Princeton Chert Research Articles

Permineralized monocotyledons from the Middle Eocene Princeton chert (Allenby Formation) of British Columbia: Alismataceae

One small monocotyledon petiole, 1.8 × 1.5 mm wide, has been recovered from the Princeton chert in the Middle Eocene Allenby Formation, British Columbia. The petiole, rectangular in transverse outline, shows approximately 36 circular to oval-shaped vascular bundles within aerenchymatous ground tissue that includes tannin cells. The epidermis is underlain by a discontinuous hypodermis of thick-walled, pitted cells. Vascular bundles are in five series: (I) a median U-shaped arc of 11 – 13 bundles; (II) an abaxial arc of 6 bundles located below the main arc; (III) two short abaxial arcs of 3 bundles each; (IV) 2 bundles just below the abaxial surface; and (V) an adaxial series of 7 bundles that show an inverse orientation to those bundles in series I–IV. Larger bundles are collateral, with a protoxylem lacuna encircled by a ring of 9 – 14 thin-walled parenchyma cells, a relatively well-developed phloem strand, and one to three thin-walled metaxylem elements. Based on bundle arrangement, orientation, and morphology, the fossil petiole most closely resembles those of the Butomaceae and Alismataceae. This new species, Heleophyton helobiaeoides Erwin and Stockey gen. et sp.nov., in the Princeton chert flora, documents the presence of the Alismataceae in the Middle Eocene of western North America and provides further evidence that the locality represents an ancient aquatic ecosystem.

Permineralized Fruits and Seeds from the Princeton Chert (Middle Eocene) of British Columbia: Nymphaeaceae

Nymphaeaceous seeds and their probable fruit remains are described from the Middle Eocene Allenby Formation of Princeton. British Columbia. These seeds are subspheroidal to ovoid, anatropous, exarillate, and operculate. Two openings in the operculum represent the position of hilum and micropyle. Cells of the operculum are radially elongate with straight walls and can be distinguished from cells of the palisade-like outer integumentary layer that have sinuous radial walls. Surface views of cells of the outer integument show sinuous walls with four to 10 undulations per cell. The inner zone of the outer integument is composed of one to two rows of tangentially elongated thin-walled parenchymatous cells. The inner integument is composed of one to two rows of tangentially elongated, thin-walled cells. Two seeds have been found with preserved perisperm. A single vascular strand runs from hilum to chalaza, and a raphal ridge is present. Comparisons to seeds of both extant and fossil Nymphaeales show that these seeds should be included in the family Nymphaeaceae. Allenbya collinsonae Cevallos-Ferriz et Stockey gen. et sp. nov. shows closest similarities to the fossil genera Dusembaya and Palaeonymphaea and may be related to the extant genus Victoria. The description of this new taxon adds to our knowledge of the four lines of nymphaeaceous seed morphology and their evolution. Evolutionary trends within the Nymphaeales include: the establishment of tubercles or small protuberances in the outer integument in one line, and the change in position of the hilum from the operculum to the seed body and development of a broad integumentary zone beneath the palisade in a second line. These fossils show that the family Nymphaeaceae was more diverse in the past and add to our knowledge of the aquatic elements in the Princeton flora.

Permineralized Fruits and Seeds from the Princeton Chert (Middle Eocene) of British Columbia: Araceae

Some of the most conspicuous fruit and seed remains from the Middle Eocene Princeton chert locality (Allenby Formation) are assignable to the Araceae. Several thousand campylotropous, reniform, spiny seeds 2.5–3.2 mm long x 1.8–2.3 mm wide have been found dispersed in the chert. Two fruits, each with a single locule and at least eight ovules, show three distinct wall layers and appear to have been fleshy. The two layers of seed integument have pitted isodiametric sclereids aligned in radial rows, the outermost being thinner walled. Spines are borne in three rows on the dorsal seed surface while the ventral side is flattened. Idioblasts that probably contained raphides are present in the outer integument. Nucellar tissue is attached to the integument for most of its length and is well-preserved in some seeds near the chalaza and in the large conical shaped area beneath the micropyle. Endosperm cells with dark contents and curved, linear, monocotyledonous embryos are present in some seeds. Sections of seeds of living araceous taxa of the subfamilies Monsteroideae (Epipremnum, Rhodospatha) and Lasioideae (Urospatha, Cyrtosperma) were made for comparison. The fossil seeds represent a new taxon most closely related to Cyrtosperma, Keratosperma allenbyensis Cevallos-Ferriz et Stockey gen. et sp. nov., Family: Araceae, Subfamily: Lasioideae, Tribe: Lasieae. The Princeton remains are the oldest described seeds of this tribe known to date and add to our knowledge of the subtropical elements of the Princeton flora.

PERMINERALIZED FRUITS AND SEEDS FROM THE PRINCETON CHERT (MIDDLE EOCENE) OF BRITISH COLUMBIA: ARACEAE

Some of the most conspicuous fruit and seed remains from the Middle Eocene Princeton chert locality (Allenby Formation) are assignable to the Araceae. Several thousand campylotropous, reniform, spiny seeds 2.5–3.2 mm long x 1.8–2.3 mm wide have been found dispersed in the chert. Two fruits, each with a single locule and at least eight ovules, show three distinct wall layers and appear to have been fleshy. The two layers of seed integument have pitted isodiametric sclereids aligned in radial rows, the outermost being thinner walled. Spines are borne in three rows on the dorsal seed surface while the ventral side is flattened. Idioblasts that probably contained raphides are present in the outer integument. Nucellar tissue is attached to the integument for most of its length and is well‐preserved in some seeds near the chalaza and in the large conical shaped area beneath the micropyle. Endosperm cells with dark contents and curved, linear, monocotyledonous embryos are present in some seeds. Sections of seeds of living araceous taxa of the subfamilies Monsteroideae (Epipremnum, Rhodospatha) and Lasioideae (Urospatha, Cyrtosperma) were made for comparison. The fossil seeds represent a new taxon most closely related to Cyrtosperma, Keratosperma allenbyensis Cevallos‐Ferriz et Stockey gen. et sp. nov., Family: Araceae, Subfamily: Lasioideae, Tribe: Lasieae. The Princeton remains are the oldest described seeds of this tribe known to date and add to our knowledge of the subtropical elements of the Princeton flora.

Permineralized fruits and Seeds from the Princeton chert (Middle Eocene) of British Columbia: Lythraceae

Two types of lythraceous fruit and seed remains are described from the Middle Eocene, Allenby Formation of Princeton, British Columbia. Small globose, pedicellate fruits with persistent sepals, four to six locules, septa complete to the fruit apex, axial placentation, a palisade-like endocarp, and pyramidal seeds are referable to the genus Decodon Gmel. Numerous dispersed anatropous seeds of the same morphology show rounded or angular comers, lack lateral wings, and have a raphe that begins near the micropyle, passing through the middle integumentary layer on the dorsal face to the seed chalaza. Integuments are three zoned and seeds bear a ventral germination valve composed of radially elongated rectangular cells similar to the outer integumentary layer. Seeds show a prominent hypostase and dicotyledonous embryos with remains of a suspensor. These fruits and seeds are described as a new species of Decodon, D. allenibyensis Cevallos-Ferriz et Stockey sp.nov., which represents the oldest known species of the genus described to date. One fruit with several large, pyramidal lythraceous seeds is described. Seeds have three integumentary zones, the outer of which appears to have been mucilaginous. These remains most closely resemble the genus Lawsonia L., but additional specimens are needed to confirm this comparison. These lythraceous remains add to our knowledge of the Princeton angiosperm flora and further support the idea that these plants lived along a lake or marsh.

A PERMINERALIZED FLOWER FROM THE MIDDLE EOCENE OF BRITISH COLUMBIA

A permineralized flower bud, two stamen clusters and one isolated stamen of similar morphology have been found in the black cherts of the Middle Eocene Allenby Formation of Princeton, British Columbia. Specimens were studied using a modified cellulose acetate peel technique and hydrofluoric acid. The single flower specimen, 4.5 mm long and 4.0 mm in diameter, represents half of a relatively mature bud of a bisexual flower with a superior ovary. The two‐loculate pistil is 2.5 mm long with a solid style and a lobed stigmatic surface. No ovules have been observed in attachment. Twenty‐two to 24 stamens are borne in three whorls or a tight helix. Pollen sacs of the anther are elongate with a thin connective while filaments are laminar. Anther walls contain rectangular cells with dark contents that also can be identified in isolated stamens or stamen clusters. Abundant stephanocolpate (pentacolpate), psilate pollen grains 20 μm in diameter have been isolated and examined using scanning and transmission electron microscopy. Grains are tectate, columellate with a broad foot layer that thins near the apertures, and an endexine of small platelets. The remains of four petals are surrounded by one large sepal, suggesting two in the whole flower. Morphological features of this flower are comparable to taxa of the Flacourtiaceae and Papaveraceae, but show closest similarities to the Eschscholziaeae of the Papaveraceae. Difficulties with reconciling the placement of this flower in the Eschscholziaeae and the known environment of deposition of the Princeton chert are discussed. The fossil material represents a new angiospermous taxon: Princetonia allenbyensis Stockey gen. et sp. nov., family Incertae sedis.

The vegetative body of Metasequoia milleri from the Middle Eocene of southern British Columbia

Anatomically preserved vegetative remains of Metasequoia milleri Rothwell and Basinger are common in the Princeton chert of the Allenby Formation in southern British Columbia. Deposition of the Allenby Formation and associated volcanics occurred during Middle Eocene time. The Princeton chert locality is in the upper strata of the Allenby Formation and is of late Middle Eocene age. The Princeton chert was formed by siliceous permineralization of marsh soil. Dissolved silicates were probably introduced by periodic influx of water from mineral springs or geysers.Anatomical features of stems, wood, and leaves are well preserved. Mature wood of the fossil resembles that of Metasequoia glyptostroboides in having traumatic resin cysts; opposite pitting on radial walls of tracheids; taxodioid cross-field pitting; tall, uniseriate rays; smooth-walled ray parenchyma; and diffuse, resinous, smooth-walled wood parenchyma. Leaves are linear, hypostomatic, and borne decussately, and have one or three resin ducts and slightly undulate to smooth epidermal cell walls. Leaves of living M. glyptostroboides differ in consistently having three resin ducts and much more pronounced undulations of epidermal cell walls. Metasequoia milleri has leaves of a generalized structure from which leaf types of many taxodiaceous genera could be derived. Roots associated with M. milleri are dimorphic. Primary tissues and secondary phloem are exceedingly well preserved. Cortex of both long and short roots contains mycorrhizal fungi.Compression remains of M. occidentalis are not distinguishable from M. glyptostroboides. Anatomical features of M. milleri, however, do reveal differences between Eocene and living Metasequoia. Wood rays of M. milleri are much higher than those of M. glyptostroboides. Other features of stem anatomy are similar in both species. Pollen cones differ in developmental and minor anatomical features. The close similarity of most organs of M. milleri to M. glyptostroboides and the dissimilarity of leaf structure indicate mosaic evolution within the genus.