Trithuria is a genus of small ephemeral aquaticherb that represent the only members of the family Hydatellaceae found in India, Australia, and New Zealand. Almost all described species of Trithuria are found in Australia, with the exception of T. inconspicua and T. konkanensis, from New Zealand and India respectively. Until DNA sequence data and a reinterpretation of morphology proved otherwise, these plants were believed to be monocots related to the grasses (Poaceae). They are unique in being the only plants besides two members of Triuridaceae (Lacandonia schizmatica and L. braziliana) in which the stamens are centred and surrounded by the pistils; in Hydatellaceae the resulting 'flowers' may instead represent condensed inflorescences or non-flowers.
These diminutive, superficially moss-like, aquatic plants are the closest living relatives of a clade comprising two closely related water-lily families Nymphaeaceae and Cabombaceae. Together, these three families compose the orderNymphaeales in the APG III system of flowering plant classification. Trithuria (Hydatellaceae) diverged from the rest of Nymphaeales soon after Nymphaeales diverged from its sister taxon, although the crown clade evolved relatively recently, in the early Miocene (~19 Ma;). The order as a whole is the sister group of all flowering plants except Amborellales.
Trithuria exhibits a remarkable similarity to Centrolepis and species of both genera were mistaken for members of the other genus.
Description
Flowering Trithuria submersaTrithuria inconspicuaBotanical illustration of Trithuria australis
Plants are submerged and emergent aquatic plants, rooted in the substrate below the water. They are tiny plants, just a few cm tall. Most species are ephemeral aquatics that flower in vernal pools when the water draws down, but several species are submerged perennials found in shallow lakes. The simple leaves are concentrated basally around a short stem. Individual species are cosexual (with several types of hermaphroditic conditions) or dioecious, and are either wind-pollinated (anemophilous) or self-pollinating (autogamous). Two predominantly apomictic species are also known. Flower-like reproductive units are composed of small collections of minute stamen- and/or pistil-like structures that may each represent very reduced individual flower, so that the reproductive units may be pseudanthia. The non-fleshy fruits are follicles or achenes. Hydatellaceae are unusual among angiosperms in allocating nutrients to the embryo before fertilization. In all other flowering plants, nutrient provisioning is known to occur only after fertilization. This trait is similar to gymnosperms, although in gymnosperms the nutrients come from the gametophyte rather than maternal tissue.
Cytology
The diploid chromosome count of Trithuria inconspicua subsp. inconspicua is 2n = c. 24.
The diploid chromosome count of Trithuria submersa is 2n = 56. The diploid chromosome count of the tetraploid species Trithuria konkanensis 2n = 40. The diploid chromosome count of Trithuria australis is 2n = 14.
Taxonomy
The genus TrithuriaHook.f. was described by Joseph Dalton Hooker in 1858 with the type species Trithuria submersaHook.f. It has two synonyms: In 1888 the genus JuncellaF.Muell. ex Hieron. was described without a type designation by Georg Hans Emmo Wolfgang Hieronymus based on previous work by Ferdinand von Mueller. Mueller had invalidly published the nomen invalidum and nomen nudumJuncella tasmanicaF.Muell. in 1854. Juncella is a nomen illegitimum. In 1904 the genus HydatellaDiels was described by Friedrich Ludwig Emil Diels without a type designation, but the lectotypeHydatella australisDiels. has been designated in 2008. The genus Hydatella was synonymised with Trithuria in 2008.
Trithuria was initially placed in the family Centrolepidaceae Endl. (now synonymous with RestionaceaeR.Br.), which is placed in the order PoalesSmall but it was separated as its own family Hydatellaceae U.Hamann by Ulrich Hamann in 1976 with HydatellaDiels as the type genus. Upon its separation, Hamann stated the new families affinity or placement were still obscure. The correct placement of the family became apparent in 2007, when it was identified as a basal angiosperm lineage. The family Hydatellaceae is now placed in the order Nymphaeales Salisb. ex Bercht. & J. Presl.. Alternatively, it is placed in a separate order Hydatellales Cronquist ex Reveal & Doweld validly published by James Lauritz Reveal and Alexander Borissowitsch Doweld in 1999 based on previous work by Arthur Cronquist (see the Cronquist system). This is however not widely accepted, as the order Hydatellales is mostly treated as a synonym of Nymphaeales.
Etymology
The generic name Trithuria is derived from the Greek words τρεις treis meaning "three", and θυρις thyris meaning "window". It references the dehiscence of the capsule fruit.
Species
Trithuria has 13 species, although species diversity in the family has probably been substantially underestimated.
Trithuria polybracteata D.A.Cooke ex D.D.Sokoloff, Remizowa, T.D.Macfarl. & Rudall - Western Australia
Trithuria submersa Hook.f. - Western Australia, South Australia, Victoria, New South Wales, Tasmania
Distribution and habitat
It is native to India, Australia, and New Zealand, where it occurs in seasonally dry aquatic environments as annuals, or more rarely in permanently inundated habitats as perennial plants.
Conservation
The IUCN conservation status of Trithuria lanterna is Least Concern (LC). Trithuria inconspicua is critically endangered.
↑Smissen, Rob D.; Ford, Kerry A.; Champion, Paul D.; Heenan, Peter B. (2019). "Genetic variation in Trithuria inconspicua and T. filamentosa (Hydatellaceae): a new subspecies and a hypothesis of apomixis arising within a predominantly selfing lineage". Australian Systematic Botany. CSIRO Publishing. doi:10.1071/sb18013. ISSN1030-1887. S2CID91261563.
↑Macfarlane, T.D.; Watson, L. & Marchant, N.G., eds. (2000). "Hydatellaceae U Hamann". FloraBase: Flora of Western Australian. Archived from the original on 14 July 2007. Retrieved 2013-11-15.
↑Sokoloff, D. D., Marques, I., Macfarlane, T. D., Remizowa, M. V., Lam, V. K. Y., Pellicer, J., … Graham, S. W. (2019). Cryptic species in an ancient flowering-plant lineage (Hydatellaceae, Nymphaeales) revealed by molecular and micromorphological data. TAXON, 68(1), 1–19. doi:10.1002/tax.12026
↑Hooker, Joseph Dalton, Fitch, W. H., & Reeve Brothers. (1844). The botany of the Antarctic voyage of H.M. discovery ships Erebus and Terror in the Years 1839-1843 :under the command of Captain Sir James Clark Ross (Vol. 3, Issue 2, pp. 78-79). Reeve Brothers. https://www.biodiversitylibrary.org/page/28467263
↑Sokoloff, Dmitry D.; Remizowa, Margarita V.; Beer, Anton S.; Yadav, Shrirang R.; Macfarlane, Terry D.; Ramsay, Margaret M.; Rudall, Paula J. (May 2013). "Impact of spatial constraints during seed germination on the evolution of angiosperm cotyledons: A case study from tropical Hydatellaceae (Nymphaeales)". American Journal of Botany. 100 (5): 824–843. doi:10.3732/ajb.1200620. PMID23613353.
↑Iles, Will; Rudall, Paula J.; Sokoloff, D. D.; Graham, Sean W (March 2012). "Molecular phylogenetics of Hydatellaceae (Nymphaeales): Sexual-system homoplasy and a new sectional classification". American Journal of Botany. 99 (4): 663–676. Bibcode:2012AmJB...99..663I. doi:10.3732/ajb.1100524. PMID22473977.