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Metzgeriales

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Metzgeriales

Metzgeriales is an order of liverworts. The group is sometimes called the simple thalloid liverworts: "thalloid" because the members lack structures resembling stems or leaves, and "simple" because their tissues are thin and relatively undifferentiated. All species in the order have a small gametophyte stage and a smaller, relatively short-lived, spore-bearing stage. Although these plants are almost entirely restricted to regions with high humidity or readily available moisture, the group as a whole is widely distributed, and occurs on every continent except Antarctica.

Members of the Metzgeriales typically are small and thin enough to be translucent, with most of the tissues only a single cell layer in thickness. Because these plants are thin and relatively undifferentiated, with little evidence of distinct tissues, the Metzgeriales are sometimes called the "simple thalloid liverworts".

There is considerable diversity in vegetative structure of the Metzgeriales. As a rule, simple thalloid liverworts do not have structures resembling leaves. However, a few genera, such as Fossombronia, and Symphyogyna, are "semileafy" and have a thallus that is very deeply lobed, thus giving the appearance of leafiness. The genus Phyllothallia has a more striking leafiness, with paired lobes of tissue spaced regularly at swollen nodes along a central, forked stem. The several semileafy groups within the Metzgeriales are not closely related to each other, and the currently accepted view is that the leafy condition evolved separately and independently in each of the groups where it occurs.

Members of the Metzgeriales also differ from the related Jungermanniales in the location of their archegonia (female reproductive structures). Whereas archegonia in the Jungermanniales develop directly from the apical cell at the tip of a fertile branch, archegonia in the Metzgeriales develop from a cell that is behind the apical cell. As a result, the female reproductive organs, and the sporophytes that develop within them, are always located on the dorsal surface of the plant. Because these structures do not develop at the apex of the branch, their development in the Metzgeriales is described as anacrogynous, from Greek ἀν- (an-, "not") + ἄκρος (akros, "tip") + γυνή (gynē, “female”). The group was accordingly known as the Anacrogynae prior to being recognized as a separate order.

Although these plants are almost entirely restricted to regions with high humidity or readily available moisture, the group as a whole is widely distributed, and occurs on every continent except Antarctica.

One simple thalloid liverwort is not photosynthetic. The species Cryptothallus mirabilis is white as a result of lacking chlorophyll, and has plastids that do not differentiate into chloroplasts. This species is a myco-heterotroph that obtains its nutrients from fungi growing among its tissues. These plants grow in bogs and are typically found under peat moss near birch trees.

The beginning of modern liverwort nomenclature is marked with the 1753 publication of Linnaeus' Species Plantarum, although this relied heavily upon the prior work of Micheli (1729) and Dillenius (1741). Linnaeus included all 25 known species of liverworts, together with mosses, algae, and fungi, within a single class Cryptogamia. Linnaeus' system was heavily revised by workers in the early nineteenth century, so that by the time Endlicher published his Enchiridion Botanicum in 1841, five orders of liverworts were defined, and the "Frondosae" were segregated as a group that is congruent with the modern concept of the Metzgeriales. Endlicher's "Frondosae" included five subgroups (Metzgerieae, Aneureae, Haplolaeneae, Diplomitrieae, and Codonieae) with no assigned taxonomic rank, but these groups were termed Familien by Dědeček in 1886. The same five subgroups of "Frondosae", without significant change, were used in the Synopsis Hepaticarum of Gottsche, Lindenberg, and Nees.

A more thorough understanding of the Metzgeriales was not achieved until the morphological and developmental work of Leitgeb in the late nineteenth century. Leitgeb was among the first to recognize and appreciate the significance of development and reproductive morphology as a guide to distinguishing liverwort groups. His careful examinations guided revisions made in the classification published from 1893 to 1895 by Schiffner in Engler and Prantl. Schiffner thus divided his "Jungermanniales" into two broad groups according to whether the archegonia were terminal on reproductive branches (Jungermanniales akrogynae) or sub-terminal (Jungermanniales anakrogynae). This latter group included what are now recognized as the Metzgeriales, Sphaerocarpales, and Haplomitriales.

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