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''Baragwanathia'' is a genus of extinct
lycopsid Lycopodiopsida is a class of vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching s ...
plants of Late
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
to Early
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
age (), fossils of which have been found in Australia, Canada, China and Czechia. The name derives from
William Baragwanath William Baragwanath (1878–1966) was an Australian surveyor, geologist and public servant. In 1922 he was appointed director of the Geological Survey of Victoria, and in 1932 Secretary for Mines. He discovered fossils of ''Baragwanathia'', a ...
who discovered the first specimens of the type species, ''Baragwanathia longifolia'', at Thomson River (Victoria, Australia).


Description

''Baragwanathia'' differed from such taxa as ''
Asteroxylon ''Asteroxylon'' ("star-shaped xylem") is an extinct genus of vascular plants of the Division Lycopodiophyta known from anatomically preserved specimens described from the famous Early Devonian Rhynie chert and Windyfield chert in Aberdeenshire, ...
'' by the presence of vascular tissue in its leaves—''Asteroxylon'' had
enations Enations are scaly leaflike structures, differing from leaves in their lack of vascular tissue. They are created by some leaf diseases and occur normally on ''Psilotum''. Enations are also found on some early plants such as '' Rhynia'', where th ...
without vascular tissue. The
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their lif ...
were borne in the
axil A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
s of the leaves, which were spirally arranged. By comparison, the closely related genus ''
Drepanophycus ''Drepanophycus'' is a genus of extinct plants of the division Lycopodiophyta of Early to Late Devonian age (around ), found in Eastern Canada and Northeast US, China, Russia, Egypt and various parts of Northern Europe and Britain. Description E ...
'' of the same period (see
Drepanophycaceae Drepanophycaceae is a family of extinct lycophytes of Late Silurian to Late Devonian age (), found in North America, China, Russia, Europe, and Australia. Description The stems are several mm to several cm in diameter and several cm to several ...
for more details) bore its sporangia on the upper surface of specialized leaves known as
sporophyll A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls (whether they are microphylls or megaphylls) bear either megasporangia and thus are called megasporophylls, or ...
s. ''Baragwanathia'' varied in size, with stems up to a few cm in diameter and up to a few metres in length. They were erect or arched, dichotomized (forked) occasionally, and had adventitious roots arising directly from prostrate stems. As in ''Asteroxylon'' the vascular bundle in the stems was an
exarch An exarch (; from Ancient Greek ἔξαρχος ''exarchos'', meaning “leader”) was the holder of any of various historical offices, some of them being political or military and others being ecclesiastical. In the late Roman Empire and e ...
actinostele, with a star-shaped arrangement of
tracheid A tracheid is a long and tapered lignified cell in the xylem of vascular plants. It is a type of conductive cell called a tracheary element. Angiosperms use another type of tracheary element, called vessel elements, to transport water through the ...
s of a primitive annular or helical type (so-called G-type). Leaves were unbranched strap-shaped
microphyll In plant anatomy and evolution a microphyll (or lycophyll) is a type of plant leaf with one single, unbranched leaf vein. Plants with microphyll leaves occur early in the fossil record, and few such plants exist today. In the classical concep ...
s (4 cm long in ''B. longifolia'') with a single prominent vascular thread, arranged spirally on the stem. The
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their lif ...
were borne in the axils of the leaves, broader than long, dehiscing by a transversely-orientated slit.
Spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s were trilete isospores. The gametophyte of ''Baragwanathia'' is currently unknown. The species ''Baragwanathia brevifolia'', described in 2017 from a single fossil specimen, had smaller microphylls than other species of ''Baragwanathia''. It had marine species (
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns and
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, wh ...
s) attached to it, and apparently growing on it, showing that at least the lower part of the plant grew in marine water. As it is one of the oldest lycophyte fossils, the implication is that land-based lycophytes evolved from aquatic precursors.


Age

The age of ''Baragwanathia'' has been uncertain because the fossils described by Lang and Cookson (1935) at first appeared to be of Late Silurian age, associated as they were with the
graptolite Graptolites are a group of colonial animals, members of the subclass Graptolithina within the class Pterobranchia. These filter-feeding organisms are known chiefly from fossils found from the Middle Cambrian (Miaolingian, Wuliuan) through the ...
genus ''
Monograptus ''Monograptus'' is a genus of graptolites in the Order Graptoloidea. This particular genus is the last stage of the graptoloid evolution before its extinction in the early Devonian. A characteristic of the genus includes one uniserial stipes ...
''. This would make the species by far the most advanced known plant of the time. However, the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
( Ludlovian) dating of the deposit in
Victoria, Australia Victoria is a state in southeastern Australia. It is the second-smallest state with a land area of , the second most populated state (after New South Wales) with a population of over 6.5 million, and the most densely populated state in A ...
which produced the type specimens of ''Baragwanathia longifolia'' was later disproved, as ''Monograptus'' (and in particular the species present at that site) was later proved to persist into the
Early Devonian The Early Devonian is the first of three epochs comprising the Devonian period, corresponding to the Lower Devonian series. It lasted from and began with the Lochkovian Stage , which was followed by the Pragian from and then by the Emsian, wh ...
. Since then, specimens from a different Victorian locality have been found that occur with veritable Late Silurian
graptolite Graptolites are a group of colonial animals, members of the subclass Graptolithina within the class Pterobranchia. These filter-feeding organisms are known chiefly from fossils found from the Middle Cambrian (Miaolingian, Wuliuan) through the ...
s., and earlier papers referred to therein. See
Yea Flora Fossil Site The Yea Flora Fossil Site is a roadside cutting on Limestone Road, Yea, Victoria, Australia. It contains fossils of genus Baragwanathia, some of the world's earliest vascular plants dating back to the begin of the Devonian period, 415 million y ...
.
The species ''Baragwanathis brevifolia'' has been dated to the lower Pridoli, about . The genus ''Baragwanathia'' persisted at least until the
Emsian The Emsian is one of three faunal stages in the Early Devonian Epoch. It lasted from 407.6 ± 2.6 million years ago to 393.3 ± 1.2 million years ago. It was preceded by the Pragian Stage and followed by the Eifelian Stage. It is named after the ...
(Late Lower Devonian) and probably had a worldwide distribution.


See also

*
Yea Flora Fossil Site The Yea Flora Fossil Site is a roadside cutting on Limestone Road, Yea, Victoria, Australia. It contains fossils of genus Baragwanathia, some of the world's earliest vascular plants dating back to the begin of the Devonian period, 415 million y ...
*
List of Early Devonian land plants The list of Early Devonian (419.2 ± 2.8 to 393.3 ± 2.5 million years ago) land plants includes currently known vascular and potentially vascular plants, along with some possibly non-vascular plants, that have been described from global Early Devon ...


Notes


References

*Hao SG and Gensel PG (2001) The Posongchang Floral Assemblages of Southeastern Yunnan, China - Diversity and Disparity in Early Devonian Plant Assemblages. In ''Plants Invade the Land. Evolutionary and Environmental Perspectives'', pp. 103–119. Eds PG Gensel and D Edwards. (Columbia University Press, New York). * * * * * {{Taxonbar, from=Q1452954 Early Devonian plants Silurian plants Drepanophycales Paleozoic life of Ontario Prehistoric lycophyte genera