Palynollogy of the Eocene Kitsilano Formation, southwest British Columbia

WILLIAMS. HOPMINS,JR. Zt~stituteofSedin7entar.y and Petroleltm Geology, Calgary, Alberta Received February 5, 1969

HOPKINS,W. S., JR. 19G9. Palynology of the Eocene Kitsilano Formation, southwest British Columbia. Can. J. Bot. 47: 1101-1131. Approximately 55 genera and 75 species of plant microfossils are illustrated and described from the Kitsilano Formation of soi~thwesternBritish Columbia. The general composition of the flora suggests a humid, warm temperate climate prevailed at time of deposition. Previous work on plant megafossils indicates a late Eocene to possibly early Oligocene age. Plant microfossils recovered in this study support a probable middle to late Eocene age for the Kitsilano Formation.

Introduction the basin is rimmed by hills and mountains This report presents the results of a Tertiary composed of older and diverse rock types. To palynological study of the Kitsilano Formation, the west the Whatcom basin is open to the southwestern British Columbia. This formation, Strait of Georgia depression but the structural probably an Eocene coastal deposit, produced and stratigraphic connections are only imper- a relatively large number of species of plant fectly understood. Further discussion of the microfossils. The work is based on analysis of general geology, plus a complete list of refer- about 40 shale and coal samples collected in ences, can be found in Hopkins (1966). The dis- a various Kitsilano outcrops and building excava- cussion of the geology of the Kitsilano and tions, as well as 14 stratigraphically located Burrard formations is based largely on the work samples from an artificial excavation known as of Johnston (1923); little additional geologic I work on either of these formations has been I the Highbury Tunnel. I done since Johnston's classic studies. I Acknowledgments- Burrard Formation Appreciation is expressed to Dr. Glenn E. The Burrard Formation was named by John- For personal use only. Rouse of the University of British Columbia, ston(l923)for a series of continental clastic rocks who supervised the Ph.D. thesis of which this that underlie the city of Vancouver and which is a part. Dr. Rouse also read this manuscript sporadically crop out along the south shore of and offered many useful suggestions. Thanks Burrard Inlet for a distance of approximately are also due to Dr. Richard Hedlund of the 15 mi. Several small outcrops occur on the Atlantic Richfield Company, who also read the north shore of Burrard Inlet along the lower manuscript at an early stage and offered numer- reaches of Capilano Canyon (Brothers Creek, ous suggestions and improvements. Text-figs. 2 and 3). The Burrard Formation is about 2000 ft thick, consisting of conglomerates, sandstones, Geology and shales and a few thin interbedded lignitic General Remarks seams. The base of the formation, at least to The Kitsilano Formation is the upper of two the north, rests unconformably upon the Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 continental sedimentary formations which granitic rocks of the Coast Range Mountains underlie the city of Vancouver, British Columbia. and is exposed only in the lower canyon of the Both formations, the lower Burrard and the Capilano River. Within the Whatcom basin, the overlying Kitsilano, dip gently (ca. 10') into the Burrard apparently rests on Cretaceous rocks Whatcom basin, a structural and topographic equivalent to the Nanaimo Group. This rela- low (Text-fig. 1). This basin is about 50 mi in tionship is not exposed but is documented in northwest-southeast diameter with its north several well sections and will be reported on in a margin along the Coast Mountains north of later paper. Beds higher up in the section are Vancouver and its south margin near Belling- exposed from Prospect Point (Stanley Park, ham, Washington. To the north, east, and south Text-fig. 2) more or less continuously along the 1102 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

west side of Stanley Park. The general attitude total of 1300 ft of strata is exposed along the of the formation appears to be a continuous west shore of Stanley Park; of this thickness, south dip of 10 to 15'. 1100 ft is sandstone and the rest shale. The base of the formation is marked by a 50- Johnston placed the top of the Burrard ft thick basal conglomerate that is composed Formation beneath a "basal conglomerate" of largely of subangular granodiorite boulders up the overlying Kitsilano Formation. This con- to 6 in. in diameter, accompanied by minor glomerate is not everywhere present and the amounts of diorite, greenstone, chert, quartzite, similar lithologies of the two formations make and schist boulders set in a sandy, ferruginous distinctions difficult. A real possibility exists matrix (Johnston 1923 ;Hughes 1946). The upper that this may simply be an interbedded con- portion of the Burrard Formation consists of a glomerate and may have no particular time coarse-grained feldspathic sandstone interbedded significance, i.e. a time break here may be with sandy shale. According to Johnston, a inconsequential (Rouse 1962). For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20

TEXT-FIG.1. Index map of southwestern British Columbia and northwestern Washington indicating area of interest. HOPKINS: EOCENE KITSILANO FORMATION 1103

A number of assemblages of fossil plants As suggested above, the evidence for a pro- have been collected and examined from the longed time break between the Burrard and Burrard Formation. J. W. Dawson (1895) Kitsilano Formations is highly uncertain at studied a collection of plant remains from the best. Indeed, even Johnston (1923) observed Stanley Park area along the south shore of ". . . since both formations are in large part Burrard Inlet and concluded the beds were composed of material deposited on land or in Eocene in age. E. W. Berry, after studying a shallow water, it is possible that the apparent plant collection made by Johnston from the break is due to contemporaneous erosion." same general site, reported that the rocks were Roddick (1965) no longer maintains the dis- middle or upper Eocene (reported in Johnston tinction between Kitsilano and Burrard forma- 1923). Berry (1926) added: "There can be no tions and considers them one unit. He adds that question of the Eocene age of these plants, . . . ." "Armstrong (report in preparation) indicates Rouse (1962), after a study of the plant micro- that the subdivision should be abandoned and fossils, concluded that the Burrard Formation a new name proposed for the assemblage." south of Burrard Inlet is middle Eocene in age. For the purpose of this report, I shall continue He reported further that the "Burrard Forma- to use the terms Burrard and Kitsilano forma- tion" of the North Shore in Capilano Canyon tions in the older sense of distinct rock units and (Brothers Creek) was considerably older, pos- in the sense they were defined by Johnston. Be- sibly Cretaceous and equivalent to part of the cause the Kitsilano dips to the south and dis- Upper Nanaimo Group (Text-fig. 3). The nature appears beneath a mantle of Pleistocene out- of the contact between the North Shore beds and wash and till, the top of the formation is not the Burrard Formation to the south of the inlet visible. From several lines of evidence, Johnston is unknown because it lies below the waters of considered a conservative estimate for the Burrard Inlet. Crickmay and Pocock (1963), thickness of the Kitsilano Formation to be using palynological techniques, suggested that 1500 ft. However, more recent data acquired the Burrard Formation was Upper Cretaceous from excavation of sewer and other tunnels, and correlated it with the plant-bearing portions as well as from water wells, suggests that a of the Nanaimo Group on Vancouver Island. minimum thickness of 2500 ft is present. But According to Johnston (1923) the "Burrard because outcrops are limited and because we Formation was deposited mostly in shallow are still uncertain about the validity of the For personal use only. water and in part sub-aerially on an alluvial Kitsilano-Burrard contact, the actual thickness plain under humid-warm climatic conditions and is still unknown. nearly at sea level. It is not a true delta deposit, The so-called Kitsilano basal conglomerate at least in the landward part, but may pass into contains pebbles and boulders ranging from 1 delta deposits." He suggests the possibility that to 10 in. in diameter. Near Second Narrows, all of the Georgia depression (the Strait of Johnston described imbricated gravels, suggesting Georgia, to the west, and now below sea level), deposition by a westerly flowing river. The rock was an alluvial plain and that the sea did not fragments are mostly granitic and apparently extend into the region. Because some 2000 ft of derived from the Coast Mountains. Also present sedimentary rock are present in the Burrard are schistose rocks and pebbles of shale, the Formation, subsidence probably was active latter presumably derived from the Burrard during the period of deposition. Formation. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Kitsilano Formation Sandstones and shales make up most of the The Kitsilano is a rather inadequately de- middle and upper parts of the Kitsilano Forma- limited unit overlying the Burrard Formation, tion. The sandstones are coarse-grained, fre- and underlying the Pleistocene sediments on quently cross-bedded on a small scale, and which the city of Vancouver is built. It crops occasionally contain thin lenses and irregular out along Kitsilano Beach, at various places masses of lignitic coals. The sand grains are along the south shore of Burrard Inlet, and generally angular, contain an abundance of east to Burnaby Mountain. Building excava- heavy minerals such as biotite, chlorite, horn- tions within the northern part of the city blende, olivine, garnet, and sphene (Thomson frequently penetrate the Kitsilano (Text-fig. 2). 1958). Thomson also added that the fresh, 1104 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

angular nature of the sand grains "indicates The Kitsilano Formation shows the same that transportation has not been great, and that attitude as the underlying Burrard Formation conditions of rock destruction were mechanical and at Kitsilano Beach dips to the south at 6 rather than chemical." The source rocks were to 9". Attitudes measured south of the Kitsilano apparently metamorphic and igneous, also sug- Beach outcrop, within artificially excavated gesting a source in the Coast Range to the tunnels are 9 to 10". To the south, for as far as north. direct measurements can be made, the southerly The shale is often blue-grey and usually sandy. dip is reasonably constant. What happens deeper Locally, both sands and shales contain remains in the Whatcom basin to the south, can only be of logs that have been altered to lignite. Stream inferred from the study of plant microfossil channelling, similar to that of the Burrard assemblages in wells and outcrops. The results Formation, is found throughout the middle and of the well investigations will be published in a upper Kitsilano. later report.

Vancouver

For personal use only. New Westminster

formation Contact (mostly concealed)

recent deposits Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20

Metamorphic and Igneous Rocks

TEXT-FIG.2. Geologic sketch map of Vancouver area showing location of Kitsilano Formation out- crops. HOPKINS: EOCENE KITSILANO FORMATION 1105

Plant microfossils and leaves are locally as Upper Eocene or Lower Oligocene." Some- abundant in the Kitsilano Formation. A col- what later (1926) Berry states "the conclusion lection of leaves from exposures at Kitsilano that the beds at Kitsilano are Eocene is strongly Beach were identified and discussed by E. W. indicated . . . Just what part of the Eocene is Berry. He concluded that "the general facies of perhaps not determinable at the present time. the Kitsilano plants is, in my judgement, I would regard it as Late Eocene . . . ." entirely Eocene and does not in the slightest Johnston, in referring to the origin of the degree suggest later Tertiary" (quoted by Kitsilano sediments, suggests they are similar to Johnston 1923). Berry, in personal communica- the Burrard, i.e. an alluvial plain deposit which tion to Johnston, goes on to say "there is no may pass into delta sediments westward and objection to considering the Burrard Inlet plants southward. Neither formation shows any evi- as Middle or Upper Eocene, and Kitsilano plants dence of marine deposition, nor are there any

NORTHWESTERN SOUTHWESTERN NORTHWESTERN

EPOCH WASHINGTON BRITISH COLUMBIA WASHINGTON

WELL SECTIONS OUTCROPS OUTCROPS For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20

UPPER NANAIMO GROUP (Vancouver Island) CRETACEOUS

TEXT-FIG.3. Stratigraphic section showing major rock u~iitsof Vancouver area. Ages and correlations are interpretations of the author based on available geologic Literature. Age revisions in the future are possible but relative stratigraphic positions are correct. 1106 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

marine fossils. Channel structures, cut and fill this unit was based on samples taken from structures, and imbricated gravels all imply an Kitsilano Beach, from transient exposures in alluvial origin. Even the shales must have been building excavations, and a series of samples deposited in shallow water because they con- taken from the north-south Highbury sewer tain abundant sand grains which probably could tunnel on Point Grey. This tunnel (Text-fig. 2), not have been carried far. which was driven from 5th and Highbury Streets due south to the north arm of the Fraser Age- River, penetrated about 1000 ft of Kitsilano General Comments m or mat ion dipping 8 to 10' due south. In the vicinity of 22nd and Highbury Streets, the tunnel The Tertiary was a period of pronounced climatic and topographic change, during which passed through the Kitsilano into the overlying Pleistocene outwash and till. floras evolved and altered their composition. Because of these changes, analysis of the floral There is little indication of a fundamental record should be, and often is, usable as an change in either age or environment within the rocks of the Kitsilano Formation. As a result, age-dating method. Unfortunately, however, the study of Tertiary I have considered the formation as a single unit. palynology in North America has not progressed The microfossil assemblage is very similar to that of the underlying Burrard as described by to the point where it can give absolute answers Rouse (1962). Some of the more important to the question of age. In fact, because of inter- pretive difficulties, some workers have assigned genera are Osmunda, Anemia, Cicatricosisporites, Lygodium, Pinus, Taxodium, Podocarpus, Salix, only Paleogene or Neogene ages to certain floras. The problem of the Pacific Coast Tertiary Alnus, Carpinus, Corylus, Castanea, Fagus, Quercus, Carya, Engelhardtia, Pterocarya, Ulmus- is particularly difficult because only enough has been done to be tantalizing. Many of the Zelkova, Tilia, Myrica, and Liliaceae. A variety of fungal spores are also widely represented. Alaskan and British Columbia floras have not been assigned absolute ages, but are themselves There are, however, several distinct and per- haps significant differences in the microfossil dated by floral similarities and differences. assemblages of the Burrard and Kitsilano For- Recently MacGinitie (1966), while discussing Eocene floras of the Middle Rocky Mountains, mations. Tsuga is not found in the Kitsilano but occurs in low frequency in the Burrard Forma-

For personal use only. commented on the significant vegetative varia- tion. Ilex is absent in the Burrard, but is com- tion present in Wyoming floras of essentially the same age. He added (p. 40) that "such monly present throughout the Kitsilano, becoming more abundant stratigraphically up- diverse vegetation types indicate that fossil floras cannot be used as accurate age indicators wards. In the stratigraphically youngest Kit- silano sample, which is a carbonaceous shale, on the basis of floral composition alone, unless Ilex reaches 27% of the total pollen count. a sequence of vegetation types is established for each separate sedimentation basin." Platycarya, while not common, is present in the Further complications of interpretation are upper half of the Kitsilano but was not reported added in that definite Paleocene, Oligocene, and from the Burrard. Finally, Pistillipollenites, Pliocene floras have not been reported in the which locally reaches several percent in the Burrard, is present only in the lower part of the North Pacific area. In short, the typical micro- fossil or leaf assemblages for these time inter- Kitsilano. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 vals have not been established. Eocene assem- The fact that Ilex is present suggests either a blages, however, are well documented from short time break between Burrard and Kitsilano various other parts of NorthAmerica,and appear times or the establishment of Ilex during the to have basically similar floral elements. early stages of Kitsilano deposition. Platycaiaya appears in the upper part of the Kitsilano, sug- Age of the Kitsilano gesting that the plant migrated into the region Rouse (1962) suggested that the Burrard during Kitsilano time. Rouse (1962 and personal Formation is most probably middle Eocene, communication) and Hills (1965) believe that which, if correct, indicates that the Kitsilano is Pistillipollenites is most representative of the middle Eocene or younger. Microfloral study of middle Eocene. However, Hedlund (1966) found HOPKINS: EOCENE KJTSILANO FORMATION 1107

Pistillipollenites in the Cenomanian of Oklahoma Paleoecological Interpretations and has since concluded (personal communica- General Considerations tion) that this genus is pre-Eocene in age. How- Ideally, plants are the most sensitive of the ever, in British Columbia it is absent in Cre- terrestrial ecological indicators. Animals can taceous rocks and has been found in radio- roam and move about if climatic conditions be- metrically dated middle Eocene rocks of come unfavorable, but plants are rooted to one Interior British Columbia. Pistillipollenites ap- spot and must tolerate the environment in pears to be an Eocene form of the North Pacific which they grow. Furthermore, the tolerance in coast. Because of the presence of Cicatricosis- their environmental requirements is less than porites and Platycarya I have concluded that the for most animals. Because the most critical Kitsilano is no younger than Eocene. The former stage in a plant's entire growth cycle is at ger- seems to have a stratigraphic range on the mination, a changing environment will allow Pacific coast which terminates in upper Eocene survival of the reproductive propagules only if or lower Oligocene, while the latter appears they fall in a favorable site. Because of this, a restricted to Eocene and older rocks through- changing climate can markedly alter the flora in out North America. The complete absence of a comparatively few years. As a consequence, exclusively Cretaceous forms suggests the Kit- analysis of fossil floras should provide data silano is no older than Eocene, which is con- on climatic conditions at the time of growth. sistent with the upper Eocene age suggested by And indeed they do, but several problems Berry (1926). loom large. In palynology, as in all paleontology, Geologic considerations also bear on the a generally accepted truism is the old saw, "the age of the Kitsilano. The lithologies of the present is the key to the past." In paleoecologic Burrard and Kitsilano are similar, and both interpretations it must be assumed that orga- units apparently were formed under the same nisms, whether plant or animal, reacted to a depositional environment. No clear-cut for- given environment in much the same way as do mational contact appears between them except their modern counterparts. In other words, an for a discontinuous conglomerate which has alder and an elm would have had the same been assumed to be a basal conglomerate. No ecologic requirement in the Miocene as they do real evidence has been presented to show that today. The practical problem with this assump- this is anything more than a local conglomeratic tion is that ecologists are not really familiar with For personal use only. lens, without regional or temporal significance. the complete ecological requirements and the The lack of a conspicuous break between the range of tolerance of most genera and species of formations; the lithologic, textural, and struc- plants. This is true with temperate species, and is tural similarities; and the floral similarities lead even more so with tropical and subtropical me to conclude that the Kitsilano is not really species. Furthermore, within any given genus the a distinct unit differentiated from the Burrard, range of variability may be (and usually is) high, but that both represent a continued period of with each species requiring slightly different con- locally discontinuous deposition. Perhaps local ditions. However, in virtually all studies where disastems occur within the Burrard-Kitsilano, plant microfossils are used to interpret paleo- but they are not of sufficient magnitude to be ecology, palynologists are not dealing with called unconformities. If this is so, the forma- natural species but only with natural genera. The tional names Burrard and Kitsilano should be species of microfossils or other plant parts are

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 discarded and an all-encompassing name applied artificial, at least in rocks older than Pliocene, to both. and are based on various morphological dif- In summary the Burrard Formation has been ferences which may or may not have significance established as probably middle Eocene age in reflecting natural (or phylogenetic) relation- (Berry 1926; Rouse 1962); the Kitsilano Forma- ships. In any event, in very few cases can they tion contains a microflora which appears to be be equated to modern species. It is usually best characteristically coastal middle to upper to take the total range of variables within a Eocene; and the two formatioils are probably genus, and use as many genera as possible to conformable. Therefore, I have concluded that interpret paleoecology. Hopefully, a large micro- the Kitsilano is middle to late Eocene in age. fossil assemblage will give a qualitative estimate CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

TABLE I Range and ecological requirements of modern genera whose fossil counterparts have been identified from the Kitsilano Formation (modified after Rouse (1962), Hills (1965) with additions from Bailey (1949), Lawrence (1951), Graham (1965), Smiley (1966), and Willis (1966)

Genus Habitat and geographic range Climate Isoetes Moist areas to aquatic, cosmopolitan Temperate to tropical Lycopodi~im Most are mesophytic, cosmopolitan Temperate to tropical Osmitnda Swamps, shaded moist woodland, mainly Temperate to tropical Northern Hemisphere Anemia Wet lowlands and rain forests; particularly Subtropical to tropical in tropical America Lygodium Swamps, shaded moist areas, Southern Subtropical to tropical Hemisphere Swamps, shaded moist areas, mainly Tropical to subtropical Southern Hemisphere Quiet lake and pond waters Warm temperate to tropical Ginkgo Open stands, well-drained areas, west China Warm temperate, 4&60 in. precipitation Cedriis Dense forests, northern Africa, southern Asia Temperate to subtropical Keteleeria Dense forests, China Warm temperate Larix Marshes to woodlands, mainly Northern Cool temperate to boreal Hemisphere Picea Moist soils, mainly Northern Hemisphere Cool temperate, generally high altitude Swamps to rocky highlands, predominantly Variable dry sites, Northern Hemisphere Associated with evergreen oak forest, Warm temperate to generally moist to swampy habitats, subtropical, 50-60 in. southeast China precipitation Metasequoia Wet ravines in mountains, central China Taxodiitr7i Swamps and flood plains of southeastern Warm temperate to United States and Mexico subtropical, 50-60 in. precipitation Podocarpits Moist woodlands and mountains of the Warm temperate Southern Hemisphere, Caribbean, and South America

For personal use only. Northern Hemisphere, mostly small trees in Temperate woodlands Magnoliaceae Trees and shrubs, some climbing, Warm temperate, cosmopolitan subtropical to tropical Salix Damp thickets, swamps, cool woods, Variable cosmopolitan Swamps, wet woods, stream margins, Variable cosmopolitan Uplands to bog and wooded swamp, Cool temperate Northern Hemisphere Carpinus Upland woodlands to coastal swamps, Cool temperate Northern Hemisphere Corylus Thickets, woodlands, Northern Hemisphere Temperate Castanea Dry woods, thickets, Northern Hemisphere Cool to warm temperate Fagus Forms homogeneous, Northern Hemisphere Temperate Quercrts Wide range of habitats, Northern Variable Hemisphere, mountains of the tropics

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Carya Variable habitats, China, southeast Asia, Cool temperate to eastern North America subtropical Woods and river terraces, southeastern Warm temperate to United States, South America, subtropical southeastern Europe, Asia Platycarya Japan and northern China Pterocarya Northern Hemisphere of old world Temperate Tilia Low slopes and along streams, Northern Temperate Hemisphere Acer Variable habitats, Northern Hemisphere Cool to warm temperate Aesclilus Woods, bottomlands and stream borders and Temperate thickets, Northern Hemisphere, and South America HOPKINS: EOCENE KITSILANO FORMATION

TABLE I (Conclrided)

Genus Habitat and geographic range Climate Ilex Bogs, moist depressions, cosmopolitan Warm temperate to subtropical Proteaceae Mostly xerophytic, restricted to Southern Tropical; most indicate Hemisphere long annual dry season Myrica Variable habitats, cosmopolitan Temperate to subtropical Sabal Lowlands, river bottoms, coastal plains, Subtropical to tropical southeastern United States, Caribbean, Columbia TYP~~ Marshes, along river banks, cosmopolitan, Temperate to tropical except south of equator in Africa Polarnogeton Herbaceous in streams and ponds, a few Cool temperate to along sea margin subtropical

of the climatic conditions at the site of deposition parative fragility may result in its absence while the oarticular flora was in existence. entirely from the pollen record. noth he; problem is whether a given sample is Factors such as these undoubtedly lead to truly representative of the extant flora at the complications of interpretation and must always time of deoosition. Several considerations must be borne in mind when arriving at ecological be borne in mind while evaluating the data. conclusions. However, if floras are taken as a Differential preservation is always a factor- whole, and if adequate consideration is given to some pollen grains survive bacterial and fungal the various modifying factors, a fairly reliable attack, oxidation, hydrolysis, and rock dia- analysis of surrounding vegetation is possible. genesis more readily than others. Certain pollen grains are thin-walled and fragile (i.e. Thuja Discussion of Kitsilano Paleoecology and Populus) and easily destroyed. Others are The Kitsilano microfossil assemblage is largely attacked by bacterial action (i.e. Acer) and may composed of genera whose modern components be underre~resented.Others. because of their are warm temperate to subtropical (see Table 1). exceptionally durable nature (i.e. fungal spores Most of the genera, such as Anemia, Taxodium, and many spores of the Pterophyta) may be Salix, Alnus, Ilex, Sabal, and Typha are now For personal use only. overrepresented. Furthermore, pollen and spores characteristic of low, moist, and poorly drained are produced in vastly different quantities by coastal areas. The presence of Azalla and different genera. For example, a 10-year-old Potamogeton, which presently inhabit ponds and branch system of beech has been estimated to (or) lakes, indicate the presence of standing produce 28 million pollen grains per year, water on this lowland. while an equivalent branch system of pine may Other microfossils which suggest a more up- produce 350 million grains (Faegri and Iverson land habitat are present. These include Pinus, 1964). This difference will obviously be reflected Picea, Podocarpus, Carya, Corylus, and Quercus. in the quantities of pollen grains obtained from How far these trees were growing from the site maceration of rock samples and used in slide of deposition is uncertain, but their low fre- counts. quency of occurrence would suggest it was some The method of pollen dispersal also reflects distance away. Pinus and Picea, whose pollen is Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 relative quantities. Wind pollinated species (i.e. produced in prolific amounts, are not common Pinus) which usually produce pollen in enormous microfossils in any of the Kitsilano rocks. The quantities will be abundant in the fossil record, few bladdered conifer grains that are present are whereas insect pollinated plants (i.e. Acer) pro- seldom well preserved, are often physically duce relatively few pollen grains. As a result, broken. and are almost inevitably corroded. The Acer may be underrepresented in a microfossil implication is that their habitat was a consider- spectrum and its importance in the assemblage able distance away, probably to the north, east, may be underrated. In the case of Acer, a pollen or south. Transport to the depositional site was grain which is also easily destroyed, the com- probably largely by streams, which accounts for bination of low relative productivity and com- the poor physical preservation. 1110 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

The lithologies, insofar as they are visible in but confusion results, and comparisons between the Kitsilano and Burrard Formations, as well microfossil assemblages become difficult. Be- as in other Eocene outcrops, bear out this sides, this leads to unnecessary and meaning- interpretation of a lowland environment. Sands less proliferation of specific names. For paly- and shales predominate, river and stream nomorphs which cannot be assigned to modern channels are common, and thin coal seams are genera I have used form generic names. At times, present, as are occasional cross-beds. Together I have suggested possible or probable botanical these characteristics indicate the existence of affiliations, but this is often only an opinion and lowlands and swamps. The picture that emerges in some cases cannot be substantiated. is similar to that proposed by Johnston (1923) A section in the taxonomy called "Frequency" of a low, swampy coastal plain, spotted with gives a rough measure of the abundance of the small lakes, ponds, and swamps, and traversed palynomorph. "Rare" indicates that the given by generally slow-moving, meandering streams palynomorph occurs as less than lyo of the flowing westward to their junction with the sea. total spore and pollen count, exclusive of fungal In summary, the Eocene in the area of the spores. "Occasional" indicates 1 to 5y0. "Com- Whatcom basin appears to have been warmer mon" indicates 6 to 10% while "Abundant" and more humid than at present, but probably refers to grain occurrence in excess of 10% of not truly subtropical. Highlands must have sur- the total count. rounded the basin of deposition, but relief was Because of the variation in application and far less than at present. Precipitation was definition of various textural and sculptural moderately high, probably 50 to 60 in. annually terms, I have followed closely the terminology and was more or less uniformly distributed suggested by Faegri and Iverson (1964). The throughout the year. plant taxonomic classification employed is the

1 system outlined in Scagel et al. (1965). I Palynology In the following section I have described I General Taxonomic Considerations microfossils representing approximately 55 gen- As every palynologist knows, the classifica- era and 75 species from the Kitsilano Formation, tion and description of plant microfossils is in whose occurrence ranges from abundant in a chaotic state with no uniform system acceptable nearly all samples to a single occurrence in one to all workers. There are, however, two broad sample. For personal use only. schemes in use, each with its advantages, i.e. a natural classification and a morphologic clas- Division EUMYCOTA sification. Class FUNGI IMPERFECTI In this work I have elected to use the system Form Genus Dyadosporonites Elsik 1968 which is commonly applied by many Tertiary Dyadosporonites sp. 1 palynologists, i.e. a combination of these two Fig. 1 broad systems. This involves identification to an DESCRIPTION: Spores bilocular, 35 to 74 p extant genus wherever possible, and in some long, elliptical, aperture characteristics question- cases assignment to a form species in that genus. able, but apical and presumably circular. Central However, I have generally been conservative in septum simple and 2-3 p thick. Wall psilate. my approach to subdividing a genus into species. REMARKS : Similar fungal spores have been It is often the practice in paleontology to sub- recorded in Upper Cretaceous, Eocene, and Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 divide or to split into ever smaller units, often Pliocene beds. on dubious grounds. If, within a given genus BOTANICAL AFFINITY : Probably spores of a there are two or more forms which are clear cut, rust. distinct, and easily recognized, a subdivision FREQUENCY : Rare. into form species is more useful in stratigraphic studies. If, however, as appears to be more Dyadosporonites sp. 2 often the case, subdivisions are based on end Figs. 2-3 products of evolution showing intergradations DESCRIPTION : Spores elliptical, biporate, and of form or on idefinite and vague criteria, little bicellular, size 55 to 75 p. Pores large, circular, HOPKINS: EOCENE KrrSILANO FORMATION 1111

slightly aspidate, some subwall thickening. Large four cells separated by three septa; furthermore, thickened septum through middle of grain, ap- no intermediate stages were recognized. Con- parently dividing grain into two cells, each sequently assignment to a separate species is with an apical pore. Wall thick (ca. 2 p) and tentative. heavy. FREQUENCY: Rare. REMARKS: This grain is referred to the genus Fungal Hyphae Dyadosporonites as redefined by Elsik (1968). Fig. 8 It differs from Dyadosporonites sp. 1 in having a DESCRIPTION: Segmented and germinating heavier and less well-defined septum as well as fungal hyphae, overall length 150 p, average larger and more distinct apical pores. thickness of hypha is 7 p. FREQUENCY : Rare. REMARKS: Although of no known diagnostic Form Genus Pleuricellaesporites Hammen ex value, this is a common fossil in the Kitsilano Clarke 1965 Formation. Of interest are the young budding hypha and widely spaced and perforate septa Pleuricellaesporites psilatus Clarke 1965 shown on Fig. 8. Figs. 4-6 FREQUENCY: Abundant. DESCRIPTION: Uniseriate fungal spores with individuals consisting of several to many cells. Division LYCOPODOPHYTA The width varies from 12 to 25 p and the length Order ISOETALES may be in excess of 100 p, depending upon the Family ISOETACEAE number of component cells. Length of in- Genus Zsoetes L. dividual cells 6 to 12 p. A 0.5 to 1.0 p diameter ?Zsoetes sp. aperture occurs in each septum. Thickenings on Figs. 9-10 one side of the septum appear as adjacent DESCRIPTION: Small (17 to 23 p) monolete triangles with the aperture occurring between microspore (pollen?). Characteristically the sul- them. Cell walls psilate. cate furrow extends about two-thirds length of REMARKS: These spores are identical in all grain, in some gaping slightly, in others tightly respects with those described by Clarke (1965). closed. Sculpture scabrate. They are particularly common in many early REMARKS: For personal use only. Assignment of this microfossil to Tertiary horizons of Arctic and Western Canada. Zsoetes remains highly tentative, but it closely They are also common in numerous outcrop resembles the illustration and descriptions given samples from Eocene and Oligocene rocks of by Erdtman (1943) for Zsoetes microspores. coastal Oregon and Washington. FREQUENCY: Rare. BOTANICAL AFFINITY : Spores of a fungus. FREQUENCY: Common. Order LYCOPODIALES Pleuricellaesporites sp. Family LYCOPODIACEAE Fig. 7 Genus Lycopodium L. DESCRIPTION: Spores 35 to 50 p long with a Lycopodium annotinioides (Krutzsch) 1ength:width ratio of about 2:1, multicellular, Martin and Rouse 1966 usually with three perforate cross walls. There is Fig. 11 a suggestion of a pore at either end, but ruptured DESCRIPTION: Trilete spore, 30 to 35 p, with Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 ends leave this unclear. Thickening on one side moderately distinct laesurae reaching from pole of the septum appears as two adjacent triangles. to equator. Subcircular in polar view; distal REMARKS: The pattern of thickened septa surface and equatorial area of proximal surface which appear triangular suggests strongly that covered with a reticulum of narrow muri and this particular fungal spore should be assigned wide lumina. Reticulum 4 to 6 p between muri; to the genus Pleuricellaesporites. Possibly this muri up to 2 p high. Exine thin. form is identical with P. psilatus but is more REMARKS: These specimens are identical with mature, developed by constriction of every those found by Martin and Rouse (1966) from third cell. However, these spores always have the Miocene or Pliocene Skonun Formation of 1112 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

the Queen Charlotte Islands, British Columbia. accounts for the wide variation in shape, rang- Their extremely thin exine makes them rather ing from prolate to spherical to irregular. inconspicuous and difficult to photograph. Observed size range is 42 to 60 p. FREQUENCY: Rare. REMARKS: These microfossils differ from 0. irregulites in the smaller and more densely Division PTEROPHYTA packed bacula. Order FILICALES BOTANICAL AFFINITY: This spore type appears Family OSMUNDACEAE almost identical with modern 0. claytonia. Genus Oa?zunda L. FREQUENCY:Common, but see "Remarks" Osrn~rnclaregalites Martin and Rouse 1966 under Osn7urzda sp. 2. Fig. 12 Ostnuncla sp. 2 DESCRIPTION: "Trilete spores, spherical in out- Figs. 18-20 line. Laesurae simple, usually gaping slightly, DESCRIPTION: Subspherical, echinate, trilete and ranging from 18 to 22 p in length. A thin spore, laesurae sharp and distinct, tapering to a but distinct margo borders the commissure. The point at the equator; wall thin, covered with wall is about 1 p thick and appears to be rigid, short spines less than 1 p high which are widely as little folding has been observed. The orna- spaced-over the entire surface. Size range 45 to mentation is characteristically rugulate with the 55 p. rugulae short and thick, and packed close to- REMARKS: This form differs from Osmunda gether. Individual rugulae range to 3 p in length sp. 1 in having much more widely spaced pro- but rarely exceed 0.5 p in height" (Martin and jections. Furthermore, the ornamentation ap- Rouse 1966). pears to be much more spiny than is char- FREQUENCY: Abundant. acteristic of the three preceding species. Oc- Osmunda irregulites Martin and Rouse 1966 casional blunt bacula are observed but on the Figs. 13-14 whole the ornamentation is echinate. Although four different species of Osmunda DESCRIPTION: "Trilete spores, sub-circular in are described here, these are in reality probably outline but usually folded or split open. Laesurae end products of a continuum. Attempts to distinct, with a thin margo, and measuring about count Osmunda were continuously frustrated by For personal use only. 17-22 p in length. The ornamentation consists uncertainty about which of the four groups of bacula 1.5-2.5 p long, 0.5-2.0 p wide, and particular specimens should be assigned. I feel spaced 1-3 p apart. The bacula are usually at this time it would be unwise to assume that straight, occasionally clavate. There are no these grains represent four different species. All rugulate thickenings subtending the bacula. The that can be said with certainty is that Osmunda most diagnostic feature is the complete irregu- was a common genus of fern throughout the larity of the width of the bacula; delicate, slender time interval represented by Kitsilano de- bacula are randomly mixed with stout-stump position. like bacula and with all grades in between. Size FREQUENCY: Occasional, but see above. range 41-57 p." (Martin and Rouse 1966). FREQUENCY: Rare, but see "Remarks'; under Family SCHIZAECEAE Osmunda sp. 2. Genus Anemia Swartz

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Osmunda sp. 1 Anemia poolensis Chandler 1955 Figs. 15-17 Figs. 21-23 DESCRIPTION: Trilete spore showing consider- DESCRIPTION: These trilete spores are identical able variation in shape. Trilete mark usually not in all respects with those described by Chandler. distinct, but laesurae extend to equator. Orna- Size range is 34 to 58 p with most approximately mentation varies slightly from definitely baculate 52 p. to weakly vermiculobaculate; bacula closely REMARKS : Kitsilano specimens are identical spaced, ranging from 1 to 3 p in height. The with those described by Rouse (1962) from the wall is comparatively thin, which probably Burrard Formation and by Hills (1965) from HOPKINS: EOCENE KITSILANO FORMATION 1113

the middle Eocene strata of interior British established species. Several specimens bear a Columbia. resemblance to G. triangula Stanley 1965; FREQUENCY:Abundant. however, for a discussion of fossil gleicheniace- Form genus Cicatricosisporites PotoniC ous spores, see Skarby (1964). and Gelletich FREQUENCY:Rare. Cicatricosisporites intersectus Rouse 1962 Family POLYPODIACEAE Figs. 24-27 Form genus Laevigatosporites (Ibrahim) REMARKS: Specimens from the Kitsilano Schopf, Wilson aild Bentall Formation are identical with those described by Rouse (1962) from the Burrard Formation. Laevigatosporites discordatus Thompson Cicatricosisporites intersectus has not been and Pflug 1953 found to date in the middle Eocene beds Figs. 31-32 from the interior of British Columbia. Appar- DESCRIPTION: Spores generally bean-shaped, ently it is an Eocene-?Oligocene species re- monolete, laesurae short, wall Iaevigate, al- stricted to coastal lowlands. This species appears though some appear to be nearly circular; size identical with C. dorogensis PotoniC and Gel- range 40 to 80 p. letich except that C. intersectus has ribbing over BOTANICAL AFFILIATION:Rouse (1962) states the entire proximal surface. "the closest affiliation appears to be with BOTANICAL AFFILIATION : Cicatrico~i~porites Dryopteris, particularly D. Iatifrons." a form genus with very definite schizaeceous FREQUENCY: Common to abundant. affinities, resembling certain species of Mohria and Anemia. Laevigatosporites albertensis Rouse 1962 Fig. 33 FREQUENCY: Abundant. DESCRIPTION : Kidney-shaped, monolete spores, Genus Lygodiunz Swartz with a weakly defined suture, always oc- Lygodium reticulosporites Rouse 1962 curring along the concave crest. Ornamenta- Fig. 28 tion weakly punctate. Size range 32 to 36 p. REMARKS:The grains of this species agree with BOTANICAL AFFILIATION : Rouse (1962) de- those described by Rouse (1962) with an ob- scribed this species from the Burrard Formation,

For personal use only. served size range of 55 to 70 p. suggesting that it is most likely related to BOTANICAL AFFILIATION: Rouse comments Dryopteris or Asplenites, both of which have (1962, p. 197) that "spores of this species re- been reported as leaves in the Burrard. semble those of the living Lygodium japoniczrm FREQUENCY: Rare. Swartz and L. kerstenii Kuhn very closely." Laevigatosporites ovatus Wilson and FREQUENCY: Rare, but present in all Kitsilano Webster 1946 samples. Not illustrated Family GLEICHENACEAE DESCRIPTION: Monolete, bean-shaped spores, Genus Gleichenia Smith length 33 to 37 p, smooth wall, simply monolete suture. Gleicheizia s p. BOTANICAL AFFINITY: See L. gracilis. Figs. 29-30 FREQUENCY: Rare.

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 DESCRIPTION: Sharply triangular, laevigate, trilete spores, 25 to 30 p in diameter. Rounded Laevigatosporites gracilis Wilson and to pointed corners, weakly to strongly concave Webster 1946 interradial areas. Wall thickens near proximal Not illustrated pole in interradial area, the thickenings extend DESCRIPTION : Bean-shaped monolete spores, toward the angles so that the trilete mark is en- laevigate, simple monolete suture on concave closed by thickenings. side, length 25 to 32 p. REMARKS: Very few specimens were found, and BOTANICAL AFFILIATION: Wilson and Webster these could not, with confidence, be placed in an (1946) remark that L. gracilis and L. ovatus 1114 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

probably belong to Thelypteris, Asplenium, Division GINKGOPHYTA Atlzyrium, Aspidium, or Blechnum. Order GINKGOALES FREQUENCY : Rare. Family GINKGOACEAE Genus Ginkgo L. Families DENNSTAEDTIACEAE-POLYPODIACEAE Figs. 34-39 ?Ginkgo sp. REMARKS: Martin and Rouse (1966) have dis- Figs. 41-43 cussed the difficulties in classifying the large DESCRIPTION: Pollen monosulcate, shape ob- reticulate-verrucate-warty monolete spores late to peroblate, sulcus broad, generally closed which are abundant in the Miocene and (or) in the middle and gaping at the ends. Sculpture Pliocene Skonun Formation. This spore pattern psilate to lightly scabrate. Length 28-34 p. occurs in several different fern genera in at REMARKS: Comparison with modern Ginkgo least two families, viz. Polypodiaceae and biloba strongly suggests affiliation with Ginkgo. Dennstaedtiaceae, and sufficient gradation of Similar grains have been reported as Cycado- ornamentation exists to defy generic identifica- pites Wodehouse 1933 and Cycadopites follicu- tion. PotoniC (1956) employed three form genera laris Wilson and Webster 1946. Various other to include these three spore types, i.e. Poly- names have been applied to similar monocolpate podiisporites, Polypodiites, and Verrucatospor- grains found in Mesozoic rocks. ites. However, the fossil spores encountered in FREQUENCY: Rare. the Skonun Formation could not be assigned Division CONIFEROPHYTA even to form genera. As a result, Martin and Order CONIFERALES Rouse elected to use "non-committal form Family PINACEAE designates" for the three main forms. Genus Cedrus Link In the present work I encountered identical problems of classifying these monolete spores- ?Cedrus sp. gradations from one morphological form to Fig. 44 another occur. As a result, I have used the DESCRIPTION: The two grains found from the system suggested by Martin and Rouse. Forms Kitsilano appear to be that of Cedrus; both are 1,2, and 3 are illustrated but not described. They about 65 p in diameter. The bladders are are identical in all respects with the descriptions moderately large and tend to enclose the body,

For personal use only. given by Martin and Rouse (1966, pp. 187-188). except at the leptoma, which slightly separates FREQUENCY: Rare to common. One, two, or the bladders. Sculpture of the body is finely three of the forms may occur in the same reticulate, whereas the bladders are somewhat sample. more coarsely reticulate. REMARKS: Only two grains of this form were Order SALVINIALES found. Preservation is only fair and many of Family SALVINIACEAE the finer details are not clear. As a result it is Genus Azolla Lamarck only tentatively assigned to Cedrus sp. on the basis of comparisons made with modern Cedrus Azolla sp. pollen. However, Dacrydium (Podocarpaceae) Fig. 40 is superficially similar in that it has a body en- DESCRIPTION: Only isolated anchor-shaped closed by bladders with a narrow cleft leptoma. glochidia of Azolla were found in this study. There have been very few reports of Cedrus Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 The massulae and microspores were not en- in North American Tertiary rocks, the most countered; the reason for their absence is not recent documented occurrence being in the known. The glochidia are about 20 p long, and Miocene/Pliocene of the Queen Charlotte 8 p across the barbs. Islands (Martin and Rouse 1966). REMARKS: AzoIla appears to be especially FREQUENCY : TWOgrains only found. common in Eocene rocks in the interior of British Columbia (Hills 1965) but has also been Genus Keteleeria Carr. reported from Upper Cretaceous and rocks of Keteleeria sp. other Tertiary epochs. Figs. 45-46 FREQUENCY : Rare. DESCRIPTION: Bisaccate grains, large, body HOPKINS: EOCENE KITSILANO FORMATION 11 15

oval, 85 by 65 p; bladders nearly subspherical, Genus Pinus L. 55 by 45 p. The body appears to be scabrate to Pitzus strobipites Wodehouse 1933 minutely punctate, cap thin. Bladders are Figs. 5 1-52 finely reticulate, attached discretely to body and REMARKS: Although preservation is invariably directed distally. Structure and dimensions of poor, there is little doubt that this is the same leptoma obscured by bladders. specimen described by Wodehouse (1933) from REMARKS:The character of this grain fits the Eocene Green River Formation. [t has also closely those of Keteleeria clclvidatzcl as illustrated been reported from the Eocene Burrard Forma- in Erdtman (1943) and Macko (1957). However, tion by Rouse (1962). Macko gives overall dimensions of up to 200 p FREQUENCY: Although more commoll than while Erdtman gives dimensions of 102 to 161 p. other saccate pollen grains P. strobipites is Because only a few of these grains were found, relatively rare. I have no satisfactory range of sizes and it may be the average size is higher. Pinus sp. haploxylon-type The only other grain which resembles this is Figs. 53-54 Pseudolarix, which is considerably smaller. DESCRIPTION:Bisaccate pollen grain with According to Wodehouse (1935) Pseudolarix bladders attached to lateral equatorial extrem- ranges from 51 to 53 p. ities of body, bladders larger than body, body At the present time Keteleericl is restricted to circular and finely reticulate; bladders moder- central China where it grows in damp lowland ately coarsely reticulate, becoming finer toward forests and lower belts of mountain forests up bladder roots. Leptoma broad, straight-mar- to 8000 ft above sea level. gined, and finely granular. Body ranges from 40 FREQUENCY : Rare. to 50 p, bladders slightly larger.

I REMARKS: Thompson and PRup (1953) refer 1 Genus Larix Miller this species to the form species Piryosporites I Larix plica~ipollenitesRouse 1962 microalatus, which, they say, belongs to the Figs. 47-48 Pinus hap10,xylon group of Rudolph (1935). FREQUENCY:Rare. REMARKS:These grains appear to be identical with those reported by Rouse (1962) from the Pinus sp.

For personal use only. Burrard Formation. However, a number are Fig. 55 smaller, with the observed size range for the DESCRIPTION: Bisaccate pollen grain with present specimens being 50 to 70 p. They bladders about equal in size or slightly smaller clearly agree in size with the L. plicatipollenites than body. The body is esseiltially spherical reported by Hills (1962) from the Princeton with exine coarsely granular proximally. Lightly Basin of interior British Columbia. Their small reticulate sculpture on both bladders and distal size suggests affiliation with Larix rather than portion of body. Cap thick. The bladder con- Pseudotsuga. Wodehousz (1935) states that nection at the proximal root is well defined, and Larix pollen ranges in size from 62.5 to 90.2 p, at the distal root the contact is sharp and dis- while Pseudotsuga lies within the comparatively tinct. Lepto~naindistinct, but smooth to fine narrow range of 90 to 100 p. granulate. Observed body diameter 50 to 55 p. FREQUENCY: Rare to common. BOTANICAL AFFILIATION: This pollen grain is

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 almost identical with that of the extant Pinus Genus Picea Dietrich strobus. Picea grandivescipites Wodehouse 1933 FREQUENCY: Rare to occasional. Figs. 49-50 Family CUPRESSACEAE REMARKS: Although poorly preserved, this Figs. 5&57 bisaccate pollen appears essentially identical DESCRIPTION : Pollen grains subspherical, al- with that described by Wodehouse (1933) and though in some cases deformed by folding. Rouse (1957, 1962). The observed range of Inaperturate, size range 20 to 40 p. Exine thin, body size is 25 to 80 p. transparent, invariably with surface folds, FREQUENCY: Rare. occasionally ruptured. Surface of exine covered 1116 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

with small flecks that are slightly more stained have been widespread in the late Mesozoic and than exine. No evidence of pores, colpae, or Tertiary of western North America. papillae. Irregular area of thinning is commonly FREQUENCY: Rare. present which may represent a leptoma. BOTANICAL AFFINITY:This pollen appears to Genus Metasequoia Miki ex Hu and Cheng be that of Juniperus. However, other genera of the Cupressaceae have similar pollen, and dif- Metasequoia papillapollenites Rouse 1962 ferentiation is difficult or impossible. As a Fig. 62 result, I have placed grains of this morphology REMARKS: The few grains of Metasequoia en- and size in the family Cupressaceae, recognizing countered appear to be identical with those de- that one or more genera may be represented. scribed by Rouse (1962) from the Burrard Forma- FREQUENCY: Common to abundant. tion and by Hills (1962) from the Eocene Family CUPRESSACEAE,TAXODIACEAE, Princeton coal fields. or PINACEAE FREQUENCY:Metasequoia is frequently dif- Figs. 58-59 ficult to distinguish from Taxodium and so the apparent frequency may be less than it is in DESCRIPTION: Usually subcircular, inapertur- reality. Definitely identifiable Metasequoia is ate, laevigate pollen grains, identical with those rare. of Larix, except smaller. Observed size range 22 to 42 p. BOTANICAL AFFINITY: These grains would be Genus Sciadopitys Siebold and Zuccarini assignable to Larix except for their small size. They also bear a marked resemblance in size ?Sciadopitys sp. and form to modern species of Thuja. At this Fig. 63 time it is not possible to classify them more DESCRIPTION : Pollen grains large (78 and 75 p), closely than probably affiliated to one or more spherical, covered with large warts, 3 to 6 p in of the above families. diameter, usually lower than broad. Although a FREQUENCY: Rare. distinct aperture is not visible, there is a distinct leptoma that is usually psilate to granulate, but Family TAXODIACEAE covered with warty projections in some speci- For personal use only. Genus Glyptostrobus Endlicher mens. Glyptostrobus vacuipites Wodehouse 1933 REMARKS: Although this grain closely re- Figs. 60-61 sembles the single extant species Sciadopitys DESCRIPTION: According to Wodehouse (1933) verticillatn, as figured and described by Erdtman "the case skins of pollen grains split into two (1934) and Van Campo (1951), it is consider- approximately equal halves. Exiile in life ably larger. S. verticillata, both in modern apparently stiff and under mechanical strain so reference slides and in the above literature, that, in separating, the two halves buckle with appears to have a size range of 28 to 44 p, with the formation of longitudinal folds. Outer sur- most being 35 to 40 p. Morphologically this face dotted with small flecks openly and irregu- grain is also very similar to S. serratzrs as de- larly spaced. Length of halves 37.6 p." scribed by Martin and Rouse (1966) but again is Martin and Rouse (1966) add that the "pollen much larger. S. serratus has an observed size Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 grains usually split into two valves, with the range of 29 to 41 p. wall folded parallel to the split edges. Size There is also a similarity to Verrucosisporites range 27-30 p, wall thickness about 0.5 p and obscurilaesuratus Pockock 1962. However, the the ornamentatioil decidedly scabrate." Kitsilano form never shows a trilete mark and is The specimens in this study correspond closely somewhat larger. to the above descriptions except that the length Although size is not always a critical criterion, of the split halves ranges from 30 to 33 p. the discrepancies mentioned here are enough to REMARKS: Although Glyptostrobus is now cast some doubt on the generic assignment. confined to southeastern China, it appears to FREQUENCY:Rare. HOPKINS: EOCENE KITSILANO FORMATION

Genus Taxodium Richard Division GNETOPHYTA Taxodium hiatipites Wodehouse 1-933 Order EPHEDRALES Fig. 64 Family GNETACEAE Genus Ephedra L. REMARKS: Some specimens appear to be identical with those described by Wodehouse ?Ephedra sp. (1933), Wilson and Webster (1946), Rouse Fig. 69 (1962), and Hills (1962). It is quite possible that DESCRIPTION:Large (80 p), elliptical pollen some grains are Metasequoia because of the grains. Exine thick, heavy, and provided with frequent difficulty in distinguishing the two broad, low, and irregular ridges that are sub- species except under conditions of exceptional parallel to the long axis of the grain. Ridges preservation. appear to bifurcate and anastomose in an FREQUENCY:Common. irregular pattern. No furrows or pores are apparent. Family PODOCARPACEAE REMARKS: It is with the greatest uncertainty Genus Podocarpus L'Heritier ex Persoon that I assign this grain to Ephedra, which, in Podocarpus sp. extant forms, ranges from 35 to 55 p in length. Figs. 65-66 It is much larger than modern members of this DESCRIPTION:Bisaccate grain, body circular, genus, but is similar in morphology to Ephedri- 28 to 33 p in diameter. Bladders large and pites Bolkhovitina 1953 as illustrated in PotoniC irregular. Body sculpture scabrate, bladders (1958). finely reticulate. FREQUENCY: Rare. REMARKS: Only a few grains referable to Division ANTHOPHYTA Podocarpus were found in this study and all are Class DICOTYLEDONAE poorly preserved. As a result, no attempt has Order MAGNOLIALES been made to assign them to particular species. Superficially these grains appear similar to Family MAGNOLIACEAE Podocarpidites microreticuloidatus Cookson. Fig. 70 Hills (1962, 1965) reported Podocarpus from the DESCRIPTION: Ellipsoidal, monocolpate grains

For personal use only. Eocene rocks of interior British Columbia. ranging from 46 to 56 p in polar length. A single Rouse (1962) found a single grain of Podocarpus furrow extends from end to end of the grain, from the middle Eocene Burrard Formation. usually gaping slightly at the ends. Margins of FREQUENCY: Rare. the furrow are closed and slightly wavy, and are not thickened along margins. Exine moderately Family TAXACEAE thick and slightly roughened. Genus Taxus L. REMARKS: Except for the roughened exine, ?Taxus sp. this pollen is identical with that of Liriodendron Figs. 67-68 psilopites reported by Wodehouse (1933) from DESCRIPTION : Spherical pollen grains, 18 to the Eocene Green River Formation. 21 p in diameter. In most specimens a leptoma FREQUENCY: Rare. appears to have ruptured to form a large Order ARALIALES

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 irregular opening. Exine psilate to faintly Family CORNACEAE scabrate. Genus Nyssa Gronovius ex L. BOTANICAL AFFINITY: Lack of definite and diagnostic features makes assignment of this ?Nyssa sp. form to Taxus somewhat tenuous. However, Fig. 71 morphologically it appears identical with grains DESCRIPTION: Large (45 to 50 p equatorial of the western yew, Taxus brevifolia. The diameter) tricolporate pollen grains. Outline Kitsilano specimens average about 5 p smaller essentially circular, colpae long and tapering, than extant pollen. becoming wider at the equator. Each colpus FREQUENCY: Rare. contains a deeply sunken, pronounced slightly 1118 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

elliptical pore. Exine finely but distinctly granu- four- and five-pored types, but are often associ- late. ated with them. Six-pored grains occasionally REMARKS: Morphologically these grains appear make up several percent of the total alder count to be affiliated with Nyssa but are generally in the Kitsilano. larger than extant pollen. Also the shoulders of the wall surrounding the colpae are sloping Genus Betulrr L. rather than right-angled as in extant pollen of Betula cf B. claripites Wodehouse 1933 Njssa. Figs. 84-85 FREQUENCY:Rare. REMARKS:Pollen grains assignable to Betula are very rare in rocks from this formation. The Order SALICALES several grains found seem to fit the description Family SALICACEAE of B. claripites as given by Wodehouse (1933). Genus Salix L. Betula pollen is a widespread and common Salix discoloripites Wodehouse 1933 element in Tertiary rocks throughout North Figs. 72-73 America. Among the numerous localities where REMARKS: The grains found in this investiga- it has been found are the Paleocene Fort Union tion are identical with those described by Formation, the Eocene Green River Forma- Wodehouse except for a more restricted size tion, the middle Eocene of interior British range of 15 to 19 p. Columbia, and the middle to late Miocene FREQUENCY:Rare. (?early Pliocene) Latah Formation. FREQUENCY: Rare. ?Snlix sp. Figs. 74-75 Genus Carpitzus Rafinesque DESCRIPTION:Tricolpate, mostly oblate with cf. Carpinus sp. colpae extending almost from pole to pole. Figs. 86-87 Furrows closed, with a slight internal marginal DESCRIPTION: A number of grains were found thickening. which suggest definite affinities with Carpirzus. Sculpture minutely reticulate to punctate, Generally they fit the description of Carpinus maintained right up to the margins of the colpae. ancipites Wodehouse 1933 but many are con- For personal use only. Reticulations about 1 p. Length of grain 25 to siderably smaller. The size range of the present 30 p, equatorial diameter 15 to 20 p. speci~nensare 20 to 37 p, whereas Wodehouse REMARKS: This grain is only tentatively reports a range of 27.4 to 44.5 p. The only assigned to Sulix because of the lack of con- modern Curpinus pollen available for reference clusive and diagnostic features. Tricolpate, was C. betul~rs,with a size averaging 35 p. reticulate pollen grains are abundant in various REMARKS:Carpinus was once much more families and their identification is always an widespread than it is at present, and appears to uncertain procedure. have been a common element in many of the FREQUENCY: Rare. North American Tertiary floras. FREQUENCY: Rare to occasional. Order FAGALES Family BETULACEAE Genus Corylus L. Genus Ab~usMiller

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Cor:ylus tripollenites Rouse 1962 Alnus verus (Potonik) Martin and Rouse 1966 Figs. 88-89 Figs. 76-83 REMARKS: Corylus encountered in this study is REMARKS: This genus and species include all morphologically identical with the forms de- the various pollen of alder described in the past scribed by Rouse, but tend to be smaller with a (see synonyms in Martin and Rouse 1966, pp. minimum size of 18 p and an average of about 196-197). Pore number can vary from three to 24 p. seven but four- and five-pored pollen is markedly Corylus has been reported locally from the more common. Burrard Formation and the Miocene or Pliocene FREQUENCY: Occasional to abundant. Six- Skonun Formation. pored pollen grains are never as common as the FREQUENCY : Rare to occasional. HOPKINS: EOCENE KITSILANO FORMATION 1119

Family FAGACEAE Genus Quercus L. Genus Castanea Miller Quercus explanata Anderson 1960 ?Crrstanea sp. Figs. 97-98 Figs. 90-91 REMARKS: This form is identical with that DESCRIPTION : Grains small (1 6-20 p), tri- described by Anderson from Paleocene rocks of colporate, with colpae extending from pole to New Mexico. pole. Small circular pore, situated in short FREQUENCY : Rare. transverse furrow, lies in the center of each Quercus sp. 1 colpus. The exine is psilate to very faintly Figs. 99-100 granular. DESCRIPTION:Prolate tricolpate pollen grains REMARKS: These specimens look much like with furrows extending nearly from pole to pole. pollen of the modern species Castanea dentaro Occasionally a weak margo flanks colpae. Pores but are slightly larger. However, pollen of the absent but the middle of the furrow sometimes genus Castrrnopsis looks much like that of gapes slightly. A tectate-collumnellate exine Crrstanea. In fact, pollen of Crrstanopsis chryso- suggests a reticulate pattern which appears phylla is virtually identical with that of Castanen slightly coarser at the poles. Polar axis varies dentata. For this reason assignment to Castanea from 33 to 40 p, width 23-30 p. is questionable. REMARKS : This grain resembles Querc~rslongi- Castanea-type pollen has been described from canalis Traverse 1955. See "Remarks" under numerous Tertiary rocks ranging in age from Quercus sp. 4. Eocene to Pliocene. Rouse (1957) and Couper (1960) have also reported Cmtanea-type pollen Quercus sp. 2 from Upper Cretaceous rocks. Figs. 101-102 FREQUENCY : Occasional. DESCRIPTION : Fairly large (29-35 p polar diameter), tricolpate grain, nearly circular in Genus Fagus L. polar and equatorial views. Colpae two-thirds Fagus grrrnulata Martin and Rouse 1966 to three-fourths length of grain, usually tightly Figs. 92-94 closed. No subexinous thickenings along colpae. Sculpture scabrate.

For personal use only. REMARKS: These specimens appear identical REMARKS : See "Remarks" under Quercus sp. 4. with those described by Martin and Rouse FREQUENCY : Rare. (1966) from the Miocene /Pliocene of the Queen Charlotte Islands. They go on to say that this Quercus sp. 3 species probably represents "either an extinct or Figs. 103-104 extant Asiatic species." DESCRIPTION:Tricolpate grain, 20 to 24 p FREQUENCY: Rare. long, prolate. Closed colpae extending from end Fagus sp. to end of grain. Weak subexinous thickenings Figs. 95-96 beneath colpae margins. Sculpture frequently scabrate. DESCRIPTION: Subspherical and tricolpate, 30 REMARKS: This grain, although somewhat to 34 p in diameter. Colpae one-half to two- smaller, superficially resembles Quercus vir- thirds total grain length. Conspicuous margo giniana Mill as described by Macko 1957. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 bordering colpae; small, slightly elliptical pores REMARKS : See "Remarks" under Quercus in center of each colpus. Exine finely scabrate, sp. 4. becoming much finer on the margo. FREQUENCY:Rare to common. REMARKS:This form differs from F. granulata mainly in having shorter colpae, but also appears Quercus sp. 4 to have consistently smaller pores. This probably Fig. 105 represents an undescribed species, but there is DESCRIPTION:Tricolpate grain with the same little doubt that it is Fagus, probably an extinct size and same morphology as Q~rercussp. 3. form. The sculpture, however, is psilate rather than FREQUENCY: Rare. scabrate. 1120 CANADIAN JOURNAL OF BOTANY. VOL. 47. 1969

REMARKS: During the Tertiary Quercus ap- most specimens there is a narrow and often in- pears to have been a major constituent of conspicuous collar, sculpture slightly scabrate. northern hemisphere floras, and now is a wide- On proximal hemisphere, between pores, there spread genus consisting of over 300 species. are three subcircular areas of exine thinning; a It seems probable that the pollen found in the broad indefinite triradiate mark is present on Kitsilano Formation represent more than four ~roximalsurface. species, but these would be impossible to sub- REMARKS: These forms appear very similar to divide meaningfully. The four main mor- those described by Simpson (1961) from Lower phological types found in the Kitsilano rocks Tertiary rocks of Scotland. See "Remarks" are illustrated but no attempt is made to assign under Engellzardtia sp. them to species. FREQUENCY: Rare. FREQUENCY: Rare, but see above. Engelhardtia sp. Figs. 112-113 Order JUGLANDALES Family JUGLANDACEAE DESCRIPTION: Pollen grains in this group are Genus Carya Nuttall similar to those of E. granulata but differ in two respects: (I) there is no area of thinning in the Carya juxtaporites (Wodehouse, 1933) interpolar areas and (2) the exine is laevigate to Rouse 1962 faintly scabrate. Figs. 106-109 REMARKS: This grain appears similar to DESCRIPTION: Circular to subtriangular tri- Engelhardtia corylipites Wodehouse 1933. porate grains, 24 to 36 p in diameter. Pores It is with some uncertainty that I assign the usually circular, 14 to 2 p in diameter, occasion- grains encountered here to two fossil species of ally slightly oval. Pores on one hemisphere, Engelhardtia. Modern Engelhardtia pollen can slightly offset from equator. Ornamentation be distinguished from Myrica and Corylus ranges widely from psilate to distinctly granulate. mainly on the basis of size, but it is only sup- REMARKS: The Kitsilano Formation speci- position that this relationship held throughout mens appear identical with the Hicoria juxta- the Tertiary. Because these forms are smaller porites Wodehouse 1933 and with Carya than typical Myrica, and because the pores are juxtaporites (Wodehouse 1933) Rouse 1962. The not set precisely on the equator, which is a

For personal use only. size range given by Rouse (23 to 29 p) is slightly characteristic of the Juglandaceae, I have con- less than that given above but in all other re- cluded that these grains are those of Engel- spects they appear identical. The average size of hardtia. the specimens encountered in this study is 30 p FREQUENCY: Rare. and the range in ornamentation is pronounced. Possibly more than one species is included Genus Platycarya Sieb. and Zucc. under this designation but the gradational nature Platycarya sp. of the ornamentation makes it difficult to draw Figs. 114-1 16 definite conclusions. Wodehouse (1933) in his DESCRIPTION: Small (14 to 17 p), triporate original description of this species included pollen grains. More or less distinctly triangular; psilate to granular forms and I have followed sharp, slit-like pores at each angle. Pores are re- the same procedure in this study. ported by several authors Wodehouse (1933) ; Carya is a common fossil, both as leaves and

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Thiergart (1940) to have a characteristic Corylus pollen, in North American Tertiary rocks. pattern. Pores are characteristically slightly off- FREQUENCY: Common. set from the equator in the Juglandaceous- Genus EngelIfardtia Leschen. ex B1. corr. B1. fashion. Exine is scabrate and marked by at least two crossed folds (splits?). Engelhardtia cf. E. granulata Simpson 1961 REMARKS: Although to my knowledge it is Figs. 110-1 11 not published, the unofficial feeling among DESCRIPTION: Pollen grains 22 to 26 p in Tertiary palynologists is that Platycarya is diameter, profile in polar view triarcuate; three restricted to Eocene rocks. However, as a pores, close to circular, slightly aspidate; on number of writers have mentioned, the dis- HOPKINS: EOCENE KITSILANO FORMATION 1121

tinction between Platycarya and Engelhardtia REMARKS: The closest species appears to be and certain members of the Betulaceae can at Juglans periporites Martin and Rouse 1966, times be difficult. I have not had access to but these have a more pronounced annulus. modern Platycarya or Engelhardtia pollen but This type of Juglans pollen from the Kitsilano have compared descriptions and illustrations and Formation is similar to that of the modern J. can see how difficulties of differentiation might nigra. occur. As a result of this potential confusion it FREQUENCY: Rare. seems possible that Platycarya might be more long ranging than commonly thought, simply Genus Pterocarya Kunth because it has not been identified. I have assigned these grains with some con- Pterocarya stellatus Martin and Rouse 1966 fidence because of their small size, their slit- Figs. 122-124 like pores, and characteristic crossfolds (splits?). DESCRIPTION: Polyporate grain with five to FREQUENCY: Rare. eight subcircular to circular pores, located at or near the equator. Pores vary from 5 to 7 p Genus Juglans L. in diameter, most grains having six pores. Juglans periporites Martin and Rouse 1966 Grains definitely angular with the number of Figs. 117-118 side dependent on the number of pores. Sub- DESCRIPTION: These forms are essentially exinous thickenings beneath the pores shallow. morphologically identical with those described Almost invariably the surface is creased with by Martin and Rouse. However, the size range one or more folds. Surface sculpture ranges can be expanded because forms from the from psilate to faintly scabrate. Size range 28 Kitsilano Formation range from 26 to 35 p. to 37 p. Martin and Rouse give an annulus diameter of REMARKS: NO obvious morphological differ- 8 to 10 p, which is somewhat higher than shown ence appears to exist between fossil and extant in their illustrations. The annulus of the present Pterocarya pollen. specimens ranges from 5 to 6 p with pore FREQUENCY: Infrequent to frequent. diameters, in different specimens, varying from 1.5 to 4 p. Order ?URTICALES REMARKS: Juglans pollen has been found in Family ?ULMACEAE For personal use only. numerous Tertiary floras, including the Eocene Form genus Monzipites Wodehouse Green River Formation (Wodehouse 1933) and Monzipites tenuipolis Anderson 1960 the Oligocene-Miocene of Alaska (Wolfe et al. Figs. 125-127 1966). Wodehouse (1933) states "Juglans is an REMARKS: These grains are identical with ancient group, with a history dating back to the those described by Anderson (1960); size range Upper Cretaceous. In the Tertiary is was repre- 14 to 17 p. sented by many different species and is a con- Stanley (1965) describes what appears to be spicuous feature of most of the Tertiary floras, M. tenuipolis from the Paleocene of South particularly those of the more northerly and Dakota, and calls it Engelhardtia n~icrofovealata. colder climates . . . ." He does not refer to Anderson's species, nor FREQUENCY: Rare to common. does he comment on polar exine thinning which Juglans sp. characterizes M. tenuipolus, but his illustrations Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Figs. 119-121 appear to show this thinning. DESCRIPTION: Polyporate pollen grain, 28 BOTANICAL AFFILIATION : Ecologically it is Un- to 35 p, subrounded. Number of pores highly fortunate that this grain cannot be assigned to variable but usually around 12. Most located the natural classification. It is locally very near the equator, the rest located on one hemi- abundant and apparently was a major consti- sphere. Pores generally small (1 to 2 p in diam- tutent of the Kitsilano flora. Possibly it belongs eter), circular to slightly elliptical, annulus to the family Juglandaceae; possibly related to weak or absent altogether. Sculpture is weakly Engelhardtia or Pterocarya. On the other hand scabrate. it may belong to the family Ulmaceae. Momipites 1122 CANADIAN JOURNAL OF BOTANY VOL. 47, 1969

shows a marked resemblance to the modern Order TILIALES genus Moinisia. Family TILIACEAE FREQUENCY:Occasional to abundant. Genus Tilia L. ?Tilia cf. T. vescipites Wodehouse 1933 Order URTICALES Figs. 131-133 Family ULMACEAE DESCRIPTION: Circular to rounded subtrian- Genus Ulmus L. or Genus Zelkova Spach. gular, 24 to 34 11, tricolporate, pores more or less Figs. 128-129 deeply sunk. Very little subexinous thickening DESCRIPTION: Square to subrectangular; 28 to around pores. This form is similar to T. vescipites 37 p in diameter. Four pores are the rule, although reticulation is somewhat coarser. although three and five have been observed; REMARKS:This particular species shows a they are somewhat elliptical, 3 to 4 p in length. variation in external form from the circular As a rule, pores are located on the angles of the (Fig. 131) to the triangular with broadly rounded grain, but as in modern Ulmus, they may be all angles (Figs. 132-133). The latter two forms on one hemisphere, adjacent to the equator. approach in morphology several genera of the Slight, subexinous thickenings do occur. Exine Bombacaceae, which with the Tiliaceae, com- characteristically laevigate with a rugulate to prise the order Tiliales. Differentiation of these reticulate pattern impressed into it. Wodehouse two families along with the Malvaceae has long (1935) has remarked that these undulations are been controversial. Erdtman (1952) expressed due to "internal thickenings." doubt about the validity of elevating Bom- REMARKS:Ulmus and Zelkova cannot be dis- bacaceae to family rank. Fuchs (1967) feels tinguished on the basis of pollen, even in sufficient evidence is now available to dis- modern pollen grains, so I have not attempted to tinguish these families. but additional work is separate them here. This form is quite different necessary before the present confusion can be from the U. granopollenites described by Rouse settled. from the Burrard Formation. It looks much FREQUENCY : Common to abundant. like the modern U. americana but according to Tilia sp. Wodehouse (1935), U. americana character- Fig. 134 istically has five pores. The Ulnzus described DESCRIPTION: Small (21 to 24 y), tricolporate

For personal use only. here is dominantly four-pored. FREQUENCY: Rare. grain, invariably subcircular. Pores circular and deeply sunken, forming pits. Subexinous thick- Genus Planera Gmel. ening around pores is considerable and always Planera sp. conspicuous. Exine is pitted to minutely reticu- Fig. 130 late. REMARKS: This grain differs from T. vescipites DESCRIPTION:Morphologically, Planera is in its smaller size, in the punctate rather than identical with Ulmus-Zelkova already described, reticulate sculpture, and in the proilounced but has the addition of arci or curved linear thickenings around the pores. thickenings reaching from pore to pore. A FREQUENCY: Rare. description of Planera given by Simpson (1961) fits these grains almost exactly. Wodehouse Family ?TILIACEAE (1935), commenting on the arci of Planera, Fig. 135 Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 states that arci are more than adequate to DESCRIPTION: Elliptical pollen grain, 22 to differentiate Plailera from Ulinus. 26 y, markedly pointed polar ends. Tricol- REMARKS: I feel there is little doubt that these porate, colpae extending from end to end of are truly Planera. The difference in these grains grain. Marked subexinous thickenings. Small to those referred to as Ulmus-Zellcova is striking circular pore located in short, shallow, trans- and consistent. Leaves of Planera have been verse furrow. Exine is fine granular with a sug- found in the Burrard Formation. gestion that alignment or ornamentation is FREQUENCY: Rare. parallel to long axis of grain. HOPKINS: EOCENE KITSILANO FORMATION 1123

REMARKS: Assignment to the Tiliaceae is PistillipoIlenites mcgregorii Rouse 1962 tentative, but the morphologic resemblance to Figs. 139-142 the modern Grewia is pronounced. See Erdtman REMARKS: This form is identical with that (1952, 1966). described by Rouse (1962). P. nwgregorii has FREQUENCY: Rare. been reported from rocks as old as Cenomanian (Hedlund 1966) and as young as Eocene (Rouse Order SAPINDALES 1962). R. W. Hedlund (personal communica- Family ACERACEAE tion) states that PistiIIipoIIenites occurs only in Genus Acer L. Paleocene and older rocks of the United States Acer sp. Gulf Coast. However, it has been found in rocks Figs. 136-137 of Eocene age in interior British Columbia DESCRIPTION: Grains predominantly oblate, which are well dated radiometrically and by 24 to 30 y long, 10 to 20 y in width. Colpae fossil mammal bones (Rouse and Mathews extend from pole to pole, usually closed; where 1961 ; Hills 1962, 1965). gaping slightly, the colpa floor is faintly gran- BOTANICAL AFFINITY : Rouse (1962 suggested) ular. No thickening of colpae margins. The affinity with the modern monotypic genus exine is always distinctly granulate; almost Rusbyanthus, a member of the family Gentian- always the granules are arranged in rows, aceae. giving a striate appearance. Frequently the FREQUENCY: Frequent in lower Kitsilano, striations are weakly developed. absent in upper Kitsilano. REMARKS: Wodehouse (1935) and Simpson (1961) both remark on the diagnostic value of Family AQUIFOLIACEAE the granular striations. Among modern forms Genus IIex L. only Acer negundo fails to show this striate IIex sp. appearance. As a consequence I have identified Figs. 143-148 as Acer only those grains which show this DESCRIPTION: Tricolpate pollen grains, sub- feature. Tricolpate grains are common in many prolate to prolate. Some grains show the colpae diverse groups in the Dicotyledonae, and I am distinctly, others with the colpae barely visible. convinced that many misidentifications are All specimens show a clavate sculptured ektexine.

For personal use only. found in the literature. As a result I have Clavae vary from 1.5 to 3.5 y in diameter, decided to err on the conservative side and markedly expanded, and rounded on distal ends. assign to Acer only those grains which show the Most specimeils have equal-sized clavae, others characteristic striations. This undoubtedly has have slight variations. No clavae present on resulted in Acer appearing in lesser percentage furrows. Size range (excluding ornaments) 25 to than it did originally. 37 p in polar length. FREQUENCY : Rare to occasional. REMARKS: At the present time there are 400 species of IIex (Willis 1966) and many of these Family HIPPOCASTANACEAE undoubtedly exhibit slight variation in pollen Genus Aesculus L. morphology. Presumably considerable multi- Aesculus sp. plicity of species was present in the Tertiary, Fig. 138 as several different forms of Zlex are reported from Tertiary floras. Differences in the pollen Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 DESCRIPTION: Prolate, tricolpate grain, 20 to 26 y in polar diameter. Colpae short, each con- of Iles in the Kitsilano Formation are slight taining a small elliptical pore in the center. and do not lend themselves to meaningful sub- Exine faintly scabrate. division. Considerable overlap of form exists, FREQUENCY: Only two grains found. and it is difficult and arbitrary to assign most of the grains to any particular species. Traverse Order ?CONTORTAE (1955) has created five fossil species, but here Family ?GENTIANACEAE this would have little value. Gradation and Form Genus Pistillipollenites Rouse overlap of morphological forms precludes 1124 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

meaningful assignment to species. As a result r REMARKS:This grain appears similar to have simply placed all forms in Ilex sp. and Proteacidites terrazus Rouse 1962 but is rather illustrated some of the minor variations present. poorly preserved. FREQUENCY: Common to abundant. FREQUENCY: Only one specimen found, prob- ably contamination from underlying Cretaceous. Order PROTEALES Family PROTEACEAE Order MYRICALES Form genus Proteacidites (Cookson) Couper Family MYRICACEAE Genus Myrica L. Proteacidites thalrnani Anderson 1960 Figs. 149-1 5 1 Myrica annulites Martin and Rouse 1966 DESCRIPTION:Triporate, triangular pollen Figs. 153-1 54 grain about 22 to 30 p in diameter. Rounded REMARKS:Triporate grains, 26 to 33 p in corners, slightly convex interradial areas. Pores diameter. Species from the Kitsilano appear at angles, somewhat variable in character, very similar to those found in the Skonun generally appear elliptical and notchlike. An- Formation of the Queen Charlotte Islands. nulus usually pronounced around pores. Exine Wodehouse (1933) has stated "Mj~ricais one of is rather coarsely reticulate near equator, be- the most abundant and widely distributed genera coming finer toward poles. of the Tertiary occurring in practically all of the REMARKS:Proteacidites is difficult to break floras of the epoch." It was also present in the into species because of the transitional nature Upper Cretaceous and is still widely distributed of the pollen grain morphology. However, the although greatly reduced in number of species. present specimens appear identical with P. FREQUENCY: Rare. thalnlanii Anderson 1960. Class MONOCOTYLEDONAE Available evidence indicates that Proteacidites Order LILIALES is restricted to Upper Cretaceous in New Family LILIACEAE Mexico (Anderson 1960). Stanley (1 965) found Form genus Liliacidites Couper Proteacidites retzrszrs Anderson (1960) restricted Liliacidites sp. to Upper Cretaceous in northwestern South Figs. 155-1 56 Dakota. Rouse (1962) states that two species, P. terrazzrs Rouse and P. n~arginusRouse 1962, DESCRIPTION: Monocolpate pollen grains, For personal use only. are restricted to Upper Cretaceous rocks in prolate to perprolate, 14 to 45 p in polar length. British Columbia. However, the stratigraphic Furrow extending to extremities of grain, usually range of Proteacidites in North America is still not gaping but well defined; some grains show a uncertain. I have seen them in various Eocene slight margo. Sculpture reticulate, becoming rocks of western Oregon and Washington with finer toward the furrow and end of grain. no especial evidence that they were reworked. Lumina irregular and angular in shape. However, they are seldom abundant in rocks REMARKS:These grains are morphologically younger than Cretaceous and the real possibility similar to those of Liliaciclites leei Anderson 1960 exists that they were reworked into the Kitsilano but many are 5 to 10 p shorter in polar diameter. rocks. They are also similar to Aponogeton but be- FREQUENCY: Rare. cause this is strictly an African genus it seems unlikely it was ever native to North America. Proteacidites sp. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 FREQUENCY: Rare. Fig. 152 PALMALES DESCRIPTION: Rather sharply triangular grain, Order abruptly rounded angles, straight to slightly Family PALMAE concave sides. Size about 28 p. Pores indistinct Genus Sabal Adams but appear notch-like, no particular thickening Sabal gratzopollenites Rouse 1962 of exine around pores. Sculpture is wart-like at Figs. 157-1 58 equator grading to reticulate and punctate at DESCRIPTION: "Pollen monocolpate, fusiform poles. in outline, coarsely granulate to weakly reticu- HOPKINS: EOCENE KITSILANO FORMATION 1125

late. The single colpae is long and narrow with species of Typha." However, detailed examina- weak margins. Size range 28 to 32 p." (Rouse tion of the reticulate ornamentation indicates 1962). that the muri of modern Sparganiurn pollen are REMARKS: Sabal granopollenites has been not always closed, while Potamogeton appears reported from Eocene Burrard Formation to always have closed muri. Furthermore, Tjyha (Rouse 1962), the middle Eocene Allenby has a much finer and more delicate reticulum Formation (Hills 1965), and the Paleocene(?) than either Sparganium or Potarnogeton and Chuckanut Formation (Griggs 1965). usually has a conspicuous germ pore. Because a FREQUENCY : Rare. germ pore is never present and the muri are always closed, T have assigned these grains to Order PANDANALES the genus Potamogeton. Family TYPHACEAE Potamogeton has been described from a Genus Typha L. number of Tertiary formations, including the Eocene Green River (Wodehouse 1933), the Typha sp. Figs. 159-1 60 middle Eocene Allenby Formation of interior British Columbia (Hills 1962), and the Miocene DESCRIPTTON:Pollen grains small (18 to 25 p), - lower Pliocene Latah Formation of the irregularly spheroidal. Single germ pore which Columbia Plateau (Chaney and Axelrod 1959). is often not distinct, but usually occurs as a FREQUENCY: Occasional. rather irregular hole. Exine is thin and covered with a fine, foam-like reticulation. REMARKS: These grains appear identical with Incertae sedis those of the extant Typha latifolia which grows Trilites solidus (PotoniC) Krutzsch 1959 I in marshes throughout temperate North Figs. 162-163 America, sometimes abundantly. DESCRIPTION:Moderate-sized (35 to 50 p) FREQUENCY: Rare. I trilete spore, subangular, rounded angles. Slightly concave to slightly convex interradial Order NAJADALES areas. Trilete mark distinct, extending from pole Family POTAMOGETONACEAE almost to equator. Ornamentation coarsely Genus Potamogeton L. warty rugulate. For personal use only. I Potamogeton llollickipites Wodehouse 1933 REMARKS: Krutzsch (1959) states this spore Fig. 161 occurs in the Eocene of Germany, but its overall stratigraphic range is unknown. DESCRIPTION:According to Wodehouse these FREQUENCY:TWO specimens only found. grains are "spheroidal, somewhat ellipsoidal or variously irregular, 16 to 27.4 p in diameter. Delroidospora sp. 1 Exine rather thin and conspicuously reticulate Fig. 164 with a coarse network of beaded ridges. Without DESCRIPTION:Small (22 to 28 p), trilete spore. pores or furrows or vestiges of them." Subtriangular, broadly rounded angles, straight REMARKS: The specimens here are similar to to slightly concave interradial areas. Trilete those described by Wodehouse from the Eocene mark weakly developed, extending about two- Green River Formation except that the upper thirds of distance toward equator. Exine very

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 size limit is 32 p. The Kitsilano specimens are. weakly and minutely punctate. identical with those described by Hills (1965) REMARKS:This spore is similar to Deltoido- from the Eocene Allenby Formation, Princeton spora microforma Rouse 1962, but these grains Basin, interior British Columbia. do not have the generally gaping laesurae which Wodehouse (1933) remarked: "It is not Rouse describes. Tn size and shape it is more like known for certain whether they (Potarnogeton D. rhytisrna Rouse 1962 but does not often have pollen) have a germ pore or not, the absence of the concave interradial areas which characterize which is the only character which distinguishes Rouse's illustrations. these grains from those of Sparganium and some FREQUENCY: Rare. 1126 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

Deltoidospora sp. 2 BOTANICAL AFFINITY: Probably fungal spore. Fig. 165 HOLOTYPE: In writers collection, slide 14-5- DESCRIPTION:Small (20 to 22 p) trilete spore. 65-3-1, Leitz ortholux coordinates 25.2 X 118.0. Subtriangular in shape, broadly rounded angles, FREQUENCY: Rare. straight to weakly concave interradial areas. Punctodiporites llarrisii Varma & Rawat 1963 Trilete mark weak, extends to spore equator. Figs. 171-172 Sculpture scabrate. REMARKS:Specimens from the Kitsilano For- REMARKS : This grain is similar to Deltoidospora taenia Rouse 1962, but is considerably smaller. mation are identical with those described by Rouse gives a size range for this species of 26 Varma and Rawat, with a size range of 50 to to 35 y. 65 y. FREQUENCY:Rare. BOTANICAL AFFILIATION:Probably a fungal or algal spore, although it bears resemblance to Monoporate A certain forms in the family Onagraceae, especi- Figs. 166-167 ally the diporate forms of Fuchsia. DESCRIPTION: Almost spherical spore, 16 to FREQUENCY : Rare. 17 p, with one very slightly aspidate pore. Exine about 3 y thick with no thickening about the Diporisporites (Hammen) Elsik 1968 pore. Exine psilate. Diporisporites sp. BOTANICAL AFFILIATION: Fungal spore. Fig. 173 FREQUENCY: Rare. DESCRIPTION : Elongate spore or pollen grain Monoporate B elliptical to fusiform in outline, 55 to 60 p in Fig. 168 length. Biporate, pores opposed, slightly aspi- DESCRIPTION: Spherical to oval, 17 to 20 p, date. Poral costae prominent with exine thinner monoporate spore. Pore very minute, sur- toward pores. Exine psilate. rounded by slight subexinous thickenings, non- BOTANICAL AFFINITY:Probably a fungal or aspidate. Exine psilate. algal spore. BOTANICAL AFFILIATION : Fungal spore. FREQUENCY: Rare. FREQUENCY: Rare. Diporate A

For personal use only. Monoporate C Fig. 174 Fig. 169 DESCRIPTION:Large (63 to 68 y), irregularly DESCRIPTION:Small (15 to 16 p), oval, mono- oval pollen(?) grain. Diporate, pores are small, porate spore. Pore located at one end, circular, circular, and surrounded by a narrow zone of showing slight thickening of the exine of pore subexinous thickening. Pores are always on the margins. Exine psilate. same surface, but not always in the same place. BOTANICAL AFFILIATION : Fungal spore. Wall is thin and always creased with a number FREQUENCY: Rare. of folds. Sculpture minutely scabrate. BOTANICAL AFFINITY:Unknown. Fus~ormisporitesmicrostriatrrs n. sp. Fig. 170 FREQUENCY:Rare in Kitsilano Formation, abundant in lignite layers of equivalent age in DIAGNOSIS: Oval spore, 42 to 49 p in length. northwest Washington. Grain divided into two cells by a septum in Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 middle of grain. Fine longitudinal ribs extend Diporate C from the poles to the equatorial septum (which Fig. 175 is very slightly constricted). Most of these ribs DESCRIPTION:Large (75 to 85 p), oval, dipo- terminate at the equator, although occasionally rate. Circular, slightly aspidate pore at each several may be continuous across it. The wall is end. Subexinous thickening around each pore. thick and appears granular. Grain divided into four cells by three moderately REMARKS: This form is placed in the genus thick but simple septa. No pores apparent in Fus$or?nisporite.s Rouse 1962 but has much septa. Walls psilate. finer ribs and grooves than F. crabbii Rouse BOTANICAL AFFINITY : Probably fungal spores. 1962. FREQUENCY: Rare. HOPKTNS: EOCENE KITSILANO FORMATION 1127

Tricolpate A Fungal Spore Figs. 176-1 77 Fig. 183 DESCRIPTION: Large (40 to 44 p), tricolpate DESCRIPTION: Uniseriate fungal spores with pollen grain. Colpae about three-fourths total individual spores consisting of several to many length of grain, somewhat wavy and tending to cells. The width varies from 12 to 25 p and the gape in middle. Exine coarsely reticulate, be- length may be in excess of 100 p, depending coming slightly coarser at poles and finer toward upon the number of component cells. Length the colpae. of individual cells 6 to 12 p. A minute, slit- BOTANICAL AFFINITY: Unknown, but I have like aperture occurs in each septum. seen this form in various Tertiary rocks of FREQUENCY:Occasional to abundant. western Washington. Asterinn sp. FREQUENCY: Rare. Fig. 184 Tricolpate B REMARKS:These structures, which range from Fig. 178 65 to 85 p in diameter, are identical with the DESCRIPTION: Prolate, tricolpate pollen grain, cylindrical plates of cells described by Dilchner 26 to 28 p in polar diameter. Colpae sharp and (1963). He found them in Eocene clay deposits tightly closed, extending about three-fourths of Tennessee and assigned them to the epiphyl- length of grain. Exine laevigate. lous fungi family Microthyraceae. Similar struc- FREQUENCY:TWO grains found. tures are illustrated but not described by Hills Tricolpate C (1965) from the Eocene of interior British Fig. 179 Columbia. Martin and Rouse (1966) found them I I in the Miocene-Pliocene of the Queen Charlotte I DESCRIPTION:Small (15 to 19 p, prolate, I tricolpate pollen grain. Colpae extending al- Islands. most from end to end of grain, always tightly Unidentified I closed and showing no subexinous thickening. Figs. 185-186 I Exine moderately thick, scabrate to minutely DESCRIPTION: Large (75 to 80 p), circular I granular. organic body of unknown origin. FREQUENCY:Rare. FREQUENCY:Rare.

For personal use only. I Tricolporate A I Fig. 180 ANDERSON,R. Y. 1960. Cretaceous-Tertiary palynology, eastern side of the San Juan basin, New Mexico. , DESCRIPTION: Tricolporate pollen grain, pro- N. Mex. Burea~~Mines Mineral Reso~~rces,Mem. 6. late, 23 to 28 p long. Three colpae extend almost AXELROD,D. I. 1949. Eocene and Oligocene formations in the Western Great Basin. Geol. Soc. Amer. Bull. from pole to pole. Circular protruding pore in 60: 1935-1936A. center of each colpae. Weak subexinous thicken- 1950. Studies in Late Tertiary paleobotany. Carnegie Inst. Wash. Publ. 590. ings along margins of colpae. Exine scabrate. BAILEY,L. H. 1949. Manual of cultivated plants. 9th ed. FREQUENCY: Rare. The Macmillan Co.. New York. BARGHOORN,E. S. 1951.Age and environment; a survey Triporate A of North American Tertiary floras in relation to Figs. 181-182 paleoecology. J. Paleontol. 25: 736-744. BARGHOORN,E. S. and SPACKMAN,W. 1950. Geological DESCRIPTION: Triporate pollen grain, 26 to and botanical study of the Brandon lignite and its 38 p in diameter. Pores moderately aspidate, but significance in coal petrology. Econ. Geol. 45: 344-357. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 BELL,W. A. 1949. Uppermost Cretaceous and Paleocene nature of exine around pores is not clear; there floras of western Alberta. Geol. Surv. Can., Bull. 13. does not appear to be any thickening of exine BERRY,E. W. 1926. Tertiary floras from British Columbia. around pores. Exine finely and irregularly Geol. Surv. Can., Bull. 42, pp. 91-116. BRADLEY,W. H. 1931. Origin of the microfoss~lsof the striate. oil shales of the Green River Formation of Colorado BOTANICAL AFFINITY: 0111~tW0 species, both and Utah. U.S. Geol. Surv. Prof. Pap. 168. illustrated, were found, both in upper part of 1964. Geology of the Green River Formation and associated Eocene rocks in southwestern Wyoming the Kitsilano Formation. These two forms may and adjacent parts of Colorado and Utah. U.S. Geol. not be the same species, botanical affiliation un- Surv. Prof. Pap. 496A, pp. A1-A86. CAMP,W. H. 1947. Distribution patterns in modern known. plants and problems of ancient dispersais. Ecol. FREQUENCY: TWOgrains only were found. Monogr. 17: 159-183. 1128 CANADIAN JOURNAL OF BOTANY. VOL. 47, 1969

CHANDLER,M. E. J. 1955. The Schizaeaceae of the south 1956a. A palynological systematic nomenclature. of England in Early Tertiary times. Bull. Brit. Mus. Bogota, Bol. Geologico, 4: 63-101. (Natur. Hist.) 2, No. 7: 291-314. --- 1956b. Description of some genera and species CHANEY,R. W. 1936. Plant distribution as a guide to of fossil pollen and spores. Bogota, Bol. Geologico, 4:

age determination. Wash. Acad. Sci. 26: 313-324. 11- A 1-1 .17 . . 1940. Tertiary forests and continental history. HEDLUND,R. W. 1966. Palynology of the Red Branch Geol. Soc. Amer. Bull. 51: 469-488. Member (Woodbine Formation). Okla. Geol. Surv. 1948. The bearing of the living Metaseq~toiaon Bull. 112. problems of Tertiary paleobotany. Nat. Acad. Sci. HILLS,L. V. 1962. Glaciation, stratigraphy. structure and Proc. 34: 503-515. micropaleontology of the Princeton coalfield, British 1951. A revision of fossil Seqlroia and Taxodiu~n Columbia. Unpublished M.Sc. Thesis, Univ. of in western North America based on the recent dis- British Columbia, Vancouver. coveries of Mernseqltoia. Amer. Phil. Soc. Trans., -- 1965. Palynology and age of early Tertiary basins, new ser., 40(3): 171-239. interior British Columbia. Unpublished Ph.D. Thesis, CHANEY,R. W. and AXELROD,D. I. 1959. Miocene floras Univ. of Alberta. of the Columbia Plateau. Pt. 1 and 2. Carnegie Inst. HOLLAND,S. S. 1964. Landforms of British Columbia-A Wash. Publ. 617. physiographic outline. Brit. Columbia Dept. Mines CLARKE,R. T. 1965. Fungal spores from Vermejo Petrol. Resources, Bull. 48. Formation coal beds (Upper Cretaceous) of central HOPKINS,W. S. 1966. Palynology of Tertiary rocks of Colorado. The Mountain Geol. 2: 85-93. the Whatcom basin, southwestern British Columbia COUPER,R. A. 1953. Upper Mesozoic and Cainozoic and northwestern Washington. Unpublished Ph.D. spores and pollen grains from New Zealand. N.Z. Thesis, Univ. of British Columbia, Vancouver. Geol. Surv., Paleontological Bull. 22. 1968. Subsurface miocene rocks, British 1960. New Zealand Mesozoic and Cainozoic Columbia-Washington, a palynological investigation. plant microfossils. N.Z. Geol. Surv., Paleontological Geol. Soc. Amer. Bull. 79: 763-768.

Bull.- -~. 32.~ - HUGHES.R. 1946. Fraser River Tertiary. Unpublished Cox, R. L. 1962. Age and correlation of the Sooke B.A. Thesis, Univ. of British Columbia, Vancouver. Formation with a section on its palynology. Un- JOHNSTON,W. A. 1923. Geology of the Fraser River published M.Sc. Thesis, Univ. of British Columbia, Delta map area. Geol. Surv. Can., Mem. 135. Vancouver. KRAJINA,V. J. 1959. Bioclimatic zones of British CRICKMAY,C. H. and POCOCK,S. A. J. 1963. Cretaceous Columbia. Univ. of British Columbia, Botanical Ser. of Vancouver, British Columbia, Canada. Amer. No. 1. Ass. Petrol. Geol. Bull. 47: 1928-1942. 1965. Biogeoclimatic zones and classification of DAWSON,J. W. 1895. On collections of Tertiary plants British Columbia. 111 Ecology of North America. Vol. from the vicinity of the city of Vancouver, B.C. Roy. 1. pp. 1-17. University of British Columbia, Dept. Soc. Can. Trans., Ser. 2, l(4): 137-161. Botany. DILCHER,D. L. 1963. Eocene epiphyllous fungi. Science, KREMP,G. 0. W. 1965. Morphologic encyclopedia of 42: 667-669. palynology. Univ. of Arizona Press, Tucson, Arizona. ELSIK,W. C. 1968. Palynology of a Paleocene Rockdale KREMP,G. 0. W. and GERH~RD,J. E. 1960. Pollen and Lignite, Milam County, Texas. 1. Morphology and spores from the Lower Tertiary of North and South taxonomy. Pollen Spores, 10: 263-314. Dakota and some taxonomic problems concerning For personal use only. ERDTMAN,G. 1943. An introduction to pollen analysis. their designation. Pa. State Univ. College Min. Chronica Botanica Co., Waltham, Mass. Industries. Contrib. 58-64, pp. 257-269. 1952. Pollen morphology and plant taxonomy; KRUT~SCFI,W. 1957. Sporen- ~~ndpollengruppen aus der angiosperms (An introduction to palynology, 1). Oberkreide und dem Tertiar Mitteleuropas und ihre Chronica Botanica Co., Waltham, Mass. stratigraphische verteilung. 2. Angew. Geol. Bd. 3, 1957. Pollen and spore morphology /plant taxon- Heft 11 112, S. 490-588. omy; gymnospermae, pteridophyta, bryophyta. The 19590. Mikropalontologische (Sporenpalaeonto- Ronald Press Co., New York. logische) ~lntersuchungen in der Braunkohle des - 1966. Pollen morphology and plant taxonomy; pep!ta!l: Akad.-Verlag-Berlin, Nr. 21-22, Jahrgang Angiosperms. Hafner Pub. Co., New York. 6. 3. 1-4L.7. EVERNDEN,J. F. and JAMES,G. T. 1964. Potassium argon 19596. Einige neue formgattungen und -arten dates and the Tertiary floras of North America. von sporen und pollen aus der Mitteleuropaischen Amer. J. Sci. 262: 945-974. Oberkreide und dem Tertiar. Paleontogr., Bd. 105, FAFGRI,K. and IVERSON,J. 1964. Textbook of pollen S. 105-157. analysis. 2nd ed. Hafner Publishing Co., New York. -- 1962. Atlas der Mittel- und Juntertiaren dis- FELIX,C. J. 1965. Neogene Tnsmnnites and leiospheres persen sporen- und pollen sowie der mikroplankton- from southern Louisiana, U.S.A. Paleontology, 8(1): formen des nordlichen Mitteleuropas (in three parts). Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 16-26. Veb Deutcher verlag der Wissenschaften, Berlin. Fuc~s,H. P. 1967. Pollen morphology of the family LAWRENCE,G. M. 1951. Taxonomy of vascular plants. Bottibncncene. Rev. Palaeobotan. Palynol. 3: 119- Macmillan Co., New York. 1'71 1 .JL. MACGINTIE,H. D. 1941. A Middle Eocene flora from GRAHAM;A. 1965. The Sucker Creek and Trout Creek the celitral Sierra Nevada. Carnegie Inst. Wash. Publ. Miocene floras of southeastern Oregon. Kent State 534. Univ. Bull., Res. Ser., 9. 1966. Some vegetation types in Eocene of Middle GRIGGS,P. H. 1965. Stratigraphic significance of fossil Rocky Mountains. Geol. Soc. Amer. abstracts for pollen and spores of the Chuckanut Formation, 1966 annual meetings at Las Vegas, Nevada, pp. northwest Washington. Unpublished M.Sc. Thesis, 40-41 A. Michigan State Univ. MACKO,S. 1957. Lower Miocene pollen flora from the HAMMEN,TH. VAN DER. 1954. El desarrolo de la flora valley of Klodnica near Gliwice (Upper Silesia). Colombiana en 10s periodos geologicos. Bogota, Bol. Trav. Soc. Sciences Lettres Wroclaw, Ser. B, Nr. 88, Geolbgico, 2: 49-106. pp. 5-314. HOPKINS: EOCENE KITSILANO FORMATION 1129

1959. Pollen grains and spores from Miocene SMILEY,C. J. 1966. Cretaceous floras from Kuk River brown coals in Lower Silesia, I. Trav. Soc. Lettres area, Alaska; stratigraphic and climatic interpreta- Wroclaw, Ser. B. Nr. 96, pp. 5-178. tions. Geol. Soc. Amer. BLIII.77: 1-14. 1963. Sporomorphs from Upper Cretaceous SNAVELY,P. D. and WAGNER,H. C. 1963. Tertiary near Opole (Silesia) and from the London clays. geologic history of western Oregon and Washington. Trav. Soc. Sciences Lettres Wroclaw, Ser. R, Nr. 106, Wash. Div. Mines Geol. Rep. Inv. 22. pp. 5-136. STANLEY,E. A. 1965. Upper Cretaceous and Paleocene MARTIN,H. A. and ROUSE,G. E. 1966. Palynology of plant microfossils and Paleocene dinoflagellates and Late Tertiary sediments from Queen Charlotte Islands, hystrichosphaerids from northwestern South Dakota. British Columbia. Can. J. Bot. 44: 171-208. Bull. Amer. Paleontol. 49: 179-384. MULLER,J. 1959. Palynology of the Recent Orinoco THIERGART,F. 1940. Die mikropalaontologie als pollen- delta and shelf sediments. Micropaleontology, 5: analyses im dienst der Braunkohlenforschung. Von 1-32. Ferdinand Enke, Stuttgart, 13 Heft. NEWMAN,K. R. 1965. Upper Cretaceous-Paleocene THOMSON,P. W. and PFLUG,H. 1953. Pollen und sporen guide palynomorphs from northwestern Colorado. des Mitteleuropaischen Tertiars. Palaeontogr. 94, Pt. Univ. of Co!orado, Ser. in Earth Sciences No. 2. B: 1-138. NOREM.W. I-. 1955. Pollen, spores, and other organic THOMSON,R. V. 1958. Petrographic and sedimentary microfossils from the Eocene of Venezuela. Micro- comparison of sandstones of the Kitsilano and Bur- paleontology, 1 : 261-268. rard Formation. Unpublished B.A. Thesis, Univ. of POTONIP.R. 1951. Pollen- und s~oreformenals leitfos- British Columbia, Vancouver. silien des Tzrtiars. ~onderdruckaus Mikroscopie, Bd. TRAVERSE.A. 1955. Pollen analysis of the Brandon 6, Heft 9/10, S. 272-283. lignite of Vermo-it. U.S. Bur. Mines Rep. Inv. 5151. -- 1956. Synopsis der gattungen der sporae dis- VAN CAMPO-DUPLAN,M. 1951. Recherches sur la persae, 1 Teil, sporites. Beih. Geol. Jahrb., Hannover, phylogenie des Taxodiacees d'apres leurs grains de Heft 23. 103 S. pollen. Trav. Lab. Forest. Univ. Toulouse, 4, Art. 2: 1958. Synopsis der gattungen der sporae dis- 1-14. persae, 2 Teil, sporites (nachtrage), saccites, aletes, VARMA,C. P. and ROWAT,M. S. 1963. A note on some praceolpates, monocolpates. Beih. Geol. Jahrb., diporate grains recovered from Tertiary horizons of Hannover, Hzft 31, S. 1-114. India and their potential marker value. Grana Paly- 1960. Synopsis der gattungen der sporae dis- nologica, 4: 130-139. persae, 3 Teil, nachtrage sporites, fortsetzung pol- WHITE,W. H. 1959. Cordilleran tectonics in British lenites mit general register zu Teil 1-3. Beih. Geol. Columbia. Amer. Ass. Petrol. Geol. 43: 60-100. Jahrb., Hannover, Heft 39, S. 1-187. WILLIS,J. C. 1966. A dictionary of the flowering plants RODDICK,J. A. 1965. Vanco~lvernorth, Coq~~itlamand and ferns. 7th ed. Cambridge Univ. Press, England. Pitt Lake map-area, British Columbia. Geol. Surv. WILSON.L. F. 1956. Composite micropaleontology and Can., Mem. 335. its application to Tertiary and near Recent strati- ROUSE,G. E. 1957. The application of a new nomen- graphy. Micropaleontology, 2: 1-6. clatural approach to Upper Cretaceous plant micro- WILSON,L. R. and WEBSTER,R. M. 1946. P!ant micro- fossils from western Canada. Can. J. Bot. 35: 349-375. fossils from a Fort Union coal of Montana. Amer. J. 1962. Plant nlicrofossils from the Burrard For- Bot 33: 271-278. mation of western British Columbia. Micropaleon- WODEITOUSE,R. P. 1933. Tertiary pollen -2; the oil

For personal use only. tology, 8: 187-218. shales of the Green River Formation. Torrey Bot.

Rous~.--- ~ G. E. and MATHEWS.W. H. 1961. Radioactive Club Bull. 60: 479-524. dating- of Tertiary plant-bearing deposits. Science, 1935. Pollen grains, their structure, identifica- 133: 10791080. tion and significance in science and medicine. McGraw- RUDOLPH,K. 1935. Mikrofloristische Untersuchungen Hill Book Co.. Inc., N.Y. Tertiarer Ablager~lngenim nordlichen Bohmen. Bech. WOLFE,J. A. and BARGHOORN,E. S. 1960. Generic change Bot. Cbl. 54: 244-328. in Tertiary floras in relation to age. Amer. J. Sci., SCAGEL..R. F., BANDONI.R. J.. ROUSE.G. E., SCHOFIELD. Bradley Vol. 258A: 388-399. W. B., STEIN,J. R., and TAYLOR,T. M. C. 1965. ~n WCLFE.J. A,. GOWER.H. D.. and VINE. J. D. 1961. Aee evolutionary survey of the plant kingdom. Wads- and correlation of the Puget Group 'in King COLI~~, worth Publishing Co., Belmont, Ca!ifornia. Washington. U.S. Geol. Surv. Prof. Pap. 424-C, Art. SIMPSON.J. B. 1961. The Tertiary pollen-flora of Mull 231, pp. C230-273. and Ardnamurchan. Trans. Roy. Soc. Edinburgh, 64. WOLFE,J. A., HOPKINS,D. M., and LEOPOLD,E. R. SKARRY,A. 1964. Revision of Gleichetziidites senonicirs 1966. Tertiary stratigraphy and paleobotany of the Ross. Acta Univ. Stockholmie~~sis,Stockholm Con- Cook lnlrt region. Alaska. U.S. Geol. Surv. Prof. trib. Geol. 11: 59-77. Pap. 398-A, p.-~1:~29. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 EXPLANATIONOF FIGS. 1-186 FIG. I. Dyadosporotzifes sp. 1 (X 1000). FIGS. 2-3. Dyadosporotzites sp. 2,(X 500). FIGS. 4-6. Pleurice/lrresporites psrlatrrs Hammen ex Clarke 1965 (X 500). FIG. 7. PlerlriceNae~~oritesSD. (X 1000). FIG. 8. Fungal hyp6ae (X 5'00). FIGS. 9-10. ?Isoetes sp. (X 1000). FIG. 11. Lycopoclilltn annotinioides (Krutzsch) Martin & Rouse 1966 (X 1000). FIG. 12. Osnzr~ndaregalires Martin & Rouse 1966 (X 1000). FIGS. 13-14. Osmutzdrr irrepirlites Martin & Rouse 1966 (X 1000). FIG. 18. Osmlrnda sp. 2'(~1'000). 1130 CANADIAN JOURNAL OF BOTANY. VOL. 47, I969

FIG. 19. Oanrrnda sp. 2 (X 500). FIG. 20. Osmundn sp. 2 (X 1000). FIGS. 21-23. Anemia poolensis Chandler 1955 (X 1000). FIG. 24. Cicntricosisporites itztersect~rsRouse 1962 (X 1000). FIG. 25. Cicatricosisporites irltersectus Rouse 1962 (X 500). FIGS. 26-27. Cicatricosisoorires intersectus Rouse 1962 (X 1000). F~G. 28. ~ygorlirmzretic~~los~orites Rouse 1962 (X 1000). FIGS. 29-30. Gleiclretrin sp. (X 1000). FIGS. 31-32. Lnevigotosporites rliscorclntrrs Thompson & Mug 1953 (X 1000). FIG. 33. Loevigntosporites albertetrsis Rouse 1962 (X 1000). FIG. 34. Derrrrsraedriaceae-Po/~vporliaceneform 1 (X 1000). FIGS. 35-36. Det~rrsraedtincene-Polvoodiaceaeform 2 (X 1000). FIGS. 37-38. ~etrtrstnedtincene-~o&~odiaceaeform 2 (X 1000). FIG. 39. De~~r~smedriacene-Polyponinceaeform 3 (X 1000). FIG. 40. Azolln glochidia (X 1000). FIGS.41-43. ?Ginkgo sp. (X 1000). FIG. 44. ?Cedrus sp. (X 1000). FIG. 45. Kereleerirr sp. (X 1000). FIG. 46. Keteleerin sp. (X 750). FIGS.47-48. Lrrrix plicatipollenites Rouse 1962 (X 1000). FIGS.49-50. Picen grrrtrrlivescipites Wodehouse 1933 (X 750). FIGS.5 1-52. Pit~rrssrrobipites Wodehouse 1933 (X 750). FIGS. 53-54. Pitlrts sp. 11nploxylor1-type(X 750). FIG. 55. Pinrrs sp. (X 750). FIGS.56-57. Cupressaceae (X 1000). FIGS. 58-59. Cupressaceae, Taxodiaceae, or Pinaceae (X 1000). FIGS. 60-61. Glyptostrobr~svncrripites Wodehouse 1933 (X 1000). FIG. 62. Metoseqrroin pnpillnpol!enites Rouse 1962 (X 1000). FIG. 63. ?Sciarlopitys sp. (X 750). FIG. 64. Tnxodirrm lrirrtipires Wodehouse 1933 (X 1000). FIGS. 65-66. Podocarpus sp. (X 1000). FIGS. 67-68. ?Tnxlrs sp. (X 1000). FIG. 69. ?Epherlrtr sp. (X 1000). FIG. 70. Mngtzoli(rcetr (X 1000). FIG. 71. ?Nys.sn sp. (X 1000). FIGS. 72-73. Snlix rliscoloripites Wodehouse 1933 (X 1000). FIGS.74-75. ?Snltx sp. (X 1000). FIGS. 76-83. Alnrrs verrts (Potonit) Martin & Rouse 1966 (X 1000). FIGS. 84-85. Betrrla cf. B. clrrripires Wodehouse 1933 (X 1000). FIGS. 86-87. cf. Cnrpitrrrs sp. (X 1000). FIGS. 88-89. Corylrrs tripolle~iitesRouse 1962 (X 1000).

For personal use only. FIGS. 90-91. ?Cnstntlea sp. (X 1000). FIGS.92-94. Frigrrs grn~rrrlntaMartin & Rouse 1966 FIGS. 95-96. Fagrrs sp. (X 1000). FIGS. 97-98. Qrrercrrs explatirrm Anderson 1960 (X 1000). FIGS. 99-100. Qrrercrrs sp. 1 (X 1000). FIGS. 100-102. Qrrercrrs sp. 2 (X 1000). FIGS, 103-104. Qrrercrrs sp. 3 (X 1000). FIG. 105. ~rrercissp. 4 (X 1000). FIGS. 106-109. Cnrya jr~stnporites (Wodehouse) Rouse 1962 (X 1000). FIGS. 110-1 1 1. Et~gell~nrrltincf. E. gr~rrrrrlataSimpson 1961 (X 1000). FIGS.112-1 13. Etlgelharrltin sp. (X 1000). FIGS. 114-1 16. P/ntycarj~osp. (X 1000). FIGS. 117-1 18. Jriglntis periporites Martin & Rouse 1966 (X 1000). FIGS. 119-121. Jrrglnns sp. (X 1000). FIGS. 122-124. Pterocorya srellntrrs Martin & Rouse 1966 (X 1000). FIGS. 125-127. Monzipiies tenrripolis Anderson 1960. FIGS. 128-129. Ulmrrs sp. or Zelkova sp. (X 1000).

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 FIG. 130. Planera sp. (X 1000). FIGS. 131-133. ?Tilin cf. T. vescipires Wodehouse 1933 (X 1000). FIG. 134 Tilin sp. (X 1000). FIG. 135. ?Tiliacene. FIGS. 136-137. Acer sp. (X 1000). FIG. 138. A~SCIIIUSsp. (X 1000). FIGS. 139-142. Pi.rtillipo!lenites mcgregorii Rouse 1962 (X 1000). FIGS. 143-148. flex sp. (X 1000). FIGS. 149-1 5 1. Protencidites tlznlnlnni Anderson 1960 (X 1000). FIG. 152. Protencidites sp. (X 1000). FIGS. 153-154. Myrica ntltlrrlites Martin & Rouse 1966 (X 1000). FIGS. 155-1 56. Lilincirlifes sp. (X 1000). FIGS. 157-158. Sabnl grnnopollenites Rouse 1962 (X 1000). FIGS. 159-160. Typhn sp. (X 1000). HOPKTNS: EOCENE KITSILANO FORMATION

FIG. 161. Potanlogeton hollickipites Wodehouse 1933 (X 1000). FIGS.162-163. Triletes solidus (Potonie) Krutzsch 1959 (X 1000). FIG. 164. Deltoidospora sp. 1 (X 1000). FIG. 165. Deltoidospora sp. 2 (X 1000). FIGS.166167. Monoporate A (X 1000). FIG. 168. Monoporate B (X 1000). FIG. 169. Monoporate C (X 1000). FIG. 170. F~isiformisporitesmicrostriatru n. sp. (X 1000). FIGS. 171-172. Plmctodiporites harrisii Varma and Rawat 1963 (X 750). FIG. 173. Diporisporites sp. (X 500). FIG. 174. Diporate A (X 1000). FIG. 175. Diporate C (X 500). FIGS. 176177. Tricolpate A (X 1000). FIG. 178. Tricolpate B (X 1000). FIG. 179. Tricolpate C (X 1000). FIG. 180. Tricolporate A (X 1000). FIGS. 181-182. Triporate A (X 1000). FIG. L83. Fungal spore (X 1000). FIG. 184. Asterha sp. (X 500). FIGS. 185-186. Unidentified (X 500). NOTE:Figs. 1-186 follow. For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. PLATEVII For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only. Can. J. Bot. Downloaded from www.nrcresearchpress.com by 66.183.244.166 on 05/15/20 For personal use only.