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.     

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.

     

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.     

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.     

Interactions between a seed‐eating neotropical rodent,the Azara's agouti (Dasyprocta azarae), and the Brazilian ‘pine’ Araucaria angustifolia

We studied the seed predation and scatter‐hoarding behaviour of Azara's agoutis Dasyprocta azarae (Rodentia: Dasyproctidae) in relation to the seeds of the Brazilian ‘pine’, Araucaria angustifolia (Araucariaceae), the rodent's main winter food source. We compared seed‐removal rates, seed‐caching rates, cache distances and recovery rates between a summer period of food abundance (with a low demand for A. angustifolia seeds and no such seeds naturally available) and a winter period of food scarcity (with a high demand for A. angustifolia seeds). We investigated whether the relative seed value affected the rodent's seed‐handling behaviour. We predicted that during the high seed‐demand period (winter): (1) cache distances would be greater; (2) fewer seeds would be stored; (3) more seeds would be recovered and the seed‐recovery time would be lower. In support of our first two predictions, the caching distances were greater in winter (mean ± SE = 15.67 ± 5.11 m) than in summer (9.40 ± 1.59 m), and agoutis hoarded >9 times more seeds in summer (55) than in winter (6). Our third prediction was not supported, and the proportion of unrecovered caches and buried seed recovery times did not differ between winter (mean ± SE = 3.00 ± 0.00 days, n = 5 seeds) and summer (11.05 ± 3.68 days, n = 20 seeds). The high resource density (during summer) rather than the density of A. angustifolia seeds likely influenced seed fate. Agoutis acted mainly as predators, leaving few intact seeds, caching a low proportion of handled seeds (? 8%) and rapidly consuming the caches. Agoutis may cache seeds to keep them safe from competitors on a short‐term basis rather than maintaining medium‐ or long‐term reserves for use during food‐scarcity periods.     

太白红杉种实数量特征

太白红杉 (Larixchinensis) 是仅分布于我国陕西秦岭境内的国家二级保护植物, 研究其生殖规律将有助于加深对其濒危原因的认识。该文用单因素方差分析的方法研究了太白红杉种实数量特征, 并用Pearson相关系数检验了不同环境条件下的变化及其与环境因子和植株生长状况的关系, 结果表明种实数量特征在不同环境条件下差异明显, 太白山国家森林公园单株种子产量最高 ;长安光头山平均每株产生的种子总重、球果总重和球果平均重量最重, 平均每株球果数目和每球果种子数也是最多。多重比较表明牛背梁种群太白红杉种子千粒重最重。其中球果的种子数在小种群中较多 ;每株的种子总重量、球果平均重和球果总重在小种群中较重。太白红杉种实数量特征与海拔、坡度和坡向呈负相关, 与植株年龄呈正相关, 球果的种子数目与年均温呈正相关 (p <0.0 5 ), 而与一月均温呈负相关 (p <0.0 5 ) 。种子大小与种子总数呈弱正相关, 种子大小和数目间不存在权衡 (Trade-ff) 。     

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.     

Seed and cone diversity and seed germination of Pinus pinea in Strofylia Site of the Natura 2000 Network

Variation in cone size, seed number per cone, seed potential, seed efficiency, seed morphology and seed germination behavior of Pinus pinea and its relation to stand conditions was analyzed. Data were collected from P. pinea forest in Strofylia, southern Greece, a forest that belongs to the Natura 2000 European network and the RAMSAR convention and is characterized by the absence of regeneration for many decades. The pine stands found in the area were distinguished into five categories according to a previous study and our observations, regarding stand age, canopy cover and the degree of stress by human pressure. The categories are: (I) young artificial, (II) closed-mature, (III) open-mature, (IV) over-mature stands and (V) highly degraded stands. Cones were collected from all stand types and their morphological characteristics as well as their seed production were measured. Seeds were extracted from the collected cones, measured and their germination behavior was tested. The findings showed that the over-mature and the high degraded stands and to a lesser extent, the closed-mature stands, produced significantly smaller cones with a lower seed potential, a lower number of filled seeds per cone, a greater number of not fully developed seeds and reduced seed morphological characteristics than the young and open-mature stands. However, the seed germination behavior of fully developed seeds was only slightly affected by the stand type.     

Enhanced seed defenses potentially relax selection by seed predators against serotiny in lodgepole pine

Serotiny, the retention of seeds in a canopy seed bank until high temperatures cause seeds to be released, is an important life history trait for many woody plants in fire‐prone habitats. Serotiny provides a competitive advantage after fire but increases vulnerability to predispersal seed predation, due to the seeds being retained in clusters in predictable locations for extended periods. This creates opposing selection pressures. Serotiny is favored in areas of high fire frequency, but is selected against by predispersal seed predators. However, predation also selects for cone traits associated with seed defense that could reduce predation on serotinous cones and thereby relax selection against serotiny. This helps explain the elevated defenses in highly serotinous species. However, whether such interactions drive variation in seed defenses within variably serotinous populations has been studied rarely. We investigated the effects of phenotypic selection exerted by red squirrel (Tamiasciurus hudsonicus) predation on Rocky Mountain lodgepole pine (Pinus contorta latifolia) seeds. Squirrels preferentially harvested cones with more and larger seeds, indicating a preference for a higher food reward. We found evidence for stronger selection on trees with serotinous cones, which presumably accounts for the elevated defenses of and lower predation on serotinous compared to non‐serotinous cones. Lower levels of predation on serotinous cones in turn lessen selection against serotiny by squirrels. This has important implications because the frequency of serotiny in lodgepole pine has profound consequences for post‐fire communities and ecosystems widespread in the Rocky Mountains.     

Germination ecology of <Emphasis Type="Italic">Scorpiurus subvillosus</Emphasis> L. seeds: the role of temperature and storage time

Scorpiurus subvillosus L., wide spread in pastures of Mediterranean basin, is disappearing in the native pastures of the Hyblean plateau (Sicily, southern Italy), because of overgrazing and intensive management techniques. Moreover, it exhibits seed coat dormancy, which delays and reduces germination preventing its diffusion. This paper represents a first attempt in order to investigate changing in germination determined by storage time and temperature on seeds of two populations of S. subvillosus. Germination of S.␣subvillosus seeds was tested in relation to four storage time (30, 130, 200 and 360 days after harvest (DAH)), eight constant temperatures (5, 10, 15, 20, 25, 30, 35 and 40°C) and two populations of different provenience (30 and 600 m above mean sea level). The experiments were conducted either on scarified and unscarified seeds. In S. subvillosus the failure of germination under favourable conditions must be attributed␣only to seed coat, since seed scarification enhanced germination percentage with values up to 100% at almost all tested temperatures. In both treatments, but with a grater incidence in unscarified, seed germination increased gradually as temperature raised, peaking at 20–25°C, then declined with further increases of temperatures. At 40°C no germination occurred. Storage time induced a softening effect, which is somewhat limited by the natural ageing of seeds occurring from about 6 months after harvest.     

CONSISTENCY AND VARIATION IN PHENOTYPIC SELECTION EXERTED BY A COMMUNITY OF SEED PREDATORS

Phenotypic selection that is sustained over time underlies both anagenesis and cladogenesis, but the conditions that lead to such selection and what causes variation in selection are not well known. We measured the selection exerted by three species of predispersal seed predators of lodgepole pine (Pinus contorta latifolia) in the South Hills, Idaho, and found that net selection on different cone and seed traits exerted by red crossbills (Loxia curvirostra) and cone borer moths (Eucosma recissoriana) over 10 years of seed crops was similar to that measured in another mountain range. We also found that the strength of selection increased as seed predation increased, which provides a mechanism for the correlation between the escalation of seed defenses and the density of seed predators. Red crossbills consume the most seeds and selection they exert accounts for much of the selection experienced by lodgepole pine, providing additional support for a coevolutionary arms race between crossbills and lodgepole pine in the South Hills. The third seed predator, hairy woodpeckers (Picoides villosus), consumed less than one‐sixth as many seeds as crossbills. Across the northern Rocky Mountains, woodpecker abundance and therefore selective impact appears limited by the elevated seed defenses of lodgepole pine.     

Coevolution between crossbills and black pine: the importance of competitors,forest area and resource stability

Studies of predator‐prey interactions have found that geographically structured coevolution has played an important role in the adaptive diversification of crossbills (Loxia spp.). We extend those studies by considering common crossbills (L. curvirostra) in the Mediterranean where they rely on seeds in the cones of black pine (Pinus nigra). On the continent, where tree squirrels (Sciurus vulgaris) are present, enhanced defenses against crossbills were most evident in larger areas of pine forest. On islands in the absence of tree squirrels, crossbills and black pine have coevolved in a predator‐prey arms race on Cyprus but not Corsica. In contrast to other conifers that island endemic crossbills rely upon, black pine does not hold seeds in its cones year round. Consequently, key to the strong crossbill–pine interaction on Cyprus is likely the presence of an alternative conifer that provides seeds during early summer when black pine seeds are scarce.     

Seed dispersal of the Korean pine,Pinus koraiensis,by the red squirrel,Sciurus vulgaris

Seed hoarding behavior of the red squirrel,Sciurus vulgaris, was studied in relation to the amount of dispersed seeds of the Korean pine,Pinus koraiensis, and the distribution of its seedlings. After removing a cone from a tree, squirrels sat on the ground and ripped off its cone scales before transporting it. A mean of 3.2 seeds were scatter-hoarded per hole. Of 7.7×10⁴ mature seeds produced in a 0.21 ha planted Korean pine forest, 22% were estimated to be directly eaten by four squirrels, 9% were hoarded by them in the pine forest and 65% were cached outside the forest. Squirrels rediscovered hoarded seeds frequently, until the ground was covered with snow, during the period from snow fall until seed germination the next spring, few hoarded seeds were utilized. Korean pine seedlings were found up to 600 m from their mother trees. Scatter-hoarding by squirrels extensively contributes to seed dispersal to places suitable for the regeneration of the Korean pine. The large size of the cone, the absciss-layer at the cone penduncle, the infrequent dehiscence of cone scales, the large and wingless seeds, and the thick seed-coats have probably all been specialized to facilitate utilization by GenusSciurus.     

Temporal variation in breeding and cone size selection by three species of crossbills Loxia spp. in a native Scots pinewood

The breeding biology and cone size selection of crossbills was studied mainly during 1995 to 2002 at Abernethy Forest, Scotland, an ancient native Scots pine Pinus sylvestris wood, where only a single crossbill species, the Scottish crossbill Loxia scotica, was assumed to occur and to be adapted to feed on seeds in Scots pine cones. However, three crossbill species (common Loxia curvirostra, Scottish and parrot crossbills Loxia pytyopsittacus) nested in some years, with the parrot crossbill being the most abundant. Most nests were in old large pines, with the three crossbill species not differing in their use of tree size or stand density for nesting. The mean clutch and brood sizes were 3.8 and 2.9, and their mean survivals were 86 and 74%, respectively, with no significant differences among species. The timing of breeding differed between species, with parrot crossbills breeding earliest (median date 21 March, including second attempts) and common crossbills breeding last (median date 21 April), probably in response to the differing accessibility of Scots pine seeds to these species. The difference in the time of breeding may reduce mixed mating. Crossbills foraged preferentially on trees with small cones when the cones were closed. Small cones had thinner scales than large cones, suggesting that the preference for small cones was related to higher feeding rates on these cones when cones are closed. Such a preference was also found for captive crossbills with the Scottish crossbill showing a more pronounced preference for smaller cones than the larger‐billed parrot crossbill. However, crossbills selected larger cones within trees and trees with larger cones once the cones opened in April. Such a shift occurred presumably because variation in scale thickness has little impact on seed accessibility once cones open, and larger cones have larger and more seeds. The greater ability of parrot crossbills to exploit seeds in closed Scots pine cones allowed parrot crossbills to start breeding earlier and to have young when seeds were most accessible. Only after the cones opened were the smaller‐billed common crossbills able to easily access seeds and to start breeding. The time of breeding of Scottish crossbills was intermediate between common and parrot crossbills, and they probably had an intermediate ability to exploit Scots pine cones. The reason why there were few Scottish crossbills nesting in Abernethy Forest remains a puzzle, considering that native pine wood is assumed to be the ancestral habitat to which the Scottish crossbill is adapted. The breeding season for all crossbills ended in June, when most of the seed from a given cone cohort was shed. This is when starved broods were found, not associated with bad weather.     

A comparative analysis of seed and cone characteristics and seed-dispersal strategies of three pines in the subsection Sabinianae

The seed-dispersal systems of Coulter pine (Pinus coulteri), gray pine (P. sabiniana), and Torrey pine (P. torreyana), all of the subsection Sabinianae, are not well understood. These pines occur in arid and semi-arid foothills and mountains of California that are subjected to frequent fires. Cone and seed traits of these three California pines are compared to those of four species of pines (sugar pine, P. lambertiana; Jeffrey pine, P. jeffreyi; ponderosa pine, P. ponderosa; and lodgepole pine, P. contorta) that occur in more mesic environments in the nearby Sierra Nevada mountains. The cones of the Sabinianae pines are large with thick, dense scales, and the scales of gray and Coulter pines are armed with sharp, recurved spines. The seeds of all three species are large, and those of gray and Torrey pines are nearly wingless. In contrast, the Sierra Nevada pines have small to medium-sized seeds with large wings that are initially dispersed by the wind. Heavy wing loading of the Sabinianae pine seeds causes them to fall rapidly, and they are not dispersed far by wind. However, animals remove the fallen seeds rapidly, and rodents and jays scatter hoarded many seeds in the soil. This caching activity results in seedling establishment. The unusual morphology of the cones and seeds of the Sabinianae pines is interpreted as a combination of traits that attract animal dispersers, thwart the foraging activities of seed predators, and promote the survival of seeds in an environment subject to frequent fires.     

The role of birds and mammals in Korean pine (Pinus koraiensis) regeneration dynamics

This study examined the interrelationships of the fall seed-foraging guild with Korean pine (Pinus koraiensis) and its regeneration. The study took place in old-growth and secondary-growth forests in Northeastern China. Foraging behavior, seed transportation distance, and cache location of various birds and mammals were observed. Regeneration characteristics of Korean pine were also analyzed using plots in various vegetation cover types, successional stages, and topographical situations. Ten species of vertebrates were able to harvest seed from the closed-cone Korean pine. Of these, the Eurasian nutcracker, Eurasian nuthatch, red squirrel, and Siberian chipmunk were found to be potential seed dispersal agents. The nutcracker was the most important dispersal agent, easily acquiring seed with its large pointed bill, carrying up to 62 seeds in one trip, carrying seed at least 4 km, and placing seed in a variety of sites 2.5–3 cm deep in the soil. The Eurasian nuthatch carried a single seed per trip at distances less than 50 m, needed nutcrackers to open the cones and expose seed before they could acquire seed, and occasionally cached seed in the soil. Red squirrels were uncommon visitors to the tree tops of Korean pine, carried cones shorter distances than nutcrackers, and were only found under forest canopies. Human harvest of cones by knocking off branches also affected squirrel behavior and reduced future cone crops. Siberian chipmunks also collected seed from cones in trees and appeared to transport seed less than 50 m. Six other species were observed in this study successfully harvesting seed from cones but were not potential seed dispersers. Natural seedling establishment was found to be over 1000 seedlings/ha except on old-growth pine-hardwood sites. Squirrels were commonest here, but few seedlings survived past the 2nd year due to the intense shading. Second-growth forest types, including aPicea plantation where nutcrackers cached seed daily, and an old-growth pine-hardwood selective-harvest site, had the greatest regeneration. In conclusion, most natural regeneration of Korean pine in this part of its range is due primarily to the Eurasian nutcracker. Nutcrackers can aid forest managers in reaching desired stocking levels after disturbance, as well as a more natural-appearing forest. Squirrels, chipmunks, and nuthatches are minor seedling establishment agents. Korean pine seed is an important food source used by at least 22 species of forest wildlife.     

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。     

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.     

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.     

Role of heat tolerance and cone protection of seeds in the response of three pine species to wildfires

The post-fire regenerative ability of Pinus halepensis, Pinus nigra and Pinus sylvestris, three of the most important pine species present in the West Mediterranean basin, has been analyzed in the light of seed tolerance to different temperatures and times of exposure, and of seed position during the fire event (seeds inside cones versus free seeds). The combination of different fire intensities and degrees of seed protection allows us to draw different scenarios during the fire event: canopy scenarios (seeds inside cones), surface scenarios (seeds on the ground surface), and soil scenarios (seeds in the top soil layers). There were interspecific differences in the pattern of cone opening under the different heat treatments: cones of P. nigra and P. sylvestris showed similar percentages of opening, but considerably higher than those of P. halepensis. In the three species, seeds inside cones showed higher percentages of germination than those that were free, emphasizing the important role of cones in the protection of pine seeds from high temperatures. The percentage of germination decreased when both the temperature and the time of exposure increased, and there was also a significant species effect: P. halepensis showed higher germination rates than P. nigra, and both were higher than P. sylvestris. The overall scores of seed germination of these three pine species under the conditions tested suggests that their regeneration after fire should come either from the soil bank, or from the canopy bank, but rarely from the ground surface. As the existence of a permanent seed bank in Mediterranean pines is probably limited or nil, pine recruitment after fire appears to be mainly controlled by the existence of a canopy seed bank. The contribution of this canopy bank to the differences in postfire regeneration success of the three pine species is discussed in the light of their seeding phenology and the effects of fire severity on cone opening. The results obtained in this study contribute to explain the successful regeneration of P. halepensis, and the failure of P. nigra and P. sylvestris after fire.