Amer. J. Bot. 73(3): 353-363. 1986.

MORPHOLOGY AND DISPERSAL POTENTIAL OF WIND-DISPERSED

DIASPORES OF NEOTROPICAL TREESI

CAROL K. AUGSPURGER
Department of Plant Biology, University of lI1inois, 505 S. Goodwin, Urbana, lI1inois 6180 I

ABSTRACT
Morphological and aerodynamic traits affecting mean potential dispersal distance are quan-
tified for wind-dispersed diaspores of tree species on Barro Colorado Island, Panama. The
sample includes 34 species in 16 families and represents six aerodynamic groups. Mass and
area (maximum cross section) each vary over six orders of magnitude among the species. In
contrast, wing-loading, defined as weight divided by area, varies over only one order of mag-
nitude, as does the rate of descent. While the regression of rate of descent on the square root
of wing-loading is significant overall, the slopes vary significantly among five aerodynamic
groups. At comparable wing-loading values, diaspores of fluffy kapok fall faster than four other
aerodynamic groups and rolling autogyros fall faster than non-rolling autogyros. Assuming the
diaspores are released from their typical tree height and experience a mean windspeed of 1.75
m sec:", the expected mean dispersal distance varies among the 34 species from 22 to 194 m.
Rate of descent is weakly correlated with shade tolerance of seedlings for a subset of 18 species;
rate of descent is more strongly correlated with the log of dry mass of seed for all 34 species.
Given these wide differences in dispersal potential, any generalizations about tropical trees that
use wind dispersal are of dubious value.

MORPHOLOGICAL DESIGNS of wind-dispersed quences of this variation for rate of descent

diaspores appear to slow their rates of descent and dispersal potential are the subject of this
and increase their chances of exposure to hor- study.
izontal or gusty winds. Rate of descent and Horizontal winds spread diaspores over a
dispersal potential have been documented for broad area and carry them away from the par-
temperate herbaceous species (Sheldon and ent. Hence, the potential dispersal distance and
Burrows, 1973; Werner and Platt, 1976; Platt area are inversely related to the rate ofdescent.
and Weis, 1977; Rabinowitz and Rapp, 1981) Because consistency ofwind direction will also
and for temperate trees of North America (Sig- affect the total area over which diaspores are
gins, 1933; Green, 1980; Guries and Nord- dispersed, many studies of wind dispersal, in-
heim, 1984) and Eucalyptus trees of Australia cluding the present one, focus on dispersal po-
(Cremer, 1977). Values for rate of descent for tential. Dispersal potential here refers specif-
temperate tree seeds have been incorporated ically to potential mean dispersal distance,
into a model of wind dispersal that predicts although the area covered is implicitly in-
the seeds' spatial distribution under specified volved as it tends to increase with increasing
wind conditions and crop size (Fields and mean distance.
Sharpe, 1980; Sharpe and Fields, 1982). In the Green (1980) found for seven species oftem-
species-rich tropical forests, there is much vari- perate trees that a diaspore's rate of descent is
ation in morphology and size of wind-dis- highly correlated with its (wing-loading) 1/2; here
persed diaspores (Ridley, 1930). The conse- wing-loading is defined as wt/area. His dia-
spores fall into two morphological groups; one
group simultaneously autorotates around the
I Received for publication 4 June 1985; revision ac-

cepted 27 September 1985.

diaspore's longitudinal axis and autogyrates
This research was supported by NSF grant BSR 8219856 around one end of the diaspore ("rolling
and was made possible by the support facilities and per- samaras"), while the other group only autogy-
sonnel of the Smithsonian Tropical Research Institute, rates ("non-rolling samaras"). For a given wing-
especially D. Windsor and the Environmental Science Pro- loading, the first group has a faster rate of de-
gram. I thank R. Beebe and his staff for use ofthe Krannert
Center for the Performing Arts, Univ. of 111., to determine scent in still air than the second. The validity
rates of descent, L. Siri Kimsey for the artwork, S. Franson of these relationships can be further tested us-
for statistical analyses, and B. Benner, S. Franson, K. Ho- ing the wider variety of morphological groups
gan, C. Kelly, and H. Michaels for assistance in deter- and sizes present among tropical species.
mining rates of descent. R. B. Foster collected the fruits
of Centrolobium sp. in Peru. K. Hogan and S. Franson
This paper presents data for 34 species of
contributed constructive comments on earlier drafts of the tropical trees found on Barro Colorado Island,
manuscript. Panama. I investigated the relationship of
353
354 AMERICAN JOURNAL OF BOTANY [Vol. 73

(wing-loading)!" to the rate of descent for six determined shortly after the time of dispersal.
diverse morphological groups to determine Diaspores with a mass ~ 100 mg were weighed
whether the groups vary in their dispersal po- on a Cahn electrobalance, those with a mass
tentials. Using representative tree ht and wind > 100 mg on a Mettler top-loading balance.
speeds, expected mean dispersal distances were Area was defined as the maximum cross sec-
then calculated for each species. Finally, using tion of the diaspore and was determined using
existing data for shade tolerance of seedlings a Licor leaf area meter. The majority of the
and dry wt of seeds of the species (Augspurger, diaspores were flat enough so they could be
1984), I tested whether rates of descent are passed directly through the meter. For dia-
related to the light requirements for seedlings spores with a pronounced third dimension, the
of these species. The probability of some di- following modifications were required. For
aspores landing in a light-gap increases as di- species 4, 5, 8, 22, and 25 (Fig. 1), the radius
aspores descend more slowly, thus traveling of the fluffy sphere was used to calculate the
greater distances and spreading over a larger maximum cross-sectional area of the sphere.
area. Hence, species requiring light-gaps for For species 21, 24, and 34 (Fig. 1), the shape
early seedling survival may have slower rates of the diaspores was traced onto paper, cut,
of descent of their diaspores. and passed through the meter. For species 3,
11, and 33 (Fig. 1), all wings were traced onto
STUDY SPECIES AND METHODS- The study in- paper, cut, and passed through the meter. For
volves 34 tree species in 16 families and in- species 6 (Fig. 1), all five wings were traced,
cludes approximately 90% oftree species using cut, and passed through the meter and mean
wind dispersal in the semi-deciduous forest on area per wing was determined; the area of this
Barro Colorado Island, Panama (Table 1, Fig. diaspore producing lift at anyone time and
1). Croat (1978) provides a detailed description used to calculate its wing-loading was consid-
of each species. Foster and Brokaw (1982) de- ered to equal the area of two wings. Finally,
scribe the vegetation on the island. The trees for species 1, 9, and 20 (Fig. 1), the diaspores
include emergent, canopy, and subcanopy trees. were magnified using a photographic enlarger;
Their crown ht range from 15-45 m, and the their magnified shadows were traced, cut, and
ht of a tree relative to the surrounding vege- passed through the meter.
tation is expected to affect wind dispersal as Wing-loading was defined as wt (mass x ac-
well. The average canopy ht of the old forest celeration) divided by area. This value was
on Barro Colorado Island is 30-40 m; it is 10 used in subsequent analyses to determine
m shorter in the young forest (Foster and Bro- whether the rate of descent was proportional
kaw, 1982). to (wing-loading)':".
Field collections of diaspores were made in The rate ofdescent in still air was determined
1982 and 1983. Because of the high species for each diaspore shortly after field collection.
diversity in this tropical forest, individual adults Each diaspore was dropped from the top of a
of a particular species are sometimes widely theater stage and the time required to fall
separated, and not all individuals of a species through 27 m of still air was determined. This
fruit in the same year. Therefore it is possible ht approximates the mean ht of reproductive
to locate isolated individual parents of these canopy trees on Barro Colorado Island. The
species. Fifteen intact diaspores were collected minute diaspores of species 1, 9, and 20 were
from the ground just outside the crown of a dropped from a ht of 4 m so that they could
specified parent tree for each species. This stan- be observed more readily. The rate of descent
dard location was selected for all species to (cm· sec:") is not precisely equivalent to the
allow interspecific comparisons. Previous terminal velocity because the diaspore requires
measurements of spatial distributions of dis- some time after release before it begins its aero-
persed diaspores indicated that this location dynamic movement and before it reaches its
was typically where the peak density of seeds terminal velocity. For most species this time
occurred (c. Augspurger, unpubl. data). Al- is a very minor portion of the total fall time;
though these samples may be slightly biased the maximum time was approximately 1.5 sec
toward diaspores with low dispersal ability, (8.4% of the total time) for those species with
such a bias is consistent across species. Also large wing-loading values. This initial free fall
given the stochastic element of dispersal and time is a small proportion of total time for
the fact that dispersal potential and distance these tall trees, but can become important for
are only loosely coupled for a given diaspore, smaller trees and shrubs (see Guries and Nord-
even diaspores with high dispersal potential heim, 1984).
may be included in the sample. The repeatability of the results for rate of
The mass and area for each diaspore were descent was determined for a subset of species.
March, 1986] AUGSPURGER-WIND-DISPERSED TROPICAL DIASPORES 355

FLOATER UNDULATOR

10 HELICOPTER

ROLLING AUTOGYRO ~05Cm~m

~C)~
II 33

\
2em 29
;;.~
30 31 ~
~ 23 3
~~
~ 13
17 ~ TUMBLER
~15~
O.5em lem
1--1 ~
--=-9
20

AUTOGYRO

~ 34 2em

~' NONCLASSI FIED

____--........24 ~7
28~
_~~~ <019 27~
--.:Jf7 21" 4:::> 18
2em
~

Fig. 1. Morphologies of the diaspores of 34 tree species using wind dispersal on Barro Colorado Island, Panama.
The number below a diaspore refers to its species number (see Table I for species identification). The label above a
cluster of diaspores refers to their aerodynamic classification (see Table 2 for a description of aerodynamic behaviors).
TABLE 1. Morphological characters oj wind-dispersed diaspores, rates of descent in still air, tree height, and expected mean dispersal distance at three wind speeds for 34 tree v.>
species on Barro Colorado Island, Panama. N = 15 for each species (except N = 7 for species 4, N = 3 Jar species 12, and N = 4 for species 17). Taxonomic nomenclature VI
0\
follows Groat (1978)

Rank
Expected mean of ex- Rank
dispersal distance (m) peeted of
(Wing- Rate of descent Tree _ _ _ _ _ _ _ _ disper- shade Wood
Type of Aerodynamic Mass (mg) Area (em') loadingj'v (em sec"} height 7 sal dis- toler- density
diaspore group x (SO) x (SO) x (SO) X (SD) (m) 1.75 3.5 m sec- 1b tance ances (g cm-1)d

1. Alseis blackiana Seed Rolling 0.181 (0.050) 0.030 (0.006) 76.2 (10.4) 65.0 (13.5) 35 94 188 377 31 0.57
(Rubiaceae) autogyro
2. Aspidosperma cruenata Seed Undu1ator 871 (95) 64.5 (3.3) 115.7(7.1) 94.8 (8.7) 35 65 129 258 23 5 0.71
(Apocynaceae)
3. Astronium graveolens Fruits Helicopter 47.5 (5.5) 1.93 (0.21) 143.7 (11.2) 132.8 (6.3) 35 46 92 184 12 0.99
(Anacardiaceae)
4. Bombacopsis quinata Seed Floater 32.9 (5.2) 13.7 (3.7) 49.2 (4.0) 75.6 (15.0) 35 81 162 324 27 0.39
(Bombacaceae) ;I>
5. Bombacopsis sessilis Seed Floater 462 (65) 38.8 (7.2) 109.0 (11.2) 198.1 (24.4) 25 22 44 88 2 0.42 s::tr1
(Bombacaceae) ::0
6. Cavanillesia platanifolia Fruit Tumbler 3,055 (569) 155 (14) 137.9 (9.9) 140.4 (16.2) 45 56 112 224 18 16 0.12 n
(Bombacaceae) ~
7. Cedrela odorata Seed Autogyro 14.1 (2.1) 2.28 (0.31) 78.3 (6.9) 59.7(11.4) 35 103 205 410 32 (0.47) '-<
(Meliaceae) o
C
8. Ceiba pentandra Seed Floater 74.1 (13.4) 18.2 (2.3) 63.6 (6.0) 100.3 (17.7) 45 78 157 314 26 14 0.24 ::0
(Born bacaceae)
9. Cespedezia macrophylla Seed Rolling 0.132 (0.047) 0.068 (0.008) 43.1 (6.4) 31.6 (7.1) 35 194 388 775 34 0.64
~
r-
(Ochnaceae) autogyro o'r1
10. Cochlospermum vitifolium Seed Floater 24.8 (1.1) 5.79 (0.39) 64.8 (2.8) 109.1 (15.9) 15 24 48 96 2 17 0.30
I:l:I
(Coch1osperrnaceae) o...,
11. Cordia alliodora Fruit- Helicopter 6.25 (0.68) 0.50 (0.05) 110.7 (9.7) 124.4 (9.9) 25 35 70 140 6 10 0.44
(Boraginaceae) ~
12. Couratari panamensis Seed Rolling 236 (25) 14.1 (1.5) 128.4 (8.2) 107.0 (12.2) 35 57 114 229 20 0.49 -<
(Lecythidaceae) autogyro
13. Dalbergia retusa Fruit Rolling 248 (23) 10.7 (0.9) 150.6 (5.1) 149.8 (17.0) 25 29 58 117 3 7 0.95
(Papi1ionoideae) autogyro
14. Jacaranda copaia Seed Undu1ator 6.50 (1.52) 4.68 (0.26) 36.7 (4.3) 39.0 (6.2) 30 135 269 538 33 0.38
(Bignoniaceae)
15. Lafoensia punicifolia Seed Rolling 38.0 (2.4) 2.46 (0.25) 123.4 (3.9) 109.3 (11.4) 30 48 96 192 15 6 0.72
(Lythraceae) autogyro
16. Lonchocarpus pentaphyllus Fruit Rolling 143 (15) 8.33 (0.81) 129.9 (4.5) 132.1 (15.0) 35 46 93 185 13 8 (0.75)
(Papilionoideae) autogyro
17. Lonchocarpus velutinus Fruit Rolling 127 (54) 8.58 (0.75) 126.0 (19.8) 102.7 (13.7) 35 60 119 239 21 (0.75)
(Papi1ionoideae) autogyro
18. Luehea seemannii Seed Autogyro 2.10 (0.45) 0.26 (0.02) 88.7 (10.3) 70.0 (7.0) 35 88 175 350 30 13 (0.50)
(Tiliaceae) '<S
19. Luehea speciosa Seed Autogyro 4.95 (0.50) 0.30 (0.04) 128.1 (13.0) 96.7 (10.9) 25 45 90 181 11 (0.50)
(Tiliaceae) -...J
v.>
TABLE 1. Continued ~
II)
'"1
o
Rank
Expected mean of ex- Rank
r-.....
dispersal
distance (m) pected of \0
(Wing- Rate of descent Tree disper- shade Wood 00
Type of Aerodynamic Mass (mg} Area (em') Ioadingj's ' (em sec") height 7 sal dis- toler- density 0\
~
diaspore group x (SO) x (SO) x (SO) .e(SO) (m) 1.75 3.5 m sec-w tance ance- (g cm- 3) d

20. Macrocnemum glabrescens Seed Rolling 0.0194 (0.0056) 0.0029 (0.0004) 80.6 (9.8) 51.5 (9.0) 25 85 170 340 29 0.60
(Rubiaceae) autogyro
21. Myroxylon balsamum Fruit Autogyro 749 (138) 14.2 (2.1) 227.1 (15.0) 154.5 (22.4) 35 40 79 159 9 1 0.78
(Papilionoideae) e>
22. Ochroma pyramidale Seed Floater 14.8 (3.8) 7.31 (2.76) 45.9 (6.5) 61.3 (7.2) 20 57 114 228 19 18 0.13 0~
(Born bacaceae) 'tI
23. Platymiscium pinnatum Fruit Rolling 462 (52) 20.0 (2.7) 151.0 (11.9) 128.4 (13.3) 35 48 95 191 14 0.83
c
it!
(Papilionoideae) autogyro 0
rn
24. Platypodium elegans Fruit Autogyro 1,640 (218) 23.5 (2.5) 261.7 (8.8) 178.2 (12.6) 35 34 69 137 4 3 0.75 it!
(Papilionoideae) I
25. Pseudobombax septenatum Seed Floater 86.9 (9.6) 15.7 (4.4) 75.3 (7.9) 125.7 (11.8) 25 35 70 139 5 11 0.14 ~
(Bombacaceae) Z
26. Pterocarpus rohrii Fruit Undulator 337 (106) 35.8 (5.9) 95.0 (10.4) 91.4 (14.3) 35 67 134 268 24 0.57 0I
(Papilionoideae) 0
27. Tabebuia guayacan Seed Nonclassified 35.7 (4.3) 4.50 (0.72) 88.8 (9.0) 109.9(15.7) 35 56 111 223 17 0.85 Vi
'tI
(Bignoniaceae) tTl
it!
28. Tabebuia rosea Seed Nonclassified 35.3 (4.7) 4.17 (0.28) 91.0 (5.0) 85.5 (8.6) 30 61 123 246 22 15 0.52 tr:
tTl
(Bignoniaceae) 0
29. Tachigalia versicolor Fruit Rolling 1,421 (163) 44.2 (4.1) 178.2 (8.7) 173.2 (33.7) 35 35 71 141 7 (0.52) ...,
(Caesalpinoideae) autogyro it!
0
30. Terminalia amazonica Fruits Rolling 4.07 (1.07) 1.01 (0.10) 62.8 (9.0) 73.2 (7.8) 35 84 167 335 28 9 0.68 'tI
(Combretaceae) autogyro n
31. Terminalia oblonga Fruits Rolling 54.3 (12.5) 5.33 (1.03) 99.6 (2.9) 88.6 (12.1) 35 69 138 276 25 12 (0.70)
>r-
(Combretaceae) autogyro 0
32. Trichospermum mexicanum Seed Floater 2.96 (0.30) 0.48 (0.05) 77.8 (5.3) 118.9 (8.0) 25 37 74 147 8 0.25 ;;
~
(Tiliaceae) 'tI
33. Triplaris cumingiana Fruits Helicopter 76.7 (13.1) 7.88 (0.95) 97.7 (7.2) 83.0 (11.6) 25 53 105 211 16 4 0.56 0
it!
(Polygonaceae) tTl
tr:
34. Vatairea erythrocarpa Fruit Autogyro 1,181 (233) 23.3 (2.5) 222.4 (19.6) 138.9 (14.1) 35 44 88 176 10 (0.64)
(Papilionoideae)
aWing-loading = wt/area; weight is expressed in terms ofmillidynes (mgcmsec") and area is in cm-.
b1.75 m sec:" = (4 mph); mean velocity over 24 h for March-April in laboratory clearing on Barro Colorado Island. 3.5 m sec" = (8 mph); mean velocity at noon for
March-April in laboratory clearing in Barro Colorado Island. 7 m sec-' = (15 mph); velocity used by Green (1980) to estimate wind dispersal of some temperate trees.
C From Augspurger (1984).
d Oven dry wt/green volume; values primarily from Chudnoff (1979); values in parentheses are not species-specific, but refer to genus level values.
e Seed mass includes ovary tissues.
(,;J
VI
-...I
358 AMERICAN JOURNAL OF BOT ANY [Vol. 73

TABLE 2. A description of the aerodynamic behavior of morphological groups of wind-dispersed diaspores. See Fig. 1
for drawings ofthe various groups

Aerodynamic group Aerodynamic behavior in still air

Floater Floats downward in a vertical line.

Rolling autogyro = Rotates on two axes simultaneously:
Flettner-Rotating Autogyro I) around diaspore's longitudinal axis;
(Vogel 1981) = 2) around one end of the diaspore in a semi-tight spiral.
"Rolling Samara" (Green 1980)
Autogyro = Rotates tightly around the seed end of the diaspore.
Lift-producing Airfoil (Vogel 1981) =
"Non-rolling Samara" (Green 1980)
Undulator Glides and undulates, but not with cumulative forward motion (not
continuous glider or fugoid oscillator).
Helicopter Spins tightly around a vertical line; similar to autogyro with added
wings.
Tumbler Tumbles but not on a consistent axis; randomly oriented tumbles;
also rotates around a vertical line, but in a very large spiral.
Nonclassified Complex and variable behavior within and between diaspores; closer
to autogyro and undulator than rolling autogyro.

Using a paired t-test it was determined that no data), it appears that mean wind speeds be-
significant differences occurred between the first tween I. 75 and 3.5 m sec:" are closest to the
and second releases of a given diaspore. mean speeds dispersing diaspores on Barro
Based upon observations of their aerody- Colorado Island during the dry season when
namic behavior in still air and their mor- the majority of these species disperse their
phology, each species was placed into one of seeds. Distances estimated using a mean wind
six categories (Table 2); the motion of species speed of 7 m- sec- 1 appear to more closely ap-
27 and 28 were too inconsistent to classify. proximate the observed maxima, not the mean
The expected mean dispersal distances were dispersal distance. Typical release height ofdi-
estimated using Cremer's (1977) formula: D = aspores was estimated for each species based
Vw(H/Vr) where D equals the horizontal dis- on general observations of these tree species
tance to which wind with a velocity of Vwwill over many years.
carry a diaspore falling at a mean velocity of
Vrfrom a height H; here Vwis the average wind RESULTS - Both mass and area, two variables
velocity between the release point and the expected to strongly affect the diaspore's rate
ground. Estimates were made using three dif- ofdescent, vary widely among the species; each
ferent wind velocities: I. 75 m- sec", the mean variable covers six orders of magnitude. Mass
velocity over 24 h in March-April in the lab- varies from 0.0194-3055 mg; area varies from
oratory clearing on Barro Colorado Island; 3.5 0.0029-155 em- (Table 1). Mass and area do
m .sec-I, the mean velocity at noon for March- not differ proportionately and, consequently,
April in the laboratory clearing on Barro Col- the species have different values for (wing-
orado Island, and 7 m- sec:", the velocity used loading)!"; they range from 36.7-261.7 (mil-
by Green (1980) to estimate dispersal distances lidynescm <)!" (Table 1). The rate of descent
by diaspores of some temperate trees. Based in still air ranges from 31. 6-19 8.1 em sec- 1
on observations ofdistances ofdiaspores from among the species (Table 1).
known parent trees (C. Augspurger, unpubl. A regression analysis of rate of descent on

TABLE 3. Regression analyses of rate of descent in still air on (wing-loading)'!' for five aerodynamic groups of wind-
dispersed diaspores. Each regression is significant (P < 0.00001). Significant differences exist among the slopes of
the five groups (F•.4U = 77.891, P < 0.00001;from Zar, 1978, pp. 230-233). Multiple comparisons of slopes are
indicated by underlinings, where separate lines indicate significant differences (P < 0.05)

Aerodynamic group Floaters Rolling autogyros Helicopters Undulators Autogyros

Sample size 97 157 45 45 90

r2 0.848 0.837 0.511 0.848 0.913
Intercept -22.61 -8.72 21.61 14.10 15.94
Slope 1.950 0.989 0.787 0.741 0.599
Multiple comparison
result
March, 1986] AUGSPURGER-WINO-DISPERSED TROPICAL DIASPORES 359

250
,
FLOATER
,,
5 '

o FLOATER
• ROLLING AUTOGYRO
o AUTOGYRO
• UNDULATOR
6. HELICOPTER
• TUMBLER
<> NONCLASSIFIED

28 56 84 112 140 168 196 224 252 280

(WI NG-LOADI NG)112
Fig. 2. The regression of rate of descent on (wing-loading)"? for individual diaspores of five aerodynamic groups
from Table 3. Dotted lines extend beyond the range of validity for each regression line for visual clarity only. The
symbols are plotted at the mean values for each species; see Table I for species' names.

(wing-loading)"? for all diaspores, irrespective area and thus increases the probability that
of species or aerodynamic group, is significant some of its diaspores land in light-gaps where
(rate of descent = 0.559 [wing-loading]"? + survival may be enhanced particularly for
44.62;N=479,r 2 = 0.503,P < 0.00001). The shade-intolerant species. Shade-tolerant species
coefficient of determination (r 2) increases are expected, in general, to have heavier seeds
greatly when regression analyses are done on than shade-intolerant species (Grime and Jef-
each of five aerodynamic groups separately fery, 1965; Ng, 1978). Earlier studies of 18 of
(Table 3). The slopes of the regression of rate these wind-dispersed species determined the
ofdescent on (wing-loading) 1/2 are not the same survival rate (slope of the regression of log
for the five groups (Table 3). At a given value number seedlings surviving on time) of seed-
for (wing-loading)!", diaspores of floaters fall lings under sun and shaded forest conditions
significantly more rapidly in still air than each (Augspurger, 1984). The survival rate in the
of the other four groups (Table 3, Fig. 2). In shade is used as the index to rank the shade
addition, the rolling autogyros fall more rap- tolerance of the species (Table 1). The species
idly than the non-rolling autogyros (Table 3; represent a continuum of shade tolerance and
Fig. 2). The sample sizes ofthe remaining two do not fit a dichotomy of shade-tolerant vs.
groups, tumblers and nonclassifieds, are in- shade-intolerant species. Wood density ofadult
sufficient for similar analysis. trees ofthese 18 species (Table 1) is positively
Rankings among the 34 species for their ex- correlated with their shade tolerance as seed-
pected mean dispersal distances are provided lings; however, shade tolerance and dry mass
in Table I. The expected mean dispersal dis- of seed are not significantly correlated (Aug-
tances vary nearly nine-fold among the 34 spurger, 1984).
species; using a windspeed of 1.75 m .sec' the Rate ofdescent shows a weak, but significant,
values range from 22-194 m (Table 1). rank correlation with the shade tolerance of
Analyses were performed to determine the 18 species (Table 4). The diaspores of the
whether rate ofdescent is correlated with shade floater group also show a significant rank cor-
tolerance of seedlings and dry mass of seeds. relation between rate of descent and shade tol-
A parent tree with diaspores that have a slow erance (Table 4). Rate of descent shows a sig-
rate ofdescent spreads its diaspores over a wide nificant correlation (r = 0.70, P < 0.00 I) with
360 AMERICAN JOURNAL OF BOTANY [Vol. 73

TABLE 4. Values of Kendall's Tau for rank correlations Given the extreme morphological diversity
of mean rate of descent with shade tolerance. Rank among the six groups, it is difficult to know
correlations were performed first for all species for
which data were available and second for the three what part of the diaspore to consistently mea-
aerodynamic groups with the most representatives sure for area to determine (wing-loading) 1/2.
How the cross-sectional area measured reflects
Shade tolerance the actual forces affecting air columns and aero-
n dynamics certainly varies among the diverse
All species 18 0.372 a groups. Further studies are needed to deter-
Floaters 5 0.800a mine the aerodynamics of the different groups
Rolling autogyros 5 00400 and to ascertain which aspects of morphology
Autogyros 5 0.333 best predict the rate of descent.
a = P < 0.05. The values for the wt/area ratio and the rate
of descent for diaspores of these 34 tropical
species do not deviate strongly from those of
the log of dry mass of seeds of the 34 species. earlier studies of temperate species. Despite a
The correlation is lower (r = 0.54, P < 0.001) range of six orders of magnitude for mass and
when values for dry mass rather than log dry area, the 34 species converge on a relatively
mass are used. This result suggests that the restricted range of values for (wing-loading)"?
relation between rate of descent and dry mass (37 to 262 millidynes-cm q-", The seven
r

of seeds is curvilinear; however, a description species of temperate trees of Green's (1980)

of that relationship is impossible because of study range from 70to 230 (millidynes-cm ")!",
the limited number of species with high mass. although their values for mass and area cover
a smaller range than those in this study. The
DISCUSSION - This study illustrates the wide rates of descent of the 34 species range from
array of morphological design found among 39 to 198 em-sec:". These values are within
species with wind-dispersed diaspores within the same order of magnitude of other studies
one tropical forest. It further documents the of trees (Green, 1980; Guries and Nordheim,
wide range in dispersal potential that occurs 1984). Slightly lower values occur in a few
among them. The most abundant morpholog- species of Compositae with a pappus (Sheldon
ical group represented is the rolling autogyro. and Burrows, 1973) and in some species of
Evolutionary convergence onto that morphol- Apocynaceae/Asclepidaceae with a parachute
ogy has occurred in six families. Evolutionary to lower their rate of descent (Platt and Weis,
divergence is illustrated by the Bombacaceae 1977). The maximum rate of descent reported
with five species of floaters and one very dif- among north temperate trees is 293 cm'sec l
ferent tumbler, and the Leguminosae repre- for Chamaecyparis thyoides (Siggins, 1933). All
sented by three autogyros, five rolling auto- 15 species of Eucalyptus studied by Cremer
gyros, and one undulator. One aerodynamic (1977) exceed the values in this study; they
group conspicuously missing from the trees on range from 210 to 554 em ·sec- I . None ofthose
Barro Colorado Island is the true glider capable species have any wings or dispersal appendages
of self-propelled forward momentum in still attached to the small seed.
air. Such a morphology occurs among the lia- On a world-wide basis the wind-dispersed
nas on the island. McCutcheon (1977) sug- diaspore with one of the most extreme values
gested that the overall rarity of gliders can be appears to be the huge autogyro of Centrolo-
explained because of their relative inferiority bium sp., a canopy tree growing in the Ama-
to autogyros and the rolling autogyros in tur- zonian lowland forests. Mean values for a sam-
bulent air. He predicted such unstable gliders ple from lowland Peru were: mass = 17.84 g,
would be found in the still air of the tropical area = 109.4 cm-, (wing-loading)"? = 407.1
rainforest. The three undulators of this study (millidynes-cm ")!", and rate of descent =
occasionally glide forward for a portion oftheir 367.6 em-sec>' (N = 14) (c. Augspurger, un-
descent in still air, but more commonly they pub. data). These values greatly exceed those
undulate back and forth in short arcs and ac- for any species in this study although the re-
complish little, if any, forward motion. Bur- lationship of rate of descent to wing-loading
rows (1975) pointed out that gliders ought to for Centrolobium sp. is within the predicted
undergo "fugoid oscillations" resulting in a range based on regression analysis of the 34
flight path in which the diaspore sweeps out species (Fig. 2).
elongated L-shaped arcs during its descent. The specific values for rate of descent re-
These arcs are necessarily undirectional with ported here are unlikely to be repeatable when
no back and forth motion. The three undula- trees in other locations are studied. Siggins
tors in this study may therefore represent a (1933) found considerable intraspecific vari-
new aerodynamic category. ation in rate ofdescent over a broad geographic
March, 1986] AUGSPURGER-WINO-DISPERSED TROPICAL DlASPORES 361

range of one species of Pinus. More locally on groups with shallow slopes. Hence the potential
Barro Colorado Island, significant variation in risk of losing dispersal distance while modi-
wt and area ofsamaras and their rate ofdescent fying wing-loading is greatest for floaters;
occurs among trees ofone species in this study, alternatively, with the same reduction in wing-
Platypodium elegans. Among 20 trees of this loading floaters can gain more dispersal dis-
species the mean values for (wing-loading)!" tance than other groups, although they still fall
range from 244 to 319 (millidynes- cm- 2) 112 and faster and therefore disperse less far than other
mean values for rate ofdescent range from 178 groups.
to 236 em- sec:" (c. Augspurger, unpubl. data). Any increase in seed wt or in the size of
Therefore, species' values reported here pro- dispersal appendages that alters rate of descent
vide only relative data for making very general represents a cost to the parent. Some morpho-
comparisons among species. logical designs are undoubtedly more costly to
Knowledge of mass and area of diaspores build or alter than others. For example, pro-
allows predictions of their rate of descent. The duction ofmore fluffy kapok ofthe floaters may
correlation between (wing-loading)"? and rate require more or fewer resources than increasing
of descent is significant in this study when all the woody wing of an auto-rotating samara,
individual diaspores ofall aerodynamic groups depending on the wing's photosynthetic con-
are combined. The correlation improves mark- tribution. In addition, for two diaspores with
edly when the analysis is restricted to any given comparable wing-loadings, but different areas,
aerodynamic group, indicating that the specific a similar decrease in wing-loading achieved by
relationship varies among aerodynamic groups. increasing area is more costly on an absolute
Except for the helicopter group (r? = 0.51), the basis for the diaspore with the larger initial
coefficient of determination is high for each area.
group (r? ranges from 0.84 to 0.91). The im- The narrow range in wing-loading values
portance of identifying these strong relation- among the 34 species, relative to the wide ranges
ships lies principally in their predictive power. in wt and area, indicates a constraint on se-
Rate of descent in still air can be accurately lection acting on the relationship between wt
predicted without experimentation for other and area. Selection for increased seed wt ap-
species, provided their wt, area, and mor- parently has been accompanied by selection
phology are known. for increased area of dispersal appendages in
The regression analyses of rate of descent on order to diminish any loss of dispersal poten-
(wing-loading)"? indicate strong differences tial. If a plant is resource limited, selection
among morphological groups in the slope de- acting to change dispersal potential is inextri-
scribing that relationship (Fig. 2). The floaters cably linked to the number of diaspores that
have the steepest slope (1.95). This group is can be made. No data are available for inter-
dominated by species of Bombacaceae; their specific comparisons in the size of fruit crops
diaspore consists of a seed surrounded by a of the study species. Species with larger dia-
sphere of silky fibers (kapok) that provides a spore areas and/or heavier seeds are predicted
drag, but no lift, component to its aerodynamic to have smaller crop sizes.
motion (M. Denny, pers. comm.). All other Knowledge of rate of descent, ht of release,
groups have both lift and drag components. and mean windspeed allows predictions of ex-
The slope for the autogyros (0.60) differs sig- pected mean dispersal distance for each species.
nificantly from that of the rolling autogyros Despite the narrow range among species in
(0.99). The rolling autogyros fall faster than wing-loading, relative to their range in wt and
autogyros, but at progressively smaller amounts in area, the species still have a nine-fold range
as the (wing-loading)"? declines. These results in predicted mean dispersal distance; they range
differ from those found for temperate trees from 22 to 194 m at a mean windspeed of 1.75
(Green, 1980). He found that rolling autogyros m- sec-I. Actual dispersal distance achieved in
and autogyros have almost identical slopes the field will be affected by a diversity of fac-
(0.52 and 0.53, respectively), but different in- tors, e.g., windspeed required for abscission,
tercepts, such that at a given (wing-loading)!", turbulence and variability in windspeed during
a rolling autogyro falls faster than an auto- descent, and the surrounding vegetation. The
gyro. expected mean distances are not likely to be
Similar evolutionary shifts in wing-loading accurate predictions. However, such values are
among morphological groups should have quite useful in providing a general idea of how the
different consequences for dispersal. Any com- various species rank in their dispersal potential
parable change in wing-loading resulting from and in illustrating the tremendous range in dis-
altering seed weight or diaspore area will have persal potential that exists within one tropical
a greater effect on rate of descent and dispersal forest. Ultimately values for rate of descent
distance for groups with steeper slopes than for can be incorporated into a model that predicts
362 AMERICAN JOURNAL OF BOT ANY [Vol. 73

the spatial distribution ofwind-dispersed seeds, study species to test the validity of the predic-
such as that developed by Fields and Sharpe tions from this study.
(1980). Before such a model can be applied to The dispersal potential of a species has im-
these tropical trees, additional measurements portant implications for other aspects of its
will be required of the variability in rate of biology, e.g., size of fruit crop, traits affecting
descent within a species (and within individual seedling survival and establishment, the spatial
trees) and ofthe wind environment of tropical relationship of juveniles and adults, and the
forests. genetic structure ofthe population due to gene
The predicted mean dispersal distance rep- flow via seed dispersal. Given the differences
resents a one-dimensional value, but seeds are among the species in their dispersal potential,
dispersed over a two-dimensional area. Be- they are expected to also differ widely in ad-
cause the area ofa circle increases by the square ditional aspects affected by their dispersal bi-
of its radius, any difference among species in ology.
mean distance will result in even greater dif- Species with a more restricted dispersal dis-
ferences among species in their area over which tance and area must have other life history
seeds are dispersed. Selection on morpholog- characters that allow seedling establishment
ical traits that results in only a small increase nearer the parent. Factors affecting such es-
in mean dispersal distance should have a sub- tablishment include shade tolerance and resis-
stantially greater effect on the area ofa parent's tance to mortality factors that act in a density-
seed distribution. This study places emphasis and/or distance-dependent manner, e.g., seed/
on mean dispersal distances. Anecdotal ob- seedling pathogens, seed predators, seedling
servations suggest extreme values in distance herbivores, and seedling competitors. This
are sometimes achieved by wind-dispersed di- study shows that rate of descent correlates
aspores (Whitmore, 1984; C. Augspurger, pers. weakly with two traits affecting seedling estab-
obs.). What is unknown is the relative impor- lishment, viz. log ofdry mass ofseed and shade
tance of mean versus extreme values of dis- tolerance of seedlings. It appears that species
persal distance for successful seedling estab- requiring light-gaps as seedlings have lighter
lishment. seed mass and/or larger area of their diaspores
Given these background data, the next step so as to lower their rate of descent. Airborne
is to determine whether these rates of descent for a longer time, they potentially disperse over
and expected dispersal distances are realized a broader area and increase the chance that
under field conditions. Previous studies con- some of parent's diaspores land in light-gaps.
firm the general predictability of dispersal dis- Dispersal potential also has implications for
tance under field conditions from lab results the genetic structure and speed of movement
of rate of descent in still air. A field study of ofnew alleles through a population. With great-
Lonchocarpus pentaphyllus, one of the species er dispersal potential, offspring of one parent
in the present study, demonstrated that fruits are more likely to encounter those of other
with 0, 1, 2, and 3 seeds have increasing values parents. This overlapping of seed distributions
for (wing-loading) 1/2 and rate of descent in still enhances the probability of outbreeding and
air. Actual dispersal distance declined among increases the movement of alleles in the pop-
the four fruit types with an increase in rate of ulation.
descent (Augspurger and Hogan, 1983). For In summary, considerable morphological
herbs Platt and Weis (1977) found a negative variation exists among the tropical tree species
correlation between rate of descent in still air with wind-dispersed diaspores on Barro Col-
and dispersal distance under field conditions. orado Island, Panama. Knowledge of their
In contrast, Rabinowitz and Rapp (1981) found mass, area, and wing-loading values allows ac-
no such relationship for grass species and curate predictions oftheir rates ofdescent, par-
stressed the importance of specific morpho- ticularly within a given morphological group.
logical features in affecting dispersal. For seeds The species differ greatly in their dispersal po-
of Melaleuca trees, Woodall (1982) found the tential. These differences are likely to have im-
dispersal distance he predicted to be shorter portant implications for their fruit crop size,
than actual distance by a factor of two. By population demography and spatial patterns of
adding a correction factor for wind turbulence, surviving seedlings, and the overall genetic
he achieved a reasonable prediction of actual structure ofthe population. The study indicates
distance based on knowledge of rate ofdescent the danger inherent in assuming that knowl-
and wind conditions. Studies on Barro Colo- edge of the general dispersal mode ofa species,
rado Island are currently underway to docu- e.g., wind dispersal, is sufficient to predict other
ment the seed distributions of many of the aspects of its biology. Generalizations about
March, 1986] AUGSPURGER - WIND-DISPERSED TROPICAL DIASPORES 363

species that use wind dispersal in tropical for- acteristics and dispersal potential of maple samaras.
ests are unlikely to be valid. Forest Sci. 30: 434-440.
MCCUTCHEN, C. W. 1977. The spinning rotation of ash
and tulip tree samaras. Science 197: 691-692.
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