Fossil records of subsection Pinus (genus Pinus,Pinaceae) from the Cenozoic in Japan

Extant pines of subsection Pinus (section Pinus, genus Pinus, Pinaceae) are predominantly distributed in Eastern Asia. However, the extent of diversification in the section has yet to be fully clarified. We reviewed fossil records of subsection Pinus from Japan and collected permineralized materials, in which anatomical details are preserved for better understanding of the diversification. Our results suggest that this subsection appeared in Japan no earlier than the Middle Eocene, with extant species (i.e., Pinus densiflora and Pinus thunbergii) appearing around the beginning of the Pleistocene. Pinus fujiii (Early Miocene to Early Pleistocene) is inferred to have a close affinity to P. thunbergii based on the medial arrangement of its leaf resin canals. Additionally, P. fujiii has a similar cone morphology to those of extant species living in China, bridging the morphological gap between P. thunbergii and Chinese relatives of P. thunbergii as inferred by molecular phylogenetic analyses. Our results also suggest that taxonomic revisions of Pinus miocenica and Pinus oligolepis are required among the Japanese fossil species reported to date.     

Pinus MUTOI (Pinaceae), a new species of permineralized seed cone from the upper Cretaceous of Hokkaido,Japan

Pinus mutoi is described as a new species on the basis of a permineralized seed cone from the Upper Cretaceous of Hokkaido, Japan. The cone is at least 20 cm long and up to 6 cm in diameter, consisting of a cone axis and numerous cone-scale complexes that are arranged helically around the axis. Two winged seeds are borne on the adaxial surface of each ovuliferous scale. Each complex receives a single trace from the vascular cylinder of the cone axis. In the scale base, all the resin canals occur abaxially to the vascular strand. The spatulate bract of the fossil is unique to the specimen among the cones of both living and fossil Pinus. The central umbo, broad sclerotic cortex of cone axis, and absence of serotinous features of the fossil cone suggest affinity with the subsection Sylvestres of the section Pinus, subgenus Pinus. This is the first record of permineralized preserved Pinus cone from the Cretaceous of Eastern Eurasia.     

Contribution to the taxonomy of Pinus uncinata (Pinaceae) based on cone characters

Eight natural populations of Pinus uncinata Ramond from the Spanish Pyrenees, five of P. mugo Turra from the Carpathians, the Sudethians and Abruzzian Apennines, three of P. sylvestris L. from the Central-European Lowland and the Iberian Peninsula and one of P. uliginosa Neumann from SW Poland have been compared biometrically in 16 cone characters. P. uncinata appears the most distant from all other taxa, differing by having the largest and most asymmetric cones and the thickest apophyses with the most prominent umbo. In cone characters, P. mugo is more similar to P. sylvestris than to P. uncinata. Nevertheless, P. mugo and P. sylvestris remain distant from each other in a number of morphological characters examined. P. uliginosa has cones intermediate between the other three taxa and is moreover very variable. The locality of P. uliginosa in the Silesian Lowland may be of a relict character and conserve genes from P. uncinata.     

Pinus baileyi (section Pinus, Pinaceae) from the Paleogene of Idaho, USA

Pinus baileyi from the Paleogene of Idaho was initially related to the bristlecone pine P. longaeva (subgen. Strobus, sect. Parrya, subsect. Balfourianae) from western North America. Unlike the centromucronate condition in P. longaeva, P. baileyi cones have raised umbos that are excentromucronate, i.e., the mucro positioned in the upper umbo field above the keel. Cone size and scale morphology shows that P. baileyi more closely resembles excentromucronate pines of subsects. Halepenses and Pinus sensu Gernandt et al. (2005, Taxon 54: 29-42), but is most similar to P. resinosa, P. kesiya, and P. massoniana of subsect. Pinus. Morphologically, P. baileyi resembles the fossil species P. princetonensis and P. arnoldii from the Eocene Princeton Chert, British Columbia, Canada. Pinus baileyi extends the western North American range of ovulate cones resembling subsect. Pinus from the middle Eocene of British Columbia, Canada and Washington, USA to the Oligocene of Idaho, USA. Pinus baileyi, and possibly P. princetonensis and P. arnoldii, indicates the presence of early populations of subsect. Pinus-type pines in the western cordillera of North America, raising the possibility that P. resinosa and P. tropicalis may have evolved from this group.     

The genus Taxodium (Cupressaceae) in the Palaeogene and Neogene of Central Europe

Fossil remains of Taxodium from upper Eocene to lower Miocene localities in Germany, the Czech Republic and Russia are investigated to reconsider taxonomic concepts. It is demonstrated that all material belongs to one species which is named Taxodium dubium (Sternberg) Heer emend. Using the whole-plant concept, T. dubium is considered as a synthetic species comprising foliage, seed cones, seeds, and male cones with pollen in situ. Shared identity of T. dubium and the wood morpho-taxon Taxodioxylon taxodii Gothan is assumed. Taxodium dubium underwent evolutionary changes through the Oligocene expressed mainly in changes of the cone scale ornamentation. To designate this intraspecific variation taxonomically two morpho-formae are distinguished: T. dubium forma heerii (Dorofeev) Kunzmann, Kva?ek, Mai et Walther stat. nov. et emend. and T. dubium forma dubium. Simultaneous intraspecific variations in leaf morphology and leaf epidermal anatomy are missing. Taxodium dubium occurs from the late Eocene to late Miocene in the (Atlantic-) Boreal province sensu Mai of Central Europe and from the late Oligocene to late Pliocene in the Transeuropean Paratethys province sensu Mai.     

Geographical cline and inter-seaside difference in cone characteristics related to climatic conditions of old planted Pinus thunbergii populations throughout Japan

Although cone morphology has been mainly utilized for taxonomy or forestry purposes, characterization of the environmental factors that influence its variation is still insufficient. Major conifers distributed over diverse climatic conditions like the Japanese archipelago could show cone morphology variation among populations related to large differences in key influential factors such as temperature, snow, irradiance and sunshine. Geographical variation in cone characteristics (cone size, seed production, seed productivity per cone and seed size) was examined across the species distribution in 24 old planted Pinus thunbergii populations (479 trees) along two major seasides in Japan (Pacific Ocean side and Japan Sea side). Variance components of cone characteristics explained by seaside were at similar levels to those for populations nested within seasides. Populations on the Japan Sea side produced cones that had larger size (length and width), higher seed production (number of filled seeds per cone and seed mass per cone) and seed productivity per cone (seed-to-ovule ratio). Analysis of covariance showed that significant latitudinal clines observed in cone size and seed productivity per cone were mostly a result of inter-seaside differences. Linear mixed model analysis detected that geographical variation in cone size, seed production and seed productivity per cone in the populations were affected significantly by low temperature, more snow, less solar irradiance and less sunshine time. Large cone sizes and high seed productivity per cone found in the populations on the Japan Sea side could be explained by a genetic or plastic response to maintain reproductive success.     

Discrimination among pinyon pine trees by Clark's Nutcrackers: effects of cone crop size and cone characters

Summary The influences of Colorado pinyon pine (Pinus edulis) cone crop size, cone and seed weight, cone length, number of seeds per cone, number of viable seeds, and percent viable seeds on the foraging behavior of avian seed dispersal agents were examined in field and laboratory settings. In the field, there was a significant positive relationship between cone number per tree and both the absolute number of cones and the percentage of the cone crop from which seeds were harvested. Cone weight and the number of viable seeds were also significantly related to seed harvest intensity. Laboratory experiments examined the relationship between crop size and cone characters on seed harvest by 18 Clark's Nutcrackers (Nucifraga columbiana). Nutcrackers were offered a choice of two tree types: one with 20 cones attached, and another with 10 cones attached. Significantly more birds chose to remove seeds first from the tree with 20 cones than the tree with 10 cones. In timed trials, they also harvested seeds from significantly more cones on the tree with the higher cone density. In the laboratory, cones chosen for seed removal by the nutcrackers had significantly more viable seeds, more seeds, and were longer compared to cones that were not chosen. Such discriminatory foraging behavior may increase avian foraging efficiency and result in differential reproductive success of pinyon pines. This behavior may therefore influence the evolution of pinyon pine reproductive traits.     

IMMUNOCHEMICAL COMPARISONS OF SEED PROTEINS FROM POPULATIONS OF PINUS RADIATA (PINACEAE)

Antisera were prepared to seed proteins from five populations of Pinus radiata and to seed proteins from six other pine species. Differences between the P. radiata populations detected with these antisera were compared to differences in sizes of cones from the same populations. The strong correlation between antigenic differences and cone size differences provides evidence that in P. radiata, both seed proteins and cone size indicate quantitative genetic relationships among the populations. The data suggest that the three California mainland populations of P. radiata are more closely related to one another than to the insular populations, and that the insular populations are more closely related to each other than to the California populations, to P. attenuata, or to P. muricata.     

Great spotted woodpeckers Dendrocopos major exert multiple forms of phenotypic selection on Scots pine Pinus sylvestris

Relatively few animal species extract seeds from closed conifer cones because of the forces required to spread apart or penetrate the woody scales. Those species that forage on seeds in closed cones tend to forage selectively, and therefore act as selective agents on cone structure. However, little is known about the foraging preferences and thus phenotypic selection that is exerted on conifers by many species that forage extensively on seeds in closed cones, including especially woodpeckers (Picidae). Great spotted woodpeckers Dendrocopos major are one of the main predators of seeds in closed cones of Scots pine Pinus sylvestris in central and eastern Europe. To estimate the cone preferences of these woodpeckers foraging on Scots pine, we contrasted traits of cones that were and were not foraged on by woodpeckers. Woodpeckers preferred to forage on shorter cones when scales were thin (smaller apophyses) but preferred cones of intermediate length when scales were thicker, providing evidence for correlational selection. The preference for intermediate‐sized cones indicates that woodpeckers exert disruptive selection on cone length when cones have thicker scales, but the overall selection on cone length across all scale types indicates directional favoring the evolution of longer cones. Woodpeckers avoided cones with thicker scales, which would lead to directional selection favoring the evolution of thicker scales. Preferences for intermediate‐sized cones have been found in tree squirrels and directional selection favoring the evolution of cones with thicker scales may be a common outcome of the foraging behavior of birds.     

Morpho-anatomical affinities and evolutionary relationships of three paleoendemic podocarp genera based on seed cone traits

Background and AimsThe three relict genera Pherosphaera, Microcachrys and Saxegothaea in Podocarpaceae produce quite distinct seed cone types in comparison with other genera and do not form a clade along with Acmopyle. The detailed seed cone morpho-anatomy of these three relict genera and affinities with other podocarps are poorly known. This study aims to understand the seed cone morpho-anatomy and affinities among these three disjunct relict genera and with other podocarps.MethodsWe comparatively analysed the seed cone morpho-anatomical traits of the three podocarps genera and used ancestral state reconstruction to understand the evolution of these traits.Key ResultsWe described the seed cone morpho-anatomical structures of the three relict genera in detail. The three genera produce aggregated multiovulate cones. Both Microcachrys and Saxegothaea have an asymmetrical free cup-like epimatium. Both species of Pherosphaera lack an epimatium. The ancestral state reconstruction implies that the presence of an epimatium is an ancestral trait in podocarps and is independently lost in Pherosphaera and Phyllocladus. The seed cones are fleshy in Microcachrys and non-fleshy in Saxegothaea and Pherosphaera. The seed cone macrofossils of both extinct and living podocarps also show the presence of an epimatium and fleshiness in podocarps.ConclusionsAltogether, the morpho-anatomy suggests that Pherosphaera, Microcachrys and Saxegothaea present affinities with each other and other podocarps, but the reconstruction of the ancestral seed cone in Podcarpaceae is quite complex due to multiple convergent evolutions of several structures. These structures (e.g. epimatium, aril and receptaculum) are of low taxonomic value but of great evolutionary and ecological significance, and are responsive adaptations to ever-changing environmental conditions.     

Influence of host log age and refuge from natural enemies on colonization and survival of Phoracantha semipunctata

The pine cone weevil, Pissodes validirostris Gyll. (Coleoptera: Curculionidae), attacks seed cones of most Eurasian pine species, except these of Swiss stone pine (Pinus cembra L.). Behavioural responses of adult weevils to cone volatile emissions of Swiss stone pine and to those of a common host, mountain pine (Pinus uncinata Ram.), were compared in an olfactometer. Weevils were significantly attracted by the volatile blend emitted by mountain pine, but Swiss stone pine volatiles elicited an inverse response, with most weevils moving in the opposite direction to the odour source. However, the majority of second instar weevil larvae that were extracted from mountain pine cones and transferred into Swiss stone pine cones were capable of developing to the adult stage. This suggests that Swiss stone pine cones do not contain strong feeding deterrents that could prevent larval development. The possible factors involved in the absence of colonization of Swiss stone pine cones by cone weevils are discussed.     

PINUS AVONENSIS,A NEW SPECIES OF PETRIFIED CONES FROM THE OLIGOCENE OF WESTERN MONTANA

Fossils from the Oligocene of western Montana described in this treatment are the first structurally preserved ovulate cones of Pinus to be reported from the Tertiary of North America. They are about 5.5 cm long and have a maximum diam of 2.5 cm. Numerous scales are arranged spirally around the axis and each scale bears two winged seeds. The bract subtending the ovuli-ferous scale is 3-4 mm long and is free from the scale throughout its length. The pith and cortex of the axis are constructed of thick-walled parenchyma cells and 18-21 resin canals occur at the inner edge of the cortex. Resin canals entering the base of the ovuliferous scale are restricted to the abaxial side with vascular tissues occupying the adaxial side. Vascular strands near the tip of the scale are strongly rounded on the adaxial or phloem side. At the abaxial side of the tip of the ovuliferous scale is a broadly rhomboidal apophysis with a raised umbo that terminates in a short spine. The fossils differ from the several Recent cones examined in having fewer resin canals and biseriate rays in the secondary xylem of the cone axis. The shape of the cone, its anatomical features, and the morphology of the tip of the cone scale indicate affinity with the subgenus Diploxylon.     

SILICIFIED PINUS REMAINS FROM THE MIOCENE OF WASHINGTON

Silicified leaves, dwarf shoots, pollen cones, and seed cones of Pinus from a Late Miocene chert bed within the Yakima Basalt Formation near Yakima, Washington are interpreted as coming from a single new species, P. foisyi. The needles and dwarf shoots are those of a three-needle pine. The needles contain two to four medial resin canals, a biform hypodermis, and endodermal cells with uniformly thickened walls. The pollen cones are ellipsoidal and about 1 cm long, and many contain bisaccate pollen grains. The seed cones are at least 6 cm long and are slightly asymmetrical. The cone axis has a broad sclerotic outer cortex, and the seed wing extends from a thick parenchymatous base. The scale apex bears a conspicuously swollen projection. The foliage and seed cones are identifiable with the Subgenus Pinus, Section Pinus, Subsection Oocarpae independently of one another, and together indicate a fossil species related to the modem Californian closed cone pines. Pinus foisyi represents one of the earliest occurrences of cone asymmetry associated with this group. However, cone serotiny characteristic of the modem species appears to have evolved after the Late Miocene.     

Yezonia,a new section ofAraucaria (Araucariaceae) based on permineralized vegetative and reproductive organs ofA. vulgaris comb. nov. from the upper cretaceous of Hokkaido,Japan

A new specimen of an araucarian cone,Araucaria nihongii, was found attached to the vegetative organs ofYezonia vulgaris, and is described asAraucaria vulgaris comb. nov. Thick branches show characteristic bark structure with lenticular patches. Secondary wood is usually araucarioid. Leaves are arranged helically on shoots, which are imbricate, appressed and fused to surface of the stem. External and anatomical features of leaves closely resembleBrachyphyllum. The seed cone is spherical with winged bracts and thin ovuliferous scales. One seed is borne per cone-scale complex. The seed coat and nucellus wall show typical araucarian structure. An araucarian plant that boreBrachyphyllum-like foliage and aEutacta-like seed cone was predicted by Harris in 1979. This reconstructed plant,Araucaria vulgaris, supports this theory and proves the presence of an extinct characteristic-form of the genus. A new section of the genus was proposed forAraucaria vulgaris. Structure and Affinities of the Petrified Plants from the Cretaceous of Northern Japan and Saghalien XV, Consecutive number from previous paper (Nishidaet al. 1993).     

Tsuga seed cones from the late Paleogene of southwestern China and their biogeographical and paleoenvironmental implications

Six Tsuga ovuliferous/seed cone impression fossils were discovered from the late Eocene (34.6 ± 0.8 Ma) Lawula Formation in Mangkang County, eastern Tibet and the early Oligocene (32 ± 1 Ma) lacustrine deposits in Lühe Basin, Nanhua County, Yunnan Province. These two fossil sites are both located in southwestern China, ∼800 km apart from each other. These fossils represent the oldest records of this genus in southwestern China, even earliest reliable macrofossil records of this genus in the world. These well-preserved seed cones provide sufficient materials for the establishment of Tsuga asiatica Wu et Zhou n. sp. to accommodate five specimens, leaving one to be assigned to T. cf. dumosa Eichler (cf. Wu et Zhou). Both qualitative and quantitative comparisons with other cone fossils and cones of all living species of the genus suggested that T. asiatica shares more similarities with one of the basal species of the genus T. heterophylla. The discovery of late Paleogene macrofossil records of Tsuga in southwestern China supports the previous hypothesis of the early disposal routes of this coniferous genus predicted by phylogenetic analysis. The elevation ranges and the climate requirements of living species that are closely related to our fossils suggest that the southeastern edge of the Qinghai-Tibetan Plateau should be much warmer, and wetter in late Paleogene than nowadays.     

Asymmetrical development of biovulate cones resulting in uniovulate cones in Ephedra rhytidosperma (Ephedraceae)

Female cone morphology in Ephedra, including the number of initiated ovules and mature seeds per cone, provides important taxonomic characters used in sectional or species delimitation within Ephedra. Recent molecular phylogenies have indicated, however, that seed number per cone has changed repeatedly during the evolution of the genus. This study reports on the development of the female cone of E. rhytidosperma, based on a large sample of dissected cones studied under SEM. All cones were initially biovulate, and in the majority of cases, both female reproductive units (FRUs) developed a micropylar tube and formed mature seeds. In a few cases, the FRU pair developed asymmetrically in a cone, with one of them eventually aborting. There was no evidence of fusion of the FRU pair. Phylogenetically, E. rhytidosperma is in a clade with E. equisetina, which has uniovulate cones, and E. gerardiana and E. minuta, which have biovulate cones that also become unispermous via abortion. The biovulate condition may thus be ancestral in this clade.     

Comparison of survey methods for monitoring Clark's Nutcrackers and predicting dispersal of whitebark pine seeds

ABSTRACT Clark's Nutcrackers (Nucifraga columbiana) disperse seeds of whitebark pines (Pinus albicaulis) in western North America by their scatter‐hoarding behavior. Because of declines in whitebark pine, resource managers are seeking an effective means of monitoring nutcracker population trends and the probability of seed dispersal by nutcrackers. We tested the reliability of four survey techniques (standard point counts, playback point counts, line transects, and Breeding Bird Survey routes) for estimating population size by conducting surveys at sites where a portion of the nutcracker population was marked with radio transmitters. The efficacy of distance sampling, based on detection rates from our unadjusted surveys, was also assessed. We conducted counts of whitebark pine cones within stands and related the probability of seed dispersal within stands to cone production and nutcracker abundance. We conducted 70 h of surveys for Clark's Nutcrackers at eight sites from July through November in 2007 and 2009 and estimated cone densities at six of these sites. Detection rates for all survey techniques were low and variable and we detected an average of 5.6 nutcrackers per 30 min of survey time. We also found no difference in detection rates among survey types, although significantly more nutcrackers were detected during surveys conducted during the peak of whitebark pine cone harvest (P < 0.0001). Nutcracker abundance was not correlated with cone density (P= 0.29) and we observed nutcrackers pouching seeds at all sites. Thus, cone density did not provide reliable information on whether seed dispersal was likely to occur. We suggest that alternate methods be considered for monitoring populations and assessing seed dispersal probability because we did not reliably detect nutcrackers using conventional survey techniques and because nutcracker abundance was not correlated with cone density.     

A NEW SCIADOPITYACEOUS SEED CONE FROM THE UPPER CRETACEOUS OF HOKKAIDO,JAPAN

A sciadopityaceous seed cone, Sciadopityostrobus kerae, gen. et sp. nov., is described on the basis of a permineralized specimen from the Upper Cretaceous (Cenomanian–Coniacian) of Hokkaido, Japan. The peel method was applied for anatomical observations. The seed cone consists of a cone axis receiving numerous cone scale complexes that are arranged helically. Each complex generally has five inverted ovules that are oriented adaxially. The cone is similar to those of living Sciadopitys verticillata with respect to its peltate cone scale complex, with free apices of both bract and ovuliferous scale, trichomes on the bract, and a trifurcated ovuliferous scale strand. In the fossil, the bract and ovuliferous scale strands fuse with each other in the basal part of the cone scale complex, while S. verticillata bract and ovuliferous scale strands are derived separately from the vascular cylinder and remain separate throughout their length. The present specimen is one of the oldest records of structurally preserved cones that can be assigned to the family Sciadopityaceae.