A dwarf conifer tree from the Triassic of Antarctica: the first fossil evidence of suppressed growth in a favorable climate?

BACKGROUND AND AIMS: The complexity of fossil forest ecosystems is difficult to reconstruct due to the fragmentary nature of the fossil record. However, detailed morpho-anatomical studies of well-preserved individual fossils can provide key information on tree growth and ecology, including in biomes with no modern analog such as the lush forests that developed in the polar regions during past greenhouse climatic episodes. METHODS: We describe an unusual-looking stem from Middle Triassic (ca 240 Ma) deposits of Antarctica with over 100 very narrow growth-rings and conspicuous persistent vascular traces through the wood. Sections of the specimen were prepared using the cellulose acetate peel technique to determine its systematic affinities and analyse its growth. KEY RESULTS: The new fossil shows similarities with the form genus Woodworthia and with conifer stems from the Triassic of Antarctica, and is assigned to the conifers. Vascular traces are interpreted as those of small branches retained on the trunk. Growth-ring analyses reveal one of the slowest growth rates reported in the fossil record, with an average of 0.2 mm/season. While the tree was growing within the Triassic polar circle, sedimentological data and growth-ring information from other fossil trees, including from the same locality, support the presence of favorable conditions in the region. CONCLUSIONS: The specimen is interpreted as a dwarf conifer tree that grew under a generally favorable regional climate but whose growth was suppressed due to stressful local site conditions. This is the first time that a tree with suppressed growth is identified as such in the fossil record, providing new insights on the structure of polar forests under greenhouse climates and, more generally, on the complexity of tree communities in deep time.

Fossil evidence of tylosis formation in Late Devonian plants.

Tyloses are swellings of parenchyma cells into adjacent water-conducting cells that develop in vascular plants as part of heartwood formation or specifically in response to embolism and pathogen infection. Here we document tyloses in Late Devonian (approximately 360 Myr ago) Callixylon wood. This discovery suggests that some of the earliest woody trees were already capable of protecting their vascular system by occluding individual conducting cells.

Keraphyton gen. nov., a new Late Devonian fern-like plant from Australia.

The first plants related to the ferns are represented by several extinct groups that emerged during the Devonian. Among them, the iridopterids are closely allied to the sphenopsids, a group represented today by the genus Equisetum. They have been documented in Middle to early Late Devonian deposits of Laurussia and the Kazakhstan plate. Their Gondwanan record is poor, with occurrences limited to Venezuela and Morocco. Here we describe a new genus from a late Late Devonian locality of New South Wales. It is represented by a single anatomically preserved large stem characterized by a star-shaped vascular system with protoxylem strands located at rib tips, and by a lack of secondary tissues. Within the first fern-like plants, this stem shares the largest number of characters with iridopterid axes but differs by the pattern of its vascular system. Keraphyton mawsoniae gen. et sp. nov. adds a new record of early fern-like plants in eastern Gondwana. It provides new insights into the anatomical diversity within this key group of plants and supports the distinctiveness of the Australian flora in the latest Devonian.

Under pressure? Epicormic shoots and traumatic growth zones in high-latitude Triassic trees from East Antarctica.

Background and Aims: Investigating the biology of trees that were growing at high latitudes during warmer geological periods is key to understanding the functioning of both past and future forest ecosystems. The aim of this study is to report the first co-occurrence of epicormic shoots and traumatic growth zones in fossil trees from the Triassic of Antarctica and to discuss their biological and environmental implications. Methods: Permineralized woods bearing scars of epicormic shoots were collected from the Triassic Fremouw Formation in Gordon Valley, Central Transantarctic Mountains, Antarctica in 2010. Samples from different portions of three specimens were prepared using standard thin section and hydrofluoric (HF) acid peel techniques, and anatomical details were studied in transmitted light. Key Results: The fossil woods represent the outer part of trunks, with at least 40 growth rings that are 0.2-4.8 mm in width. Anatomical comparisons suggest that they represent a new tree taxon for the Triassic of Antarctica. Numerous small epicormic shoots can be seen crossing the wood almost horizontally and are locally branched. Each specimen also contains several occurrences of traumatic growth zones located in the early wood, in the cells produced either at the very start of the growing season or slightly later. Conclusions: This is the first report of epicormic shoots and traumatic growth zones in the wood of a Triassic tree from Antarctica. Their co-occurrence indicates that these trees from Gordon Valley were subjected to environmental stresses not seen in Triassic trees previously described from this region. This suggests that they had a different biology and/or were growing in a different habitat, which offers a new glimpse into the diversity of high-latitude trees in the Triassic greenhouse climate.

A Callixylon (Archaeopteridales, Progymnospermopsida) trunk with preserved secondary phloem from the Late Devonian of Morocco.

PREMISE OF THE STUDY: During the Devonian, the evolution of secondary phloem produced by a bifacial vascular cambium was a key innovation that increased the ability of plants within the lignophyte clade to redistribute photosynthates and other organic compounds throughout their body. Unraveling the secondary phloem anatomy of the first arborescent lignophytes is crucial to understand the evolution of this tissue and the physiology of early trees. METHODS: A 10 cm wide stem of Callixylon with preserved secondary phloem from the Famennian of Morocco is described using thin-sections. KEY RESULTS: The secondary phloem of this Callixylon zalesskyi-type of stem is composed of fibers, sclereids, rays, axial parenchyma, and putative sieve cells. Fibers differentiate early and are relatively abundant in the inner phloem. In the older phloem, fibers are arranged in tangential bands alternating with extensive layers of axial parenchyma. Changes between the young and old phloem involve the periclinal division and radial elongation of the axial parenchyma cells. The presence of fibers in the inner, presumably functional phloem, combined with evidence for rhythmicity in the production of different phloem cell types are documented for the first time in detail in an archaeopteridalean progymnosperm. No periderm was observed within the preserved seven millimeters of bark tissues. CONCLUSIONS: The secondary phloem anatomy supports a close affinity of archaeopteridalean progymnosperms with both aneurophytalean progymnosperms and seed plants. The production of secondary phloem might have provided an advantage to these first arborescent lignophytes over other types of Devonian early trees, especially in dry conditions.

Morphological and functional stasis in mycorrhizal root nodules as exhibited by a Triassic conifer.

Mycorrhizal root nodules occur in the conifer families Araucariaceae, Podocarpaceae, and Sciadopityaceae. Although the fossil record of these families can be traced back into the early Mesozoic, the oldest fossil evidence of root nodules previously came from the Cretaceous. Here we report on cellularly preserved root nodules of the early conifer Notophytum from Middle Triassic permineralized peat of Antarctica. These fossil root nodules contain fungal arbuscules, hyphal coils, and vesicles in their cortex. Numerous glomoid-type spores are found in the peat matrix surrounding the nodules. This discovery indicates that mutualistic associations between conifer root nodules and arbuscular mycorrhizal fungi date back to at least the early Mesozoic, the period during which most of the modern conifer families first appeared. Notophytum root nodules predate the next known appearance of this association by 100 million years, indicating that this specialized form of mycorrhizal symbiosis has ancient origins.

Root suckering in a Triassic conifer from Antarctica: paleoecological and evolutionary implications.

PREMISE OF THE STUDY: Although root suckering and other types of sprouting are well studied in extant woody plants, little is known about the distribution of these traits at a macroevolutionary scale. Anatomically preserved fossil plants represent an excellent but understudied source of information of the distribution of sprouting behavior through time and across taxa. METHODS: A block of silicified peat collected in the Middle Triassic Fremouw Formation at the Fremouw Peak locality, Central Transantarctic Mountains, Antarctica, contains a group of anatomically preserved roots of the fossil conifer Notophytum krauselii that bear young shoots. The specimen was prepared using the standard acetate peel technique and studied in reflected and transmitted light. KEY RESULTS: Young sucker shoots bearing well-preserved leaves are produced in groups in some areas of the Notophytum roots. CONCLUSIONS: The production of root suckers in Notophytum indicates that some of the trees growing in polar forests during the Triassic could respond to environmental stresses by regenerating their vegetative structures and had the potential to reproduce vegetatively. The specimens also represent the first anatomical evidence of root suckering in any fossil seed plant, and its occurrence in an early putative podocarp supports the idea that this trait might be ancestral in at least some extant conifer families.