TRIPLOCHITIOXYLON (STERCULIACEAE): A NEW GENUS OF WOOD FROM THE EOCENE OF OREGON AND ITS BEARING ON XYLEM EVOLUTION IN THE EXTANT GENUS TRIPLOCHITON

A new sterculiaceous wood, Triplochitioxylon oregonensis gen. et sp. n., was collected from a Middle Eocene locality in the Clarno Formation of Oregon. Anatomical data indicate a close natural relationship between T. oregonensis and the living species of Triplochiton, a genus endemic to tropical Africa. The fossil is believed to represent a population of the group or complex from which Triplochiton evolved. The basic differences in the xylem organizations of the two genera are explained by a significant reduction of fusiform initial length and by a complete suppression of post-cambial parenchyma strand elongation in the extant genus. Paleobotanical and biogeographical evidence suggest that the xylem evolution has been strongly influenced by the increasing aridity of the African continent.     

FOSSIL CERATOPHYLLUM (CERATOPHYLLACEAE) FROM THE TERTIARY OF NORTH AMERICA

Fossil fruits and a vegetative axis assignable to the extant genus Ceratophyllum are described from four North American Tertiary localities. Fossil fruits assignable to the extant species C. muricatum and C. echinatum are reported from the Eocene Green River and Claiborne formations, and the Miocene Esmerelda Formation, respectively. An extinct species, C. furcatispinum, is described from the Paleocene Fort Union Formation and represents the oldest published report of Ceratophyllum in the fossil record. The existence of extant angiosperm species in the Eocene is very unusual and may be attributable in this case to slow evolutionary rates and unusual evolutionary properties associated with hydrophily in the genus Ceratophyllum.     

EVOLUTION OF THE GENUS CHRYSOCOCCYX

Modern geological ideas on ocean-floor spreading are briefly reviewed. Pangea began to break up at the end of the Trias, but Africa, Antarctica and Australia remained together or close to each other till the end of the Cretaceous. The position of western New Guinea at the start of the Miocene could have been approximately where Arnhem Land is now, and at the start of the Pliocene somewhat north of the present-day Aru Islands. Its size until the end of the Pliocene was much smaller than it is today. Friedmann's proposal for the evolutionary spread of Chrysococcyx therefore demands that the whole process occurred since about the start of the Pliocene. There may not have been enough time in these seven million years for the evolutionary dispersal of a genus of parasitic cuckoos halfway round the world. His proposal also regards C. osculans as an awkward throw-back, and leaves a gap between species in New Guinea and southeastern Asia that is not bridged by intermediates. If a stock of cuckoos had been in Gondwanaland before it broke up, that stock could have given rise to the genus Cacomantis and the forerunners of C. osculans. The lineage of osculans would have quickly given rise to a lineage of glossy cuckoos that then divided into two branches. One could have penetrated Africa, south of where Madagascar then was, produced the species klaas, cupreus, caprius and flavigularis (an aberrant end-product), and much later, after Madagascar had drifted south from India (having been separate from Africa since the Cretaceous), colonised Asia where maculatus and xanthorhynchus would have differentiated. The other line could have differentiated, perhaps more slowly, in Australia into basalts, lucidus, ruficollis and malayanut (minutillus). When Australia had drifted near enough to the Malay Archipelago and as New Guinea grew, ruficollis and minutillus could have moved forward to colonise the islands, where minutillus would have produced the many races oimalayanus and meyerii differentiated as an aberrant end-product. This proposal overcomes some of the objections to Friedmann's theory because it arranges events to accord better with geological developments and avoids the evolutionary discontinuities of his proposal. The migratory habits of the cuckoos, thought by Friedmann to be significant, are discounted as evidence for evolutionary history. The parasitic habits are re-interpreted. Better data are presented for the parasitic behaviour of basalis and lucidus; they suggest that both species are sophisticated and probably host-specific parasites. Jensen & Jensen (1969) have already given evidence that some African glossy cuckoos are highly host-specific. There seems to be a trend of decreasing parasitic sophistication in Australian species, correlated with the possible age of the species. In Africa parasitism seems to be far in advance of that in Australia, probably because the opportunities for parasitism are far better in Africa. These trends and differences in parasitic behaviour are compatible with an evolutionary spread from Gondwanaland. The crucial question is whether the stock of Cacomantis and Chrysococcyx could have existed before the break-up of Gondwanaland, i.e. before the early Eocene. The present fossil record suggests that this is unlikely, but the paucity of fossils and the difficulties of palaeoclimates do not seem to be insuperable and it is suggested that a southern origin for these cuckoos should be considered seriously.     

ATTACHED REPRODUCTIVE AND VEGETATIVE REMAINS OF THE EXTINCT AMERICAN-EUROPEAN GENUS CEDRELOSPERMUM (ULMACEAE) FROM THE EARLY TERTIARY OF UTAH AND COLORADO

Fossil twigs with attached foliage, fruits, and flowers from the middle Eocene of the Green River Formation in northeastern Utah and northwestern Colorado and from the early Oligocene Florissant beds of central Colorado provide a firm basis for reconstructing two species of an extinct ulmaceous genus that was widely distributed in the Tertiary of midlatitude western North America and Europe. The fruits are samaras of Cedrelospermum Saporta, a genus previously known only from isolated specimens. The distichously arranged, slender, pinnate-veined leaves vary from serrate with simple teeth to, less commonly, entire-margined. Corresponding isolated leaves in the Green River, Florissant, and other Eocene to Oligocene localities of western North America are now excluded from Zelkova and Myrica, to which they were previously misidentified. The anthers of the staminate flowers contain 3–5 porate pollen with rugulate sculpture. Based upon combined characters of phyllotaxy, and leaf, flower, fruit, and pollen morphology, Cedrelospermum can be referred to the extant subfamily Ulmoideae, and is similar to Phyllostylon, Zelkova, and Hemiptelea. The abundance of Cedrelospermum in lake sediments of volcanic areas, together with its production of numerous small winged fruits, suggest that it was an early successional colonizer of open habitats.     

THE LEPIDOPHYTIC AFFINITIES OF THE GENUS CHINLEA AND OSMUNDITES WALKERI

Chinlea campii Daugherty and Osmundites walkeri Daugherty are species of petrified stems from the Upper Triassic Chinle Formation of Arizona that were described as members of the fern family Osmundaceae. Investigation of additional material indicates that the two species are conspecific and belong to the Lepidophyta. The stems are radially symmetric and have an ectophloic siphonostele in which the xylem cylinder is thick and deeply furrowed. Internal pressure against the xylem cylinder caused by the lateral expansion of the pith in some stems produces what appears in transverse section to be a ring of up to 60 separate xylem strands. Leaf traces are small, terete, collateral and have exarch xylem. They are arranged in a tight spiral. Adventitious roots, secondary xylem, and secondary cortex are lacking. The stems are classified under the binomial Chinlea campii, and other axes that have similar cortical anatomy but in which all vascular tissues have decayed are treated as Chinlea sp. Both types of stems are interpreted as ephemeral aerial shoots of an herbaceous plant. Of the known fossil Lepidophyta, Chinlea is most similar to Pleuromeia and Nathorstiana, but it differs from each of these genera in a number of respects and is therefore included in Lepidophyta incertae sedis.     

INTERCONNECTED REPRODUCTIVE AND VEGETATIVE REMAINS OF POPULUS (SALICACEAE) FROM THE MIDDLE EOCENE GREEN RIVER FORMATION,NORTHEASTERN UTAH

A new specimen from the Middle Eocene Evacuation Creek Member of the Green River Formation in northeastern Utah shows a twig with several leaves of Populus wilmattae Cockrell and a fruiting raceme attached. This specimen establishes for the first time the type of fruits borne by P. wilmattae and provides additional characters with which to assess its taxonomic and evolutionary status. An associated seed shows attached placental hairs like those of extant species of Populus. The Green River fossil differs from extant Populus species in having basically palmate leaf venation and in bearing its fruiting axis on a young twig. In other aspects, the fossil species is remarkably similar to the extant species Populus mexicana.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A CATKIN WITH JUGLANDACEOUS AFFINITIES

Three specimens of one type of fossil catkin from the Middle Eocene of Tennessee are excellently preserved and have been investigated morphologically. The flowers on these catkins are subtended by elongate, three-lobed bracts, are exclusively staminate, and have three conspicuous, obovate, perianth parts that bear large peltate scales. The stamens are well preserved and contain triporate pollen grains that are equivalent to the dispersed pollen genus Momipites. Floral morphology, cuticular features, and pollen indicate close affinities with the extant genera Engelhardia, Oreomunnea, and Alfaroa of the Juglandaceae; but because the fossil catkins are distinct and are a dispersed plant organ, they are placed in a new form genus: Eokachyra. These fossil flowers represent a rare opportunity to correlate the micro- and macrofossil record and to compare the relative rates of evolution of these features. The fossil catkins also demonstrate that much structural information may be gained from the study of fossil angiosperm flowers. The similarities between the staminate flowers of the fossil catkins and the staminate flowers of Engelhardia, Oreomunnea, and Alfaroa confirm the idea that this complex has had a long evolutionary history and suggest that the pollination system of certain extant genera was well developed during Middle Eocene times.     

PROTEOKALON A NEW GENUS OF PROGYMNOSPERMS FROM THE DEVONIAN OF NEW YORK STATE AND ITS BEARING ON PHYLOGENETIC TRENDS IN THE GROUP

Proteokalon gen. nov. is described from the Upper Devonian Catskill deposits of New York. Two orders of branching and ultimate appendages are preserved' by petrifaction and by compression. The first order bears branches decussately and has a skewed four-armed protostele that occasionally dichotomizes. Second-order branches dichotomize rarely and most have T-shaped or three-armed protosteles. They bear ultimate appendages alternately, either in lateral pairs, or singly from the abaxial side. These appendages divide several times in one plane. Their vascular strand is terete. Maturation of the primary xylem is mesarch, and it consists of tracheids and parenchyma. Secondary xylem and phloem and a periderm are present. The outer cortex has a system of hypodermal fibers. Proteokalon is most similar to Tetraxylopteris and Triloboxylon of the Aneurophytales. A comparison of the stratigraphic occurrence of Protopteridium, Aneurophyton, Tetraxylopterism, Sphenoxylon, Triloboxylon, and Proteokalon suggests some evolutionary trends among the Aneurophytales.     

EQUISETUM CLARNOI,A NEW SPECIES BASED ON PETRIFACTIONS FROM THE EOCENE OF OREGON

Equisetum clarnoi is described from four silicified stem fragments and numerous small roots from the Eocene Clarno Chert of Jefferson County, Oregon. Stems are up to 8.0 mm in diam and have sunken stomata arranged vertically in a single line flanking each of the external biangulate stem ridges, features that clearly ally this species with the subgenus Hippochaete. External stem ridges are equal in number to the carinal hypodermal bands. The hypodermis is composed of fibers and has prominent carinal bands up to 0.75 mm long and shorter vallecular bands. Cortical parenchyma cells enclose prominent vallecular canals which are lined by specialized thick-walled parenchyma cells. The double, common endodermis has prominent casparian strips. Vascular bundles are composed of four to seven metaxylem tracheids flanking each side of the phloem and protoxylem tracheids which occur singly on the internal surface of the small carinal canals. Leaf sheaths in cross section have an adaxial fibrous layer and an external or near external fibrous bundle. Roots are up to 2.0 mm in diam and have paired cuboidal epidermal cells from which root hairs arise. The stele of the root is central and shows exarch primary xylem maturation. Equisetum clarnoi most closely resembles the extant Equisetum hyemale var. affine.     

EOCENE FLORAL EVIDENCE OF LAURACEAE: CORROBORATION OF THE NORTH AMERICAN MEGAFOSSIL RECORD

Numerous megafossils of Lauraceae have been reported from the early Tertiary of North America, but the subfamilial affinities are usually not well understood due to the great morphological variability found in extant taxa. The flowers of Androglandula tennessensis gen. et sp. nov. Taylor, from the Middle Eocene Claiborne Formation, are six-parted, pedicellate, bracteate, and have stamens with paired basal staminal glands. The flowers have ethereal oil cells and paracytic stomates throughout. The fossil species has affinities with the subtribe Cinnamomineae, and this supports suggestions that the Middle Eocene climate of the southeastern U.S. was subtropical. The existence of this fossil, and reports of the subtribe from the Eocene of Europe, indicate a South American-North American-European-southeast Asian paleodistribution suggesting that extinction in North America and Europe was the cause of the tribe's current disjunct distribution.     

CAESALPINIA SUBGENUS MEZONEURON (LEGUMINOSAE,CAESALPINIOIDEAE) FROM THE TERTIARY OF NORTH AMERICA

Legume fruits from the Eocene of Tennessee and Wyoming and the Miocene of Idaho are described and assigned to Caesalpinia subgenus Mezoneuron (Caesalpinioideae), an extant Paleotropical taxon that does not occur in North or South America today. Morphological and anatomical details of the fruits are used in evaluating their systematic relationships. The features of the fossil fruits are accommodated only within this extant subgenus. These fossils represent the only reliable known occurrence of C. subgenus Mezoneuron in the paleobotanical record. These fossils suggest that subgenus Mezoneuron was distinct from subgenus Caesalpinia by the Middle Eocene. Further, they document the widespread occurrence of this currently Paleotropical group for at least 30 million years in North America.     

STUDIES OF PALEOZOIC FERNS: TUBICAULIS STEWARTII SP. NOV. AND EVOLUTIONARY TRENDS IN THE GENUS

Eggert , Donald A. (Yale U., New Haven, Conn.) Studies of Paleozoic ferns: Tubicaulis stewartii sp. nov. and evolutionary trends in the genus. Amer. Jour. Bot. 46(8): 594–602. Illus. 1959.—Tubicaulis stewartii, a new species of the order Coenopteridales is described. The specimen was derived from the Upper Pennsylvanian of Berryville, Illinois, and is characterized by having a lacunar middle cortex, a well-developed integumentary system bearing uniseriate hairs, and xylem parenchyma organized into vertically anastomosing strands. In addition, multiseriate (somewhat transitional to reticulate) bordered pitting is present in the petiolar metaxylem elements, while those of the stem stele are multiseriate scalariform. The habit is intermediate between that of a form such as Osmunda and a tree fern, having an upright tapering stem which gives off prominently decurrent petioles in a 2/5 divergence. A reinvestigation of the type specimen of the most closely allied species, T. multiscalariformis, of Upper-Middle Pennsylvanian age, has shown that it has similar features in the cortex, metaxylem, and integumentary layers. Tubicaulis multiscalariformis and T. stewartii form a distinct group in the 6 species now known, whose evolution has most likely involved the retention of a more primitive form of pitting (multiseriate scalariform) with parenchymatization of the xylem. The remaining species of the genus have not developed xylem parenchyma but have developed circular bordered pitting. The relationships of the genus to other genera in the Coenopteridales remain obscure.     

THE PHLOEM OF ETAPTERIS LECLERCQII AND BOTRYOPTERIS TRIDENTATA

The phloem of Etapteris leclercqii and Botryopteris tridentata petioles is described from Lower Pennsylvanian coal balls. Petioles of B. tridentata are characterized in transverse section by an omega-shaped xylem trace, a phloem zone which extends from 2-10 cells in width, and 2-parted cortex. Etapteris leclercqii petioles exhibit a 4–9 cell-wide phloem zone surrounding the central clepsydroid xylem mass, and a 3-parted cortex. In both taxa a 1–2 cell layer parenchyma sheath separates the xylem from the extra-xylary tissues. The phloem of both species consists of sieve elements that average about 20 μm in diam by 200 μm in length in Botryopteris, and 100 μm in length in Etapteris, with horizontal-slightly oblique end walls. In transmitted light, the radial walls of the sieve elements form an irregular reticulate pattern enclosing elliptical lighter areas. With the scanning electron microscope, these areas appear as horizontal-slightly oblique furrows on the cell wall, with many small indentations lining the furrows. These indentations, because of their regular occurrence and size (from a few fractions of a micron up to 1.0 μm in diam), are interpreted as sieve pores, and the elliptical areas that enclose them as sieve areas. The phloem of E. leclercqii and B. tridentata is compared with that described for other fossil genera and with that of extant ferns.     

MORPHOGRAM: A NOVEL DIAGRAM TO ORGANIZE THE TRANSITIVE SECONDARY XYLEM ELEMENTS OF BASAL TOMATO (SOLANUM LYCOPERSICUM) INTERNODES

In the internodal secondary xylem of tomato (Solanum lycopersicum L. cv. Moneymaker) there are five main groups of secondary xylem elements which can be distinguished in wood macerates of basal internodes of the plant. Their top-frequency lengths are distinct: ray parenchyma cells (80 μm); fusiform parenchyma cells and strands (250 μm); vessel members (290 μm); tracheids (350 μm); and fibrous elements (590 μm), although there is overlap in length and morphology. The imperforate axial elements are strikingly diverse and morphologically intergrading, precluding ready classification according to traditional wood anatomical standards. In a novel, more appropriate, flexible categorization the variability in imperforate axial elements is depicted in a morphogram in which pit shape is plotted against cell length and cell morphology. This morphogram organizes the various elements without imposing an absolute classification. It is concluded that the appreciable variation in tomato wood samples results from developmental plasticity. The morphogram elucidates the extent and nature of element variation at the morphological level. Thus, it can be used to record developmental plasticity of wood tissues to assess interplant variation in wood tissue development, as well as intraplant plasticity.     

MUSOPSIS N. GEN.: A BANANA-LIKE LEAF GENUS FROM THE EARLY TERTIARY OF EASTERN NORTH GREENLAND

A fossil flora from the Late Paleocene-Early Eocene Thyra Ø Formation of eastern North Greenland (paleolatitude 77° N) has yielded monocotyledon leaf impressions with characters seen only in the closely related modem species in the families of Heliconiaceae, Musaceae, and Strelitziaceae. The combination of large costae widths and parallel, nonanastomosing, lateral veins that depart at right angles from the costae in the fossil material are features present only in leaves of extant species from these families. Three basic venation patterns also are recognized in the modem species of these families, but except for the genera Strelitzia and Phenakospermum, none of these patterns are present exclusively in any one family. Musopsis n. gen. is created for the fossil material from Greenland, but it is considered a form genus due to the lack of gross morphological features that can be used for separating leaves of the modem genera in Heliconiaceae, Musaceae, and Strelitiziaceae. It is the first known Arctic occurrence of fossil leaf material resembling this modem group of taxa.     

PINUS WOLFEI,A NEW PETRIFIED CONE FROM THE EOCENE OF WASHINGTON

A silicified cone from the Late Eocene of Washington is described as a new fossil species of Pinus. The cone was probably 9–10 cm long and 3–5 cm at its widest diam in the living condition and is peculiar in having abundant resin canals in the secondary xylem of the axis arranged in three concentric rings near the cone base. The bract of the fossil is also unusual in having resin canals of distinctly unequal sizes and a vascular strand that is adaxially concave. In the absence of external features of the scale tips, these anatomical conditions along with the construction of the outer cortex of the axis of thick-walled cells suggest closest affinity of the new species with the subsections Contortae, Oocarpae, and Sylvestres of the section Pinus, subgenus Pinus.     

WOOD OF GINKGO IN THE TERTIARY OF WESTERN NORTH AMERICA

Scott , R. A., E. S. Barghoorn , and U. Prakash . (U.S. Geol. Sur., Denver, Colo.) Wood of Ginkgo in the Tertiary of western North America . Amer. Jour. Bot. 49(10): 1095–1101. Illus. 1962. —Woods of Ginkgo and extinct related genera are very rare in the fossil record in contrast to the numerous ginkgoalean leaves. Ginkgo wood may be distinguished from other gymnosperms by a combination of anatomical features herein described. Ginkgo wood from beds of Miocene age at Vantage, Washington, first identified by Beck, is assigned to a new species, G. beckii. Ginkgo wood from the upper Eocene Clarno Formation, John Day Basin, Oregon, is described as G. bonesii sp. nov. Scarcity of fossil ginkgoalean woods may reflect unusual susceptibility to degradation of their cell walls in contrast to the greater chemical resistance to degradation which features many coniferous woods.     

RELATION OF DWARFMISTLETOE (ARCEUTHOBIUM) TO THE XYLEM TISSUE OF CONIFERS. II. EFFECT OF THE PARASITE ON THE XYLEM ANATOMY OF THE HOST12

Srivastava , L. M., and K. Esau , (U. California, Davis.) Relation of dwarfmistletoe (Arceuthobium) to the xylem tissue of conifers. II. Effect of the parasite on the xylem anatomy of the host. Amer. Jour. Bot. (48(3): 209–215. Illus. 1961.—The changes in the xylem anatomy induced by dwarfmistletoe infection were studied in 7 coniferous species. The most pronounced abnormalities are observed in the shape and size of the infected rays. Because of the presence of parasite tissue, the rays assume a hypertrophied appearance; moreover, they fuse to form large composite rays. The union of rays involves intrusive growth of ray cells and displacement of fusiform initials. Some division of fusiform initials also occurs. Rays may increase in number and they may contain more host cells than normal rays. Axial tracheids in infected host woods differ more or less strongly from those of noninfected woods. They may be shorter, wider, and more irregular in shape than the axial tracheids in healthy wood. The samples of xylem from infected pines had a larger number of resin canals than those from healthy trees. Resin canals were also found in infected Tsuga, which normally lacks these structures.