Effect of defoliation on grass growth. A quantitative review

This study describes the seasonal patterns of seed and cone abortion in natural Pinus halepensis trees and assesses the effects of pollen quantity, tree size and cone location on seed and cone abortion. The cone-set ratio was similar to the expected value, based on literature on woody perennials. The seed-ovule ratio, however, was almost twice as high as its expected value. The investment in protective cone elements was high and did not vary with the seed-ovule ratio of a cone. Pollination factors influenced cone abortion, as indicated by the high abortion rate at the end of the pollination period. Furthermore, abortive cones had lower pollination rates and lower pollen loads than well developed cones. Effects of resource availability were assessed as effects of tree size and cone position on twigs. Small trees aborted more cones than big trees and cone abortion was higher in apical cones than in basal cones. Abortion in P. halepensis is selective, mediated by both pollen quantity and resource availability. The relative importance of pollen and resources is suggested to be flexible, probably varying between trees and years. The high seed-ovule ratio of P. halepensis is the result of high pollination rate and selective abortion. The selective abortion might be due to the high allocation to protective cone elements relative to the allocation to seeds.     

Interactions among moths, crossbills, squirrels, and lodgepole pine in a geographic selection mosaic

Repeated patterns among biological communities suggest similar evolutionary and ecological forces are acting on the communities. Conversely, the lack of such patterns suggests that similar forces are absent or additional ones are present. Coevolution between a seed predator, the red crossbill (Loxia curvirostra complex), and lodgepole pine (Pinus contorta var. latifolia) exemplifies the ecological and evolutionary predictions for coevolving systems. In the absence of another seed predator and preemptive competitor (pine squirrels Tamiasciurus hudsonicus), natural selection by crossbills results in the evolution of larger cones with thicker distal scales, while relaxation of selection by squirrels results in the evolution of cones with more seeds and a greater ratio of seed mass to cone mass. However, in one range, the Little Rocky Mountains, distal scale thickness has diverged as expected but cone size has not. In these mountains seed predation by lodgepole pine cone borer moths (Eucosma recissoriana) was about 10 times greater than in other ranges lacking squirrels. We quantified moth predation and cone traits and found that moths select for smaller cones with fewer seeds. Thus, selection by moths in the Little Rocky Mountains counters both selection by crossbills for large cone size and relaxation of selection by squirrels favoring more seeds per cone and accounts for the relatively small and few-seeded cones in these mountains. It is also apparent that selection by crossbills changes seed defenses in a manner that favors seed predation by moths, whereas selection by squirrels likely reduces such predation. These results demonstrate the importance of considering the evolutionary consequences of community context in locally evolved (coevolved) traits and interactions.     

濒危植物梵净山冷杉的球果与种子性状研究北大核心CSCD

梵净山冷杉(Abies fanjingshanensis)为国家一级保护濒危植物。为了揭示梵净山冷杉球果发育和成熟过程的性状特征,以确定成熟球果的适宜采种期。该研究对梵净山冷杉自然生长区内成年结实母树的球果进行跟踪观测和定期采集,通过物理解剖和形态学参数测定,比较分析不同采种期梵净山冷杉的球果、种鳞和种子的性状差异。结果显示:(1)梵净山冷杉的球果每年7月中旬开始形成,10月中旬开始成熟,发育早期至发育后期球果的长度和宽度显著增加,发育后期至成熟期形态和颜色均无明显变化。(2)成熟球果平均长、宽、鲜重、干重和相对含水量分别为7.18 cm、3.84 cm、36.98 g、20.33 g和45.06%。(3)成熟球果的平均种鳞层数为30.76层,种鳞总数为250.67片,平均出种量436.67粒,平均种子饱满率82.49%。(4)成熟球果基部、中部、上部的种子性状不同,且球果中部的种子性状参数最大,饱满率最高,种子平均长、宽、厚分别为9.14、2.30、2.37 mm,千粒重11.44 g。研究表明,梵净山冷杉的球果从形成至成熟过程约3个月,成熟球果的最佳采种时间为10月下旬,发育后期和成熟球果的形态和颜色差异不明显,不能以颜色和形态作为球果成熟的判定依据;种鳞与球果的发育和成熟同步,球果中部的种鳞能够完全发育,且形状参数最大,孕育的种子最饱满;梵净山冷杉成熟球果出种量大、饱满率高,但是种子小、重量轻。     

THE IMPACT OF TREE SQUIRRELS (TAMIASCIURUS) ON LIMBER PINE SEED DISPERSAL ADAPTATIONS

Tree squirrels (Tamiasciurus) are important selective agents on conifer reproductive strategies (Smith 1970, 1975). Although this is well established for wind-dispersed pines, the impact of tree squirrels on bird-dispersed pines has been largely ignored. I assessed the impact of tree squirrels on the allocation of reproductive energy in the bird-dispersed limber pine (Pinus flexilis) by comparing its cone and seed traits from three sites in the Rocky Mountains where tree squirrels (Tamiasciurus) are present to those from three mountain ranges in the Great Basin where tree squirrels are absent. As predicted, differences between the two regions in individual cone and seed traits are consistent with the hypothesis that tree squirrels are important selective agents on these traits. In the absence of tree squirrels, limber pine allocates more than twice as much energy to kernel compared with that invested in putative seed defenses (cone, resin, and seed coat) as does limber pine where tree squirrels are present. Such a large difference is particularly striking, because tree squirrels may have become extinct in the Great Basin in only the last 12,000 yr. Although many factors influence the allocation of energy to cones and seeds, no single factor other than the presence of tree squirrels is compatible with the large and consistent differences between limber pine in the Rocky Mountains and Great Basin. These results show that tree squirrels are an important constraint on the evolution of cone and seed traits that promote the dispersal of seeds by birds.     

Measuring the impact of Leptoglossus occidentalis (Heteroptera: Coreidae) on seed production in lodgepole pine using an antibody-based assay

We measured the impact of Leptoglossus occidentalis on seed production in lodgepole pine, Pinus contorta variety latifolia Engelmann, using an antibody marker developed to detect residual saliva in fed-on seeds. Nymphs, adult females, and adult males were caged on cones during early, mid- and late season cone development. Individual analysis of 12,887 seeds extracted from 365 cones revealed that 37.3% seeds tested positive for seed bug saliva. The antibody assay was 38 times more effective than radiography at detecting seed bug damage. Radiography can detect partially emptied seed but cannot discriminate between aborted seeds and those emptied by seed bugs. The antibody marker was least sensitive in detecting early season damage compared with mid- and late season damage. We hypothesize that residual saliva in seeds fed on early in the season was either absorbed by the damaged seed or degraded over time. Early season feeding resulted in the greatest number of seeds fused to cone scales and the extraction efficiency for cones exposed to feeding during this time was reduced by 64% compared with control cones. Adding fused seeds to antibody-positive seeds raised the proportion of damaged seeds to 48.3%. At all stages of cone development, adult females were the most destructive life stage, damaging up to two seeds per day late in the season. When seed losses were adjusted to damage per degree-day, female damage was greatest early in the season, while males caused the same amount of damage regardless of cone development period. The results of the antibody assay provide baseline data for developing damage prediction formulae, and establish L. occidentalis as a potentially serious pest in lodgepole pine seed orchards.     

Oviposition Sites of the Cypress Seed Bug <Emphasis Type="Italic">Orsillus maculatus</Emphasis> and Response of the Egg Parasitoid <Emphasis Type="Italic">Telenomus</Emphasis> gr. <Emphasis Type="Italic">Floridanus</Emphasis>

Phytophagous insects have many strategies to escape parasitoids, for example by hiding eggs into plant tissues, but oviposition in holes made by another insect is rather scarce. The cypress seed bug Orsillus maculatus Fieber (Heteroptera: Lygaeidae) is strictly dependent on the availability of cones of Cupressus sempervirens L. to oviposit. Females lay eggs either in exit holes cut through the cone scale by emerging adults of the cypress seed chalcid, Megastigmus wachtli Seitner (Hymenoptera: Torymidae), or on the inner side of scales of partly open cones. A recently discovered egg parasitoid belonging to the genus Telenomus has been shown to attack bug eggs in both oviposition sites. In this paper we investigated the parasitoid performance according to oviposition sites. Field samplings were conducted in two evergreen cypress orchards located in the south of France. The distribution and condition of the egg patches were compared between the two locations and oviposition sites. Seed bugs preferred to oviposit in emergence holes of M. wachtli, and parasitoid performance was higher in eggs laid on cone scales. The chalcid emergence holes seemed to ensure bug eggs with enemy-free space. Oviposition site selection could be an adaptive strategy to escape parasitoid attack.     

Seed cone anatomy of Cheirolepidiaceae (Coniferales): Reinterpreting Pararaucaria patagonica Wieland

? Premise of the study: Seed cone morphology and anatomy reflect some of the most important changes in the phylogeny and evolutionary biology of conifers. Reexamination of the enigmatic Jurassic seed cone Pararaucaria patagonica reveals previously unknown systematically informative characters that demonstrate affinities with the Cheirolepidiaceae. This paper documents, for the first time, internal anatomy for seed cones of this important extinct Mesozoic conifer family, which may represent the ghost lineage leading to modern Pinaceae. ? Methods: Morphology and anatomy of cones from the Jurassic La Matilde Formation in Patagonia are described from a combination of polished wafers and thin section preparations. New photographic techniques are employed to reveal histological details of thin sections in which organic cell wall remains are not preserved. Specific terminology for conifer seed cones is proposed to help clarify hypotheses of homology for the various structures of the cones. ? Key results: Specimens are demonstrated to have trilobed ovuliferous scale tips along with a seed enclosing pocket of ovuliferous scale tissue. Originally thought to represent a seed wing in P. patagonica, this pocket-forming tissue is comparable to the flap of tissue covering seeds of compressed cheirolepidiaceous cones and is probably the most diagnostic character for seed cones of the family. ? Conclusions: Pararaucaria patagonica is assigned to Cheirolepidiaceae, documenting anatomical features for seed cones of the family and providing evidence for the antiquity of pinoid conifers leading to the origin of Pinaceae. A list of key morphological and anatomical characters for seed cones of Cheirolepidiaceae is developed to facilitate assignment of a much broader range of fossil remains to the family. This confirms the presence of Cheirolepidiaceae in the Jurassic of the Southern Hemisphere, which was previously suspected from palynological records.     

Cones structure and seed traits of four species of large‐seeded pines: Adaptation to animal‐mediated dispersal

Seed dispersal selection pressures may cause morphological differences in cone structure and seed traits of large‐seeded pine trees. We investigated the cone, seed, and scale traits of four species of animal‐dispersed pine trees to explore the adaptations of morphological structures to different dispersers. The four focal pines analyzed in this study were Chinese white pine (Pinus armandi), Korean pine (P. koraiensis), Siberian dwarf pine (P. pumila), and Dabieshan white pine (P. dabeshanensis). There are significant differences in the traits of the cones and seeds of these four animal‐dispersed pines. The scales of Korean pine and Siberian dwarf pine are somewhat opened after cone maturity, the seeds are closely combined with scales, and the seed coat and scales are thick. The cones of Chinese white pine and Dabieshan white pine are open after ripening, the seeds fall easily from the cones, and the seed coat and seed scales are relatively thin. The results showed that the cone structure of Chinese white pine is similar to that of Dabieshan white pine, whereas Korean pine and Siberian dwarf pine are significantly different from the other two pines and vary significantly from each other. This suggests that species with similar seed dispersal strategies exhibit similar morphological adaptions. Accordingly, we predicted three possible seed dispersal paradigms for animal‐dispersed pines: the first, as represented by Chinese white pine and Dabieshan white pine, relies upon small forest rodents for seed dispersal; the second, represented by Korean pine, relies primarily on birds and squirrels to disperse the seeds; and the third, represented by Siberian dwarf pine, relies primarily on birds for seed dispersal. Our study highlights the significance of animal seed dispersal in shaping cone morphology, and our predictions provide a theoretical framework for research investigating the coevolution of large‐seeded pines and their seed dispersers.     

Anatomic basis and insulation of serotinous cones in Pinus halepensis Mill

Pinus halepensis Mill., a widespread, low elevation conifer common in Mediterranean Basin, shows a dual reproductive strategy: post-fire obligate seeder (from serotinous cones) and an early coloniser (from non-serotinous cones). Release of seeds encased in serotinous cones is induced either by fire (pyriscence, serotiny or bradychory) or by drying (xeriscence). Morphological differences in serotinous and non-serotinous cones in natural populations of P. halepensis in Southeastern Italy were analyzed. Relationships between tree size (diameter class) and serotiny were checked by counting and sampling serotinous and non-serotinous cones. The macro and microscopic characteristics that could affect cones’ opening were measured in sampled cones. Protection against high temperatures offered by wood scales was also evaluated by applying different temperatures and time exposures, and following the inner thermal raise. Results showed that non-serotinous cones had bigger resin ducts and more separate scales. Also it was highlighted that ovuliferous scales of serotinous cones were bigger and thicker. These scales had more lamellated (multilayered) sclereid cells, and were significantly thinner with a shorter lumen diameter. Continuous temperature-monitoring heat tests inside cones showed that temperatures close to the cone axis were rather low, so seed germination was not influenced. Results confirm that serotinous cones are more compact, rigid and consistent than non-serotinous cones. These characteristics explain the lower insulation, seed protection and the ease opening of non-serotinous cones as well. In conclusion, opening mechanism of pinecone scales under the effect of fire or dry conditions seem related to anatomic differences and it provides seeds with an efficient protection against heat.     

Variation in seed packaging of a fleshy‐fruited conifer provides insights into the ecology and evolution of multi‐seeded fruits

• The study of intraspecific seed packaging (i.e. seed size/number strategy) variation across different populations may allow better understanding of the ecological forces that drive seed evolution in plants. Juniperus thurifera (Cupressaceae) provides a good model to study this due to the existence of two subspecies differentiated by phenotypic traits, such as seed size and cone seediness (number of seeds inside a cone), across its range.
• The aim of this study was to analyse seed packaging (seed mass and cone seediness) variation at different scales (subspecies, populations and individuals) and the relationship between cone and seed traits in European and African J. thurifera populations.
• After opening more than 5300 cones and measuring 3600 seeds, we found that seed packaging traits followed different patterns of variation. Large‐scale effects (region and population) significantly contributed to cone seediness variance, while most of the seed mass variance occurred within individuals. Seed packaging differed between the two sides of the Mediterranean Sea, with African cones bearing fewer but larger seeds than the European ones. However, no differences in seed mass were found between populations when taking into account cone seediness. Larger cones contained more pulp and seeds and displayed a larger variation in individual seed mass.
• We validated previous reports on the intraspecific differences in J. thurifera seed packaging, although both subspecies followed the same seed size/number trade‐off. The higher seediness and variation in seed mass found in larger cones reveals that the positive relationship between seed and cone sizes may not be straightforward.We hypothesise that the large variation of seed size found within cones and individuals in J. thurifera, but also in other fleshy‐fruited species, could represent a bet‐hedging strategy for dispersal.

     

Branching habit and the allocation of reproductive resources in conifers

Background and Aims Correlated relationships between branch thickness, branch density, and twig and leaf size have been used extensively to study the evolution of plant canopy architecture, but fewer studies have explored the impact of these relationships on the allocation of reproductive resources. This study quantifies pollen cone production in conifers, which have similar basic reproductive biology but vary dramatically in branching habit, in order to test how differences in branch diameter influence pollen cone size and the density with which they are deployed in the canopy. Methods Measurements of canopy branch density, the number of cones per branch and cone size were used to estimate the amount of pollen cone tissues produced by 16 species in three major conifer clades. The number of pollen grains produced was also estimated using direct counts from individual pollen cones. Key Results The total amount of pollen cone tissues in the conifer canopy varied little among species and clades, although vegetative traits such as branch thickness, branch density and pollen cone size varied over several orders of magnitude. However, branching habit controls the way these tissues are deployed: taxa with small branches produce small pollen cones at a high density, while taxa with large branches produce large cones relatively sparsely. Conclusions Conifers appear to invest similar amounts of energy in pollen production independent of branching habit. However, similar associations between branch thickness, branch density and pollen cone size are seen across conifers, including members of living and extinct groups not directly studied here. This suggests that reproductive features relating to pollen cone size are in large part a function of the evolution of vegetative morphology and branching habit.     

Patterns of exploitation of annually varying Pinus sylvestris cone crops by seed‐eaters of differing dispersal ability

Food consumption by animals depends on functional and numerical responses, and particularly the latter for food specialists. Some birds and mammals are conifer seed specialists and are likely to exploit the varying seed production in different ways according to differences in dispersal capabilities. This in turn may determine which species is more likely to affect the evolution of cone morphology. This study examined the inter‐annual pattern of foraging on Scots pine Pinus sylvestris cones by crossbills Loxia spp. (highly dispersive) and red squirrels Sciurus vulgaris (weakly dispersive) over 16 yr in three stands of ancient native pines at Abernethy Forest in Highland Scotland. There were synchronous annual variations in cone production across the stands, and an indication of a three‐year cycle. The number of cones taken by crossbills was correlated with cone production, indicating a numerical response by the birds, up to a certain limit of cone production. By contrast, the number of cones taken by red squirrels was not correlated with cone production. Rather, the percentage of cones taken by squirrels was high when cone production was low, and low when production was high. There was also a long‐term decline in the number of cones taken by squirrels, suggesting a decline in squirrel numbers. Although a high percentage of cones was removed from some cohorts on some trees, either by crossbills (maximum of 94.8%) or red squirrels (100%), the mean percentage of cones taken by crossbills from trees was small, ranging from 3.7 to 17.1% across all cone cohorts. For red squirrels, mean values ranged from 0.1 to 46.1% across all cohorts. However, given that crossbills can track changes in cone production by rapid numerical responses (i.e. through migration), and take a larger percentage of cones from cohorts of high production (10.7%), compared with red squirrels (3.6%), crossbills may be more important in driving the evolution of cone morphology because future trees are more likely to come from cohorts of high cone production.     

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.     

秦岭山地太白红杉种群种实性状的生态可塑性研究

研究了太白红杉6个种群中13个种实性状的变异情况.结果表明,太白红杉种实性状在种群内和种群间均存在广泛的变异,除种子宽和种鳞宽的差异在种群间不显著外,其它11个性状在种群间差异均极显著;种群间种鳞长、种鳞宽、球果长宽比、种翅长和种翅宽的变异程度比种群内大,其它8个性状在种群内的变异程度比种群间大.海拔对太白红杉种实性状的影响比较明显,其中种子宽、种鳞长、种翅宽、种鳞宽、球果长、球果宽和种鳞数与海拔呈负相关,种子宽和球果宽相关性显著,种鳞长、种翅宽、种鳞宽、球果长和种鳞数相关性极显著;坡度与种鳞长宽比呈负相关;年降水与球果宽呈正相关.种子长宽比和种翅长宽比与海拔呈负相关.     

The seed cone Eathiestrobus gen. nov.: Fossil evidence for a Jurassic origin of Pinaceae

? Premise of the study: Pinaceae and nonpinoid species are sister groups within the conifer clade as inferred from molecular systematic comparisons of living species and therefore should have comparable geological ages. However, the fossil record for the nonpinoid lineage of extant conifer families is Triassic, nearly 100 million years older than the oldest widely accepted Lower Cretaceous record for Pinaceae. An anatomically preserved fossil conifer seed cone described here extends the stratigraphic range of Pinaceae nearly 30 million years, thus reducing the apparent discrepancy between evidence from the fossil record and inferences from systematic studies of living species. ? Methods: Material was prepared as serial thin sections by the cellulose acetate peel technique, mounted on microscope slides, and viewed and photographed using transmitted light. ? Key results: A large cylindrical cone consisting of bract-scale complexes that diverge from the cone axis in a helical phyllotaxis has bracts and scales that separate from each other in the midregion and are of equal length and of nearly equal width. The cone has two inverted and winged seeds that are attached to the adaxial surface of each cone scale and, thus, represents an early member of the Pinaceae. ? Conclusions: Eathiestrobus mackenziei gen. et sp. nov. extends the fossil record for well-documented members of the family Pinaceae from the Lower Cretaceous to the Kimmeridgian Stage of the Upper Jurassic. This species also clarifies the set of characters that are diagnostic for seed cones of Pinaceae and reveals possible plesiomorphic characters for seed cones of the family.     

Impact of Leptoglossus occidentalis (Hemiptera: Coreidae) on Douglas-fir seed production

We investigated the effect of feeding by the western conifer-seed bug, Leptoglossus occidentalis Heidemann, on seed production in developing cones of coastal Douglas-fir, Pseudostuga menziesii (Mirb.) Franco, with respect to seed bug life stage and sex (nymphs, adult females, and adult males) and timing of feeding (early, mid-, and late season cone development). Feeding by females on caged cones for a 2-wk period during late season cone development reduced the proportion of full seeds in cones by approximately 70% compared with caged control cones. There was no significant difference among nymphs, adult females, and adult males with respect to the proportion of empty or partially fed-upon seeds produced during the same feeding period. Feeding by nymphs for 2 wk early in the season resulted in a threefold increase in the number of unextractable seeds fused to cones compared with the control. Weight measurements of harvested seeds indicated that radiography is an accurate tool to distinguish among Douglas-fir seeds that have sustained light, moderate, or severe damage. Determining the full impact of L. occidentalis on conifer seed production will require the development of a reliable method to distinguish between naturally aborted seeds and seeds emptied through feeding by seed bugs.     

The influence of a competitor on the geographic mosaic of coevolution between crossbills and lodgepole pine

The geographic mosaic theory of coevolution posits that the form of selection between interacting species varies across a landscape with coevolution important and active in some locations (i.e., coevolutionary hotspots) but not in others (i.e., coevolutionary coldspots). We tested the hypothesis that the presence of red squirrels (Tamiasciurus hudsonicus) affects the occurrence of coevolution between red crossbills (Loxia curvirostra complex) and Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia) and thereby provides a mechanism giving rise to a geographic mosaic of selection. Red squirrels are the predominant predispersal seed predator and selective agent on lodgepole pine cones. However, in four isolated mountain ranges east and west of the Rocky Mountains, red squirrels are absent and red crossbills are the main predispersal seed predator. These isolated populations of pine have apparently evolved without Tamiasciurus for about 10,000 to 12,000 years. Based on published morphological, genetic, and paleobotanical studies, we infer that cone traits in these isolated populations that show parallel differences from cones in the Rocky Mountains have changed in parallel. We used data on crossbill and conifer cone morphology and feeding preferences and efficiency to detect whether red crossbills and lodgepole pine exhibit reciprocal adaptations, which would imply coevolution. Cone traits that act to deter Tamiasciurus and result in high ratios of cone mass to seed mass were less developed in the isolated populations. Cone traits that act to deter crossbills include larger and thicker scales and perhaps increased overlap between successive scales and were enhanced in the isolated populations. In the larger, isolated mountain ranges crossbills have evolved deeper, shorter, and therefore more decurved bills to exploit these cones. This provides crossbills with higher feeding rates, and the change in bill shape has improved efficiency by reducing the concomitant increases in body mass and daily energy expenditures that would have resulted if only bill size had increased. These parallel adaptations and counter adaptations in red crossbills and lodgepole pine are interpreted as reciprocal adaptations and imply that these crossbills and pine are in coevolutionary arms races where red squirrels are absent (i.e., coevolutionary hotspots) but not where red squirrels are present (i.e., coevolutionary cold-spots).     

The effect of spruce cone insects on seed production in Switzerland

Abstract: In 1989 and 1990, spruce cones were harvested at 29 sites distributed over the five main geographic regions of Switzerland, i.e. Jura, Central Plateau, North, Central and South Alps. The presence of insects and the species-specific consumption of seeds was determined for each cone by examination of the median longitudinal section. Seven seed-feeding species ( Assara terebrella , Cydia strobilella , Dioryctria abietella , Eupithecia abietaria , Megastigmus strobilobius , Plemeliella abietina , Strobilomyia anthracina ) and the spruce cone gall midge Kaltenbachiola strobi were found in the samples. The proportions of infested cones ranged from 36 to 100%. The regions did not show significant differences in terms of infestation rates, whereas differences between sites within the regions were highly significant. Seed loss ranged from 2 to 100%, its geographical distribution exhibiting a similar pattern to the infestation rates. Considering the number of infested cones, C. strobilella was the most abundant species, followed by K. strobi and P. abietina . Cones infested by conospermatophagous species ( C. strobilella , D. abietella , E. abietaria ) showed higher seed losses than those infested by spermatophages ( M. strobilobius , P. abietina ). Due to the ubiquitous occurrence of C. strobilella , its high infestation rates and the resulting seed losses caused by its offspring, this species has to be considered as the most important insect seed predator in Switzerland. A multiple linear regression analysis was carried out for each region to simultaneously explore the impact of site and tree variables as well as the effect of insect attack on the yield of viable seeds. The results indicate that site and tree conditions contributed more to the variability of seed yield than insect presence in a cone. The model predicted significant effects of insect attack mainly for cones with total seed numbers that were higher than the average.     

Frenelopsis alata and its microsporangiate and ovuliferous reproductive structures from the Cenomanian of Bohemia (Czech Republic, Central Europe)

The conifer, Frenelopsis alata (K. Feistmantel) E. Knobloch (Cheirolepidiaceae), occurring mostly in the Cenomanian of Europe, is revised on the basis of the type material. Its comparison with relevant species of Frenelopsis is discussed.The ovuliferous cone associated with the genus Frenelopsis is recorded for the first time. For the associated ovuliferous cones of Frenelopsis, a new genus, Alvinia, is introduced in a new combination for the type: Alvinia bohemica (Velenovsky) comb. n. Its association with Frenelopsis alata is based on the presence of Classopollis pollen adhering to ovuliferous cone scales, and the same type of pollen found in the microsporangiate cone of F. alata, the same cuticle pattern present on ovuliferous cones, sterile twigs and microsporangiate cones of F. alata, and also the co-occurrence of ovuliferous cones or their scales and sterile twigs of F. alata.Large ovuliferous cones of Alvinia bohemica are formed by helically arranged ovuliferous scales subtended by bracts. Each ovuliferous cone scale displays one or two seeds covered by a covering flap, and three appendages, which form distally a funnel-like structure lined in its inner part by long trichomes. Numerous pollen grains of Classopollis adhere to the trichomes, and the structure is considered to function as a protostigmatic area.The ovuliferous cones of Alvinia differ from similar cones of the Cheirolepidiaceae, Hirmeriella and Tomaxellia, mainly in a high state of unification of the ovuliferous cone scale, reduction of appendages and in a presence of the protostigmatic funnel-like structure.The ovuliferous cones, Alvinia bohemica, rarely occur intact, so it is assumed that they disintegrate when mature. It seems likely that they were not woody. This assumption is supported by the flattened appearance of cones and their cone scales in the sediment, their flexibility and the absence of massive coaly matter known from cones of the Taxodiaceae and Cupressaceae. It is proposed that this type of ovuliferous cone scale indicates a specialized type of pollination. In addition, it is suggested that cone scales enclosing seeds play an important role in propagation.     

高山松种实性状与生殖适应性

高山松(Pinus densata)是云南松(P. yunnanensis)与油松(P. tabulaeformis)自然杂交产生的二倍体杂种,是植物同倍体杂交物种形成的典型案例。高山松分布于青藏高原东南缘,占据了两个亲本种都不能正常生长的高海拔地带。良好生殖适应性的产生是物种形成的关键,但对高山松在高海拔生境下生殖适应性的变异与杂交起源遗传背景关系的研究尚未见报道。该研究通过对来自该种分布区内6个代表居群样本、13个反映高山松生殖适应性的种实性状的分析,探讨了其生殖性状在居群水平的变异模式,揭示了具有不同遗传组成和起源历史的高山松居群的生殖特性。研究结果表明:6个研究居群在种实性状上存在显著差异;可育种鳞数、总种鳞数、总种子数、可育种鳞率和可育种鳞密度等与纬度呈显著负相关关系;球果结种率与经度和生态梯度值呈显著的正相关关系;胚珠败育率则与经度和生态梯度值呈显著的负相关关系;球果长度也呈现了与经度的显著负相关。高山松球果结种率很高,平均为74%;球果的结种数、种子长和种翅长与云南松、油松及其它松属双维管亚属多数种接近;球果长度、球果总种鳞数接近油松的报道。这些结果表明,高山松在亲本种不能正常生长、繁衍的高原环境下具备正常的生殖能力;其种实特征在居群间分化显著并呈现一定的地理梯度变化。居群间的显著差异反映了其杂种起源特性,与高山松居群的遗传背景有关,也反映了处在异质生态环境中的高山松居群具有各自的生殖生态进化趋向。