Identification of the selective forces on plant dispersal engenders theoretical argument, empirical study, and speculation. We separate evidence, testable hypotheses, and conjecture surrounding two major questions in dispersal ecology. The first asks what ecological, and ultimately evolutionary, advantages exist in seed dispersal. Astonishingly little is known about the advantages to a parent plant that are actually conferred by investment in dispersal structures. Does dispersal enable seeds and ultimately seedlings to escape mortality near the parent? Is continual recolonization of unstable habitats the primary advantage? Must seeds find rare microhabitats suitable for reestablishment? Such issues are addressed through joint consideration of dispersal and establishment-those stages both mediated by parental provisioning and subject to the highest mortality in the life of a plant. The second broad question asks what general and explicit environmental forces influence the timing and mode of dispersal. Do climates or seasons favor one dispersal mode over another? Do differences in number, size, morphology, or nutritional quality of fruits influence frugivore choice, and consequently differential dispersal of species or individuals within species? Studies of dispersal process and mode should be intimately connected. A general objective of this paper is to explore the degree to which they are integrated and, in so doing, to catalyze their union. We emphasize topics most in need of critical attention: the evolutionary ecology of dispersal process and mode. Excellent recent reviews consider such related topics as dispersal mechanism (131, 184), seed dormancy (1, 30), phytogeography (11, 115, 146), masting and predator satiation (105, 156), and succession (68, 69, 189).

https://www.jstor.org/stable/info/2097067

Ecology of Seed Dispersal

Gene dispersal (flow, or migration) within and between plant populations has been of continuous interest to plant breeders and seed producers for many decades. Economic considerations have stimulated studies of gene flow as a function of distance, breeding system, pollinating agent, and planting design in numerous domestic plants. Only during the past two decades have a large body of plant evolutionists become interested in information accruing from these studies, and in the rates of gene flow in wild populations. Their efforts have concentrated primarily on related problems such as adaptations for and mechanics of pollen and seed (or fruit) dispersal, plant-pollinator coevolution, adaptive radiation in pollination and seed dispersal mechanisms, and colonization and the alteration of species boundaries. Early in this century, anecdoctal evidence on the movement of pollen and seed vectors, dispersal of pollen by wind, and the range extensions of weed species led to the casual assumption that gene flow must be extensive, and that it must play a major role in the cohesion of populations and population systems. This view eroded as more information became available and was more critically interpreted (e.g., Grant, 1958, 1971; Ehrlich and Raven, 1969; Stebbins, 1970a; Bradshaw, 1972).

Gene Flow in Seed Plants

Recent work on flapping hawkmoth models has demonstrated the importance of a spiral 'leading-edge vortex' created by dynamic stall, and maintained by some aspect of spanwise flow, for creating the lift required during flight. This study uses propeller models to investigate further the forces acting on model hawkmoth wings in 'propeller-like' rotation ('revolution'). Steadily revolving model hawkmoth wings produce high vertical ( approximately lift) and horizontal ( approximately profile drag) force coefficients because of the presence of a leading-edge vortex. Both horizontal and vertical forces, at relevant angles of attack, are dominated by the pressure difference between the upper and lower surfaces; separation at the leading edge prevents 'leading-edge suction'. This allows a simple geometric relationship between vertical and horizontal forces and the geometric angle of attack to be derived for thin, flat wings. Force coefficients are remarkably unaffected by considerable variations in leading-edge detail, twist and camber. Traditional accounts of the adaptive functions of twist and camber are based on conventional attached-flow aerodynamics and are not supported. Attempts to derive conventional profile drag and lift coefficients from 'steady' propeller coefficients are relatively successful for angles of incidence up to 50 degrees and, hence, for the angles normally applicable to insect flight.

The aerodynamics of revolving wings I. Model hawkmoth wings.

Is there necessarily a trade-off between seed size (mass) and dispersal capacity for wind-dispersed diaspores? Within three families (Pinaceae, Aceraceae, and Leguminosae) with asymmetric samaras, shape is maintained (isometry) despite size change. Consequently, within these three families, equilibrium descent velocity is proportional to samara mass raised to the 1/6 power and, necessarily, larger samaras are more poorly dispersed

Seed mass and dispersal capacity in wind-dispersed diaspores

As they descend, the autorotating seeds of maples and some other trees generate unexpectedly high lift, but how they attain this elevated performance is unknown. To elucidate the mechanisms responsible, we measured the three-dimensional flow around dynamically scaled models of maple and hornbeam seeds. Our results indicate that these seeds attain high lift by generating a stable leading-edge vortex (LEV) as they descend. The compact LEV, which we verified on real specimens, allows maple seeds to remain in the air more effectively than do a variety of nonautorotating seeds. LEVs also explain the high lift generated by hovering insects, bats, and possibly birds, suggesting that the use of LEVs represents a convergent aerodynamic solution in the evolution of flight performance in both animals and plants.

https://science.sciencemag.org/content/sci/324/5933/1438.full.pdf

Leading-Edge Vortices Elevate Lift of Autorotating Plant Seeds

SUMMARY

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

R. Åke Norberg

Papers

Autorotation, self-stability, and structure of single-winged fruits and seeds (samaras) with comparative remarks on animal flight