被子植物翅果的多样性及演化

翅果能够依靠风力进行传播, 可能是被子植物快速散布和物种分化的一个重要因素。狭义的翅果是指果皮延伸成翅且不开裂的干果; 广义的翅果则包括果皮、花被片或苞片形成果翅的所有果实。根据果翅形态及其生长方式的不同, 广义的翅果可分为单侧翅果、周位翅果(圆翅果与蝶翅果)、棱翅果、披针翅果、翼状萼翅果、叶状苞翅果6种类型, 其空中运动方式有自旋式(单侧翅果、翼状萼翅果)、波浪式(周位翅果、叶状苞翅果)、翻滚自旋式(周位翅果)、直升机式(披针翅果、翼状萼翅果)和滚筒式(棱翅果)。棱翅果与圆翅果在被子植物基部类群樟目就有发生, 并同时出现在单子叶植物和双子叶植物中, 可能是最早出现的翅果类型。翅果的演化过程呈现出果翅数量增加、果翅偏向单侧和果翅负荷(果实质量与果翅面积之比)降低的趋势, 以利于适应较小的风并增加传播距离。果翅除了促进果实与种子的风力传播外, 还具有物理防御、调节种子萌发和促进二次传播等作用。泛热带分布的金虎尾科有着极其丰富的翅果类型, 与其多次跨洋长距离扩散密切相关, 可以作为研究翅果适应与演化的一个模式类群。结合生态和演化-发育生物学方法, 研究不同类型翅果在适应风力传播方面的差异、萼片或苞片发育成翅的分子与遗传机制、翅果不同类型的演化历史及其对被子植物物种多样性的影响等是今后值得探讨的重要问题。     

绒毛白蜡二态型果实的扩散特征

多态型果实或种子的出现对植物种群的扩散具有重要的意义。绒毛白蜡(Fraxinus velutina)的果实具有二态型特征,主要表现在果翅数量上不同,分别定义为二翅型和三翅型果实,为了比较两类果实在风传扩散时的差异,研究了两类果实的形态、果翅结构和扩散距离及扩散时长。在大型封闭地下室内,以电扇在不同速度档位产生的气流作为风源,分别从2、1.5、1m处手动释放果实,对风速为0、4.6、6.5、7.3m/s时的果实扩散距离及扩散时长进行了比较;并在此基础上对果实的形态特征与扩散特征进行了线性相关分析。结果表明:在同一高度及相同风速下,三翅型果实的水平扩散距离都极显著的大于二翅型,但其相应的扩散时长都小于二翅型。在相同情况下,三翅型的果实沉降速度显著高于二翅型。两类果实随着释放高度的增加,其扩散距离和扩散时长都相应的增加;随着风速的升高,其扩散距离及扩散时长都相应的增加。三翅型果实质量显著高于二翅型,相反,三翅型果翅长与宽都显著小于二翅型。两种翅型的果翅细胞结构都一样,细胞内部都呈现气囊状,果翅表面沿纵轴方向有流线型的纵棱。通过直线相关分析发现,翅型是对扩散距离和扩散时长影响最显著的形态特征;与果实释放高度相比,风速是影响绒毛白蜡果实扩散距离与扩散时长最显著的环境因素。绒毛白蜡三翅型果实比二翅型果实传播的远,关键在于其具有三翅,三翅对阵风瞬间响应,使得沉降速度较二翅型高,可以在瞬时风的作用下,快速传播到较远的距离。三翅型与二翅型在扩散方式上的结合增强了绒毛白蜡的生存与定殖机会。     

Identification of asymmetrically winged samaras from the Western Hemisphere

A key is presented for use in identifying asymmetrically winged fruits (samaras) with either proximal or distal locules. It aids identification based on dispersed fuit morphology and can be used to identify undetermined extant herbarium specimens or fossil fruits to the correct extant family and genus. The 39 genera from 11 families (Aceraceae, Anacardiaceae, Fabaceae, Malpighiaceae, Phytolaccaceae, Polygalaceae, Polygonaceae, Rutaceae, Sapindaceae, Trigoniaceae, Ulmaceae) are distinguished on the basis of wing venation, size of fruit, presence and position of attachment surface, presence and type of subsidiary wings on the ovary wall. ornamentation, size and shape of the ovary, locule position, shape of locule cross section, style position and ornamentation, distinction between ovary wall and wing, and angle of attachment between individual samaras. The developmental origins of some of these features are discussed.     

Secondary dispersal mechanisms of winged seeds: a review

Winged seeds, or samaras, are believed to promote the long‐distance dispersal and invasive potential of wind‐dispersed trees, but the full dispersive potential of these seeds has not been well characterised. Previous research on the ecology of winged seeds has largely focussed on the initial abscission and primary dispersal of the samara, despite it being known that the primary wind dispersal of samaras is often over short distances, with only rare escapes to longer distance dispersal. Secondary dispersal, or the movement of the seeds from the initial dispersal area to the site of germination, has been largely ignored despite offering a likely important mechanism for the dispersal of samaras to microhabitats suitable for establishment. Herein, we synthesise what is known on the predation and secondary dispersal of winged seeds by multiple dispersive vectors, highlighting gaps in knowledge and offering suggestions for future research. Both hydrochory and zoochory offer the chance for samaroid seeds to disperse over longer distances than anemochory alone, but the effects of the wing structure on these dispersal mechanisms have not been well characterised. Furthermore, although some studies have investigated secondary dispersal in samaroid species, such studies are scarce and only rarely track seeds from source to seedling. Future research must be directed to studying the secondary dispersal of samaras by various vectors, in order to elucidate fully the invasive and colonisation potential of samaroid trees.     

The effect of samara wing presence on predation of Acer pseudoplatanus (Sapindaceae) seeds on the ground

The key selective pressure shaping the morphology of samaras is seen as enhancing primary wind-borne dispersal from the parent plant to the ground. However, the consequences of the samara wing of primarily wind-dispersed tree species for post-dispersal processes has not been well studied. We explored whether the presence of this wing in Acer pseudoplatanus either deters or promotes predation after dispersal, either by increasing the time and energy required to predate the seed or by increasing the seed's visibility to predators. We found that wing-removed fruits were preferred, suggesting that the presence of samaras makes seed handling more expensive for granivores. Further, we found that fewer seeds were consumed from treatments that contained the most winged seeds, thus there was no evidence of the samaras making seed finding easier for granivores. We conclude that the presence of the wing may offer an anti-predatory benefit as well as aiding primary dispersal.     

Gone with the wind and the stream: Dispersal in the invasive species Ailanthus altissima

Dispersal is a key process in plant invasions and is strongly related to diaspore morphology. Often, dispersal comprises more than one step, and morphologies adapted to a primary dispersal mechanism can aid or detract from a secondary one. The aim of this work was to assess the relationship between primary wind dispersal and secondary water dispersal in Ailanthus altissima, an invasive tree species. Wind and water dispersal potential and their association with the morphological characteristics of samaras were assessed under controlled conditions to ensure the repeatability of the measurements. We found a direct positive relationship between primary wind and secondary water dispersal in A. altissima. The main morphological characteristics of the samara that affected the success of the two types of dispersal were side perimeter and mass. However, a possibility of dispersal specialisation exists, as one morphological characteristic (samara width) affects wind dispersal negatively but water dispersal positively, and dispersal potential and samara morphology have been shown to differ across individuals.     

Seed size and dispersal potential of Acer rubrum (Aceraceae) samaras produced by populations in early and late successional environments

In order to determine if red maple dispersal potential or seed size change during secondary succession, samaras were collected from five populations located in early successional environments and five populations located in late successional environments. Wing loading ratios (samara mass—mg/samara area—cm²), which are inversely proportional to dispersal ability, were computed for all samaras, and seeds were excised from each samara and weighed. Samaras from the early successional red maples showed slightly but significantly lower wing loading ratios than those from the late successional environments. This result corresponds with the conclusions reached by several theoretical investigations of seed dispersal evolution that predict that recently founded populations will show greater dispersal abilities than more established populations. The earlier successional populations had slightly heavier seeds than the later successional populations, which suggests that the changes in community composition and dynamics that occur during this successional sequence do not select for heavier seeds in older red maple populations. Coefficients of variation for wing loading and seed size showed no consistent trends with successional stage, which indicates that variation in these characters does not decrease as succession proceeds.     

THE TERMINAL VELOCITY AND DISPERSAL OF SPINNING SAMARAS

Strobe photographs were taken of over 200 spinning samaras from seven species of trees. These were used to measure the rate of descent, angular velocity, orientation, and other parameters of the samaras as they fell. These data were then used to compare the aerodynamic behavior of samaras, helicopters, and theoretical ideal rotors. Plotting morphological data for each samara against its rate of descent showed that this rate was highly correlated with the square root of the samara's wing loading (samara weight divided by wing-surface area). This plot demonstrated the existence of two distinct groups of samaras, distinguished by their morphology, spinning motion, and rate of descent. These results are of greatest use in characterizing local dispersal patterns.     

From falling to flying: the path to powered flight of a robotic samara nano air vehicle

This paper details the development of a nano-scale (>15 cm) robotic samara, or winged seed. The design of prototypes inspired by naturally occurring geometries is presented along with a detailed experimental process which elucidates similarities between mechanical and robotic samara flight dynamics. The helical trajectories of a samara in flight were observed to differ in-flight path and descent velocity. The body roll and pitch angular rates for the differing trajectories were observed to be coupled to variations in wing pitch, and thus provide a means of control. Inspired by the flight modalities of the bio-inspired samaras, a robotic device has been created that mimics the autorotative capability of the samara, whilst providing the ability to hover, climb and translate. A high-speed camera-based motion capture system is used to observe the flight dynamics of the mechanical and robotic samara. Similarities in the flight dynamics are compared and discussed as it relates to the design of the robotic samara.     

长白山9种槭树的翅果扩散及种子萌发研究

翅果的风媒传播是槭属植物的主要扩散方式之一,且与种子萌发有着密切关联,但具体机理一直还并不明确。以分布于长白山的9种槭树为对象,探讨翅果的形态特征,测定它们在空气中的垂直沉降速度、不同风速下的水平扩散距离以及在扩散距离上的种子萌发率,进而比较并分析翅果的形态性状与沉降速度、水平扩散距离的相关性以及萌发率在不同扩散距离上的差异性。结果表明：(1) 9种槭树的翅果长、宽和面积与沉降速度、水平扩散距离均呈负相关;尽管如此,翅果形态并不是风传播物种的最佳分类指标,而翅载力能较好地反应物种的风传播能力;(2)翅果垂直沉降速度和水平扩散距离间存在显著负相关,表明沉降速度越小,翅果在空气中停留的时间越长,水平方向上扩散距离越远,且强风有助于提高翅果的扩散能力;(3)沉降速度最慢的花楷槭在不同风速下的水平扩散距离均最远,而沉降速度最快的拧筋槭水平扩散距离最短;(4)种子萌发率随扩散距离的增加呈下降趋势。上述结果不仅为深入理解翅果的风力传播机制以及种子萌发对水平扩散距离的响应机制提供科学依据,还可为种群实生更新方面的理论研究提供参考。     

Intraspecific variation in samara morphology and flight behavior in acer saccharinum (Aceraceae)

We studied intraspecific variation in samara morphology and flight behavior within and among parent trees of Acer saccharinum (silver maple), with a particular focus on the effect of samara shape. Samara mass, area, wing loading, and descent rate from a 4.5-m indoor balcony were measured for 50 undamaged mature samaras from each of six parents. We found significant differences among parental types for all morphological variables and descent rate. These differences yielded a 50% range in mean dispersal potential among the six parents. There was a strong linear correlation between descent rate and square root of wing loading when mean values were plotted for each of the six parental types. But there was considerable within-parent variation for all measured variables, including substantial nonallometric variation in wing loading caused in part by poor correlations between wing area and fruit weight. Parents also differed widely in the relationship between square root of wing loading and descent rate (linear r² = 0.150-0.788), with one parental type showing no significant relationship. Fruits from the same parent with similar values of the square root of wing loading showed as much as a 75-100% difference in descent rate. The usefulness of mass : area indices such as wing loading is limited by its exclusion of aerodynamically important factors such as mass distribution and wing shape, which in our case caused the six parents to behave aerodynamically almost as if they were six separate species.     

Ecological role of distinct fruit‐wing perianth color in synchronization of seed germination in Haloxylon salicornicum

Haloxylon salicornicum is a perennial shrub that grows in sandy and stony desert areas. This species produces two types of dispersal unit with yellow and pink winged perianths that are wind dispersed. This species has been considered as an excellent sand binder and can tolerate extreme temperature, drought and salinity. We compared the germination characteristics of both colored dispersal units (with and without a wing) under various light, temperature and salinity treatments. It was found that wing color, presence of wings, temperature and light have a significant effect on germination percentage and germination rate. The presence of a winged perianth significantly reduced the germination percentage and germination rate in both yellow and pink dispersal units. De‐winged yellow dispersal units were able to tolerate higher salinity and, therefore, their germination was higher in all the tested concentrations of NaCl compared with pink de‐winged dispersal units. However, salinity tolerance in both pink and yellow dispersal units decreased with increasing salinity concentration. The germination‐recovery percentage was also higher for yellow compared with pink dispersal units. The present study has demonstrated that both morphological and physiological differences exist in different colored dispersal units, which might represent the combination of different complementary adaptive strategies and have ecological significance for Haloxylon salicornicum's successful survival in harsh desert environmental conditions.     

果实类型和种子传播的进化:系统发育和生态学简论

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angiosperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

Large-scale phylogenetic analyses reveal fagalean diversification promoted by the interplay of diaspores and environments in the Paleogene

The order Fagales (ca. 1325 species, 32 genera) is one of the most important orders of woody angiosperms in both tropical and temperate forests. Fagalean plants have diverse diaspores and dispersal modes, and have abundant macrofossil records. Here, we present, to our knowledge, the most comprehensive phylogenetic analysis of Fagales to date based on five plastid loci. We reconstructed the phylogenetic relationships within Fagales using parsimony, likelihood and Bayesian approaches. We inferred the evolutionary shifts of diaspore types, dispersal modes, habitats, and pollination syndromes, and estimated divergence times and rates of diversification. Fossil fruit records of Fagales were also reviewed. Our results suggest that fagalean families are all monophyletic and 29 of the 32 genera are monophyletic. The majority of inter-familial and inter-generic relationships were well resolved. Our evolutionary reconstructions indicate that winged diaspore is synapomorphic, and animal dispersal system is symplesiomorphic in Fagales. Within the order, the families diverged in the mid-Cretaceous but mainly diversified after the Cretaceous–Paleogene (K–Pg) boundary. The overwhelming majority of winged and wingless fruited genera diverged or diversified during the Paleogene, reflecting adaptation to wind and animal dispersals, respectively. Correlated evolution analyses strongly supported the correlated transitions between dispersal mode and habitat. The winged fruited groups often inhabit open habitats and are commonly dispersed abiotically, whereas the wingless fruits usually inhabit closed habitats and have a vertebrate-dispersal mode. Environmental changes triggered increased diversification of Fagales in the Paleogene together with evolution of diaspores and dispersal modes. Additionally, our paleobotanically calibrated time-scale for Fagales may be useful for ecological and physiological studies.     

The seed dimorphism of Spergularia marina in relation to dispersal by wind and water

Summary The seeds of the halophyte Spergularia marina differ both within and between individuals in that they either possess or lack a membranaceous border. This paper presents a morphological study of the length, weight and area of the seed types, and their dispersal characteristics under experimental conditions of wind and water dispersal. The winged seeds are shown to be larger both by length and by weight. Their rate of descent increases with wing loading. If the wing is lacking, however, the rate of descent increases with weight only. The distance of dispersal is equal for both seed types except at low wind speeds, when the winged seeds disperse farther. If the seed wing is removed, the excised seeds have shorter dispersal distances. When dispersed by water, a difference in the distance seeds are dispersed can only be detected in the presence of vegetation. The winged seeds are more frequently trapped in the vegetation as compared to the unwinged seeds. The hypothesis that the seed dimorphism is an adaptation for differential dispersal distances is discussed.     

Increased seed dispersal potential towards geographic range limits in a Pacific coast dune plant

Dispersal may be favoured at geographic range edges by unstable population and metapopulation dynamics. However, dispersal may also evolve in response to geographic variation in other life-history traits, especially the mating system. Here, increased dispersal at range margins was tested for with a range-wide analysis of seed dispersal and mating system traits in Abronia umbellata, a plant endemic to Pacific coastal dunes of North America. Seeds disperse within winged anthocarps. Anthocarps from 34 populations varied widely in wing size (mass-corrected wing index). Wing index correlated negatively with threshold wind velocity for dispersal in wind tunnel tests, suggesting that wings facilitate tumbling over open sandy substrate. As predicted, wing index increased and threshold velocity decreased towards both range limits. Flower size, herkogamy and self-incompatibility declined towards range limits, indicating a shift to self-fertilization, and flower size and wing index correlated negatively. However, the increase in wing index towards range limits remained after statistically controlling flower size. These results are consistent with selection favouring dispersal at range margins. The evolutionary lability of dispersal across the range may affect the interaction between selection and gene flow in the establishment and maintenance of geographic range limits.     

AUTOROTATION, SELF-STABILITY, AND STRUCTURE OF SINGLE-WINGED FRUITS AND SEEDS (SAMARAS) WITH COMPARATIVE REMARKS ON ANIMAL FLIGHT

• 1 A samara is a winged fruit or seed that autorotates when falling, thereby reducing the sinking speed of the diaspore and increasing the distance it may be transported by winds. Samaras have evolved independently in a large number of plants.
• 2 Aerodynamical, mechanical, and structural properties crucial for the inherent self-stability are analysed, and formulae for calculation of performance data are given.
• 3 The momentum theorem is applied to samaras to calculate induced air velocities. As a basis for blade element analysis, and for directional stability analysis, various velocity components are put together into resultant relative air velocities normal to the blade's span axis for a samara in vertical autorotation and also in autorotation with side-slip.
• 4 When falling, a samara is free to move in any sense, but in autorotation it possesses static and dynamic stability. Mainly qualitative aspects on static stability are pre sented. Simple experiments on flat plates at Reynolds numbers about 2000 as in samaras, showed that pitch stability prevails when the C. M. (centre of mass) is located 27–35 % of the chord behind the leading edge. The aerodynamic c.p. (centre of pressure) moves forward upon a decrease of the angle of attack, backward upon an increase. In samara blades the c.m. lies ca. one-third chord behind the leading edge, and hence the aerodynamic and centrifugal forces interact so as to give pitch stability, involving stability of the angles of attack and gliding angles.
• 5 Photographs show that the centre of rotation of the samara approximately coincides with its c.m.
• 6 The coning angle (blade angle to tip path plane) taken up by the samara is determined by opposing moments set up by the centrifugal and aerodynamic forces. It is essentially the centrifugal moment (being a tangent function of the coning angle, which is small) that changes upon a change of coning angle, until the centrifugal and aerodynamic moments cancel out at the equilibrium coning angle.
• 7 Directional stability is maintained by keeping the tip path plane horizontal whereby a vertical descent path relative to the ambient air is maintained. Tilting of the tip path plane results in side-slip. Side-slip leads to an increased relative air speed at the blade when advancing, a reduced speed when retreating. The correspondingly fluctuating aerodynamic force and the gyroscopic action of the samara lead to restoring moments that bring the tip path plane back to the horizontal.
• 8 Entrance into autorotation is due to interaction between aerodynamic forces, the force of gravity, and inertial forces (when the blade accelerates towards a trailing position behind the c.m. of the samara).
• 9 The mass distribution must be such that the c.m. lies 0–30 % of the span from one end. In Acer and Plcea samaras the C.M. lies 10–20% from one end, thereby making the disk area swept by the blade large and the sinking speed low.
• 10 The blade plan-form is discussed in relation to aerodynamics. The width is largest far out on the blade where the relative air velocities are large. The large width of the blade contributes to a high Re number and thus probably to a better L/D (lift/drag) ratio and a slower descent.
• 11 The concentration of vascular bundles at the leading edge of the blade and the tapering of the blade thickness towards the trailing edge are essential for a proper chord wise mass distribution.
• 12 Data are given for samaras of Acer and Plcea, and calculations of performance are made by means of the formulae given in the paper. Some figures for an Acer samara are: sinking speed 0.9 m/sec, tip path inclination 15°, average total force coefficient 1.7 (which is discussed), and a L/D ratio of the blade approximately 3.
• 13 The performances of samaras are compared with those of insects, birds, bats, a flat plate, and a parachute. They show the samara to be a relatively very efficient structure in braking the sinking speed of the diaspore.
• 14 In samaras the mass, aerodynamic, and torsion axes coincide, whereas in insect wings the torsicn axis often lies ahead of the other two. Location of the torsion axis in front of the aerodynamic axis in insects tends towards passive wing twisting and passive adjustment of the angles of attack relative to the incident air stream, the direction of which varies along the wing because of wing flapping.
• 15 Location of the mass axis behind the torsion axis may lead to unfavourable

     

Effects of monsoon on distribution patterns of tropical plants in Asia

Comparing with other regions, Asia is mostly dominated by the monsoon climate and tropical plants can be found at the furthest places away from the equator. Understanding the role of monsoon in the dispersal and evolution of tropical plants is helpful for exploring the distribution patterns of vegetation and mechanisms underlying the origin and maintenance of biodiversity in Asia. In summer, there are three types of monsoon in Asia, i.e. East Asia Monsoon, South Asia Monsoon, North-west Pacific Ocean Monsoon. The summer monsoon climate in Asia originated at about 40 Ma, when the early angiosperm evolved and started its diversification in Southeast Asia and South China. It suggested that the monsoon may facilitate the quick speciation and spread of early angiosperm. Monsoon climate facilitates the northward spread of Asia’s tropical plants and some tropical plants can be found even at Yarlung Zangbo River and the boundaries of Guizhou-Guangxi-Yunnan. Such effetcs largely change distribution patterns of zonal vegetation and even causes local vegetation types in some places with unusual topography such as tropical seasonal rainforests, monsoon rainforests, savanna and grassland along dry-hot valley in Southwest China, coastal savanna in West Hainan Island. The three summer monsoons interact at Southwest China and Indo-China Peninsula and these regions are dominated by limestone landscapes and high mountains with big rivers. Some Asia-endemic tropical taxa even formed a diversification and endemism center at this region, which may be a reason for the formation and maintenance of Indo-Burma biodiversity hotspots with global warming, the monsoon may further promote the northward spread of tropical plants and may have fundamental effects on biodiversity and flora evolution in South China.     

The fossil record of Eucommia (Eucommiaceae) in North America

Reproductive and vegetative remains of Eucommia from 25 localities in the United States, Canada, and Mexico document the wide distribution of this genus in North America during the Cenozoic. Autofluorescent elastic latex filaments bearing capitate termini are preserved in nearly all of the remains and provide conclusive evidence of their affinity to Eucommia. Four species of Eucommia are recognized on the basis of the characteristic samaras: E. eocenica from middle Eocene strata of the Mississippi Embayment in Missouri, Tennessee and Mississippi; E. montana from early Eocene to early Oligocene localities in British Columbia, Washington, Oregon, Utah, Colorado, and Montana; E. constans from Neogene rocks in central Mexico; and E. jeffersonensis n. sp. from the latest Eocene or earliest Oligocene John Day Formation of Oregon. Atypical specimens of E. montana and E. eocenica are the first records of two-seeded fruits for the genus. Eucommia leaves from Eocene localities in British Columbia and Mississippi are the first records of Eucommia foliage in North America whose identifications are confirmed by the presence of capitate latex strands. These leaves and a specimen from Oregon are referred to E. rolandii n. sp. Fruit evolution in Eucommia may have involved increases in samara size and symmetry, and reduction in seed number from two to one, perhaps as adaptations for wind dispersal. All fossil Eucommia samaras from North America are smaller and less symmetrical than those of the living species, E. ulmoides. Preliminary flight tests of E. ulmoides samaras and of models of the fossils suggest that E. ulmoides fruits are aerodynamically better suited for wind dispersal than the fossils.     

The taxonomic value of fruit wing types in the order Apiales

This study shows that structural data, when carefully examined, can provide valuable characters for delimiting monophyletic groups and can complement DNA with observable features to recognize and circumscribe taxa. In the angiosperm order Apiales, traditional classification has relied heavily (often exclusively) on fruit characters. Recent molecular systematic studies, however, provided a radically different picture of relationships, calling into question the utility of fruit characters. We have studied fruit anatomy from 18 genera (Annesorhiza, Asteriscium, Astrotricha, Choritaenia, Dasispermum, Elaeoselinum, Heptaptera, Hermas, Heteromorpha, Laretia, Molopospermum, Myodocarpus, Pachypleurum, Peucedanum, Polemanniopsis, Polylophium, Rouya, and Tordylium) that represent all major taxonomic groups of Apiales characterized by winged fruits and the full range of wing types. Fruit anatomy closely corresponded with the phylogenetic position of these genera, as suggested by molecular studies. Fruit features of taxonomic importance include developmental origin of the wings, carpel shape, presence of vittae, woodiness of the endocarp, position of crystals, and type of carpophores. Despite the long history of recognizing umbellifers as a "natural group," few studies have been able to provide structural characters to help circumscribe the clades identified by molecular data. The interpretations presented are an important step toward erecting a stable system of classification for this difficult family.