* ''Gonocarpus'' (36)
* ''Haloragis'' (24)
* ''Haloragodendron'' (6)
* ''Laurembergia'' (4)
* ''Myriophyllum'' (60)
* ''Proserpinaca'' (2)
* ''Meionectes'' (2)
* ''Trihaloragis'' (1)
{{div col end}}


== Distribution and habitat ==
== Distribution and habitat ==


The distribution of the family is nearly worldwide.{{sfn|Moody|Les|2007}} The center of species diversity is in [[Australia]] where all genera are found excepting ''Proserpinaca'' and ''Laurembergia''. Habitats vary from arid desert regions to freshwater lakes. The terrestrial genera (''Glischrocaryon'', ''Gonocarpus'', ''Haloragis'', ''Haloragodendron'', ''Trihaloragis'' ) are primarily limited to the [[southern hemisphere]]. ''Meionectes'', ''Meziella'', ''Myriophyllum'' and ''Proserpinaca'' are aquatic, while ''Laurembergia'' are semiaquatic. ''Glischrocaryon'', ''Haloragodendron'', ''Meionectes'' and ''Trihaloragis'' are Australian [[endemic]]s, where about 70% of all species are found.{{sfn|Moody|2004}}{{sfn|Moody|Les|2007}}
The distribution of the family is nearly worldwide.{{sfn|Moody|Les|2007}} The center of species diversity is in [[Australia]] where all genera are found excepting ''Proserpinaca'' and ''Laurembergia''. Habitats vary from arid desert regions to freshwater lakes. The terrestrial genera (''Glischrocaryon'', ''Gonocarpus'', ''Haloragis'', ''Haloragodendron'', ''Trihaloragis'' ) are primarily limited to the [[southern hemisphere]]. ''Meionectes'', ''Meziella'', ''Myriophyllum'' and ''Proserpinaca'' are aquatic, while ''Laurembergia'' are semiaquatic. ''Glischrocaryon'', ''Haloragodendron'', ''Meionectes'' and ''Trihaloragis'' are Australian [[endemic]]s, where about 70% of all species are found.{{sfn|Moody|2004}}{{sfn|Moody|Les|2007}} For detailed maps of the distribution of each genus, see {{harvtxt|Chen et al|2014}} ''Figure 1''.


==References==
==References==
Line 47: Line 140:


== Bibliography ==
== Bibliography ==
{{refbegin|30em}}
{{Refbegin|30em}}

=== Books and theses ===
=== Books and theses ===


* {{cite book|last1=Byng|first1=James W.|authorlink=James W. Byng|title=The Flowering Plants Handbook: A practical guide to families and genera of the world|date=2014|publisher=Plant Gateway Ltd.|isbn=978-0-9929993-1-5|url=https://books.google.com/books?id=yoLaBAAAQBAJ|chapter=Haloragaceae|page=165|ref=harv}}
* {{cite book|last1=Byng|first1=James W.|author-link=James W. Byng|title=The Flowering Plants Handbook: A practical guide to families and genera of the world|date=2014|publisher=Plant Gateway Ltd.|isbn=978-0-9929993-1-5|url=https://books.google.com/books?id=yoLaBAAAQBAJ|chapter=Haloragaceae|page=165}}
* {{cite book|last1=Christenhusz|first1=Maarten J. M.|last2=Fay|first2=Michael F.|last3=Chase|first3=Mark W.|authorlink1=Maarten Christenhusz|authorlink2=Michael F. Fay|authorlink3=Mark W. Chase|title=Plants of the World: An Illustrated Encyclopedia of Vascular Plants|url=https://books.google.com/books?id=LLo7DwAAQBAJ|date= 2017|publisher=[[University of Chicago Press]]|isbn=978-0-226-52292-0|chapter=Saxifragales|pages=231–244|ref={{harvid|Christenhusz et al|2017}}}}
* {{cite book|last1=Christenhusz|first1=Maarten J. M.|last2=Fay|first2=Michael F.|last3=Chase|first3=Mark W.|author-link1=Maarten Christenhusz|author-link2=Michael F. Fay|author-link3=Mark W. Chase|title=Plants of the World: An Illustrated Encyclopedia of Vascular Plants|url=https://books.google.com/books?id=LLo7DwAAQBAJ|date= 2017|publisher=[[University of Chicago Press]]|isbn=978-0-226-52292-0|chapter=Saxifragales|pages=231–244|ref={{harvid|Christenhusz et al|2017}}}}
* {{cite book |editor-first=Klaus |editor-last=Kubitzki |editor-link=Klaus Kubitzki |title=Flowering Plants. Eudicots: Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae Alliance, Passifloraceae Alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae |series=The Families and Genera of Vascular Plants |url=https://books.google.com/books?id=PdSL7jBNX9EC |date=2007 |publisher=Springer |volume=IX |isbn=978-3-540-32219-1 |chapter=|pages=|ref=harv}}
* {{cite book |editor-first=Klaus |editor-last=Kubitzki |editor-link=Klaus Kubitzki |title=Flowering Plants. Eudicots: Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae Alliance, Passifloraceae Alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae |series=The Families and Genera of Vascular Plants |url=https://books.google.com/books?id=PdSL7jBNX9EC |date=2007 |publisher=Springer |volume=IX |isbn=978-3-540-32219-1 }}
** {{cite book |first=Klaus |last=Kubitzki |title=Haloragaceae |pages=184–190|date=2007a|ref=harv }}, in {{harvtxt|Kubitzki|2007}}
** {{cite book |first=Klaus |last=Kubitzki |title=Haloragaceae |pages=184–190|date=2007a}}, in {{harvtxt|Kubitzki|2007}}
* {{cite book|last=Les|first=Donald H.|title=Aquatic Dicotyledons of North America: Ecology, Life History, and Systematics|url=https://books.google.com/books?id=w6QzDwAAQBAJ|date= 2017|publisher=[[CRC Press]]|isbn=978-1-351-64440-2}}
* {{cite thesis |last1=Moody |first1=Michael Lee |title=Systematics of the angiosperm family Haloragaceae R. Br. emphasizing the aquatic genus Myriophyllum: Phylogeny, hybridization and character evolution |date=2004 |publisher=[[University of Connecticut]] |url=https://opencommons.uconn.edu/dissertations/AAI3166004/|type=[[PhD thesis]]|ref=harv}}
* {{cite thesis |last1=Orchard |first1=Anthony Edward |title=Taxonomic revisions in the family Haloragaceae R. Br. |date=1972 |publisher=Department of Botany, [[University of Adelaide]] |url=https://digital.library.adelaide.edu.au/dspace/handle/2440/19634 |language=en|type=[[PhD thesis]]|ref=harv}}
* {{cite thesis |last1=Moody |first1=Michael Lee |title=Systematics of the angiosperm family Haloragaceae R. Br. emphasizing the aquatic genus Myriophyllum: Phylogeny, hybridization and character evolution |date=2004 |publisher=[[University of Connecticut]] |url=https://opencommons.uconn.edu/dissertations/AAI3166004/|type=[[PhD thesis]]}}
* {{cite thesis |last1=Orchard |first1=Anthony Edward |title=Taxonomic revisions in the family Haloragaceae R. Br. |date=1972 |publisher=Department of Botany, [[University of Adelaide]] |url=https://digital.library.adelaide.edu.au/dspace/handle/2440/19634 |language=en|type=[[PhD thesis]]}}
* {{cite book|editor1-last=Engler|editor1-first=Adolf|editorlink=Adolf Engler|last=Schindler|first=A.K.|authorlink=Anton Karl Schindler|title=Das Pflanzenreich: regni vegetablilis conspectus|publisher=W. Engelmann|location=Leipzig|year=1905|volume=23|chapter=Halorrhagaceae|url=https://www.biodiversitylibrary.org/item/68180#page/5/mode/1up|pages=|language=la, de|ref=harv}}
* {{cite book|editor1-last=Engler|editor1-first=Adolf|editor-link=Adolf Engler|last=Schindler|first=A.K.|author-link=Anton Karl Schindler|title=Das Pflanzenreich: regni vegetablilis conspectus|publisher=W. Engelmann|location=Leipzig|year=1905|volume=23|chapter=Halorrhagaceae|chapter-url=https://www.biodiversitylibrary.org/item/68180#page/5/mode/1up|language=la, de}}
* {{cite book|last1=Willis|first1=John Christopher|last2=Shaw|first2=Herbert Kenneth Airy|title=A Dictionary of the Flowering Plants and Ferns|url=https://books.google.com/books?id=RuYNAAAAMAAJ|year=1966|edition=7th|publisher=[[Cambridge University Press|Cambridge U.P.]]|ref=harv}}
* {{cite book|last1=Willis|first1=John Christopher|last2=Shaw|first2=Herbert Kenneth Airy|title=A Dictionary of the Flowering Plants and Ferns|url=https://books.google.com/books?id=RuYNAAAAMAAJ|year=1966|edition=7th|publisher=[[Cambridge University Press|Cambridge U.P.]]}}


=== Articles ===
=== Articles ===


* {{Cite journal |author=Angiosperm Phylogeny Group IV|authorlink=Angiosperm Phylogeny Group|year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=[[Botanical Journal of the Linnean Society]] |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385|ref={{harvid|APG IV|2016}} }}
* {{Cite journal |author=Angiosperm Phylogeny Group IV|author-link=Angiosperm Phylogeny Group|year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=[[Botanical Journal of the Linnean Society]] |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385|ref={{harvid|APG IV|2016}} |doi-access=free }}
* {{cite journal |last1=Chen |first1=Ling-Yun |last2=Zhao |first2=Shu-Ying |last3=Mao |first3=Kang-Shan |last4=Les |first4=Donald H. |last5=Wang |first5=Qing-Feng |last6=Moody |first6=Michael L. |title=Historical biogeography of Haloragaceae: An out-of-Australia hypothesis with multiple intercontinental dispersals |url=http://www.heimbiotop.de/Molec_Phylo_Evol_78(2014).pdf|journal=[[Molecular Phylogenetics and Evolution]] |date=September 2014 |volume=78 |pages=87–95 |doi=10.1016/j.ympev.2014.04.030|pmid=24841538 |ref={{harvid|Chen et al|2014}}}}
* {{cite journal |last1=Christenhusz|first1=Maarten JM|last2=Byng|first2= J. W. |authorlink1=Maarten Christenhusz|authorlink2=James W. Byng|lastauthoramp=yes | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = [[Phytotaxa]] | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press |ref=harv}}
* {{cite journal |last1=Christenhusz|first1=Maarten JM|last2=Byng|first2= J. W. |author-link1=Maarten Christenhusz|author-link2=James W. Byng|name-list-style=amp | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = [[Phytotaxa]] | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press | doi-access = free }}
* {{cite journal |last1=Moody |first1=Michael L. |last2=Les |first2=Donald H. |title=Phylogenetic Systematics and Character Evolution in the Angiosperm Family Haloragaceae |url=http://hydrodictyon.eeb.uconn.edu/people/les/Manuscript_Files/AJB_94%282005%29%5B2007%5D.pdf|journal=[[American Journal of Botany]] |date=2007 |volume=94 |issue=12 |pages=2005–2025 |jstor=27733378 |issn=0002-9122|ref=harv}} [https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.3732%2Fajb.94.12.2005&file=ajb22005-sup-0001.pdf Supplement]
* {{cite journal |last1=Jian|first1=Shuguang |author2-link=Pamela S. Soltis |first2=Pamela S. |last2=Soltis |first3=Matthew A. |last3=Gitzendanner |first4=Michael J. |last4=Moore |first5=Ruiqi |last5=Li |first6=Tory A. |last6=Hendry |first7=Yin-Long |last7=Qiu |first8=Amit |last8=Dhingra |first9=Charles D. |last9=Bell |author10-link=Douglas E. Soltis |first10=Douglas E. |last10=Soltis
* {{cite journal|last=Orchard|first=A.E.|title=Taxonomic revisions in the family Haloragaceae. 1. The genera ''Haloragis'', ''Haloragodendron'', ''Glischrocaryon'', ''Meziella'' and ''Gonocarpus'' .|journal=Bulletin of the Auckland Institute and Museum|year=1975|volume=10|pages=1–299|ref=harv}}
|doi=10.1080/10635150801888871 | pmid =18275001 | volume=57 | issue=1 | title=Resolving an ancient, rapid radiation in Saxifragales | date=February 2008 | pages=38–57 | journal=[[Systematic Biology]] |ref={{harvid|Jian et al|2008}}|doi-access=free }}
*{{ cite journal |last1=Orchard |first1=A E |title=Taxonomic revisions in the family Haloragaceae. 2. Further notes on Haloragus, Haloragodendron and Gonocarpus |journal=[[Nuytsia (journal)|Nuytsia]]: journal of the Western Australian Herbarium |date=1977 |volume=2 |issue=3 |pages=126–144 |url=https://www.biodiversitylibrary.org/part/228649#/summary|ref=harv}}
* {{cite journal |last1=Moody |first1=Michael L. |last2=Les |first2=Donald H. |title=Phylogenetic Systematics and Character Evolution in the Angiosperm Family Haloragaceae |url=http://hydrodictyon.eeb.uconn.edu/people/les/Manuscript_Files/AJB_94%282005%29%5B2007%5D.pdf|journal=[[American Journal of Botany]] |date=2007 |volume=94 |issue=12 |pages=2005–2025 |jstor=27733378 |issn=0002-9122|doi=10.3732/ajb.94.12.2005 |pmid=21636395 |doi-access=free }} [https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.3732%2Fajb.94.12.2005&file=ajb22005-sup-0001.pdf Supplement] {{Webarchive|url=https://web.archive.org/web/20220305210956/https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.3732%2Fajb.94.12.2005&file=ajb22005-sup-0001.pdf |date=2022-03-05 }}
* {{cite journal|last=Orchard|first=A. E.|title=''Myriophyllum'' (Haloragaceae) in Australasia. I. New Zealand: A Revision of the Genus and a Synopsis of the Family|url=http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.492.3964&rep=rep1&type=pdf|journal=[[Brunonia (journal)|Brunonia]]|year=1979|volume=2|pages=247–287|doi=10.1071/bru9850173|ref=harv}}
* {{cite journal |last1=Orchard |first1=A E |title=New taxa in Gonocarpus and Haloragis (Haloragaceae) |journal=[[Nuytsia (journal)|Nuytsia]]: journal of the Western Australian Herbarium |date=1986 |volume=5 |issue=3 |pages=327–340 |url=https://www.biodiversitylibrary.org/part/229627#/summary}}
* {{cite journal|last1=Moody|first1=M.L.|last2=Les|first2=D. H.|title=Systematics of the Aquatic Angiosperm genus ''Myriophyllum'' (Haloragaceae)|url=http://hydrodictyon.eeb.uconn.edu/people/les/Manuscript_Files/Syst_Bot_35(121)%5B2010%5D.pdf|journal=[[Systematic Botany]]|year=2010|volume=35|issue=1|pages=121–139|doi=10.1600/036364410790862470|s2cid=52999042}}
* {{cite journal|last=Orchard|first=A.E.|title=Taxonomic revisions in the family Haloragaceae. 1. The genera ''Haloragis'', ''Haloragodendron'', ''Glischrocaryon'', ''Meziella'' and ''Gonocarpus'' .|journal=Bulletin of the Auckland Institute and Museum|year=1975|volume=10|pages=1–299}}
*{{ cite journal |last1=Orchard |first1=A E |title=Taxonomic revisions in the family Haloragaceae. 2. Further notes on Haloragus, Haloragodendron and Gonocarpus |journal=[[Nuytsia (journal)|Nuytsia]] |date=1977 |volume=2 |issue=3 |pages=126–144 |url=https://www.biodiversitylibrary.org/part/228649#/summary}}
* {{cite journal|last=Orchard|first=A. E.|title=''Myriophyllum'' (Haloragaceae) in Australasia. I. New Zealand: A Revision of the Genus and a Synopsis of the Family|journal=[[Brunonia (journal)|Brunonia]]|year=1979|volume=2|pages=247–287|doi=10.1071/bru9850173|citeseerx=10.1.1.492.3964}}
* {{cite journal |last1=Orchard |first1=A E |title=New taxa in Gonocarpus and Haloragis (Haloragaceae) |journal=[[Nuytsia (journal)|Nuytsia]] |date=1986 |volume=5 |issue=3 |pages=327–340 |url=https://www.biodiversitylibrary.org/part/229627#/summary}}


=== Websites ===
=== Websites ===


* {{cite web |last1=Orchard |first1=A E |editor-last=Kodela|editor-first=Phillip|title= Haloragaceae R.Br.|url=https://profiles.ala.org.au/opus/foa/profile/Haloragaceae|website=[[Flora of Australia (series)|Flora of Australia]] Online vol.18 |publisher=[[Australian Biological Resources Study |ABRS]] |accessdate=11 November 2019 |date=2019|ref=harv}}
* {{cite web |last1=Orchard |first1=A E |editor-last=Kodela|editor-first=Phillip|title= Haloragaceae R.Br.|url=https://profiles.ala.org.au/opus/foa/profile/Haloragaceae|website=[[Flora of Australia (series)|Flora of Australia]] Online vol.18 |publisher=[[Australian Biological Resources Study|ABRS]] |access-date=11 November 2019 |date=2019}}
* {{cite web |last=POWO|authorlink=Plants of the World Online|title=Haloragaceae R.Br. |url= http://plantsoftheworldonline.org/taxon/urn:lsid:ipni.org:names:77126650-1|website=Plants of the World Online |publisher=[[Royal Botanic Gardens, Kew]]| date=2019|accessdate=12 September 2019 |ref=harv}}
* {{cite web |last=POWO|author-link=Plants of the World Online|title=Haloragaceae R.Br. |url= https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:77126650-1|website=Plants of the World Online |publisher=[[Royal Botanic Gardens, Kew]]| date=2019|access-date=12 September 2019 }}
* {{cite web |title=Haloragaceae |url=https://vicflora.rbg.vic.gov.au/flora/taxon/6a71053e-aaf0-462d-8026-be78df712e07 |website=VicFlora: Flora of Victoria |publisher=[[Royal Botanic Gardens Victoria]] |accessdate=13 November 2019|ref=harv}}
* {{cite web |title=Haloragaceae |url=https://vicflora.rbg.vic.gov.au/flora/taxon/6a71053e-aaf0-462d-8026-be78df712e07 |website=VicFlora: Flora of Victoria |publisher=[[Royal Botanic Gardens Victoria]] |access-date=13 November 2019}}
* {{cite web |title=Haloragaceae R.Br |url=https://florabase.dpaw.wa.gov.au/browse/family/22882 |website=[[FloraBase]]—the Western Australian Flora |publisher=[[Western Australian Herbarium]] |accessdate=13 November 2019 |ref=harv}}
* {{cite web |title=Haloragaceae R.Br |url=https://florabase.dpaw.wa.gov.au/browse/family/22882 |website=[[FloraBase]]—the Western Australian Flora |publisher=[[Western Australian Herbarium]] |access-date=13 November 2019 }}
* {{cite web |title=Family Haloragaceae |url=http://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=fm&name=Haloragaceae |website=PlantNET - FloraOnline |publisher=[[Royal Botanic Garden, Sydney|The Royal Botanic Garden, Sydney]] |accessdate=13 November 2019}}
* {{cite web |title=Family Haloragaceae |url=http://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=fm&name=Haloragaceae |website=PlantNET - FloraOnline |publisher=[[Royal Botanic Garden, Sydney|The Royal Botanic Garden, Sydney]] |access-date=13 November 2019}}
* {{cite web |last=Stevens |first=P.F. |author-link=Peter F. Stevens|date=2019|orig-year= 2001 |title= Saxifragales|website=[[Angiosperm Phylogeny Website|AP Web v. 14]]|publisher=[[Missouri Botanical Garden]]|url=http://www.mobot.org/MOBOT/Research/APweb/orders/saxifragalesweb.htm#Saxifragales|access-date=25 September 2019}} (''see also'' [[Angiosperm Phylogeny Website]])
* [http://florabase.calm.wa.gov.au/browse/family?f=276&level=f&id=276 Taxa of the family in Western Australia]
* {{cite web |last1= Chen |first1=Jiarui |last2=Funston |first2=Michele | title= Haloragaceae|url=http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=129989 |pages=427–428|access-date=6 December 2019|year=2004}}, in [[Flora of China (series)|Flora of China online]] vol. 13
* [http://farrer.riv.csu.edu.au/ASGAP/APOL11/sep98-5.html Article about ''Haloragodendron lucasii''.]
{{Refend}}
* [http://farrer.riv.csu.edu.au/ASGAP/h-luc.html Article about ''Haloragodendron''.]
* [http://greif.uni-greifswald.de/floragreif/?fam=Haloragaceae&gen=&spec=&flora_search=taxon Haloragaceae of Mongolia in FloraGREIF]

{{refend}}


== External links ==
== External links ==
*{{Commonscat inline|Haloragaceae}}
*{{Commons category-inline|Haloragaceae}}
*{{wikispecies inline}}
*{{Wikispecies-inline}}


{{Angiosperm families}}
{{Taxonbar|from=Q157237}}
{{Taxonbar|from=Q157237}}
{{Authority control}}


[[Category:Haloragaceae| ]]
[[Category:Haloragaceae| ]]
[[Category:Saxifragales families]]
[[Category:Saxifragales families]]


{{Saxifragales-stub}}

Latest revision as of 13:34, 17 October 2023

Haloragaceae
Myriophyllum spicatum
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Saxifragales
Family: Haloragaceae
R.Br.[1]
Genera

See text

Floral formula
✶ or

Haloragaceae (the watermilfoil family) is a eudicot flowering plant family in the order Saxifragales, based on the phylogenetic APG system. In the Cronquist system, it was included in the order Haloragales.

Description[edit]

The Haloragaceae ( alternate spelling Halorrhagidaceae) are very diverse in habit, including both small trees and submerged aquatics. Most members of the Haloragaceae are herbaceous, and most of those in turn are perennials, though some species are annuals. In contrast however, members of the genus Haloragodendron are woody. Most species of Myriophyllum are monoecious while most other taxa have hermaphrodite flowers. The flowers are usually small and inconspicuous, but some genera can have more "showy" conspicuous flowers (Haloragodendron, Glischrocaryon). Flowers are usually radially symmetrical, and unusual for core eudicots, merosity is (2-3)-4 parted. Petals are usually keeled or hooded when present. In Myriophyllum female flowers usually lack a perianth. They have (2-)4-8 stamens and an inferior ovary of (2-)4 carpels. In Myriophyllum the fruit is a schizocarp of 1-seeded 'nutlets' other genera can have nuts or drupes that can be winged or inflated.[2][3] The genus Gunnera, formerly included here are now the separate family Gunneraceae.

Taxonomy[edit]

Phylogeny[edit]

Molecular phylogenetic studies, in particular, the APG system, placed the Haloragaceae within the core eudicot order, Saxifragales.[1] Earlier versions of the APG had allowed either the broader circumscription (Haloragaceae s.l.) or a narrower Haloragaceae s.s..[4]

Cladogram of Saxifragales families[5][6][1]
Saxifragales

Peridiscaceae (4)

 97 

Paeonia (Paeoniaceae)

 woody clade 

Liquidambar (Altingiaceae)

 69 
 98 

Hamamelidaceae (27)

 95 

Cercidiphyllum (Cercidiphyllaceae)

Daphniphyllum (Daphniphyllaceae)

 core Saxifragales 

Crassulaceae (34)

 Haloragaceae s.l.

Aphanopetalum (Aphanopetalaceae)

Tetracarpaea (Tetracarpaeaceae)

Penthorum (Penthoraceae)

Haloragaceae s.s. (8)

 Saxifragaceae alliance 

Iteaceae (including Pterostemonaceae) (2)

Ribes (Grossulariaceae)

Saxifragaceae (33)

Cynomorium (Cynomoriaceae) remains unplaced within this tree

Subdivision[edit]

History[edit]

Historically, the Haloragaceae included many disparate genera, since segregated. A major circumscription was carried out by Schindler in 1905, dividing the "Halorrhagaceae" into two subfamilies (Halorrhagoideae and Gunneroideae) and the former into two tribes (Halorrhageae and Myriophylleae), with a total of seven genera. He removed some of the disparate genera and merged Gonocarpus and Meionectes into Haloragis.[7] This classification long remained the standard till Shaw (1966) removed Gunnera (into its own family Gunneraceae, within the order Gunnerales), the sole genus in Gunneroideae, leaving six genera.[8] This situation remained until the monograph of Orchard (1975). Orchard restored Gonocarpus and split Haloragodendron from Haloragis, leaving 8 genera.[2][3]
List of genera, habitat, distribution (Number of species)

Molecular era[edit]

A molecular study resolved the infrafamilial relationships among the genera, resulting in some taxonomic revision, including redistribution of species. In addition, Meionectes was reinstated, separating two species from Haloragis and creating a new monotypic genus, Trihaloragis by segregating Gonocarpus hexandrus. Consequently the number of genera is increased to ten, with the addition of:[3]

  • Meionectes R.Br. (2)
  • Trihaloragis Moody & Les (1)

Glischrocaryon-Haloragodendron is resolved as the basal node, sister to the remaining family. While monophyly of this group is well supported, monophyly of the two separate genera is less well supported, and suggests some paraphyly. Thus the generic limits remain unresolved.[3]

A subsequent, more detailed study of Myriophyllum demonstrated that the monotypic genus Meziella was embedded within it, leading to its submersion within the former as Myriophyllum subgenus Meziella, thereby reducing the number of genera within the family to 9.[9]

Species[edit]

As of 2014, the family has 138 species, distributed among the nine genera as follows:[4]

  • Glischrocaryon (4)
  • Gonocarpus (36)
  • Haloragis (24)
  • Haloragodendron (6)
  • Laurembergia (4)
  • Myriophyllum (60)
  • Proserpinaca (2)
  • Meionectes (2)
  • Trihaloragis (1)

Distribution and habitat[edit]

The distribution of the family is nearly worldwide.[3] The center of species diversity is in Australia where all genera are found excepting Proserpinaca and Laurembergia. Habitats vary from arid desert regions to freshwater lakes. The terrestrial genera (Glischrocaryon, Gonocarpus, Haloragis, Haloragodendron, Trihaloragis ) are primarily limited to the southern hemisphere. Meionectes, Meziella, Myriophyllum and Proserpinaca are aquatic, while Laurembergia are semiaquatic. Glischrocaryon, Haloragodendron, Meionectes and Trihaloragis are Australian endemics, where about 70% of all species are found.[10][3] For detailed maps of the distribution of each genus, see Chen et al (2014) Figure 1.

References[edit]

  1. ^ a b c APG IV 2016.
  2. ^ a b Orchard 1975.
  3. ^ a b c d e f Moody & Les 2007.
  4. ^ a b Chen et al 2014.
  5. ^ Jian et al 2008.
  6. ^ Stevens 2019.
  7. ^ Schindler 1905.
  8. ^ Willis & Shaw 1966.
  9. ^ Moody & Les 2010.
  10. ^ Moody 2004.

Bibliography[edit]

Books and theses[edit]

Articles[edit]

Websites[edit]

External links[edit]

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