Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

The interaction of Saccharomyces paradoxus with its natural competitors on oak bark

The natural history of the model yeast Saccharomyces cerevisiae is poorly understood and confounded by domestication. In nature, S. cerevisiae and its undomesticated relative S. paradoxus are usually found on the bark of oak trees, a habitat very different from wine or other human fermentations. It is unclear whether the oak trees are really the primary habitat for wild yeast, or whether this apparent association is due to biased sampling. We use culturing and high‐throughput environmental sequencing to show that S. paradoxus is a very rare member of the oak bark microbial community. We find that S. paradoxus can grow well on sterile medium made from oak bark, but that its growth is strongly suppressed when the other members of the community are present. We purified a set of twelve common fungal and bacterial species from the oak bark community and tested how each affected the growth of S. paradoxus in direct competition on oak bark medium at summer and winter temperatures, identifying both positive and negative interactions. One Pseudomonas species produces a diffusible toxin that suppresses S. paradoxus as effectively as either the whole set of twelve species together or the complete community present in nonsterilized oak medium. Conversely, one of the twelve species, Mucilaginibacter sp., had the opposite effect and promoted S. paradoxus growth at low temperatures. We conclude that, in its natural oak tree habitat, S. paradoxus is a rare species whose success depends on the much more abundant microbial species surrounding it.     

Historical introgression among the American live oaks and the comparative nature of tests for introgression

Introgressive hybridization challenges the concepts we use to define species and infer phylogenetic relationships. Methods for inferring historical introgression from the genomes of extant species, such as ABBA‐BABA tests, are widely used, however, their results can be easily misinterpreted. Because these tests are inherently comparative, they are sensitive to the effects of missing data (unsampled species) and nonindependence (hierarchical relationships among species). We demonstrate this using genomic RADseq data sampled from all extant species in the American live oaks (Quercus series Virentes), a group notorious for hybridization. By considering all species and their phylogenetic relationships, we were able to distinguish true hybridizing lineages from those that falsely appear admixed. Six of seven species show evidence of admixture, often with multiple other species, but which is explained by introgression among a few related lineages occurring in close proximity. We identify the Cuban oak as the most admixed lineage and test alternative scenarios for its origin. The live oaks form a continuous ring‐like distribution around the Gulf of Mexico, connected in Cuba, across which they could effectively exchange alleles. However, introgression appears highly localized, suggesting that oak species boundaries and their geographic ranges have remained relatively stable over evolutionary time.     

Leaf morphological analyses in four European oak species (Quercus) and their hybrids: A comparison of traditional and geometric morphometric methods

Abstract

In this study, leaf morphology was assessed in a mixed oak stand (western France) using two geometric morphometric (landmark and outline) datasets and one dataset of 19 leaf measures. Adult oaks (817 oaks), comprising four white oak species (Quercus petraea, Q. robur, Q. pubescens and Q. pyrenaica), were sampled for DNA extraction and genetic analysis (nuclear microsatellites). Leaf morphology was assessed on 336 oaks, comprising pure species and hybrids as determined by genetic assignment. This comparative study of oak leaf morphology, based on the use of two free size geometric morphometric methods and a set of leaf measurements, combined with the genetic assignment of individuals to pure species or hybrids, provided information about the differences among species and the intermediate leaf morphology of their hybrids.     

Long‐term and interactive effects of different mammalian consumers on growth,survival, and recruitment of dominant tree species

Throughout the world, numerous tree species are reported to be in decline, either due to increased mortality of established trees or reduced recruitment. The situation appears especially acute for oaks, which are dominant features of many landscapes in the northern hemisphere. Although numerous factors have been hypothesized to explain reductions in tree performance, vertebrate herbivores and granivores may serve as important drivers of these changes. Here, using data from 8‐ and 14‐year‐old exclosure experiments, we evaluated the individual and interactive effects of large and small mammalian herbivores on the performance of three widespread oak species in California—coast live oak (Quercus agrifolia), California black oak (Q. kelloggii), and Oregon white oak (Q. garryana). Although impacts varied somewhat by species and experiment, herbivory by black‐tailed deer (Odocoileus hemionus columbianus) reduced the height and survival of juvenile coast live oaks and altered their architecture, as well as reduced the abundance of black oak seedlings, the richness of woody species and the cover of nonoak woody species. Small mammals (Microtus californicus and Peromyscus maniculatus) had even more widespread effects, reducing the abundance of black oak seedlings and the height and cover of all three oak species. We also detected numerous interactions between small mammals and deer, with one herbivore having positive or negative effects on oak abundance and cover when the other herbivore was either present or absent. For example, deer often had negative effects on seedling abundance only when, or even more so when, small mammals were present. In summary, mammalian consumers play crucial roles in limiting oak recruitment by reducing seedling abundance, maintaining trees in stunted states, and preventing them from reaching sapling stages and becoming reproductive. Interactions between large and small mammals can also alter the intensity and direction of their effects on trees.     

Responses of three bottomland species with different flood tolerance capabilities to various flooding regimes

Seedlings of baldcypress (Taxodium distichum), nuttall oak (Quercus nuttalli), and cherrybark oak (Quercus falcata var.pagodaefolia) were subjected to four flooding treatments: control, continuously flooded, intermittently flooded, and partially flooded for 70 days in a greenhouse. The treatments imposed various durations and intensities of soil redox potential (Eh) conditions representing a range encountered by plants in their habitats. Morphological changes and gas exchange responses to the treatments differed among the study species. Rapid development of adventitious root and hypertrophied lenticels were observed in baldcypress and nuttall oak under all flooded treatments. Cherrybark oak had the highest percentage reduction in net photosynthesis ranging from 65–87%, whereas reductions in nuttall oak ranged between 35–68% and in baldcypress between 6–21% in response to various treatments. Recovery of gas exchange was noted in baldcypress but no significant recovery was found in oaks. The recovery in baldcypress contributed to the continued growth and biomass accumulation under various treatments. Little evidence of consistent changes in biomass allocation patterns in response to the treatments was found in baldcypress but total biomass decreased significantly under the continuously flooded treatment. In oaks, total biomass decreased significantly in all flooded treatments. The present findings demonstrated that physiological functions are adversely affected by low soil Eh conditions and the extent of such effects are dependent on the intensity and duration of soil reduction as well as the species' capability to respond to such conditions rapidly. Management plans concerned with regeneration of bottomland forested ecosystems should consider the species flood response capabilities at seedling stages as well as the timing, durations, and intensities of soil reduction at the specific site.     

Suitability of native and ornamental oak species in California for Agrilus auroguttatus

Goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), is a new invasive species in southern California, USA. The extent of the host range of this insect is not known, but this knowledge will have a major impact on assessment of the risks that this pest poses to oaks [Quercus spp. (Fagaceae)]. We conducted laboratory tests to determine the potential suitability of native and ornamental oak species for larvae and adults of A. auroguttatus. We infested 179 cut logs (from 163 different trees) with eggs or larvae, measured neonate survival and, after 5 months, counted feeding galleries, and noted the proportion of galleries with late instars. Initial larval survival was generally high when larvae penetrated the phloem (range: 62–98% among oak species), and low by the time larvae began to feed at the phloem/xylem interface (range: 0–25% among oak species). The majority of larvae that established a visible feeding gallery survived to the fourth instar (total of 73% for all oak species). Larval galleries were established with greater frequency in red oaks (Section Lobatae) compared with other oaks (19 vs. 7 or 4%). All red oaks tested (Q. agrifolia Née, Q. kelloggii Newberry, and Q. wislizeni A. DC.) were likely suitable hosts for larvae. Larvae were apparently able to feed on some of the other oaks (Q. chrysolepis Leibmann, Q. suber L., Q. lobata Née, and Q. douglasii Hook & Arn), although it remains unclear whether these species would be preferred hosts under natural conditions. Adult longevity and fecundity varied little by species of oak foliage fed to adults. The host range of A. auroguttatus is likely limited by suitability of oak species for the larval rather than the adult life stage. Results support published field observations that red oaks are more suitable hosts than white oaks.     

POLLEN MORPHOLOGY AND PLANT TAXONOMY OF WHITE OAKS IN EASTERN NORTH AMERICA

An evaluation of possible approaches to fossil oak pollen identification utilized scanning electron microscopy to examine exine-surface features of 171 collections, representing 16 Quercus subgenus Lepidobalanus species and varieties of eastern North America. Twenty qualitative pollen morphological characters were defined and tabulated for each of 217 pollen grains. The data were subjected to cluster analysis and cluster diagrams were compared with published white oak taxonomy. Pollen morphology and plant taxonomy compared well in series of the subgenus Lepidobalanus due primarily to consistency of character presence and absence within species and varieties. Pollen morphology of white oaks appears to reflect plant systematics above the species level. Use of routine SEM analysis to identify series of white oaks among fossil pollen grains likely will yield valid results.     

桂西地区南亚热带落叶栎林群落组成及物种多样性研究北大核心CSCD

落叶栎林是桂西地区南亚热带的典型次生林,也是该区域落叶阔叶林的重要群系组。该研究采用典型样方法对桂西地区落叶栎林群落进行调查,分析了该区域落叶栎林群落的物种组成、区系成分、物种多样性特征及其与地形因子的关系,为桂西地区南亚热带植物多样性保护与恢复提供依据。结果显示:(1)研究区落叶栎林群落维管束植物共计269种,隶属80科178属。(2)种子植物区系以热带成分为主,同时表现出一定程度的温带过渡性质。(3)聚类分析表明,调查的落叶栎林群落可分为云南波罗栎林、栓皮栎林、白栎林、麻栎林4种林分类型,其中以白栎林群落的物种多样性最高,且灌木层的物种多样性显著高于乔木层和草本层。(4)RDA分析显示,落叶栎林群落不同层次物种多样性的差异受地形因子的影响,多样性指标与经度、纬度、海拔之间具有明显的相关性(P<0.05),其中乔木层物种多样性主要与经度、纬度呈显著的相关性,灌木层物种多样性与纬度、海拔相关,草本层物种多样性与经度、纬度、海拔之间均有相关性。     

Associations Among Breeding Birds and Gambel Oak in Southwestern Ponderosa Pine Forests

Abstract: Ponderosa pine (Pinus ponderosa) forests with Gambel oak (Quercus gambelii) are associated with higher bird abundance and diversity than are ponderosa pine forests lacking Gambel oak. Little is known, however, about specific structural characteristics of Gambel oak trees, clumps, and stands that may be important to birds in ponderosa pine-Gambel oak (hereafter pine-oak) forests. We examined associations among breeding birds and structural characteristics of Gambel oak at a scale similar in size to individual bird territories in pine-oak forests in northern Arizona and western New Mexico, USA. Avian species richness and occurrence of some bird species were associated with specific growth forms of Gambel oak. Estimated probability of Virginia's warblers (Vermivora virginiae), black-headed grosbeaks (Pheucticus melanocephalus), and red-faced warblers (Cardellina rubrifrons) occurring at points increased with increasing density of pole-sized Gambel oak 7–15 cm in diameter at breast height. We also found evidence that large Gambel oak trees (≥23 cm dbh) were associated with increased occurrence of yellow-rumped warblers (Dendroica coronata) at points. Some avian associations with oak were influenced by characteristics of ponderosa pines. For example, bird species richness was positively associated with the abundance of large Gambel oak when density of large pine trees ≥23 cm in diameter at breast height was low. Because large oak trees are rare and their numbers are thought to be declining, efforts should be made to retain and promote growth of additional oaks in this size class. Forest management practices that maintain forest openings, such as prescribed burning, could promote growth of pole-sized Gambel oak, which appears important to some bird species in pine-oak forests.     

Radial growth and climate responses of white oak (Quercus alba) and northern red oak (Quercus rubra) at the northern distribution limit of white oak in Quebec, Canada

Aim The objectives of this study were: (1) to compare radial growth patterns between white oak (Quercus alba L.) and northern red oak (Quercus rubra L.) growing at the northern distribution limit of white oak; and (2) to assess if the radial growth of white oak at its northern distribution limit is controlled by cold temperature. Location The study was conducted in three regions of the Ottawa valley in southern Québec. All stands selected were located at the northern limit of distribution of Q. alba. Methods Twelve mixed red and white oak stands were sampled and increment cores were extracted for radial growth analyses. For each oak species, 12 chronologies were derived from tree‐ring measurement (residual chronologies). Principal components analysis and redundancy analysis were used to highlight the difference between radial growth in both species and to determine their radial growth–climate association. Results There was little difference between the radial growth of each species; Q. alba, however, exhibits more year‐to‐year variation in growth than Q. rubra. More than 65% of the variance in radial growth was shared among sites and species. Both species showed a similar response to climate, which suggested that the limit of distribution of Q. alba might not be determined by effects on growth. Both species had a classic response to climate and drought in the early growing season. Main conclusions The northern distribution limit of Q. alba does not appear to be directly controlled by effects on growth processes as indicated by the similarities in radial growth and response to climate between the two species. The location of the stands on southern aspects suggested that cold temperature could have been a major factor controlling the distribution limit of Q. alba. However, it is speculated that stands growing on southern aspects may be more prone to forest fires or to drought, which would favour the maintenance and establishment of oaks, and of Q. alba in particular. Models relating the northern distribution limits of species to broad climate parameters like annual mean temperature will need to be reviewed to incorporate more biologically relevant information. Such assessments will in turn provide better estimates of the effect of climate changes on species distribution.     

Weed control and overstory reduction improve survival and growth of under‐planted oak and hickory seedlings

Weed control and overstory reduction are important silvicultural treatments for improving survival and growth of under‐planted oak and hickory seedlings. Mast‐producing trees in the bottomland forests of the blackland prairie and Post Oak Savannah ecoregions of Texas have declined in abundance. Oaks and hickories have been replaced by more shade‐tolerant species, including green ash (Fraxinus pennsylvanica Marshall) and sugarberry (Celtis laevigata Willd.), which do not produce significant hard mast for priority wildlife species. A split‐plot experiment design was installed on three sites at Richland Creek Wildlife Management Area in Freestone County, Texas, studying the effects of canopy coverage and competition control on survival and growth of bur oak (Quercus macrocarpa Michx.), Shumard oak (Quercus shumardii Buckl.), and pecan (Carya illinoinensis (Wagenh.) K. Koch) seedlings. Uprooting by hogs shortly after planting resulted in greater than 90% mortality of pecan on the two lower elevation sites. Year one survival of Shumard oak was significantly higher than bur oak. However, bur oak was more preferred by hogs than Shumard oak. Year one growth of bur oak was significantly greater than Shumard oak. Severe flooding during the second growing season caused complete mortality on the lower two sites. None of the species were well suited to such prolonged (3–4 months) inundation as seedlings. On the remaining site, density reduction and weed‐barrier mats improved growth and survival while herbaceous weed control with herbicides actually reduced both growth and survival.     

Polymorphic microsatellite loci in Eurytoma brunniventris,a generalist parasitoid in oak cynipid galls

Eurytoma brunniventris is a parasitic wasp with a wide range of host species. We have developed primers for nine polymorphic microsatellite loci to allow examination of intraspecific population subdivision associated with two aspects of the biology of their hosts: host species (oak gallwasps) and the tree on which the gallwasp develops (different oak species). All nine loci amplified well across individuals collected from a range of gallwasp species and across two oak taxa. These microsatellite loci are potentially of value in the study of closely related economic pests such as seed predators of almonds (E. amygdali) and pistachios (E. plotnikovi).     

Adaptation to oak and other fibrous,phenolic-rich foliage by a small mammal,Neotoma fuscipes

Summary Neotoma fuscipes, a small mammalian herbivore with apparently generalized food habits, was laboratory tested to determine its degree of dietary specialization. Woodrats from both oak woodland and coastal sage communities preferred Quercus agrifolia leaves (containing 40% phenolics and about 16% condensed tannin) over foliage from other dominant species. Approximately one-third of the oak phenolics and less than 10% of the oak condensed tannin remained in the feces. Their performance on pure oak leaves was comparable to that on a mixed diet of Quercus, Salvia, Eriogonum, and Rhus, with respect to weight maintenance, digestive efficiency and total amount ingested. Digestive efficiency was low on the oak diet (55%) relative to Salvia (77%), and to achieve similar weight levels, approximately twice as much oak as Salvia was ingested. Woodrats retained more nitrogen as oak consumption increased. Intake of oak and other foods increased with each experimental day. A sympatric species, N. lepida, was unable to maintain weight on oak leaves, although its digestive and polyphenolic-degrading capabilities, and nitrogen retention efficiency were equivalent to those of N. fuscipes. On a weight-adjusted basis, N. lepida ate about half as much oak per day as N. fuscipes. Oak intake may have been reduced by an inability to rapidly degrade fiber, which constitutes about 30% of the oak diet. In natural populations, N. fuscipes selectively feeds on evergreen sclerophyll vegetation high in fiber, tannins and related polyphenolics. Individuals ingest 2–3 plant types at a time, with a single species (oak when available) constituting most of the material consumed. Neotoma lepida diets are also dominated by a single species. The diversity of plant types eaten by different populations of N. lepida suggests that local dietary specializations may be developmentally acquired.     

Regeneration patterns in relation to canopy species composition and site variables in mixed oak forests in the Sierra de Manantlán Biosphere Reserve, Mexico

Regeneration patterns in relation to canopy species composition and site variables were analyzed in mixed oak forests of the Sierra de Manantlán in western Mexico with the aim of establishing an ecological basis for the design of management alternatives. Using ordination (canonical correspondence analysis) and classification (two-way indicator species analysis) methods, five different canopy types and three different seedling associations were revealed according to species composition, all of them dominated by one or more oak species. Red–far red ratio, slope, altitude, topography, canopy type and grazing intensity were the main variables that explained differences in species composition among the seedling associations. Oak seedlings were relatively scarce in the sampling plots, with the lowest frequency values of all species recorded except for those of Quercus crassipes Humb. & Bonpl., and also the lowest density values. The presence of a particular oak seedling species was strongly associated with a particular percentage of canopy openness; Quercus candicans Née, Quercus laurina Humb. & Bonpl. and Quercus rugosa Née were present in the plots with the least-open canopy (6.4%, 2.9 and 6.2%, respectively), while Quercus castanea Née and Quercus crassipes Humb. & Bonpl. were present in the plots with the most-open canopy (13 and 8.1%, respectively). Every oak seedling species was more frequent, although not dependent, on the canopy type where the same oak species dominated. Because of the great heterogeneity in species composition and the physiographical factors of mixed oak forests in the Sierra de Manantlán, we concluded that management alternatives must be prescribed for each ecological situation where the different oak species are growing.     

Radiative properties of hardwood leaves to ultraviolet irradiation

Spectral reflectance and transmittance of leaves to ultraviolet irradiation were determined under laboratory conditions for seven species of hardwood trees, namely red oak (Quercus rubra, L), black oak (Q. velutina, Lamarch), white oak (Q. alba, L.), sugar maple (Acer saccharum), Norway maple (A. plantanoides), hickory (Carya tomemtosa), sweetgum (Liquidambar styraciflua), and black oak litter. The experimental system consisted of a solar simulator, an integrating sphere, and a spectroradiometer. Experiments were repeated three to five times for both adaxial and abaxial surfaces of fresh leaves chosen at randomly. The spectral distributions and simple averages of the radiative properties in the wavelength ranges of ultraviolet-B (UV-B, 280–320 nm) and ultraviolet-A (UV-A, 320–400 nm) were determined. The spectral distributions of reflectance were similar between adaxial and abaxial surfaces, although the magnitude varied among tree species. Leaf reflectance was very low for the ultraviolet spectrum in general and varied among species and between adaxial and abaxial surfaces. It was generally higher over the UV-A waveband compared to UV-B, and higher on the abaxial than adaxial surface. The broadband reflectance in the UV-A range (over all species) was 5.0 and 3.9% for abaxial and adaxial surface, respectively, compared to 3.5 and 2.8% in UV-B. The transmittance through leaves was extremely small in the UV-B (<0.1%) and nearly zero in the UV-A spectral range. Consequently, the absorptance of ultraviolet radiation by leaves, as determined from the measured reflectance and transmittance, was quite high, being more than 90% for all the combinations of species and wavebands examined. The reported results are useful for studies requiring spectral radiative properties of the examined leaves with respect to ultraviolet irradiation.     

Acorn production prediction models for five common oak species of the eastern United States

Acorn production varies considerably among oak (Quercus) species, individual trees, years, and locations, which directly affects oak regeneration and populations of wildlife species that depend on acorns for food. Hard mast indices provide a relative ranking and basis for comparison of within- and between-year acorn crop size at a broad scale, but do not provide an estimate of actual acorn yield—the number of acorns that can potentially be produced on a given land area unit based on the species, number, and diameter at breast height (dbh) of oak trees present. We used 10 years of acorn production data from 475 oak trees to develop predictive models of potential average annual hard mast production by five common eastern oak species, based on tree diameter and estimated crown area. We found a weak (R² = 0.08–0.28) relationship between tree dbh and acorn production per unit crown area for most species. The relationship between tree dbh and acorn production per tree was stronger (R² = 0.33–0.57). However, this is because larger-dbh trees generally have larger crowns, not because they have a greater capacity to produce more acorns per unit crown area. Acorn production is highly variable among individual trees. We estimated that dbh of at least 60 dominant or codominant oak trees per species should be randomly sampled to obtain an adequate representation of the range of dbhs (≥12.7 cm dbh) in a given forest area, and achieve precise estimates when using these equations to predict potential acorn production. Our predictive models provide a tool for estimating potential acorn production that land managers and forest planners can apply to oak inventory data to tailor estimates of potential average annual acorn production to different forest management scenarios and multiple spatial scales. © 2011 The Wildlife Society.     

Spatial pattern of Quercus ilex and Quercus pubescens recruitment in Pinus halepensis dominated woodlands

Abstract. European Mediterranean landscapes have undergone changes in structure in recent years as a result of widespread agricultural land abandonment and cessation of silvicultural regimes. Studies concerning the regeneration dynamics of dominant forest species have become critical to the prediction of future landscape trends in these changing forest stands. Quercus ilex (holm oak) and Q. pubescens (downy oak) are considered to be the terminal point of secondary succession in extensive areas of the Mediterranean region. Recent studies, however, have suggested the existence of recruitment bottlenecks in oak genet populations as a result of current management regimes. In this study, we present evidence of the successful establishment of Q. ilex and Q. pubescens in Pinus halepensis (Aleppo pine) woodlands. We investigate the distribution patterns and spatial relationships among oak recruits and resident pines. Established P. halepensis is randomly distributed throughout the study area. Oak seedlings are positively associated with pine trees, suggesting that P. halepensis individuals provide safe sites for oak genet recruitment. We show that spatial patterns of recruitment are in agreement with the general model of spatial segregation described for other Mediterranean plant communities, with seeder species colonizing large openings after disturbance, followed by a more aggregated recruitment of resprouter species.     

Acorns containing deeper plumule survive better: how white oaks counter embryo excision by rodents

Several squirrel species excise the embryo of acorns of most white oak species to arrest germination for long‐term storage. However, it is not clear how these acorns counter embryo excision and survive in the arms race of coevolution. In this study, we simulated the embryo excision behavior of squirrels by removing 4 mm of cotyledon from the apical end of white oak acorns differing in embryo depths to investigate the effects of embryo excision on acorn germination and seedling performance of white oak species. The embryo depth in the cotyledons was significantly different among white oak acorns, with Quercus mongolica containing the embryo most deeply in the acorns. We found that artificial embryo excision significantly decreased acorn germination rates of Quercus variabilis, Quercus acutissima, Quercus aliena, Quercus aliena var. acutiserrata, Quercus serrata. var. brevipetiolata but not Q. mongolica. Artificial embryo excision exerted significant negative impacts on seedling performance of all oak species except Quercus aliena. Our study demonstrates the role of embryo depth of acorns in countering embryo excision by squirrels and may explain the fact that squirrels do not perform embryo excision in acorns of Q. mongolica with deeper embryos. This apparent adaptation of acorns sheds light on the coevolutionary dynamics between oaks and their seed predators.