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.

     

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.     

High resolution analysis of mating systems: inbreeding in natural populations of Pinus radiata

Pinus radiata has a history of population bottlenecks and is currently restricted to five relatively small populations, three in mainland California, and two on islands off the coast of Baja California. Using highly polymorphic microsatellite markers and a newly developed statistical approach, we were able to estimate individual inbreeding coefficients and can thus analyse the mating system with high resolution. We find a bimodal distribution of inbreeding coefficients: most individuals result from selfing whereas few (in the mainland populations) to a modest number (in the island populations) are likely selfed. In most other pine species and presumably in the ancestral P. radiata population, occurrence of mature selfed individuals would be impossible because of the high genetic load. We therefore conclude that inbreeding depression has been purged in P. radiata and that the mating system has changed as a consequence.     

PINUS × CRITCHFIELDII,A LATE PLEISTOCENE HYBRID PINE FROM COASTAL SOUTHERN CALIFORNIA

Fossil cones buried in an old landslide at San Clemente are dated at 30,000–34,000 BP (years before present). They generally resemble cones of the natural hybrid Pinus radiata × P. attenuata at Año Nuevo, and also the artificial hybrid P. × attenuradiata. But differences in cone shape, size, and apophyses suggest that the fossil P. × critchfieldii Axelrod and T. G. Hill represents a hybrid of populations of P. radiata and P. attenuata then in coastal southern California. This agrees with population differences between these species in central and southern California during the Pleistocene. Regional comparisons of Late Pleistocene floras indicate that at San Clemente mean annual rainfall was about 580–635 mm, or twice that of today, mean annual temperature was about 2.5 C cooler (13.5 vs. 16.0 C), the mean monthly range was lower (5.9 vs. 7.7 C), the growing season was about 40 days shorter (170 vs. 210 days), and equability was higher (M 72 vs. M 67). Forest was replaced by coastal sage scrub and grassland along the San Clemente coast as drier, warmer climate developed after 12,000 BP.     

Seed quality in hybrid swarm populations of Pinus mugo Turra and P. sylvestris L.

Crop characteristics involving cone size, average number of seeds per cone, and seed germinability have been studied for three hybrid swarm populations of Pinus mugo and P. sylvestris, two pure P. mugo populations and one pure P. sylvestris population, in northern Slovakia. Generally, reproduction capability characteristics were significantly reduced in hybrid swarms in relation to the control populations. Although there were no consistent differences between hybrid swarms and control populations in the average number of seeds per cone or cone size, seed germination rates and full seed production in hybrid swarms were lower than for the control population of P. sylvestris and one of the pure populations of P. mugo. Based on these data a conclusion has been drawn postulating partial rather than full fertility of the hybrid swarm populations of P. mugo and P. sylvestris.     

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。     

红松种子园种群表型多样性研究

为揭示红松（Pinus koraiensis Sieb.et Zucc.）不同种群的表型分化程度和变异规律,以吉林省露水河红松种子园6个种群红松为研究对象,采用巢式方差分析、多重比较、变异系数和聚类分析等方法对红松的叶片、球果和种子共15个表型性状进行了系统分析。结果表明：（1）红松15个表型性状在种群间和种群内均存在着极显著的差异,红松种群遗变异比较丰富,在松属植物中属于中等水平,其中纬度最低的露水河种群在其中10个表型性状均值上表现出最高值;（2）红松种群间的表型分化系数（Vst）均值为12.39%,种群内的变异（87.61%）大于种群间的变异（12.39%）,种群内的变异是主要变异来源;（3）各表型性状平均变异系数为13.28%,变幅为6.16%-31.48%,叶片、球果、种子的平均变异系数依次为：球果17.86% > 针叶11.19% > 种子9.87%,种子性状最小,成为最稳定的表型性状,带岭和丰林种群表型性状遗传多样性要高于其他种群;（4）利用欧氏平均距离对红松种群进行UPGMA聚类分析,红松种群的表型性状按照地理距离而聚类,可将红松种子园6个种群分为4类,其表型性状跟地形（东北山脉）有一定的契合。红松种群具有中等水平的表型遗传多样性,种群间和种群内均存在丰富的表型变异,研究结果为顺利开展红松种质资源收集、保存,以及良种选育等工作提供依据。     

GEOGRAPHICAL CORRELATION OF MORPHOLOGICAL VARIATION IN SILVETIA COMPRESSA (FUCACEAE,FUCALES, PHAEOPHYCEAE)

Silvetia compressa ( J. Agardh) Serrão et al. is a common member of the upper intertidal fucoid community on the Pacific coast of America from Humboldt County, California, to Punta Baja, Baja California, Mexico. A relatively narrow range of morphological variability is exhibited by most mainland populations, regardless of latitude, but some mainland populations and all insular populations participate in a complex pattern that we have attempted to analyze. A few populations on the Monterey Peninsula in which the fronds are atypically delicate were described by Setchell & Gardner as f. gracilis, to which was assigned a population from Santa Catalina Island. After comparing populations from various parts of the range of the species, including all of the Channel Islands, we conclude that two subspecies may be recognized. In subsp. compressa, which includes f. gracilis as a growth form and occurs chiefly on the mainland, the frond is robust with long tapered receptacles. In the variant subspecies, which is chiefly insular but also occurs on the coast of northern Baja California, the typical frond has slender axes as in f. gracilis, but is more densely branched and has short ellipsoidal receptacles. Comparison of nucleotide sequences from the ITS regions of rDNA revealed an identical pattern for subsp. compressa from Baja California and central California, including populations assignable to f. gracilis. By contrast, the pattern for the variant subspecies differed by 2 bp (0.3%) from that of subsp. compressa.     

GEOGRAPHIC VARIATION IN REPRODUCTIVE BEHAVIOR AND SIZE STRUCTURE OF THE AUSTRALIAN CYCAD MACROZAMIA COMMUNIS (ZAMIACEAE)

Macrozamia communis is a common cycad that occurs in eastern New South Wales, Australia, where it forms an understory in various forest or woodland communities. Leaf number of reproductive individuals as well as of nonreproductive ones, taken as an indication of size class, varies clinally from north to south, with reproductive individuals in southern populations having mean leaf numbers about 12 times greater than those at the northern end of the range of this species. Female cone size and mature ovule weight appear to vary also along this gradient. Using leaf number as a predictor of sexual maturity, from 37 to 100% of the individuals present in a population were judged to be mature. The percent of mature individuals participating in the 1987 coning episode ranged from none in three southern populations to 68% elsewhere. In 9 of 10 samples more males than females participated in the 1987 coning episode, with a collective sex ratio slightly male biased for all populations sampled. In three population samples average cone numbers for male and female plants were equal. In two samples males produced more cones per plant than females. Leaf numbers of males and females in individual populations were equal. Few individuals producing cones in 1987 showed evidence of recent prior cone production, suggesting that successive cone production by both sexes is separated by an interval of several years. The periodicity with which fire is used as a fire suppression method may affect the frequency of cone production in some populations, perhaps preventing it entirely. The apparent absence of recruitment in some populations may result from long-term seed collection by humans for horticultural purposes as well as the very poor seed dispersal that appears to characterize this cycad throughout its range. The weevil Tranes internatus was observed occupying the internal tissues of female cones; this insect may be involved in the pollination of M. communis.     

Reproductive traits of two closely related species-pairs on adjacent,different soil types in South African Fynbos

High levels of edaphic endemism and soil-related beta-diversity in Agulhas Plain fynbos communities suggest that reproductive traits of plants growing on different fynbos soils would be related to differences in soil regime. We investigated reproductive traits in two closely related Proteaceae species-pairs growing on adjacent soil types: Protea obtusifolia and Leucadendron meridianum occurring in shallow pockets of limestone-derived soils, and P. susannae and L. coniferum on the adjacent, uniformly deep colluvial sands.We found that species growing on the limestone soil comprised smaller plants, with fewer cones and seeds per plant, than species on the colluvial sands. These differences suggest that the small soil pockets of limestone soil limit plant size, in turn limiting the number of reproductive structures. Annual variation in cones and seed was the same in all species. There were no consistent trends in degree of serotiny, or sex allocation across soil types. The higher cone and seed predation levels of both limestone species than the colluvial sands species were ascribed to the higher plant densities of the former leading to lower insect search times. It was concluded that there were no overall patterns in reproductive traits that could be ascribed to differences in soil regime, other than through size-related effects. Fire regime is likely to have played a more important role in determining reproductive traits.Department of Statistical Sciences, University of Cape Towm     

Biochemical genetics of some seed proteins of Pinus radiata

In a high-salt soluble fraction of the total protein from single seeds of Pinus radiata, up to 45 polypeptides were resolved on SDS-polyacrylamide gels. At least one-fifth of these polypeptides showed variation between seeds. In the 27,000–29,000 dalton region, two polypeptides were inherited as codominant alleles at a single locus and were shown to assort independently of another seed protein locus and three allozyme loci. A survey of 120 individuals from the five known native populations of P. radiata in California detected only the 27K and 29K alleles at the locus. In all populations, the 29K allele predominated, and the two island populations were monomorphic for the 29K allele. The 27 and 29 kdalton polypeptides were shown to have very similar amino acid sequences, and the allelic difference at this locus is most probably in the gene sequence for the polypeptide.     

祁连山青海云杉天然群体的种实性状表型多样性

 以系统揭示其表型变异程度和变异规律为目的,对祁连山青海云杉(Picea crassifolia )天然分布区的10个群体的8个种实性状进行了比较分析。采用方差分析、多重比较,相关分析、聚类分析等多种分析方法,对群体间和群体内的表型多样性进行了讨论。方差分析结果表明,各种实性状在群体间都存在极显著差异,除球果干质量和球果长/球果径外,其余性状在群体内都存在着极显著的差异;表型分化系数即群体间变异为27.18%,小于群体内的变异(72.82%);球果长、球果径、球果干质量、球果形状指数、种子长、种子宽、千粒重和种子形状指数的变异系数分别为：10.08%、5.80%、19.29%、9.66%、8.38%、15.34%、6.52%和13.94%;8个种实性状间多数呈极显著或显著的正相关,球果干质量、种子长、千粒重、球果长和球果径为青海云杉易测定和重要的种实性状;种实性状呈现出以经度变异为主的梯度规律性;通过表型性状的聚类分析可以将青海云杉10个群体划分为4类。     

Potential for higher invasiveness of the alien Oxalis pes-caprae on islands than on the mainland

The higher vulnerability of islands to invasions compared to mainland areas has been partially attributed to a simplification of island communities, with lower levels of natural enemies and competitors on islands conferring vacant niches for invaders to establish and proliferate. However, differences in invader life-history traits between populations have received less attention. We conducted a broad geographical analysis (i.e. 1050 km wide transect) of plant traits comparing insular and mainland populations to test the hypothesis that alien plants from insular populations have the potential for higher invasiveness than their alien mainland counterparts. For this purpose plants of the annual geophyte Oxalis pes-caprae were grown from bulbs collected in the Balearic islands and the Spanish mainland under common greenhouse conditions. There were no significant differences in bulb emergence and plant survival between descendants from insular and mainland populations. However, Oxalis descendants from insular populations produced 20% more bulbs without reducing allocation to bulb size, above-ground biomass or flowering than descendants from mainland populations. Based on the lack of sexual reproduction in Oxalis and the dependence of invasion on bulb production, our study suggests that the higher occurrence of Oxalis in the Balearic islands than in the Spanish mainland can partially be explained by genetically based higher propagation potential of insular populations compared to mainland populations.     

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.     

ECOLOGICAL CONSEQUENCES AND PHENOTYPIC CORRELATES OF PETAL SIZE VARIATION IN WILD RADISH,RAPHANUS SATIVUS (BRASSICACEAE)

In this paper we examine some ecological consequences and phenotypic correlates of flower size variation in wild radish, Raphanus sativus. Mean corolla diameter varied significantly among individuals within natural populations of R. sativus in California. On the average, almost 40% of flower biomass was allocated to corolla tissue. In field experiments, pollinator visitation increased significantly with corolla size. Large flowers also accumulated more nectar when pollinators were excluded from plants. In three populations, corolla size was positively correlated with allocation to pollen per flower (either anther weight or pollen grain number), but there was usually no phenotypic relationship between corolla size and several measures of female allocation (ovule number per flower, proportion fruit set, and total seed mass per fruit). Plants growing in the field produced fewer large flowers per unit of stem, and stem biomass was negatively related to corolla size for plants grown under controlled greenhouse conditions. Male and female fitness may covary differently with allocation to attractive floral features in species such as R. sativus, where seed production is often limited by resources rather than by pollen.     

Invasive species grows faster,competes better,and shows greater evolution toward increased seed size and growth than exotic non-invasive congeners

Comparisons of introduced exotics that invade and those that do not can yield important insights into the ecology of invasions. Centaurea solstitialis, C. calcitrapa, and C. sulphurea are closely related, share a similar life history and were each introduced to western North America from Southern Europe ~100–200 years ago. However, of these three species, only C. solstitialis has become invasive. We collected seeds from different populations for each of the three species both in the native range of Spain and the non-native range of California, measured individual seed mass, and grew plants from these seeds in a greenhouse experiment in Montana. The invasive C. solstitialis had the smallest seeds and seedlings of the three congeners. However, in contrast to its non-invasive congeners, C. solstitialis had the highest relative growth rates when grown in competition. C. solstitialis was also the only species to show significant differences in traits between populations from different ranges, with plants from the non-native range of California demonstrating greater competitive resistance, larger seed size, and larger seedling mass than plants from the native range in Spain. This suggests that C. solstitialis may be evolving toward larger seed and seedling sizes in this non-native range. Relative growth rate showed no inter-regional variation for any species, but was higher for C. solstitialis than its congeners when in competition, and thus may interact with the evolution of larger seeds and plant mass in ways that contribute to the extraordinary invasive success of this species.     

A PHYLOGENETIC ANALYSIS OF BODY SIZE EVOLUTION AND BIOGEOGRAPHY IN CHUCKWALLAS (SAUROMALUS) AND OTHER IGUANINES

The evolution of body size was reconstructed in chuckwallas (genus Sauromalus), large herbivorous lizards of southwest North America, using a phylogeny derived from sequence variation in the mitochondrial cytochrome b gene. The body mass of two endemic island species (S. hispidus and S. varius) is typically fivefold larger than mainland species. We tested the hypothesis that large body size has evolved on these islands in response to local ecological conditions against the alternative hypothesis that large size is simply retained from large iguanine ancestors. The most parsimonious tree topology depicts the insular gigantic Sauromalus as monophyletic, having diverged from a common ancestor on the Baja California peninsula after the radiation of smaller bodied clades. In a robustness analysis of this topology, we found general support for this tree over alternative topologies representing minimum evolution hypotheses that imply large body size is retained from large iguanine ancestors. The most parsimonious reconstruction of body size evolution implies a change from large to small size after the Sauromalus ancestor diverged from Iguana, and one reversal back to large size within Sauromalus. The large size increase in the gigantic clade contrasts with evolutionary stasis of small body size (for an iguanine) in mainland populations. The gigantic species show 3–4% total sequence divergence from S. obesus populations on the nearby Baja California peninsula, and mainland populations of S. obesus obesus show similar levels of divergence from each other. An analysis of character transitions and comparative behavior implicates predation, and its relaxation on isolated islands, as a strong selective force in Sauromalus. Patterns of genetic differentiation in Sauromalus and biogeographic implications are discussed.     

A FURTHER CONTRIBUTION TO CLOSED-CONE PINE (OOCARPAE) HISTORY

A new record of a fossil Monterey pine (Pinus radiata D. Don) is recorded from rocks of the Plio-Pleistocene age, southern California, and the affinities of other presumed fossil closed-cone pines are revised. This has led to the recognition of two new pine species, Pinus storeyana Axelrod of subsect. Oocarpae and Pinus verdiana Axelrod of subsect. Ponderosae. Geologic evidence indicates that forerunners of the present California and Mexican species of Oocarpae were in proximity in Mexico during the Miocene. They were isolated following opening of the Gulf of California, the northward movement of Baja and Alta California on the San Andreas fault system, and the spread of desert climate. New populations of Pinus radiata developed during the Pliocene (5–2 million years) and later as environmental diversity increased. This also enabled Pinus remorala Mason and P. muricata D. Don, which the fossil record suggests were distinct into the Late Pleistocene, to hybridize in the new topographic-climatic Holocene environments.     

Early postfire seed dispersal, seedling establishment and seedling mortality of Pinus coulteri (D. Don) in central coastal California, USA

Seedling recruitment in many highly serotinous populations of Pinus coulteri on California's central coast depends almost entirely on periodic, stand-replacing fire. Compared to serotinous pines of the Mediterranean Basin, little detailed information is available on the postfire demography of California closed-cone pines, including P. coulteri. In September 1996 a wildfire burned the 760-ha American Canyon Research Natural Area (RNA). Using aerial photography, we mapped burn severity of P. coulteri-chaparral woodlands and forests within the RNA. From May to September of 1997, we also quantified seedling establishment and mortality in relation to biophysical site characteristics including fire severity. Seventy-six percent of P. coulteri forests and woodlands experienced high-severity burns, 9% moderate-severity burns, and 15% low-severity or unburned. Of the 53 plots used for seedling counts, 70% were high-severity, 26% moderate-severity, and 4% low-severity. Seedling densities 13 months postfire were low (0.21 m^–2), but seedling mortality also was low (8.4%). Aerial seed bank size increased from north-facing to south-facing slopes and from high-severity to low-severity burns. Seedling recruitment was unrelated to burn severity and increased with the size of the canopy seed bank (cone density). Many seedlings established from rodent seed caches; 23% of the seedlings established in clumps from seeds cached by Dipodomys agilis, Chaetodipus californicus and Peromyscus maniculatus. Pinus coulteri seeds have low potential for dispersal by wind, but secondary dispersal by rodents moves seeds away from source trees and into neighboring chaparral. We discuss the potential importance of rodent seed caching to postfire demography of California and Mediterranean serotinous pines.     

EFFECT OF SEED DIMORPHISM ON THE DENSITY-DEPENDENT DYNAMICS OF EXPERIMENTAL POPULATIONS OF ATRIPLEX TRIANGULARIS (CHENOPODIACEAE)

The importance of seed size and density in determining individual plant performance and plant population dynamics in experimental populations of the halophyte Atriplex triangularis was studied. Two distinct seed morphs—large, light seeds and small, dark seeds—are produced by individual A. triangularis plants. Experimental populations consisting of seed size monocultures (large or small seeds) and seed size mixtures were established at three different densities, and the time of germination, plant size, plant survivorship, and plant fecundity were monitored. Marked variation in time of germination was observed among treatments and between seed sizes, but germination within any given treatment occurred over a five- to ten-day period. Large seeds produced larger plants than small seeds did, and this dichotomy was maintained over the course of the entire experiment. Germination date and seed size interacted such that larger plants grew from seeds which germinated earlier than those which germinated later, regardless of seed size. Germination date had a more pronounced effect than seed size did on plant mortality in high density populations. At high density, large seed monocultures experienced greater mortality than small seed monocultures did, but in seed size mixtures, the mortality was evenly distributed between plants from the two seed sizes. Regardless of density conditions and parentage, large and small seeds were produced in equal proportion by the plants. Total seed production, however, was dramatically affected by plant density, and to a lesser degree by germination date. Although seed size effects alone did not appear to affect directly final plant biomass and fecundity, effects of seed size early in ontogeny may have contributed to differences in fecundity.