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Anatomically preserved fossil gleicheniaceous fern remains in carbonate marine concretions from Vancouver Island, British Columbia, Canada support the development of a whole plant concept for an Eocene species of Gleichenia , and provide data to develop the first organismal concept for an extinct species of Gleichenia from the Cenozoic fossil record. New information for completing the plant reconstruction includes anatomically preserved pinnule fragments with attached radial exindusinate sori of six to seven annulate sporangia containing trilete spores. While the overall pattern of evolution within Gleicheniaceae remains incompletely understood because well-documented fossils from pre-Cretaceous deposits are inconsistently preserved, this organismal plant concept strengthens evidence that all diagnostic features of the family Gleicheniaceae were present, and that essentially modern species were well-established and diversifying during the Cretaceous and Paleogene. • First whole plant reconstruction of a Cenozoic species of Gleichenia. • Characterized by both external morphology and internal anatomy. • Evidence of stems, frond fragments, and sori of sporangia containing spores. • Essentially modern species of Gleicheniaceae evolved no later than the Paleogene. • Family Gleicheniaceae is inconclusively present in pre-Cretaceous deposits. [ABSTRACT FROM AUTHOR]

PREMISE OF THE STUDY: Mosses, very diverse in modern ecosystems, are currently underrepresented in the fossil record. For the pre-Cenozoic, fossil mosses are known almost exclusively from compression fossils, while anatomical preservation, which is much more taxonomically informative, is rare. The Lower Cretaceous of Vancouver Island (British Columbia, Canada) hosts a diverse anatomically preserved flora at Apple Bay. While the vascular plant component of the Apple Bay flora has received much attention, the numerous bryophytes identified at the locality have yet to be characterized. METHODS: Fossil moss gametophytes in more than 20 carbonate concretions collected from the Apple Bay locality on Vancouver Island were studied in serial sections prepared using the cellulose acetate peel technique. KEY RESULTS: We describe Tricosta plicata gen. et sp. nov., a pleurocarpous moss with much-branched gametophytes, tricostate plicate leaves, rhizoid-bearing bases, and delicate gametangia (antheridia and archegonia) borne on specialized branches. A new family of hypnanaean mosses, Tricostaceae fam. nov., is recognized based on the novel combination of characters of T. plicata. CONCLUSIONS: Tricosta plicata reveals pleurocarpous moss diversity unaccounted for in extant floras. This new moss adds the first bryophyte component to an already diverse assemblage of vascular plants described from the Early Cretaceous at Apple Bay and, as the oldest representative of the Hypnanae, provides a hard minimum age for the group (136 Ma). [ABSTRACT FROM AUTHOR]

More than 800 permineralized fagaceous fruits have been studied from the Eocene Appian Way locality of Vancouver Island, British Columbia. These cupulate nuts are the most common fruits preserved at the locality. They occur in calcareous concretions and were studied using the cellulose acetate peel technique. Cupules are borne on a spiny stalk and are broadly ovate in both longitudinal and transverse section. Cupules are evalvate and have both branching and simple spines. A single, ovoid, sclerotic nut is enclosed entirely by the cupule, except at the apex, where a stylar protrusion is free from any surrounding tissues. The nut is bilocular with a glabrous endocarp lining. At maturity, a single seed occupies the locular cavity. The embryo is straight and no endosperm is evident. The single-fruited, spiny cupule is most similar to fruits of Fagaceae subfamily Castaneoideae. Bicarpellate fruits and a glabrous endocarp place them within the fossil genus Cascadiacarpa; however, they differ from Cascadiacarpa spinosa in nut wall anatomy, cupule ornamentation, shape, and size. Fruits of Cascadiacarpa exilis sp. nov. are compared with spiny, small compression and impression fossil fruits from the Eocene Taneum Creek locality of Washington State. The characters of the Washington State fossils overlap with those observed in the anatomically preserved Appian Way cupulate fruits and, as such, may represent compression and impression fossils of the same genus. The permineralized fruits document that evalvate, spiny, cupulate nuts of Fagaceae were present and common in the Eocene of western North America. [ABSTRACT FROM AUTHOR]

A fossil seed cone with characters that have been hypothesized as transitional to the origin of crown group gnetophytes has been discovered in Lower Cretaceous deposits on Vancouver Island, British Columbia, Canada. This cone, described as Protoephedrites eamesii gen. et sp. nov., provides the first anatomically preserved fossil evidence for the evolution of gnetophyte cones with seeds. The cone is a compound shoot system consisting of a primary axis with nodes that bear bracts and axillary fertile shoots in an opposite/decussate arrangement. The secondary fertile shoot axis produces one or two pairs of diminutive bracteoles and a pair of erect ovules in an opposite/decussate pattern. In contrast to crown group gnetophytes, bracteoles subtend rather than surround and enclose the ovules. Ovules have a single multiseriate integument that is adnate to the nucellus in the basal region and free distally. The micropylar tube is short with a thick integument, but it displays distinctive cells of the inner integumentary epidermis that are characteristic of gnetophyte seeds. Each seed produces a large pollen chamber with a uniseriate wall. Paired erect ovules conform to the pleisomorphic morphology previously hypothesized for Ephedra seed cones, thus supporting the proposal that ancestral gnetophyte seeds are borne terminally on oppositely arranged sporophylls. The combination of gnetophyte synapomorphies and putative pleisomorphic characters displayed by Protoephedrites broadens the known range of morphologies for the most ancient gnetophytes. Detailed comparisons to several species of Bennettitales confirm that there are fundamental structural differences separating the seeds of Gnetales from those of Bennettitales, support the hypothesis that the outer seed envelope evolved within the gnetophyte clade, and suggest that Bennettitales are not as closely related to Gnetales as hypothesized by some authors. [ABSTRACT FROM AUTHOR]

The Eocene Princeton Chert locality of southern British Columbia, Canada, provides data to develop organismal concepts for several species of fossil plants, including the first extinct species of Pinaceae, Pinus arnoldii Miller . This new species concept is based on a combination of interconnected organs, common histological features, and patterns of association among isolated plant parts. These data expand our knowledge of P. arnoldii from a morphospecies of ovulate cones to an extinct species of pine that produces woody stems with five-needled fascicles of leaves; simple pollen cones with two abaxial pollen sacs per microsporophyll, containing bisaccate pollen grains; seed cones with inflated scale apices, dorsal umbos that do not bear mucros, and a thick-walled sclerotic outer cortex; and coralloid roots with ectomycorrhizal associations. Organs preserved at various stages of development provide evidence for numerous aspects of growth and reproductive biology, including pollination biology and seed cone development. Cladistic analyses of morphological and molecular characters demonstrate that the novel combination of characters displayed by P. arnoldii causes this extinct species to fall near the base of the Pinus L. tree, either as the sister to all living species of the subgenus Pinus or outside the subgenera that have been defined by extant species of Pinus. [ABSTRACT FROM AUTHOR]

Abstract: This paper provides a whole plant concept for a new species, Emporia cryptica sp. nov. Hernandez-Castillo, Stockey, Rothwell & Mapes (Emporiaceae: Voltziales), the fourth conifer to be reconstructed from the rich fossil biota at the Late Pennsylvanian, Hamilton Quarry, Kansas. E. cryptica has an orthotropic stem, lateral plagiotropic branches with simple leaves, simple pollen cones, and compound ovulate cones. Branches have an endarch eustele with dense wood surrounding a parenchymatous pith with sclerotic nests/plates, and secondary xylem tracheids with multiseriate hexagonal bordered pits. Leaves on both penultimate and ultimate branches are simple and amphistomatic with two adaxial stomatal bands, monocyclic and dicyclic stomata, and two narrow abaxial rows of stomata with numerous trichome bases. Pollen cones are simple with helically arranged microsporophylls and adaxial pollen sacs. Prepollen is monolete, eusaccate, and monosaccate (Potonieisporites Bharadwaj). Ovulate cones are compound with bilaterally symmetrical dwarf shoots in the axils of helically arranged forked bracts. Axillary dwarf shoots bear numerous sterile scales interspersed with two megasporophylls, each bearing a single inverted terminal ovule. This plant displays morphological and cuticular characters similar to several morphogenera of Paleozoic walchian conifers but is most comparable to the fossil plant species Emporia lockardii and Hanskerpia hamiltonensis. E. cryptica is the only walchian conifer where ovules and seeds with megagametophytes, immature embryos and mature embryos have been documented, demonstrating that the most ancient conifers possessed seed dormancy and polycotyledonary embryos. This reproductive biology is similar to that of many Mesozoic and extant conifers with saccate pollen except for the presence of prepollen grains that are common among Paleozoic walchian conifers. [Copyright &y& Elsevier]

Abstract: Cunninghamia R. Brown ex. Richard is a cupressaceous arborescent conifer known from the Early Cretaceous to the Recent. Twelve specimens of permineralized leaves of Cunninghamia were identified from Late Cretaceous (Campanian) sediments on Hornby Island, British Columbia. The fossils, preserved in calcareous concretions, were examined using the cellulose acetate peel technique. The specimens are distinct from other Cunninghamia and Cunninghamia-like plants in that they appear to have a smooth leaf margin and are mostly epistomatic. The fossil leaves closely resemble the two extant species of Cunninghamia, C. lanceolata (Lambert) Hooker and C. konishii Hayata, in that they are helically arranged, some with incurved tips, and have similar leaf and stem anatomy. The Hornby Island leaves are compared to previously described extant and fossil Cunninghamia and leaves of Cunninghamiostrobus Stopes et Fujii with respect to internal anatomy. The Hornby specimens show a unique combination of characters and are described as a new species of Cunninghamia, C. hornbyensis. The presence of Cunninghamia-like plants in British Columbia demonstrates that these conifers were much more widespread in the Cretaceous than previously recognized. [Copyright &y& Elsevier]

Investigations of the Middle Eocene Princeton Chert reveal evidence for the connection of lauraceous flowers to fruits through a developmental series. Youngest fruits are found with attached floral remnants. Later stages show receptacle enlargement, fruit wall thickening, and the development of abundant sclereid clusters. Mature fruits are borne on a shallow receptacle and have an endocarp palisade layer of radially elongate cells with stellate outlines, an inner mesocarp layer of radiately arranged sclereid clusters, and a fleshy outer mesocarp layer containing numerous idioblasts with contents. Each mature fruit bears a single seed retaining the outer integument with an innermost radially elongate transfusion cell layer. Mature seeds contain a cellular embryo bearing idioblasts. Fruits are distinguishable from previously described anatomically preserved fossil taxa. This study represents the only documented developmental reconstruction of fossil fruits of Lauraceae and that self-pruning evolved prior to the Eocene. Anatomical modifications over the developmental sequence indicate that different stages of maturity preserved together, may be erroneously identified as several taxa at a fossil locality. Fossil morphotypes typically underestimate species number, but this study suggests that the number of inferred species based on fruit types may be inflated for Lauraceae, potentially exaggerating the tropical interpretation of the paleoenvironment. [ABSTRACT FROM AUTHOR]

A diverse assemblage of fossil conifer leaves was identified in calcareous marine concretions from the Lower Cretaceous (Valanginian–Hauterivian) Apple Bay locality, Vancouver Island. Of the hundreds of isolated leaf fragments, most show affinities to Pinaceae. Leaves with closest similarities to Picea (spruce) were studied using cellulose acetate peels. Picea-like leaves vary in cross-section from rhomboidal, pentagonal, triangular, to ovoid. One fused vascular bundle with a centrally located ray and abaxial sclerenchyma is surrounded by a circular endodermis and transfusion tissue. Mesophyll is plicate containing two lateral external resin canals surrounded by a sclerenchyma sheath. Hypodermal fibres are one to three layers thick, except in areas of stomata. Leaves are amphistomatic, with sunken guard cells. Vascular bundles are identical anatomically to Picea; however, plicate mesophyll is similar to that in leaves of Pinus. Extensive sclerenchyma in the hypodermis and surrounding resin canals differs from that in most extant Picea. The major difference between these leaves and those of Picea is leaf shape. These fossil leaves probably belong to an extinct pinaceous conifer, and are described as Midoriphyllum piceoides gen. et sp. nov. Similar evidence from Cretaceous seed cones suggests that like the angiosperms, the Pinaceae were undergoing rapid mosaic evolution during the Lower Cretaceous. [ABSTRACT FROM AUTHOR]

The fossil record of aquatic flowering plants broadens our understanding of their former diversity and origins from terrestrial ancestors. This paper describes a floating aquatic monocot from 71 whole plants and several isolated leaf fragments from Upper Cretaceous oxbow lake sediments in the Dinosaur Park Formation, Alberta, Canada. The new material is represented by rosettes of leaves and roots attached to short stems that are interconnected by stolons and corresponds to the fossil aroid originally described as Pistia corrugata Lesquereux. Up to six plants have been found interconnected on a single slab suggesting that these plants grew in extensive floating mats covering lakes and calm stretches of rivers. Stems have up to six leaves and large numbers of branched aquatic roots. The leaf is trumpet-shaped with an elongate clasping petiole, large aerenchymatous base, and a nearly circular blade rim. Leaf bases are often filled with sediment giving the leaf the appearance of having a basal pouch. Petioles have 6-9 veins that divide into an upper and lower set, and veins converge at an apical notch. A submarginal collective vein and at least two marginal veins with branching veins form the leaf rim. A series of dichotomizing and anastomosing veins characterize the adaxial leaf surface. Tertiary and quatemary veins form polygonal areolae. Leaf surfaces are covered in trichomes that, like those in Pistia stratiotes, probably aided in buoyancy. A reconstruction of the plant is presented. Based on unique leaf morphology, these fossil plants are clearly not assignable to the genus Pistia and are described as Cobbania corrugata (Lesquereux) Stockey, Rothwell et Johnson gen. et comb. nov. Recent systematic analyses using molecular characters resolve two separate origins of floating aquatic aroids included in the duckweeds and the genus Pistia. This new fossil genus increases our understanding of colonization of aquatic habitats by revealing a third possible origin of the floating aquatic habit within Araceae. [ABSTRACT FROM AUTHOR]

Trawetsia princetonensis Smith, Stockey, Nishida & Rothwell gen. et sp. nov. is described from several permineralized rhizomes, stipes, and higher-order frond members of a blechnoid fern that have been identified from the Middle Eocene Princeton Chert of British Columbia. Rhizomes bear stipes arising from all sides of the stem as well as diarch adventitious roots. The dictyostelic rhizomes are up to 16 mm across and have five to seven amphiphloic cauline bundles that are capped internally and externally by sclerenchyma. Ground tissue, where well preserved, is transitional from arenchymatous toparenchymatous, with abundant small sclerotic nests found throughout the pith and cortex. Multicellular scales are borne on the rhizome but are sparse and do not imbricate. At least three orders of frond material are preserved: stipes, primary pinnae, and laminar pinnules. Stipes have an adaxial groove, two adaxial hippocampiform bundles, and seven or fewer small, circular bundles arranged in an abaxial U-shaped arc. In distal sections of the rachis, the vascular bundles are reduced to three or fewer. Ground tissue of the stipe has sclerenchyma nests near the base but distally becomes completely aerenchymatous, further evidence that the Princeton Chert preserves an ancient wetland environment. This fossil material expands our knowledge of fern diversity from the Princeton Chert locality, which also includes osmundaceous, dennstaedtioid, and dryopterid ferns, and it contributes to the understanding of the timing and evolution of derived Filicales. While the vegetative anatomy of this fern allies it with the Blechnaceae, the lack of attached fertile fronds precludes the possible recognition of T. princetonensis as a fossil representative of a modern genus or perhaps even a living species. [ABSTRACT FROM AUTHOR]

Several new broad-leaved monocots were identified in gray siltstones and fine-grained sandstones from the Upper Cretaceous (Campanian-Maastrichtian) St. Mary River Formation near Cardston, Alberta, Canada. Specimens are compression/impressions of long-petiolate aquatic plants that were probably buried in situ. Leaf blades are entire, ovate to elliptic, with deeply cordate bases; leaf blades are 5-12 cm long and 3.5-8.5 cm wide. Petioles are at least 4 mm wide with five to seven primary veins that enter the leaf blade. Venation is campylodromous, resulting in 23-27 primary veins with three medial veins that remain unbranched to the apex. Major and minor secondary veins (ABAB pattern) diverge at angles of 45°-65° near the midveins and 900 near the leaf margin. Tertiary veins are usually unbranched, but occasional dichotomies and anastomoses occur. No freely ending veinlets are visible. The Cardston specimens are compared with extant leaves of Alismatales and show closest similarities to those of Lirnnocharis L., Hydrocleys Rich., and Butomopsis Kunth. These leaves are also similar to fossil leaves of Haemanthophyllum Budantsev, in particular H. cordatum Golovneva from the Maastrichtian-Danian deposits of the Koryak Highlands, Russia. A reexamination of the genus Haemanthophyllum, the generitype (H. kamtschaticum Budantsev), the holotype for H. cordatum, and the Cardston specimens results in the description of a new genus, Cardstonia tolmanii gen. et sp. nov. (Limnocharitaceae). This study points to the need for reexamination of the remaining species in the genus Haemanthophyllum, which appear to represent a diverse assemblage of leaves of varying morphology that are probably not a natural group. [ABSTRACT FROM AUTHOR]

Studies a species of the osmundaceous fern Millerocaulis from the Early Cretaceous, Upper Chickaballu Member of the Budden Canyon Formation in California. Comparison to living and fossil ferns in the family Osmundaceae; Stem characteristics; Root traces; Genera with the closest similarity to the specimen.

Focuses on the resolution of the relationships of the Lemnaceae using morphological characters of fossils and extant species. Interconnection of rosettes by stolons; Characterization of the leaves; Description of the pollen wall; Advantages of the completeness of Limnobiophyllum scutatum.

Reevaluates the hypothesis that identifies the Eorhiza as semiaquatic dicotyledonous rhizomes. Presentation of a hypothesis concerning the growth habit of E. arnoldii. Rhizome anatomy and morphology; Occurrence of horizontal rhizome-type of rooting in most stems; Reconstruction of the growth habit in rhizome systems.

Provides information on a study which examined the fossil seeds and seedlings of a Metasequoia-like taxodiaceous conifer which occur in Paleocene at the Munce's Hill and Gao Mine localities of central Alberta, Canada. Description of the sediments in Gao Mine; Information on the mature organs of Metasequoia-like plant; Other findings of the study.

Premise of the study: Two ovulate conifer cones, one of which is attached terminally to a short leafy shoot, reveal the presence of a new species of Hughmillerites in the Early Cretaceous Apple Bay flora of Vancouver Island, British Columbia, Canada. This ancient conifer expands the diversity of Cupressaceae in the Mesozoic and reveals details about the evolution of Subfamily: Cunninghamioideae. • Methods: Specimens were studied from anatomical sections prepared using the cellulose acetate peel technique. • Key results: Vegetative shoots have helically arranged leaves that are CunninghamiaAike. Seed cones have many helically arranged bract/scale complexes in which the bract is larger than the ovuliferous scale. Each ovuliferous scale has three free tips that separate from the bract immediately distal to an inverted seed. Several ovuliferous scales show interseminal ridges between seeds. • Conclusions: This study documents a new extinct species of cunninghamioid conifers, Hughmillerites vancouverensis, expanding the record of the genus from the Late Jurassic to the Early Cretaceous. This new extinct species emphasizes the important role that conifers from subfamily Cunninghamioideae played in the initial evolutionary radiation of Cupressaceae. In light of recent findings in conifer regulatory genetics, we use H. vancouverensis to hypothesize that variations of expression in certain gene homologues played an important role in the evolution of the cupressaceous ovuliferous scale. [ABSTRACT FROM AUTHOR]

Early Cretaceous fossilized leaves assignable to the extinct seed plant order Bennettitales occur within an exceptionally diverse Early Cretaceous (Valanginian) flora of anatomically preserved plant fossils at Apple Bay on Vancouver Island, British Columbia, Canada. One of the bennettitalean leaf types has an entire margin, with laminae that are attached near the adaxial surface of the midvein. Leaves are 10-15 mm wide with an adaxial surface that shows distinct corrugations, and a midrib that is exposed adaxially. The vascular system of the midrib consists of a crescent-shaped ring of collateral bundles. Lateral veins diverge from midrib at ∼90°, are typically simple but occasionally branch at base of the lamina. Vein density is 12-15 per cm. Leaves are hypostomatic with syndetocheilic stomata that are randomly oriented between veins. Internal anatomy consists of an adaxial hypodermis of closely-spaced isodiametric cells, mesophyll that is differentiated into adaxial palisade and abaxial spongy zones, and collateral bundles that show a distinct bundle sheath with bundle sheath extensions. This novel combination of characters represents the diverse bennettitalean genus Nilssoniopteris. Nilssoniopteris corrugata new species is only the second species of the genus for which internal anatomy is preserved. This species increases the known variation of bennettitalean leaf anatomy, and reinforces our appreciation for the global distribution of bennettitalean seed plants during the Mesozoic. [ABSTRACT FROM AUTHOR]

Premise of research. The occurrence of six ovulate cones and six leafy branching systems, two of which show attachment of the ovulate cone, reveals a new cunninghamioid fossil conifer from the Cretaceous Apple Bay locality of Vancouver Island, Canada. This anatomically preserved plant expands our understanding of basal Cupressaceae in the fossil record. Methodology. Specimens were studied from anatomical sections prepared by the classic cellulose acetate peel technique. Pivotal results. Vegetative shoots have helically arranged Cunninghamia-like leaves with a single vascular bundle and one to three resin canals. Ovulate cones consist of numerous helically arranged bract/scale complexes with a large bract and a small ovuliferous scale with three separate tips. There are three inverted seeds/ ovules attached adaxially, immediately proximal to the free scale tips. Conclusions. This discovery reveals the presence of a new genus and species of basal Cupressaceae, Hubbardiastrobus cunninghamioides, in the Early Cretaceous (Valanginian; ~136 Ma) of the Northern Hemisphere. These data expand the species richness of fossil Cupressaceae and further document the Cretaceous evolutionary radiation of subfamily Cunninghamioideae. [ABSTRACT FROM AUTHOR]

Permineralized cyatheaceous sori occur among remains of conifers, fungi, and other plants in newly discovered calcareous concretions from Early Cretaceous (Barremian) marine sediments of Vancouver Island, British Columbia, Canada. Sori are superficially attached in two rows to narrow pinnules and display a globose sphaeropteroid indusium. Annulate sporangia with multicellular stalks diverge from a basal, vascularized receptacle. The nearly vertical uniseriate annulus is not interrupted by the stalk. The sporangia bear 64 trilete spores with perispore sculpturing that ranges from irregular granulate/echinate to prominent rodlets. These specimens, described as Cyathea cranhamii sp. nov., are the first anatomically preserved tree fern sori from the fossil record. They represent the most ancient evidence for fertile structures of the Cyatheaceae and demonstrate that essentially modern species of cyatheaceous tree ferns had evolved by the Early Cretaceous. [ABSTRACT FROM AUTHOR]

Reports on fossil ectomycorrhizae from the British Columbia. Effects of hormones produced by living ectomycorrhizal fungi; Evolution of mycorrhizal associations; Features of the Princeton chert fossils; Characterizations of the mycorrhizal rootlets.

Premise of research. Humiriaceae occupy predominantly Neotropical lowland rainforests, with only a single Old World species in western Africa. Molecular divergence time estimates suggest that the family might have originated during the middle Cretaceous; however, fossil occurrences are lacking prior to the Paleocene. Here we provide new fossil evidence that expands our understanding of the paleobiogeographic history and evolution of this family. Methodology. Fossil endocarps and wood were compared with extant relatives of the family. Transverse and longitudinal sections of fossil and modern fruits were studied anatomically and morphologically. Pivotal results. Fruits of the new species, Duckesia berryi sp. n., from the Oligocene of Pacific coastal Peru (ca. 30-28.5 Ma), provide the earliest fossil evidence of Duckesia--the genus is now confined to Amazonia. We also document the earliest fossil fruit record for Sacoglottis tertiaria, from the early Oligocene (ca. 33.9-28.4 Ma) of Puerto Rico, and a new occurrence of Vantanea cipaconensis fruits, from the late Miocene of Panama (ca. 9-8.5 Ma). The new fossil wood, Humiriaceoxylon ocuensis gen. et. sp. n., from the late Eocene (ca. 37.2-33.9 Ma) of Ocú, Panama, confirms that this family was represented by large trees anatomically consistent with the extant genera. Conclusions. Humiriaceae originated in the Neotropics, with the oldest-known occurrences being fruits of Lacunofructus cuatrecasana and wood of H. ocuensis from the late Eocene of Panama. The family was generically diverse and widely distributed geographically across northern South America and Central America by the early Miocene. The new fossils also provide further evidence of regional extinctions within the Neotropics; extirpation of particular genera from Central America, the Caribbean, and coastal Peru indicate that Neogene orogenic and climatic events had an important effect on the modern-day distribution of the family. [ABSTRACT FROM AUTHOR]