New Ptiyiol.

(198S), 108, 357 36S

Mass allocation, moisture content, and

dispersal capacity of wind-dispersed
tropical diaspores

BY C. K. A U G S P U R G E R

Department of Plant Biology, Universitv of Illinois, 505 S. Goodwin, Urbana, IL 61801,

U.S.A.
{Received 22 December 1986; accepted 30 October 1987)

SUMMARY
Among 34 wind-dispersed tree species on Harro Colorado Island, Panama, the wet mass of diaspores ranges over
six orders of magnitude. The seed mass as a percentage of diaspore mass iS/D) varies greatly among species from
14 to 94",, with a mean value of 61 "o. The mean percent moisture of diaspores is 10 "„ ; no consistent differences
occur between seed and non-seed components of diaspores in percent moisture. Wing-loading (weight/area) and
hence dispersal capacity varies over one order of magnitude among these species and is correlated more highly with
S/D than with percent moisture of diaspores. Compared to fruit diaspores, seed diaspores have less mass and a
greater S/D, and are slightly more dehydrated. As a result, seed diaspores have lower wing-loading and greater
dispersal capacity than fruit diaspores. The morphological/aerodynamic feactures of a diaspore also have a
significant eflect on the mass variables. Overall, the type of diaspore (fruit or seed) is of major importance in
explaining mass differences of diaspores among these wind-dispersed species; morphological features and
dehydration ability contribute secondarily.

Key words: Dispersal capacity, fruits vs. seeds, mass, moisture content, wind-dispersed diaspores

I N T R O i:> II C T I O N
A diaspore is the dispersal unit that is moved from
tbe parent plant and establishes the offspring as an
independent seedling. For wind-dispersed species,
added diaspore mass increases wing-loading
(weight/area) and thus causes a decrease in dispersal
capacity. Natural selection of species with wind-
dispersed diaspores might be expected to minimize
any components of mass tbat result in loss of
dispersal and/or seedling establishment. However,
minimization of mass may be constrained evolu-
tionarily by the type and amount of tissues com-
prising the diaspore.
Seed components necessary for seedling establish-
ment, i.e. embryo and cotyledons (or endosperm),
are the first determinants of diaspore mass. Two
additional factors affect the total mass of a wind-
dispersed diaspore: (1) tissue derived from ovary
and/or seed coat (hereafter termed non-seed com-
ponents) and (2) moisture content of all tissues.
The amount of diaspore mass from non-seed
components depends on two factors. First, the
diaspore may be a fruit or a seed. For seed diaspores
the added non-seed mass accrues only from enlarge-
ment of the seed coat as dispersal appendages. For
fruit diaspores it accrues from the seed coat and from
ovarian tissues that enlarge, so increasing appendage
area for photosynthesis (Bazzaz, Carlson & Harper,
1979) and later for dispersal, and that surround the
seed to protect it from premature dehydration,
pathogens and predators (Janzen, 1971). A fruit
diaspore also has added mass if it is multi-seeded; a
seed diaspore, by definition, is always single-seeded.
Thus, relative to seed diaspores, fruit diaspores, on
average, are expected to have a greater total mass,
but a lower percent of total diaspore mass devoted to
those seed components necessary for seedling estab-
lishment.
Second, the morphological/aerodynamic features
of a diaspore will affect the amount of added mass
due to appendages (Fig. 1). Species vary in their
degree of morphological elaboration, e.g. area and
number oi wings, and nattire of the construction
material, e.g. woody, membranous, or hair-hke
fibres.
Diaspore mass also depends on the amount of
tissue dehydration that occurs before dispersal. It
would be expected tbat wind-dispersed diaspores
would lose moisture, so lowering mass during flight.
358 C. K. Augspurger
Aerodynamic grt)up Representative morphology
Fi.t)ATER ,^'' '^.^^
ROLLING AirrocYRt)
AUTOGYRO
UNDUI.ATOR
HnLICOPTER
TUMBI.ER
UNCLASSIFIED ^I ^>
Figure 1. A description of the aerodynamic behaviour of seven groups of wind-dispersed diaspores and
representative drawings of their morphology (reprinted with permission from The American Journal of Botany).
See Augspurger (1986a) for drawings of all 34 species.
In general, mature seeds bave considerably lower
water content (5-20%) compared to most plant
tissues in an active state (80-95 "/i,) (Street & Opik,
1975). Tbe decline in moisture content during seed
developinent is due primarily to an increase in dry
mass of nutrient reserves. Only in tbe final days of
maturatit)n does a simple drying process occur
(Pollock & Rt)os, 1972). In general, tbe moisture
content is maintained at low levels by impermeable
seed coats (Mayer & Poljakt)fT-Mayber, 1975). In
legumes tbere is a bygroscopic one-way valve of the
hilum that opens only wben the seed is surrounded
by even drier air (Hyde, 1954).
I)ebydratit)n causes the metabolic rate of seeds to
be extremely low and greatly increases seed long-
evity; in general, eacb 1 "„ decrease in seed moisture
doubles tbe longevity t)f crt>p seeds during storage
Aerodynamic behaviour in still air
Floats dt)wnward in a vertical line.
Rotates t)n two axes simultanet)usly ;
(1) arout-id diaspt)re's lot-igitudinal axis;
(2) around one end t)f the diaspt)rc in a
semi-tight circle.
Rt)tates tightly around the seed end t^f
the diaspore.
Glides and undulates, but not with
cumulative forward motion (not
continuous glider).
Spins tightly around a vertical line;
similar to autogyro with added wings.
Tumbles hut not on a ct)nsistent axis;
randomly oriented tumbles; also rotates
around a vertical line, but in a very
large circle.
Ct)n-iplex and variable behavit)ur within
and between diaspores; closer to
autogyro and undulator than rolling
autogyro.
(Harringtt)n, 1973). At values above 10-13% moist-
ure, seed viability decreases due to damage from
insect reproduction, fungal invasion, or heating from
microbial respiration (Cbristensen & Kaufmann,
1969; Harrington, 1972, 1973). Tbere are lower
limits to seed debydratitjn ; at values less tban 5 %
mt)isture, deterioratit)n occurs, probably due tt) lipid
autoxidation (Harrington, 1973).
Given tbis background, two questit)ns about
diaspore moisture arise. Do non-seed ct)mponents of
wind-dispersed diaspores have an equivalant cap-
acity ft>r dehydration as seed components ? Do fruit
diaspores, due to their thicker ovarian-derived
tissues, bave prt)portionately a greater mt)isture
content tban seed diaspores.'
Few data exist on tbe ct)mparison of different mass
components tjf wind-dispersed diaspores (but see
 Mass and moisture of diaspores
Sheldon & Burrows, 1973). The present study of 34
wind-dispersed tree species on Barro Colorado
Island, Panama, was designed to measure the wet
and dry mass of diaspores and their component parts
(seed and non-seed). Those data were used to
compute ; (1) seed mass as a percent of diaspore mass
(hereafter referred to as S/D) and (2) percent
moisture of diaspores and their component parts
(seed and non-seed). The goal was to determine
whether these mass variables vary significantly
between fruit and seed diaspores, betweet-i morpho-
logical categories, ar-id between seed and non-seed
compotients of diaspores. Tbe species vary widely
in wing-loading^ (range = 3-7-26-2 m Pa), rate of
descent in still air (range = 32-198 cm s~'), esti-
mated mean dispersal distance (range = 44—388 m at
mean wind velocity of 3-5 m s ' ) (Augspurger,
1986 a), and in shade tolerance of seedlings (Augs-
purger, 1984). Using published values, I determined
how mass factors correlate with those affecting
diaspore dispersal and seedling establishment. The
overall objective was to identify factors related to
mass that explain, in part, the wide disparity in
dispersal capacity among wind-dispersed species.
METHODS
Field collections of diaspores of the 34 wind-
dispersed tree species were made from 1981-4 on
Barro Colorado Island, Panama. Descriptions and
nomenclature of the species occur in Croat (1978) and
drawings of the diaspores in Augspurger (1986 a).
The species differ in whether the diaspore is a fruit
(« = 15) or a seed (n = 19) (Appendix 1), and in
morphological and aerodynamic properties (Fig. 1).
All are generally one-seeded diaspores.
During the dispersal period 15 diaspores of one
tree of each species were collected from the ground
immediately beyond the canopy of the tree. Excep-
tions occurred for very small seed diaspores that
were collected directly from dehisced fruits of one
tree. Small sample sizes arose because of the
difficulty of locating viable seeds of many of the
species which are rare, reproduce infrequently, and
produce low crop sizes of filled, undamaged seeds.
Diaspores with a mass greater than 100 mg were
measured on a Mettler top-loading balance, tbose
less than 100 mg on a Cahn electrobalance. Wet (i.e.
fresh) mass was determined immediately after col-
lection. Values for wet mass were not substantially
modified by weatber because collections were made
in the extended dry season with low humidity.
Diaspores were dried to constant mass at 105 °C and
remeasured to obtain values of dry mass. Values for
percent moisture.
dry mass\
wet mass/
359
were correct within 1-2 "„, because they were not
determined at standardized relative humidity and
temperature (Harrington, 1972).
To obtain values for seed mass as a percent of
diaspore mass (S/D)
/ seed wet mass
xlOO ,
ydiaspore wet mass
required definitions that separated those parts of
the diaspore necessary for seedling establisbment
(i.e. 'seed') from all other components; these
definitions for seed and fruit diaspores differed as
follows. The seed component of a seed diaspore
excluded the part of the seed coat modified into
appendages, i.e. wings, membranes, hairs, or fibres,
while the seed component of a fruit diaspore included
the entire seed coat. Exceptions for fruit diaspores
occurred for five species with ovary and seed coat
tissues that were fused and inseparable. For these
species, {Aslronium graveolens, Cordia alliodora,
Terminalia atnazoniea, Terminalia oblonga, and Trip-
laris cumingiana), ovary tissues directly surrounding
the seed were included as part of the seed component,
thus leading to overestimates of seed mass. A seed
diaspore thus included seed (as above) plus append-
ages; a fruit diaspore included seed (as above) plus
appendages (all ovary-derived tissue dispersed with
tbe seed).
In total 11 variables were computed for each
species (see Appendices 1 and 2). All variables,
includmg S/D and percent moisture, were deter-
mined first for individual diaspores prior to statistical
analyses. Statistics for percentages were done on
arcsine-transformed data. After mean values were
determined for each species, species were ranked for
each variable before any non-parametric rank corre-
lations were made.
RESULTS
Seed mass us. diaspore mass
For the 34 species, the mean wet mass of the diaspore
at the time of dispersal is 394 mg; values range over
six orders of magnitude. The mean wet mass of the
seed component of the diaspore is 134mg; values
range over five orders of magnitude (Fig. 2, Appen-
dix 1). Wet and dry mass measurements in Appendix
1 are highly correlated with one another (Kendall's
T = 0-986, n = 34, P < 0-001).
Values for S/D vary widely among species from
14-2% for Cavanillesia platanifolia to 93-7 "i, for
Tabebuia rosea\ the mean value for all species is
60-6% (Appendix 2).
Relative to fruit diaspores, seed diaspores have 1)
lower wet and dry mass and 2) greater values tor S/
D (Table 1). Seed diaspores also bave lower wing-
loading, lower rate of descent in still air, and greater
estimated dispersal distance than fruit diaspores
360 C. K. Augspurger
10
Wet mass of seed (g)
Figure 2. Mean wet mass of seed (without dispersal
appendages or tivarian-derived tissues) as compared with
the mean wet mass of the diaspore (entire dispersal unit)
for 34 tree species dispersed by vvit-id on Barrt) Ct)lorad()
Island, Panama. Note the log-log scale. The I : 1 line is
added for visual interpretation. Some diaspores are fruits
(O), other seeds ( • ) . For species' identification, see
Appendix 1.
(Table 1). Wing-loading increases more as wet mass
increases for fruit diaspores (T = 0-543, n = 15, P <
0-005) tban for seed diaspores (T = 0-240, n= 19,
P = 0-152). Tbis result suggests tbat fruit diaspores
do not bave sufficient area to balance tbe added mass
of ovarian-derived tissues. As a result, tbeir wing-
loading increases as diaspore mass increases. Tbe
above ct)mparist)ns bt)ld for all species witbout
regard to aerodynamic category and alst) within an
aert)dynamic category (e.g. rt)lling autogyrt)s t)r
autogyros).
Tbe 34 species represent seven morpbological/
aerodynamic categories (Fig. 1). Wben tbese cat-
egories are compared, witbout regard to fruit vs. seed
diaspores, significant differences exist among cat-
egories for eacb tjf tbe 11 mass variables (Table 2).
Particularly striking is tbe one tumbler, Cavanillesia
platanifolia\ it is tbe beaviest diaspore, bas tbe lt)west
S/D, and bas a very bigb percent moisture (see
belt)w). Its large fruit contains a gelatinous hygro-
scopic core (Garwt)t)d, 1985) and a very large
embryo witb relatively moist leafy ct)tyledotis. Tbe
second heaviest group, the auttjgyrt)s, includes three
species with particularly large legume fruits witb
heavy protective ovarian walls surrounding the seed.
Tbe two Tabebuia species (unclassified) and tbe
floaters (largely species in tbe Hombacaceae) have tbe
bigbest S/D. Both grtiups have only seed diaspores;
their appendages of thin membranes and fibres,
respectively, add little tt) ttjtal diaspore mass. Htjw-
ever, tbe fibres prt)vide tbe diaspore w itb only a drag,
and no lift, component to lower its rate of descent
(Augspurger, 1986o).
When botb tbe diaspore type (fruit or seed) and
the mt)rpholt)gical category are included in a 2-w-ay
analysis of variance, two generalities emerge. First,
for all 1 1 mass variables in Appendices 1 and 2, tbere
are significant effects due to diaspore type, morpho-
logical category, and their interaction. Sect)nd, ft)r all
mass variables, except percent moisture of diaspores
and non-seed ct)mptjnents, tbe diaspore type bas a
larger F-value tban eitber mt)rpht)logical categt)ry or
the interactit)n term. The /^-values are particularly
large for the wet and dry variables t>f S/D.
Wet mass directly determines the diasporc's wing-
loading value that, in turn, effects its dispersal
capacity. Wing-lt)ading t)f the 34 species is positively
correlated witb their rate t)f descent in still air (T =
0-604, P < 0-001) and negatively ct)rrelated with
their estimated mean dispersal distance (T = 0-441,
P < 0-001) (values ft)r the 34 species taken from
Augspurger, 1986a). Wing-loading is negatively
correlated with S/D (T = 0-358, n = 34, P < 0-003).
Species with high S/D values thus have lower
dispersal capacity as their diaspore makeup has a
large emphasis on the seed ct)mponent and less
emphasis t)n nt)n-seed ct)mponcnts, including dis-
persal appendages.
Previous work on tbese species demonstrated that
tbe more sbade-tolerant species tend to bave a faster
rate of descent and therefbre are mt)re likely tt) be
dispersed in shady areas closer to the parent tree
(Augspurger, 1984). It would be expected tbat these
mt)re shadc-tt)lerant species wt)uld invest more in
seed mass than in dispersal appendages and have
bigber S/D values, relative to more sbade-intolerant
species. Ht)wcvcr, fbr a subset t)f 18 species, sbade
tolerance t)f seedlings is not significantly ct)rrelated
witb S/D (7- = - 0 - 2 0 3 , « = 1 8 , P>(}-10; values
taken from Augspurger, 1984). Tbis lack of correla-
tion may reflect conlbunding effects due tt) variation
amt)ng families in amount of t)varian tissue (see
below) and tbe compositional differences of seed
versus fruit diaspores. Wbilc seed diaspores bave
greater values for S/D tban do fruit diaspores (Table
1), shade-tolerance values for seed diaspores (/( = 9)
do not differ from those for fruit diaspt)res (n = 9)
Mann Whitney U, U = 56, n.s.).
Percent moisture
For the 34 species, the mean percent mt)isture is
10% for tbe diaspore, as well as for seed and non-
seed components separately (Appendix 2). No con-
sistent differences occur between seed and non-seed
components in percent moisture (letter Z in Appen-
dix 2). l-'ercent mt)isturc is not significantly ctirre-
lated witb wet mass of the diaspore (T = 0-196,
n = 34, P> 0-10). Relative to fruit diaspores, seed
 Mass and moistt(re of diaspc 361

T a b l e 1. Cotnpartson of mass variables a?id dispersal capacitv betzveen 15

zvind-dispersed species zvith fruit diaspores and 19 species zvith seed
diaspores on Barro Colorado Island, Panama

Seed diaspore Eruit diaspore

(n = 273) (n= 199) t(P)
Wet mass (mg)
Diaspore 87-48 (11-02) 815-33 (74-86) 9-62***
Seed 62-00 (8-21) 232-80 (18-61) 8-39***
Non-seed 25-48 (3-29) 582-53 (60-03) 9-27***
Dry mass (mg)
Diaspore 80-45 (10-35) 680-20 (60-61) 9-76***
Seed 57-60 (7-83) 186-24 (14-82) 7-68***
Non-seed 22-85 (2-92) 486-39 (49-13) 9.42***
S/D(" )t
Wet 73-71 (0-91) 42-53 (1-34) 19-62***
Dry 73-98 (0-91) 42-59 (1-38) 19.49***
Moisture ("n)J
Diaspore 9-08 (0-26) 12-37 (0-45) 6-97***
Seed 8-72 (0-28) 13-02 (0-67) 6-15***
Non-seed 9-78 (0-64) 12-03 (0-60) 4-05***
(Wing-loading)^ (mPa)§ 8-2 (0-66) 14-6 (1-43) 4-07***
Rate of descent (cm s ' ) | | 89-95 (8-70) 126-11 (8-33) 2.94**
Estimated dispersal 143-5 (18-8) 99-3 (7-8) 2-17*
distance (in)l

Values represent mean (SE). Statistics for variables that are percentages are
done on arcsin-transformed data.
* P < 0-05 ; ** P < 0-01 ; *** P < 0-001.
f S/D = values for seed mass as a percent of diaspore mass, i.e.
/ Seed n-iass ,
\Diaspore mas? X 100 .

X Values for percent moisture = 1 —:—'- |100

Wet massj
§ Wing-loading = (mass X acceleration)/projected area; values fron-i .Aiigs-
spurgcr (1986(3).
II Based on rate of descent through 27 m of still air; values from AXugspurger
(I986«).
1 Based on average tree height, rate of descent in still air, and a mean velocity
of 3-5 m s ' (8 mph); values from Augspurger (1986rt).

diaspores have slightly lower percent moisture
( T a b l e 1).
T b e morphological categories difl'er slightly in
percent moisture of botb seed and non-seed c o m -
p o n e n t s (Table 2). T h i s result suggests tbat con-
struction materials in the various structures difier in
tbeir abilities to be dehydrated before dispersal.
Highest moisture content occurs in the tumbler,
Cavanillesia platanifolia (sec above) and the heli-
copter category with species that exhibit enlarged
calyces as dispersal appendages and bave fused
ovarian and seed tissues. Such diverse morphological
categories as floaters, rolling autogyros, and auto-
gyros do not differ in their moisture content.
D I S C U .S S I O N
Seed mass vs. diaspttre tnass
The mass of diaspores of wind-dispersed species
accrues from both seed and non-seed components.
An increase in seed n-iass generally promotes seedling
survival (Perry, 1976; Howe & Richter, 1982), but
diminishes dispersal capacity, unless accompanied
by an increase in area (and hence mass) of append-
ages to maintain wing-loading area (.'Kugspurger &
Franson, 1987). Among tbe tropical species in tbe
current study, a wide and 'continuous range in
cotyledon reserves and shade tolerance exists (.Augs-
purger, 1984). l^beir seed mass ranges widely over
five orders of magnitude. This extremely wide range
of mass dispels the notion that wind-dispersed
species necessarily are small-seeded, shade-intol-
erant species. Probably tbe most extreme mass for
any wind-dispersed species is 17-8 g for the diaspore
of Centrolt)biuni from lowland Amazonia (Augs-
purger, 1986fl); it is fi\-e times heavier than the
heaviest species in the current study. However, seed
mass is generally less for wind-dispersed than
animal-dispersed species. In the same forest in
Panama the seed mass of a small percentage of
animal-dispersed species exceeds that of tbese wind-
dispersed species (Foster, 1982).
The diaspore mass may represent a compromise
between the importance of seed mass for seedling
362 C. K. Augspurger

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 Mass and moisture of diaspores
establishment, and appendages and other mass
necessary for photosynthesis, dispersal, and seed
protection. Mass from any source increases wing-
loading at-id lowers diaspore dispersal capacity.
Among the species, the values for S/D range widely
frotn 14 to 94",,, indicating how different the speeies
are in their relative balanee of diaspore mass used for
seedling establishment versus dispersal and/or seed
protection, l^hese wide differences in S/D values do
not correspond with speeies' difFerenees in shade
tolerance of seedlings and indieate the complexity of
factors that determine diaspore mass.
A primary faetor in accounting for the differences
in balanee between seed and non-seed components is
whether the diaspore is a seed or a fruit. The n-iean
S/D is 74 "o for seed diaspores compared to 43 ",, for
fruit diaspores. Both diaspore types have dispersal
appet-idages; fruit diaspores have additional n-iass in
ovarian tissues surrounding the seed. Ovary tissues
surrounding seeds are a particularly large eon-iponent
of diaspore mass in legume fruits; the mean S/D is
only 33",, for legume fruits in contrast to 54",, for
non-legume fruits. For fruits of 24 herbaceous
species of Compositae, the S/D ranges from 39 to
85 "o (Fenner, 1983); these values more closely
overlap those for seed diaspores in the current study.
Thus the distinction drawn in this study between
fruit and seed diaspores in their mass allocation to
seed and non-seed components depends in part on
tbe family of the species.
Use of the fruit or the seed as tbe diaspore is
usually a family trait and thus has a long evolutionary
history that is only partially related to dispersal per
se. Pollination, abortion, and seed predation all
affect the evolution of packaging of seeds into
dispersal units (Casper & Wiens, 1981, Willson,
1983). In this study all species with multi-seeded
fruits are dehiscent and have seed diaspores; those
witb single-seeded fruits are indehiscent and have
fruit diaspores, although some of the legume species
produce a small proportioti of multi-seeded fruit
diaspores (Augspurger & Hogan, 1983 ; Augspurger,
1986/)),
A secondary faetor explaining mass diflerences
among these species is their n-iorpbology at-id, by
inference, the type and amount of construction
material used for non-seed compot-ients. ll-iick,
woody wings add n-iore mass per unit area than
membranes or fibres. Only seed diaspores have the
thin membranes or fibres of lower det-isity; fruit
diaspores often have thick, somewhat woody
wings.
Son-ie morphological categories include both truit
and seed diaspores and species fron-i a diversity of
families. Evolution can affect the mass of a given
construction material, but it is probably more
constrained by developmental pathways from switch-
ing to a new construction material ancl converting
from one morphology to a grossly different morph-
363
ology. Among morphological categories the relation
between wing-loading and rate of deseent varies
significantly (Augspurger, 1986a). Categories with
only aerodynamic drag, such as floating diaspores
with fibres, fall faster, at a given wing-loading, that-|
do those witb aerodynamic lift and drag, such as
spinning autogyros, helicopters, and rolling auto-
gyros.
Percent moisture
A third factor affecting mass is dehydration of
diaspore tissues before dispersal, l^he 34 species
have a mean of 10",, moisture in their wit-id-
dispersed diaspores. Because tbe species differ
significantly in tbeir diaspore moisture content, this
is another factor that explains, in part, why wind-
dispersed species vary in tbeir wing-loading and
dispersal capacity.
Non-seed components of wind-dispersed dia-
spores are dehydrated, on average, to the same extent
as seed components ; botb a\ erage 10-11 "„ moisture
at tbe time of dispersal. In contrast, non-seed
components of wind-dispersed diaspores are mark-
edly drier than the flesby, moist rewards of animal-
dispersed diaspores that have moisture contents
from 42 to 99",, (Wheelwright et al., 1984; Johnson
et al., 1985; White & Stiles, 1985), This dehydration
lowers considerably tbe total diaspore mass, par-
ticularly for those species with a large value for seed
mass as a per cent of diaspore mass (e.g. up to
86 "o ii-i Cavanillesia platanifolia).
Fruit diaspores have a slightly higher percent
moisture than seed diaspores, although tbis differ-
ence is exaggerated by two outlier species, both
legumes with a high percent moisture in their seeds.
Overall, fruit diaspores have greater mass than seed
diaspores because of their extra ovary tissues and/or
because of the use of tissues of higher density, rather
than because of a higher moisture content. Wing-
loading values of species are significantly (negatively)
correlated witb the seed mass as a percent of diaspore
mass and not witb percent moisture.
The moisture content of the seed component of
these diaspores averages 10",,; this value is withiti
the publisbed range of 5-18% for crop species
(Harrington, 1972, 1973). The majority of species
range from 6 to 14",, moisture, although extreme
species range from 4—35 %. Only values for Myroxy-
ton balsamum (23 ",,) and Platypodium elegans (35 ",,)
are above the level at which viability of temperate
crop species rapidly declines (Harrington, 1973).
How seed moisture affects viability of tropical species
is generally unknown. Platypodium elegans germ-
inates within 1-2 months after dispersal; any delay
in germination, by artiiicial storage, lowers its vi-
ability considerably (Augspurger, unpublished data).
Comparable data of seed moisture for non-crop
species are unavailable. For domesticated species.
364 C. K. Augspurger
seed moisture depends particularly on tbe cbemical
compt)sition of tbe seed; moisture is higher in seeds
with bigb starcb t)r protein tban in t)ily seeds witb
hydropbobic lipids (Harrington, 1972). Wbetber a
seed is in a flesby t)r dry fruit apparently dt)es nt)t
affect dehydration; tomatt) seeds mature in an
aqueous environment, but tbey undergo tbe same
changes in seed moisture as seeds tbat mature in air
on otber species (McIIratb, Abrt)l & Heiligman,
1963).
Given tbe limited ct)mparative data base, it
appears that seeds of wind-dispersed diaspores have
moisture contents within tbe publisbed range of
seeds in general. Tbeir mass is not minimized by
extremely low levels of moisture. Non-seed and
components sbow equivalent capacities for debydra-
tit)n; nt)n-seed ct)mponents sbow extreme levels of
dehydration, relative to their animal-dispersed
counterparts.
A t - K N O W L I-: D G E M E N T S
This research was supported hy NSF grant BSR 8219856
and made possible with support from the Smithsonian
'l'ropical Research Institute. I thank Colleen Kelly and
Sue Franst:)n for assistance with weight measurements.
Sue Franson ft)r statistical analyses, and Carol Kelly,
Kevin Hogan, Flizabeth Lacey, and Mike Melainpy for
contributing constructive comments on earlier drafts of
the manuscript.
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