Fossil woods from the Beaufort Formation (Tertiary), northwestern Banks Island, Canada

Fossil woods from the Beaufort Formation (Tertiary), northwestern Banks Island, Canada

S . K. ROY Department of Botany, University of Alberta, Edmonton, Alberta

AND

L. V. HILLS Department of Geology, University of Calgary, Calgary, Alberta

Received April 12, 1972

ROY, S. K., and L. V. HILLS. 1972. Fossil woods from the Beaufort Formation (Tertiary), northwestern Banks Island, Canada. Can. J. Bot. 50: 2637-2648.

Fossil woods belonging to the Pinaceae (Pinoxylon albicauloides Greguss, 1954; Picea beaufortense sp. nov.; Laricioxylon occidentaloides sp. nov.; Abietoxylon koreanoides sp. nov.; A. traumatiductus sp. nov.) and a single representative of the Eleagnaceae (EIeagnaceoxylon shepherdioides gen. et sp. nov.) are described and illustrated. Two main conclusions are derived from this investigation. They are (1) the fossil woods of the Beaufort Formation are very similar to the woods of some extant arborescent species of northern North America and northeastern Asia and (2) the fossil flora indicates a temperate climate similar to that of the present-day boreal forest.

ROY, S. K., et L. V. HILLS. 1972. Fossil woods from the Beaufort Formation (Tertiary), northwestern Banks Island, Canada. Can. J. Bot. 50: 2637-2648.

Les auteurs dkrivent et illustrent des bois fossiles appartenant aux Pinaceae (Pinoxylon albicalrloides Gre~uss . 1954: Picea beaufortense SD. nov.: Laricioxvlon occidentaloides SD. nov.: Abietoxvlon koreanoides sp. iov.; A. trauma ti duct;^ sp. no;.) et un seul representant des ~leagnaceae (~1eagnac;oxylon shepherdioides gen. et sp. nov.). Deux conclusions gkntrales se dtgagent de ce travail. D'abord, (1) les bois fossiles de la Formation de Beaufort ressernblent beaucoup 5. ceux d'espkes arborescentes actuelles dans le nord de 1'Amtrique du Nord et le nord-est de I'Asie, et (2) cette flore fossile est indicatrice d'un climat temptrt sernblable 5. celui qui existe actuellernent dans la forkt boriale. [Traduit par le journal]

Introduction The Beaufort Formation is a fluvio-lacustrine

deposit that occurs along the western fringe of the Arctic Archipelago. The formation has been recorded from Meighen Island on the north to the southern part of Banks Island on the south (Craig and Fyles 1960).

Hills (1969) subdivided the Beaufort Forma- tion into lower unit (X), composed of fine- to medium-grained sand, clay, and lignite, and an upper unit (Y), which is composad of gravel and sand with a few 1- to 2-in. interbeds of clay. Uncompressed logs are distributed throughout the upper unit of the Beaufort Formation.

A piece of wood described as Pinus strobus (Heer 1868) is the first record of a plant fossil from the Beaufort Formation. Since then palynomorphs assignable to Picea, Tsuga, Erica- ceae (tetrads), and Larix, and bryophytic spores have been recorded (Hills and Ogilvie 1970); Terasmae (in Tozer and Thorsteinsson 1964) identified pollen of Corylus, Carpinus, and Populus. Hills (1969) recorded cones of Picea,

Larix, Pinus, and Alnus from the formation on northwestern Banks Island. Hills and Ogilvie (1970), on the basis of about 500 cones, described a new fossil spruce species (Picea banksii). Kuc and Hills (1971) reported on fossil mosses from these strata. Hills and Sweet (1971) described a fossil Juglans similar to J. cinerea from the Beaufort Formation of southwestern Banks Island.

The Beaufort Formation has been assigned to the Miocene (Heer 1868), whereas Tozer (1956) assigned a Tertiary or Quaternary age or both to it. Hills (in Hills and Ogilvie 1970), on the basis of palynologic similarities, correlated the Beaufort Formation with the Homerian-Clam- gulchian sequence of Alaska (Wolfe et al. 1966). These local stages were assigned, on the basis of paleobotany and palynology, a late Miocene - early Pliocene age. Recent work by Hopkins el al. (1971) indicates that the extant species Picea glauca was already in existence 5.7 million years ago (K-AR). If, as suspected by Hills and Ogilvie (1970), the extinct spruce P. banksii is an ancestor

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2638 CANADlAN JOURNAL OF BOTANY. VOL. 50, 1972

of the modern species then the Beaufort Forma- tion where this species occurs cannot be younger than early Pliocene.

The aim of our present study is to describe the anatomy and determine the natural affinities and paleoclimatic significance of fossil woods collected from the Beaufort Formation on northwestern Banks Island. Figure 42 indicates the geographic location of the collection sites.

All specimens are stored in the Paleontologic Collection, Geological Survey of Canada, 601 Booth Street, Ottawa, Ontario.

Description Coniferales

FAMILY Pinaceae GENUS Pinuxylon Gothan, 1905 emend.

Pinuxylon albicaliloides Greguss, 1954 Figs. 1-6 Topography Growth rings are distinct, wide, and straight

(Fig. 1). The transition from early to late wood is gradual (Fig. 2). No appreciable differences are seen in the shapes and sizes between the early and late wood tracheids. Late wood tracheids are two to three cells wide, slightly flattened and rectangular in cross section (Fig. 2). The tracheids in the early wood are circular, oval, or rectangular in shape, mostly thick- walled with only those near the growth boundary with slightly thinner walls and with wide lumina (Fig. 2). Early wood is 40 to 50 cells wide. Sparse, large, vertical resin ducts with six to seven thin- walled epithelial cells occur only in the early wood. The rays are separated by 1 to 10 rows of tracheids. Xylem parenchyma is absent.

The rays are heterogeneous (Figs. 4 and 6), 2 to 15 cells high, mostly consisting of parenchyma cells and a marginal row of thin-walled ray tracheids with small circular bordered pits. Rays are mostly uniseriate, those with horizontal resin ducts are fusiform, multiseriate in the middle with uniseriate extensions at both the ends (Fig. 3). Horizontal resin ducts with four or five thin-walled epithelial cells are present. Ray cells are oval or elliptical in cross section. Tangential walls of the vertical tracheids are always with closely placed, oblique, spiral checkings. No bordered pits were observed on the tangential walls of the tracheids.

Radial walls of tracheids always have spiral checkings similar to those of the tangential wall. Bordered pits are mostly uniseriate, occasionally

in part biseriate (Fig. 5), when biseriate they are opposite. Pits are more apparent on the early wood tracheids. Pit contours and pores are circular. Transverse tracheids are present only at the extremities of the rays, their horizontal walls are perfectly smooth, thin-walled and without any projections or dentations on their inner surface (Figs. 4 and 6). Cross-field pits are typically pinoid, oval or circular in shape; one or two in each field. Those in the late wood appear eye-shaped and obliquely disposed (Fig. 6). Horizontal walls of the ray cells are thin- walled and smooth (Fig. 4). Crassulae are occasionally present between the bordered pits.

Collections and Type Specitnen G.S.C.

G.S.C. specimen Type location No. No.

Voucher specimen C-12672 (Fig. 42) 27926 Voucher specimen C-12672 (Fig. 42) 27949 Voucher specimen C-3790 (Fig. 42) 27968

The Family Pinaceae consists of 10 genera in the extant flora (Pinus, Picea, Larix, Pseudotsuga, Cedrus, Tsuga, Abies, Keteleeria, Pseudolarix, and Catlzaya). Xylotomically the family exhibits a fairlv wide variation in characters. On the basis of the nature of rays and the presence and absence of normal horizontal and vertical resin ducts, they can be divided into four groups (Greguss 1955; Watari 1956): (1) those with heterogeneous rays having both horizontal and vertical resin ducts, e.g., Pinus, Picea, Larix, and Pseudouuga ; (2) rays heterogeneous without normal resin ducts (Cedrus and Tsliga); (3) rays homogeneous with only vertical resin ducts, e.g., ~eteleer ia ; and (4) rays homogeneous and resin ducts absent (Pseudolarix and Abies). Since the structure of the wood of Cathaya (Chun and Kuang 1958) is not well known, it is not possible here to refer it to any of the above four groups.

Because the fossil wood herein described has heterogeneous rays with both vertical and horizontal resin ducts, it is referable to either Pinus, Picea, Larix, or Pseudotsuga. Among these four, Pinus is the only genus with typically "pinoid" cross-field pitting in which the epithelial cells of the resin ducts are thin-walled (Greguss 1955; Phillips 1941). These two features are present in the fossil wood and therefore it

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ROY AND HILLS: FOSSIL W 'OOD FROM BANKS ISLAND 2639

undoubtedly belongs to Pinus. On the basis of the structure of the inner surface of the horizontal walls of the transverse tracheids, species of Pinus are classified into two types: (1) the haploxylon type, in which the horizontal walls of the transverse tracheids are smooth; (2) the diploxylon type, in which the horizontal walls of the transverse tracheids are dentate or reticulate (Greguss 1955). The presence of smooth horizontal walls on the transverse tracheids indicates that the fossil belongs to the haploxylon type. Within this type, the "strobus" group exhibits one or two "pinoid" cross-field pits disposed side by side. This is also a characteristic of the fossil (Fig. 4). Detailed comparison with species of the "strobus" group, e.g., Pinus albicaulis, P. koraiensis, P. lambertiatza, P. parviflara, and P. strobus, for the characters of the growth rings in cross-sectional view, height of the rays, number of epithelial cells in resin ducts, nature of pitting on the tracheid walls, and the presence of spiral checkings, shows that the fossil wood cannot be distinguished from the extant species Pinus albicaulis Engelm. However, Hills (1971) recorded the presence of cones comparable to those of P. strobus from these deposits and therefore affinities of this fossil with this species should not be overlooked.

Fossil woods resembling that of Pinus are referred to the genus Pinuxylon Gothan (1905). So far about 25 species have been described within this genus, from Europe, North America, and the Arctic (Krausel 1949; Greguss 1967).

Among these, the nearest comparable form is Pinuxylon albicauloides Greguss (1954) from the Miocene of Hungary and therefore the fossil wood is referred here to the same species.

Krausel (1949) gave the following diagnosis of the genus Pinuxylon Gothan 1905. "Hori- zontal tracheids often with teeth or serrations, cross field pits + Eipore, often very big. Hori- zontal and tangential walls of parenchymatous ray cells, smooth, vertical and horizontal normal resin ducts with thin-walled epithelial cells" (Krausel 1949).

This diagnosis does not include all the structural variability of Pinus wood. Specifically, it excludes those species of Pit~us which have horizontal tracheids with -imooth walls (haploxylon groups of Pinus species). Therefore, to make this generic name more meaningful an emended diagnosis is presented here.

Emended Diagnosis Pinu;iylon (Gothan 1905)-Growth rings dis-

tinct, transition from early to late wood abrupt or gradual. Vertical and horizontal resin ducts present; epithelial cells of the resin ducts thin- walled. Tangential and radial walls of tracheids smooth or with spiral checkings. Bordered pits on the radial walls of tracheids, generally uniseriate, occasionally biseriate and opposite. Crassulae present or absent. Tangential walls of tracheids with or without pits. Rays markedly heterogeneous, ray parenchyma thin- or thick- walled. Transverse tracheids fringing rays or in the main body. Inner face of the horizontal walls of transverse tracheids smooth or toothed or serrated or reticulate. Cross-field pits generally pinoid, fenestriform, or occasionally piceoid. Xylem parenchyma generally absent.

GENUS Piceoxylon Gothan, 1905 emend. Picenxylnn beaufortense sp. nov. Figs. 7-12

Topography Growth rings are distinct, of variable breadth,

and slightly wavy. Transition from early to late wood is abrupt. Late wood is sometimes wider than the early wood, whereas early wood tracheids are rectangular or square in shape, thin-walled, and with wide lumina (Fig. 7). The late wood tracheids are thick-walled, oval o r circular, and with narrow lumina. The vertical resin ducts are generally located in the late wood with six to eight thick-walled epithelial cells (Fig. 7). The rays are separated by two to eight rows of tracheids. Xylem parenchyma is absent.

Rays are markedly heterogeneous (Figs. 9 and 10); uniseriate, those with resin ducts are fusiform multiseriate around the resin ducts (Figs. 8 and 12). Horizontal resin ducts have thick- walled epithelial cells (Fig. 12). In late wood the tangential walls of vertical tracheids possess spiral striations or thickenings (Fig. 8). Bordered pits are present on the tangential walls of early wood tracheids (Fig. 12).

Radial walls of longitudinal tracheids always have spiral checkings similar to those of the tangential walls in late wood (Fig. 8). Bordered pits on the radial walls are uniseriate, occasionally biseriate in part in the wider tracheids; biseriate pits are opposite or subopposite, discrete, circular, never crowded, and angular (Fig. 11). Transverse tracheids are only in one row, generally at the end of the rays, their horizontal

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2640 CANADIAN JOURNAL OF BOTANY. VOL. 50, 1972

walls are dentate (Fig. 10). Horizontal walls of the ray cells are also smooth (Figs. 9 and 10). Cross-field pits are minute, piceoid, sometimes tending to be taxodioid, four or five per field, generally placed at the corners (Figs. 9 and 10). Tangential walls of the ray cells have beaded thickenings (Fig. 10). Crassulae present (Fig. 1 I).

AFFINITIES AND COMPARISONS

The presence of vertical and horizontal resin ducts in the normal wood, heterogeneous rays, piceoid cross-field pits, and thick-walled epithelial cells of the resin ducts show the wood to have characters of Picea, Larix, and Pseudotsuga (Greguss 1955; Phillips 1941 ; Tang 1933). In Pseudotsuga, closely placed spiral thickenings are generally present in the transverse tracheids, wood parenchyma occurs frequently, and the horizontal walls of transverse tracheids are wavy. These features are not present in the fossil; therefore, this genus is excluded from further consideration.

There are many overlapping xylotomical characters between Larix and Picea and it is often difficult to distinguish the wood of these two genera unless all features are well preserved. Greguss (1955) suggested that the diagnostic features for Picea are 8 to 10 epithelial cells lining a resin duct, the rare occurrence of paired pits on the radial walls of the tracheids, walls of transverse tracheids mostly dentate, and two to six generally piceoid cross-field pits per field. As opposed to this, Larix has 10 to 14 epithelial cells lining a resin duct, frequent paired pits and crassulae on the radial walls of the tracheids, horizontal walls of the transverse tracheids generally smooth and 2 to 6 (8-10) piceoid or taxodioid cross-field pits per field which are arranged in two horizontal rows. Taking all these characters into consideration, the fossil appears to belong to Picea.

Gothan (1905) erected the genus Piceoxylon, which circumscribes the characteristics of the wood of the extant genera Picea, Larix, and Pseudotsuga. Krausel (1919, 1949), Bannan and Fry (1957), and Prakash (1968) used this genus in the sense of Gothan (1905). Watari (1956) and Greguss (1967) believe that Picea can be separated from Larix on xylotomical characters. We, however, are of the view that although

Pseudotsuga can be generally distinguished from Picea and Larix on the basis of wood anatomy, the latter two genera are really difficult to distinguish on the characters of the wood alone. Therefore, the generic name Piceoxylon Gothan (1905) is used here in a more restricted sense to include the structural variability of the wood of Picea and Larix and accordingly the generic diagnosis of Piceoxylon Gothan is here emended.

The similarities in the xylotomical characters in the different species of Picea are of such limited magnitude that it is almost impossible to segregate them on this basis alone. If we consider, however, such features as the number of cross-field pits per field, their arrangement, the height of rays, etc. to be reliable criteria then the fossils can be assigned to Picea. If these criteria are taken into account then the fossil under consideration is similar to Picea glauca or Picea sitchensis. Neither of these two species, however, agrees totally in wood characters with the fossils. Hills and Ogilvie (1970) described spruce cones (Picea banksii) which closely resemble P. glauca from localities where the wood was collected. Although these cones were not attached to branches so that the anatomy of the wood could be checked it is probable that they belong to the same species.

~ u m e r o u s species of Piceoxylon Gothan (1905) have been described from Mesozoic and Tertiary formations of Europe, North America, Asia, and the Arctic (Krausel 1949; Schonfeld 1956; Bannan and Fry 1957; Shilkina 1958; and Prakash 1968). Watari (1956) described three fossil woods from Japan as Picea palaeo- nzaximowiekzii, Picea wakimizui, and Picea cf. jezoensis. All these species can be broadly placed into two groups: (1) those with spiral thickenings or checkings in the tracheid walls and (2) those without any thickenings or striations on the tracheid walls. The wood from the Beaufort Formation falls within the first group. Because of the presence of spiral thickenings or checkings in the late wood only and the absence of xylem parenchyma, the wood is comparable to Piceoxylon piceae Rossler 1937. However, in P. piceae the tracheids have spiral thickenings as opposed to spiral checkings in the wood described here. The present wood is distinct from all the species of Piceoxylon described so far and, therefore, it is described as new.

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ROY AND HILLS: FOSSIL WOOD FROM BANKS ISLAND 264 1

GENUS Piceoxylon Gothan, 1905 emend. Einenderl Diagnosis

Growth rings distinct, transition from early to late wood gradual or abrupt. Vertical and horizontal resin ducts present in normal wood; epithelial cells of the resin ducts thick-walled. Tangential and radial walls of vertical tracheids with spiral thickenings or checkings or smooth. Bordered pits on the tangential walls present or absent, radial walls with uniseriate or biseriate and opposite bordered pits. Crassulae absent or present. Rays markedly heterogeneous, ray parenchyma thin-walled, walls of the transverse tracheids smooth or dentate. Cross-field pits generally piceoid, occasionally taxodioid. Xylem parenchyma generally absent; if present, restricted to growth boundaries.

Piceoxylon beaufortense sp. nov. Diagnosis Growth rings distinct; transition from early

to late wood abrupt. Width of the rings variable. Vertical resin ducts located in the late wood. Epithelial cells of resin ducts thick-walled and six to eight in number. Xylem parenchyma absent. Rays markedly heterogeneous, uniseriate, those with resin ducts fusiform, multiseriate around the resin ducts, 2 to 12 cells high. Spiral striations present in late wood tracheids. Bordered pits present in tangential and radial walls of early wood tracheids. Pits uniseriate, occasionally biseriate in part, when biseriate opposite or subopposite; circular and discrete. Horizontal walls of ray parenchyma smooth, those of ray tracheids dentate. Cross-field pits four or five per field, placed at the corners. Tangential walls of rays beaded. Crassulae present.

Collection and Type Specimen G.S.C.


Holotype C-12673 (Fig. 42) 27976 Paratype C-12673 (Fig. 42) 27939

GENUS Laricioxylon Greguss, 1967 Laricioxylon occidentaloides sp. nov. Figs. 13-1 9

Topography The growth rings are distinct and narrow (Figs.

13, 14). Transition from early to late wood is abrupt. Early wood zone is wider than late wood.

Early wood is four to six cells wide, tracheids are thin-walled, rectangular with round corners, their radial diameter is more than the tangential diameter. Lumina are wide. Late wood zone is narrow, three or four cells wide with thick- walled, irregular-shaped tracheids (Fig. 14). Tangential dimension greater than the radial dimension. Resin ducts are located only in the late wood zone (Figs. 13 and 14), epithelial cells of the resin ducts are thick-walled, 8 to 10 in number. Rays are separated by four to six rows of tracheids (Figs. 14, 15).

Rays are markedly heterogeneous (Figs. 18, 19), uniseriate, 2 to 10 cells high (Figs. 15, 16), whereas those with resin ducts are multiseriate in the middle and up to 20 cells high. Resin ducts generally have thick-walled epithelial cells, occasionally with tylosoids (Fig. 16). Ray cells in cross section oval or rectangular. Tangential walls of tracheids are without any pits or spiral thickenings or checkings. Radial walls of tracheids frequently have biseriate bordered pits with crassulae between them (Fig. 17) ; bordered pits are sometimes uniseriate, pits when biseriate are, as a rule, opposite, pores circular. Trans- verse tracheids are present in the body of the rays or at their extremities (Figs. 18, 19). Each tracheid has one to three small circular bordered pits (Figs. 18, 19). Horizontal walls of ray tracheids and ray parenchyma are smooth (Fig. 18). Cross-field pits are piceoid, mostly six or sometimes even eight in number per field, arranged in two horizontal rows in the field; aperture obliquely orientated, extends beyond the border. Xylem parenchyma absent.


The presence of horizontal and vertical resin ducts in the normal wood, markedly heterogeneous rays, piceoid cross-field pits, and thick- walled epithelial cells of the resin ducts relate the fossil wood to Picea, Larix, and Pseudotsuga. The absence of longitudinal parenchyma and spiral thickenings in longitudinal and transverse tracheids preclude its relationship with Pseudo- tsuga.

Picea and Larix are generally difficult to distinguish from each other on the basis of wood anatomy alone. However, some species of Larix have certain characteristics by which they can be segregated from species of Picea.

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The frequent presence of paired bordered pits with crassulae on the radial walls of the tracheids, typically piceoid cross-field pits arranged in two horizontal rows, and smooth-walled transverse tracheids seem to suggest a relationship with Larix (Beck 1945; Greguss 1955; Phillips 1941 ; Panshin and de Zeeuw 1968). Detailed comparison with the wood of extant species shows that the wood cannot be differentiated from Larix occide~ztalis Nutt.

Fossil woods resembling the wood of Larix have been described either as Piceoxylon Gothan 1905 (Krausel 1919; Bannan and Fry 1957) or Laricioxylon Greguss (1967). Among the species of Piceoxylon which are supposed to be related to Larix are Piceoxylon arcticum (Schmalhausen) Krausel (1949) and P. laricinoides Hareg (1932). According to the current practice of determina- tion, this fossil wood could also be placed in the more inclusive genus Piceoxylon Gothan (1905). Since the available anatomical characters of the fossil are most like those of the extant Larix wood, it is placed in the genus Laricioxylon Greguss (1 967).

The fossil wood under consideration resembles Laricioxylon nogradense, described by Greguss (1967) from the Miocene of Hungary, in number and arrangement of cross-field pitting and height of rays. In the Beaufort wood, however, the pattern of growth rings and the arrangement of the resin ducts are different. It differs from Piceoxylon arcticum Krausel(1949) in the absence of pitting on the tangential walls of the tracheids and in the frequent presence of two rows of bordered pits on the radial walls. Piceoxylon laricinoides (Hareg 1932) differs from the present fossil in the absence of horizontal resin ducts and in the presence of xylem parenchyma. Thus the fossil wood from the Beaufort Formation is different from previously described woods of Larix.

Lauicinxylon occidentaloides sp. nov. Spec& Diagnosis Growth rings distinct. Width of the rings

uniform. Transition from early to late wood abrupt. Resin ducts present only in the late wood. Epithelial cells of resin ducts thick- walled and 8 to 10 in number.

Rays heterogeneous, mostly uniseriate, 2 to 10 cells high; those with resin ducts fusiform, up

to 20 cells high. Tylosoids present in the resin ducts. Spiral thickenings or checkings absent. Frequent biseriate bordered pits with crassulae on the radial walls of tracheids. Pits opposite. Transverse tracheids present in the body of rays or at the extremities. Horizontal walls of ray tracheids and ray parenchyma smooth. Cross- field pits piceoid, four to six per field arranged in two horizontal rows. Pit apertures oblique. Xylem parenchyma absent.

Collections and Type Specimens

Type Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paraty pe

G.S.C. locality No.

C-12672 (Fig. 42) C-12671 (Fig. 42) C-3790 (Fig. 42) C-5552 (Fig. 42) C-12672 (Fig. 42) C-3790 (Fig. 42) C-12670 (Fig. 42) C-3790 (Fig. 42)

G.S.C. specimen

No.

27945 27940 27944 27945 27948 27972 27978 27981

GENUS Abietoxylon Houlbert, 19 10 Abietoxylon koueanoides sp. nov. Figs. 20-26

Topography Growth rings are distinct, consisting of 12 to

20, thick-walled cells which are rectangular in cross section with narrow lumina. Early wood tracheids are oval to square in shape, thin- walled, and with wide lumina. Rays are separated by three to eight rows of tracheids. Xylem parenchyma is scattered, consisting of solitary resiniferous cells near the growth ring boundaries (Fig. 20).

Rays are always homogeneous (Figs. 21 and 22); generally of low height, 2 to 8 (10) cells high and uniseriate. Ray cells in cross section are oval or elliptical, with sieve-like pitting on the tangential walls (Fig. 22). Tangential walls of the tracheids are smooth, without bordered pits. Radial walls of the tracheids have uniseriate, circular, discrete, bordered pits with a distinct torus (Fig. 25). Pits most common on walls of early wood tracheids. Horizontal walls of ray parenchyma distinctly pitted and thick- or thin-walled (Fig. 25). Tangential walls of ray parenchyma in radial section beaded or dentate (Figs. 25, 26), slightly oblique or vertical in

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ROY AND HILLS: FOSSIL W( 30D FROM BANKS ISLAND 2643

relation to the horizontal walls. Cross-field pits are elliptical taxodioid with oblique aper- Fres, or piceoid. Each field with two pits side by side or three or four pits placed at the corners. Borders of the pits are narrow and elliptical (Fig. 26).


The characteristic features of the fossil are the presence of distinct growth rings, xylem parenchyma, homogeneous, uniseriate rays composed of ray parenchyma with pitted horizontal walls and beaded or dentate tangential wall, sjeve-like tangential ray pitting and oblique taxodioid or piceoid cross-field pits. Such a combination of xylotomical features is found only in Abies, Keteleeria, and Pseudolarix (Greguss 1955) in the subfamily Abietoideae. Among these three genera, Keteleeria possesses vertical resin ducts in the normal wood (Watari 1956). Vertical resin ducts are absent in the fossil wood, thus precluding the possibility of its relationship to Keteleeria. Abies wood is distinguished from Pseudolarix on the basis of the presence of more parenchyma and calcium oxalate crystals in the former genus (Tang 1933, 1936; Kanehira 1926). Neither of these characters can be used to assign the fossil wood to either Abies or Pseudolarix because the calcium oxalate crystals are not likely to be preserved in fossils and the occurrence of xylem parenchyma has also been reported in Pseudolarix (Phillips 1941 ; Greguss 1955).

A detailed comparison of the fossil wood with species of Abies and Pseudolarix in the distribution of xylem parenchyma, height and width of rays, nature of pitting on the radial walls of the tracheids, the number of cross-field pits per field and their type, indicates that the fossil is closely comparable to Abies koreana Wilson. However, twigs of Pseudolarix have been found in the Beaufort Formation on northwestern Banks Island. Therefore, it is possible that this wood belongs to this genus rather than to Abies (Hills, unpublished data).

Fossil woods belonging to Abies have been described either as Abietoxylon Houlbert (1910), or as Abies. The fossil wood under consideration differs from Abiesfirmoides Watari (1956) and Abietoxylon faulense Houlbert (1910) in the

absence of partially biseriate rays and bordered pits on the tangential walls of the tracheids. Since the wood appears to be closely related to the extant species Abies koreana Wilson we have used a new specific epithet, koreanoides, to indicate this similarity.

Abietoxylon koreanoides sp. nov. Specific Diagnosis Growth rings distinct. Transition from early

to late wood gradual. Xylem parenchyma resiniferous, scattered near the growth rings. Rays homogeneous, uniseriate, and 2 to 8 (10) cells high. Sieve-like pitting present on the tangential walls of the rays. Bordered pits absent on the tangential walls of tracheids. Radial walls of tracheids with discrete, uniseriate bordered pits with distinct torus. Horizontal walls of rays pitted, tangential walls beaded or dentate. Cross-, field pits two to four per field; piceoid or oblique taxodioid. Pits placed side by side or at the corners. Pit borders elliptical with wide apertures.

Collections and Type Specimens G.S.C.


Holotype C-12671 (Fig. 42) 27938 Paratype C-12671 (Fig. 42) 27936

Abietoxylon trarrmatid~ctzis sp. nov. Figs. 27-32 Topography Growth rings are distinct (Fig. 27), 10 to 20

cells wide with a gradual transition from early to late wood (Fig. 28). Early wood tracheids are square to rectangular in shape, thin-walled, and with wide lumina. Late wood tracheids are thick- walled, four to six cells wide, with very narrow lumina. Xylem parenchyma occurs as resiniferous cells in some rings, whereas they are absent in others. Traumatic resin ducts and proliferated parenchyma cells occur in some growth rings (Fig. 27). Rays are separated by four to nine rows of tracheids.

Rays 2 to 12 (15) cells high, homogeneous, uniseriate, and composed of ray parenchyma. Tangential walls of the rays in tangential view have sieve-like pits (Fig. 31). Tangential walls of the early wood tracheids occasionally have spiral checkings (Fig. 31). Horizontal walls of xylem parenchyma are smooth.

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Radial walls of tracheids possess circular, discrete, uniseriate bordered pits (Fig. 30). The pits possess a distinct, unscalloped torus. Horizontal walls of rays are sparsely pitted (Figs. 30-32), whereas their tangential walls are always beaded or dentate. Cross-field pits in the early wood are mostly two per field, placed side by side, or up to four in the marginal cells and placed at the corners (Figs. 30-32). Pits are taxodioid or circular with obliquely oriented, wide apertures and narrow elliptical, indistinct margins.


It is obvious from the above description that the fossil under investigation is a member of Abietoideae with homogeneous rays and is closely related to Abies. It agrees, in general, with Abietoxylon koreanoides sp. nov., differing from it, however, in possessing traumatic vertical resin ducts, spiral checkings in the tracheid walls, fewer xylem parenchyma cells, and in the absence of any piceoid cross-field pits. Traumatic resin ducts occur in some of the Abies species (Pan- shin and de Zeeuw 1968). The new specific epithet traumatiductus is given because of the presence of traumatic resin ducts in the wood.

Abietoxylon traumatiductus sp. nov. Specific Diagnosis Growth rings distinct. Transition from early

to late wood gradual. Xylem parenchyma very rare. Traumatic resin ducts and proliferated parenchyma cells present. Rays homogeneous, composed of ray parenchyma; uniseriate, 2 to 12 (15) cells high. Sieve-like pitting present on the tangential walls of rays. Spiral checkings are present on early wood tracheids. Horizontal walls of rays very sparsely pitted whereas the tangential walls are beaded or dentate. Cross- field pits two to four, generally circular or taxodioid with oblique apertures.

Collection and Type Specirnerz G.S.C.


Holotype C-12672 (Fig. 42) 27953 60 ft above base Unit Y (Hills 1969)

Dicotyledoneae Rhamnales

FAMILY Elaeagnaceae GENUS Elaeagnaceoxylon gen. nov.

Elaeagnaceoxylon shepherdioides sp. nov. Figs. 33-41

Topography Wood distinctly ring-porous (Fig. 33). Tran-

sition from early to late wood vessels is abrupt. Late wood vessels are inconspicuous and small. Vessels in both the zones are almost solitary (Fig. 34). Early wood vessels are circular or angular in cross section and crowded near the growth ring boundaries, with a mean tangential diameter of 100 microns (p) or less. Vessels in the early wood are frequently plugged with a dark gummy substance (Figs. 33, 34). End walls of the vessels are oblique, without a tail-end (Figs. 37, 38). Perforation plates are always simple (Fig. 37). Intervessel pits are minute, alternate or subopposite, circular, or angular due to crowding (Figs. 38, 39). Spiral thickenings are present on the vessel walls (Fig. 38). Xylem parenchyma is very sparse and not readily discernible.

Rays are homogeneous (Kribs's type II), composed of rectangular or square procumbent cells (Fig. 41), up to 3 cells wide and 4 to 15 cells high. Uniseriate rays are few and far between (Fig. 35). Pitting in the procumbent cells is fairly common (Fig. 41).

Imperforate tracheary elements are generally tracheids or fiber tracheids (Figs. 35, 36, and 40). Tracheids have alternately arranged uni-, bi-, or tri-seriate bordered pits (Fig. 36). Pits are circular or slightly elongated in outline. Spiral thickenings are commonly present on the fiber tracheid walls (Fig. 40).


The important xylotomical characters which help in determining the affinity of the fossil are the distinct ring-porous character, tracheids and fiber tracheids with bordered pits, exclusively simple perforation plates, scarcity of xylem parenchyma, and the presence of spiral thickenings in the vessel. Such a combination of wood anatomical features is found only in the members of Anonaceae, Cistaceae, Elaeagnaceae, Eucommiaceae, Koeberliniaceae, Moraceae,

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ROY AND HILLS: FOSSIL WI

Polygalaceae, Rosaceae, Solanaceae, and Thy- meliaceae (Metcalfe and Chalk 1950; Greguss 1959). Among these, the spiral thickenings in the fiber tracheids occur only in Elaeagnaceae and Rosaceae (Record 1925). The exclusive presence of solitary pores in the spring and summer wood, heavy concentration of spring wood vessels near the growth rings, low height of the rays, and the occurrence of solid deposits in the vessels suggest that the fossil wood belongs to the Elaeagnaceae (Kanehira 1921 ; Yamabayashi 1939). Within this family Shep- herdia has rays up to 15 cells high and very sparse xylem parenchyma. It is our opinion, however, that some species of Elaeagnus cannot be distinguished on the basis of these two characters from Shepherdia. Moreover, some botanists (Nelson 1935) treat Elaeagnaceae as a monogeneric family, i.e. Elaeagnus.

Therefore the fossil wood under consideration is best referred to the family only.

Elaeagnus semiannulipora Watari (1952) from the Miocene of Japan is the only record of eleagnaceous wood known so far. The wood of E. semiar~nulipora is semi-ring-porous, the growth rings faintly indistinct. The vessels in the late wood are in groups and the rays are heterogeneous. In contrast to this, the fossil wood from the Beaufort Formation is distinctly ring- porous, the vessels are always solitary, and the rays are homogeneous. The wood is therefore given a new specific epithet, shepherdioides, to denote its relationship to the extant genus Shepherclia.

EZaeagnaceoxyZnn gen. nov.

Generic Diagnosis Wood distinctly ring-porous. Transition from

spring to summer wood abrupt. Vessels always solitary. Spiral thickenings present. Early wood vessels crowded near growth ring boundaries. End walls of vessels oblique. Perforations simple. Intervessel pitting alternate to subopposite, circular or angular. Xylem parenchyma very scanty. Rays homogeneous (Kribs's type TI), heterogeneous (Kribs's type I1 A), up to 20 cells high, one to nine are seriate. Imperforate tracheary elements with one to three seriate bordered pits and spiral thickenings.

Type species E. shepkerdioides.

OOD FROM BANKS ISLAND 2645

Elaeagnaeeoxylon shepherdioides sp. nov. Diagnosis Wood markedly ring-porous. Early wood

vessels concentrated near the growth ring boundary. End walls oblique, without a tail end. Vessels always solitary. Spiral thickenings present. Perforations simple. Intervessel pits alternate to subopposite, discrete, circular or crowded and angular. Xylem parenchyma not discernible. Rays homogeneous (Kribs's type 11), 1 to 3 cells wide and up to 15 cells high. Procumbent cells commonly pitted; rectangular to square. Im- perforate tracheary elements are fiber tracheids or tracheids, with bordered pits and spiral thickenings.

Collection and Type Specimens G.S.C.


Holotype C-12672 (Fig. 42) 27924

Discussion and Conclusions

The Beaufort Formation flora contains five genera and six species of fossil woods. These woods are referable to Pinaceae (Pinuxylon albicauloides Greguss (1 954), Piceoxylon beaufortense sp. nov., Laricioxylon occidentaloides sp. nov., and Abietoxylon trautnatiductus sp. nov.) and Elaeagnaceae (Elaeagnaceoxylon shepherdioides sp. nov.). A survey of the living equivalents of these woods indicates that all these genera now occur in the Canadian boreal forest. Since all but one (Abietoxj~lon traumatiductus sp. nov.) fossil species closely resemble living representatives, the assumption that their environmental requirements were essentially the same as their living counterparts seems valid. Hence in the discussion of the ecological re- lations a direct reference is made to the following modern equivalents.

Fossil species Modern equivalents

Pinuxylon albicauloicles Pinus albicaulis Greguss 1954 Engelmn. or

Pinus strobus L. Piceoxylon beaujortense Picea glnuca

sp. nov. (Moench) Voss. and Picea sitchensis

(Bong.) Carr.

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Laricioxylon Larix occi&nralis occirlrntaloides sp. Nutt. nov.

Abietoxylorz Abies koreona Wilson koreanoides sp. nov.

Elaeagnaceoxylon Shepherdia cana(lensis sl~epherdioides sp. Nutt. or nov. Elaeagnus canadensis

(Nutt.) Nelson

Pin~rs albicaulis is an alpine species restricted to the western Cordillera whereas P. strobus occurs in the Appalachian - Great Lakes region of North America and extends into the southern boreal forest in this same area (Harlow and Harrar 1958). Picea sitchensis is restricted to coastal areas of California to Alaska, whereas P. glauca ranges throughout the boreal forest. Shepherdia canadensis has a wide distribution in North America, occurring from Newfoundland to Alaska and south into South Dakota. In fact, in the modern flora of Canadian boreal forest these genera are common associates (Hosie 1969; Grimm 1966). The occurrence of fossil woods resembling these common boreal forest elements in the Beaufort Formation sug- gests that northwestern Banks Island was also covered by such a forest during the late Miocene or early Pliocene times. Similar conclusions were arrived at earlier by Hills (1969), Hills and Ogilvie (1970), Kuc and Hills (1971), and Hills (1971).

Pinuxylon albicauloides sp. nov., Laricioxylon occidentaloides sp. nov., Abietoxylon koreanoides sp. nov., and Elaeagnaceox~~lon shepherdioide~ sp. nov. cannot be distinguished from their modern equivalents on the basis of the xylotomical characters alone. It appears that the n~orphological features of woods of these species have not changed since the late Miocene or early Pliocene times.

Among these, Abies koreana Wilson, an alpine species, is confined to the volcanic island of Quelpaert in Korea, whereas Picea glauca, P. strobus, Larix occidentalis, and Shepherdia canadensis are present in at least some parts of the boreal forest of North America. Picea sitchetwis is found along the northwest coast of North America and Pinus albica~rlis is characteristic of alpine areas of the western Cordillera of North America. If we assume that the cones of Picea banksii and Pillus cf. P. strobus, although

not found attached to the wood, represent the same species as the fossil wood, we can ignore Picea sitchensis and Pinus albicaulus for paleo- ecologic interpretations and consider only the present distribution and ecology of Pinus strobus, Picea glauca, Larix occidentalis, and Shepherdia canadensis. All of these species are characteris$ of the Appalachian region of the United Stat+? (Harlow and Harrar 1958) and extend into the southern parts of the boreal forest of souther8 Ontario. Assuming that the environmental requirements of this flora were the same as they are today, it can be concluded that conditions on northern Banks Island in late Miocene or early Pliocene times were similar to those of the southern part of the boreal forest in Ontario today.

Based on this study it appears that members of the Pinaceae were the dominant elements of th: flora. Of the previously described (Chaney and Elias 1938; Dorf 1936, 1938; Carlton 19381 Axelrod 1938, 1950, 1956; Hollick 1930; Mart$ and Rouse 1966; Wolfe et al. 1966; Wolfe and Leopold 1967), late Tertiary deposits of Nort) America, the fossil assemblage most closery corresponds to the late Homerian or Clamghl- chian floras of Alaska (Wolfe et al. 1966; Wolre and Leopold 1967).

Acknowledgments

The authors express their sincere thanks to Dr. Wilson N. Stewart for critically going through the manuscript and giving useful sug- gestions. The research leading to this paper wa: supported financially by the National Research Council of Canada in a grant to W. N. Stewart, and by a Geological Survey of Canada Grant to L. V. Hills. The fossils herein described were collected by the junior author while in the em- ploy of the Division of Quaternary Research and Geomorphology, Geological Survey of Canada, Ottawa. The Geological Survey of Canada granted permission to publish this manuscript. Polar Continental Shelf provided field transport for the junior author.

AXELROD, D. I. 1938. A Pliocene flora from the Mount Eden beds, Southern California. Carnegie Inst. Wash. Publ., No. 476. pp. 125-183.

195C. Studies in the late Tertiary paleobotany. Carnegie Inst. Wash. Publ., No. 590. pp. 1-320.

1956. Mio-Pliocene floras from West-Central Nevada. Univ. Calif. Publ. Geol. Sci. 33: 1-322.

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ROY AND HILLS: FOSSIL WOOD FROM BANKS ISLAND 2647

BANNAN, M. W., and W. L. FRY. 1957. Three Cretaceous woods from the Canadian Arctic. Can. J. Bot. 35: 327-337.

BECK, G. F. 1945. Tertiary coniferous woods of western North America. Northwest Sci. 19(3-4): 67-102.

CARLTON, C. 1938. The San Pablo flora of west central California. -.- Carnegie Inst. Wash. Publ., No. 476. pp. 21 1-268.

CHANEY, R. W., and M. K. ELIAS. 1938. Late Tertiary floras from the high plains. With a chapter on the Lower Pliocene vertebrate fossils from Ogallala Formation (Lavern zone) of Beaver County, Oklahoma. Carnegie Inst. Wash. Publ., No. 476. pp. 1-72.

CHUN, W.-Y., and K.-Z. KUANG. 1958. A new genus of Pinaceae. Catl~aya Chun et Kuang gen. nov., from the southern and western China. (Genus novum pina- cearum ex Sina australi et occidentali.) Bot. Zh. 43: 461-470.

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DORF, E. 1936. The late Tertiary flora from southwestern Idaho. Carnegie Inst. Wash. Publ., No. 476. pp. 73-124.

-- 1959. Climatic changes of the past and present. Mus. Palaeontol. Univ. Mich. 13(8): 181-210.

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GREGUSS, P. 1954. Ax ipolylarnoci alsomiocen famara- dvanyok. (Wood remains from the lower Miocene of Ipolytarnoc.) Foldt. Kozl. 84: 91-1 10.

1955. Identification of living gymnosperms on the basis of xylotomy. Akademiai Kiado, Budapest.

1959. Holzanatomie der Europaischen Laub- holzer und Straucher. Akademiai Kiado, Budapest.

1967. Fossil gymnosperm woods in Hungary. Akademiai Kiado, Budapest.

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HARLOW, W. M., and E. S. HARRAR. 1958. Textbook of dendrology. McGraw-Hill Book Co.

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HILLS. L. V. 1969. Beaufort Formation. Northwestern ~ a n k s Island, district Franklin. Geol. L r v . Can., Pap. 69-1. pp. 204-207.

1971. Paleoclimatic interpretation of the Beau- fort flora (Late Tertiary), Banks Island, Arctic Canada. Am. J. Bot. 58(5): 2. 469 (Abstr.).

HILLS, L. V., and A: R. SWEET. 1971: Juglanscf. J . cinerea from the Beaufort Formation (Tertiarv). South- western Banks ~s land, Arctic ~ a n a d a . ~ m : J. Bot. 58(5): 2. 470 (Abstr.).

HILLS, L. V., and R. T. OG~LVIE. 1970. Picea banksii n. sp. Beaufort Formation (Tertiary), northwestern Banks Island. Arctic Canada. Can. J. Bot. 48(3): 457-464.

HBEG, 0. A. 1932. The fossil wood from'the Tertiary at Myggbukta, East Greenland. Medd. Norg. Svalbard. Uncres., 14.

HOLL~CK, A. 1936. The Tertiary floras of Alaska. U.S. Geol. Surv. Prof. Pap. 182: 1-185.

HOPKINS, D. M., J. V. MATHEWS, J. A. WOLFE, and M. L. SILBERMAN. 1971. A Pliocene flora and fauna from the Bering Strait Region. Palaeogeogr., Palaeoclimatol., Palaeoecol. 9: 21 1-231.

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FIGS. 1-6. Pinuxjllon albica~rloirles Greguss 1954. All are from G.S.C. specimen No. 27968. Fig. 1. Cross section of the wood to show growth rings. 40 X. Slide 2796814. Fig. 2. Cross section illustrating the thin- walled epithelial cells of the resin ducts and the shape and size of the tracheids. 100 X. Slide 2796813. Fig. 3.

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Tangential section, showing the ray with a radial resin duct, spiral checkings on the tracheid walls and the height and width of the rays. 250 X . Slide 2796814. Fig. 4. Radial section showing the pinoid cross-field pits, horizontal tracheids with bordered pit and smooth horizontal wall of ray tracheids and ray parenchyma. 300 X. Slide 2796813. Fig. 5. Radial section. Note the biseriate, opposite, and uniseriate bordered pits on the tracheid wall. 300 X. Slide 2796813. Fig. 6. Radial section through the late wood showing obliquely disposed, eye-shaped cross-field pits and spiral checkings on the tracheid wall. 250 X . Slide 2796813.

FIGS. 7-12. Piceo.~yIon beaufortense sp. nov. Figs. 7-9, 11, holotype, G.S.C. specimen No. 27976; Figs. 10 and 12, G.S.C. specimen No. 27939. Fig. 7. Cross section of the wood showing the nature of growth rings, the shape and size of the early and late wood tracheids, and the position of the resin ducts. 100 X. Slide 2797612. Fig. 8. Tangential section, showing the horizontal resin ducts in the ray, height, width, and distribution of rays, and the spirals on the tracheids. 65 X . Slide 2797611. Fig. 9. Radial section to show the ray tracheids with bordered pits, piceoid cross-field pits, and smooth horizontal wall of ray parenchyma. 250 X. Slide 2797611. Fig. 10. Radial section, showing cross-field pits and slightly dentate horizontal wall of ray tracheid. 250 X. Slide 2793919. Fig. 11. Partially biseriate bordered pits on the radial wall of the tracheids. Note the distinct torus. 350 X. Slide 2797613. Fig. 12. Tangential section. Note the uniseriate bordered pits on tracheid wall and horizontal resin duct with thick-walled epithelial cells. 250 X. Slide 2793911.

FIGS. 13-19. Lnvicioxylon occidentnloides sp. nov. Figs. 13, 14, holotype, G.S.C. No. 27954; Fig. 16, paratype, G.S.C. specimen No. 27944; Figs. 17-19, paratype, G.S.C. specimen No. 27945. Fig. 13. Cross section in low magnification showing distinct spring and summer wood, variable width of the growth rings and the resin ducts in the late wood. 40 X . Slide 2795413. Fig. 14. Cross section further magnified to show thick-walled epithelial cells of the resin ducts and the shape of tracheids. 100 X. Slide 2795414. Fig. 15. Tangential section. Note the height and width of rays and the horizontal resin ducts in a ray. 65 X. Slide 2795412. Fig. 16. Tangential section showing an exceptionally large horizontal resin duct with tylosoids. 100 X. Slide 2794411. Fig. 17. Radial section showing biseriate bordered pits on tracheids. Note the crassulae and distinct torus in the pits. 350 X. Slide 2794512. Figs. 18, 19. Radial section. Note the ray tracheids with bordered pits in the main body of the rays, piceoid cross-field pits in two horizontal rows with oblique apertures and smooth horizontal wall of the ray tracheids and ray parenchyma. 350 X. Slide 2794512.

FIGS. 20-26. Abietoxylotl koreatloides sp. nov. Fig. 20, holotype, G.S.C. specimen No. 2793814; Figs. 21- 26, paratype, G.S.C. specimen No. 27936. Fig. 20. Cross section illustrating the nature of growth rings and the distribution of axial parenchyma. 40 X . Slide 2793614. Fig. 21. Tangential section showing the uniseriate rays of low height. 100 X . Slide 2793811. Fig. 22. Tangential section. Note the sieve-like pitting on the tangential wall of the ray parenchyma. 250 X . Slide 3793813. Fig. 23. Cross section more highly magnified. 100 X. Slide 2793811. Fig. 24. Radial section. Note the uniseriate bordered pits on the tracheid wall. 300 X. Slide 27938/1. Fig. 25. Radial section. Note the piciform cross-field pits with oblique apertures and homogeneous rays composed of ray parenchyma. 350 X . Slide 2793811. Fig. 26. Radial section more highly magnified showing details of cross-field pits. 570 X. Slide 2793811.

FIGS. 27-32. Abietoxylon traumatidrrct~rs sp. nov. Figs. 27-32, holotype, G.S.C. specimen No. 27953. Fig. 27. Cross section illustrating the growth rings and traumatic resin ducts. 40 X . Slide 2795312. Fig. 28. Cross section more highly magnified. Note sporadic occurrence of resiniferous parenchyma. 100 X. Slide 2795413. Fig. 29. Tangential section illustrating the distribution, height, and width of the rays. 100 X. Slide 2795312. Fig. 30. Radial section; note the taxodioid cross-field pits, homogeneous rays, and dentate tangential wall of the ray parenchyma. 570 X . Slide 2795313. Fig. 31. Tangential section illustrating sieve-like pitting on the tangential wall of the ray cells. 250 X . Slide 2795313. Fig. 32. Radial section. Note the cross- field pitting in the early and late wood zones, homogeneous nature of the rays, and beaded tangential wall of the rays. 300 X. Slide 2795313.

FIGS. 33-41. Elaeagnaceoxylon shepi~erdioides sp. nov. Figs. 33-41, holotype, G.S.C. specimen No. 27924. Fig. 33. Cross section illustrating the distinct ring-porous wood. 100 X. Slide 2792415. Fig. 34. Same, more highly magnified to show the gummy deposits and solitary crowded vessels in the early wood. 250 X. Slide 2792415. Fig. 35. Tangential section showing biseriate low rays. 200 X . Slide 2792415. Fig. 36. Tangential section showing bordered pits on the tracheid walls. 250 X . Slide 2792413. Fig. 37. Radial section. Note the short vessels with oblique end wall and simple perforations. 250 X. Slide 2792415. Fig. 38. Tangential section showing bordered pits and spiral thickenings on the vessel wall. 250 X . Slide 2792411. Fig. 39. Highly magnified view of the bordered pits to show their angular nature due to crowding and circular aperture. 570 X. Slide 2792413. Fig. 40. Spiral thickenings in the fiber tracheids. 250 X. Slide 2792413. Fig. 41. Radial section illustrating the procumbent cells. 250 X. Slide 2792416.

FIG. 42. Index map showing collecting localities: G.S.C. location C-5552, north side of small tributary on the west side of Ballast Brook, 7 mi south of M'Clure Strait (lat. 740211, long. 123"8') 230 ft stratigraphic above Ballast Brook, measured up tributary; G.S.C. location C-12670, north side of small tributary on west side of Ballast Brook, 7 mi south of M'Clure Strait (lat. 74'21f, long. 123"7'30") 120 ft stratigraphic above Ballast Brook; G.S.C. location C-12671, north side of Ballast Brook, about 8.5 mi south of M'Clure Strait (lat. 74'211, long. 123"6'), about 50 ft above Ballast Brook (Unit Y, Hills, 1969); G.S.C. location C- 12672, north side of Ballast Brook, 9 mi south-southeast of M'Clure Strait (lat. 74°19'30", long. 123"5'), 150- 160 ft stratigraphic above Ballast Brook; G.S.C. location C-3790, in prominent gulley on north side of Ballast Brook, 9.75 mi S-SE of M'Clure Strait (lat. 74'19', long. 23"4'). Collecting localities are on the ridge in- dicated by the arrow and at the head of the gully imniediately to the left of the arrow. G.S.C. location C- 12673, north side of the north fork of Ballast Brook, 7 mi south and 5.75 mi east of the mouth of Ballast Brook (lat. 74°20'30", long. 122"4S1). Permission to publish Air Photo A17654-64 was granted by the National Air Photo Library, Department of Energy, Mines and Resources.

NOTE: Figs. 1-42 follow.

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Documents

Fossil woods from the Beaufort Formation (Tertiary), northwestern Banks Island, Canada