Seed Predation, Disease and Germination on Landslides in Neotropical Lower
Montane Wet Forest
Randall W. Myster
Journal of Vegetation Science, Vol. 8, No. 1. (Feb., 1997), pp. 55-64.
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.Iournal of \.e,qc.tation Science 8 55-64. 1997
O IAl'S; Oplrllrs PI-ess U11)sala PI-intedin Swederi
Seed predation, disease and germination on landslides in Neotropical lower montane wet forest Myster, Randall W.
Institure,for T~.opiculEcosystem Studies, U i z i ~ ~ e rofs iPlierto
~
Rico, P.O. Bos 363682, Sun .Tlian, PR 00936. USA;
Te1. + 1 787 767 0371 ;Fus
+ 1 787 758 0815; E-mail ~_mystc~r@lrp~.l
.~q~~:cl~t.edlr
Abstract. Seed mortality (caused by predators and pathogens)
and germination were compared between Puerto Rico and
CostaRica on landslides in lower montane wet forest. Seeds of
six common species on five Puerto Rican landslides and four
common species on two Costa Rican landslides were used
with a Cc~cropiaspecies and a GonzaIa,yutlia species included
at both sites. In the Puerto Rican experiments Cec,ropia
schl.eher.ianu was the only species to show significant seed
predation (which was due to insects), pathogens grew from all
species and fewer seeds were lost to predators than pathogens.
Also in Puerto Rico mean germination across all species was
57 C/c before dispersal (filled seeds collected while still on the
tree) and 7 1 C/c after, with Phxto1acc.u r.ir,inoide.s seeds germinating most abundantly, followed in descending order by
Ocorea Ieuco.uxlon, Cec,r.opiu spec., Micotlia I-ac,emosu,
Pa1ic~our.e~
ripu/.za and Gonzula~utliaspic,ata. In the Costa
Rican experiments three species had significant predation:
Cec,l.opia poly~~hlehia
and Ur-el-acar~ac~a.sunu
(both due to
insects) and Withcr.itl,yia c,occ,olohoides (due to mammals):
pathogenic disease caused more seed loss than predation, and
germination was high (61 % pre-dispersal, 69 C/c post-dispersal).
Similarities between these island and mainland sites included ( I ) percentage of seeds lost to predation and percentage
lost to pathogens (all in the 5 - 15 % range), (2) generalist
pathogens which claimed more seeds than predators and (3)
majority germination with a general increase after dispersal.
Finally sites were dissimilar only in the number of species
with significant predation loss and whether it was b) insects
'
or
mammals, casting doubt on the traditional islandlmainland
dichotomy.
Keywords: Costa Rica: Disturbance: Island vs. Mainland;
Montane forest: Puerto Rico; Regeneration; Seed dynamics.
Nomenclature: Liogier & Martorell (1982); Haber (1991)
Introduction
Biotic responses to disturbance, particularly to severe disturbance (Garwood et al. 1979; van der Maarel
1993),help determine ecosystem structure (Watt 1947;
Brokaw 1985;Pickett &White 1985; Brokaw &Walker
199 1). Landslides are among the most severe neotropic
disturbances (Waide & Lugo 1992), removing organic
matter and topsoil, exposing bare rock and shallow
subsoils, and generating extreme environmental gradients (Fernindez & Myster 1995; Myster & Fernindez
1995). On a larger landscape scale, landslides may help
maintain biodiversity, for instance by creating regeneration sites for rare species and ferns (Walker 1994) or
acting to restock less severe but more frequently disturbed areas such as hurricanes (Willig & Camilo 1991)
and influence nutrient cycling and material flowlredistribution (Swanson et al. 1982; Guariguata 1990).
Understanding island ecosystem structure has always included comparisons with the mainland and often
stressed species richness differences due to ( 1 ) habitat
diversity, ( 2 )island size and distance to and diversity o f
the mainland, ( 3 ) immigration and extinction rates, or
( 4 )island 'samples' o f the mainland (Martinet al. 1995).
In addition the strength o f species interactions has been
part o f this comparison (Janzen 1973; Kitayama &
Mueller-Dombois 1995).However islandlmainland differences in disturbance regime has been missing in these
studies and may be critical to understanding ecosystem
structure on both (Sugden 1992). Because neotropic
islands differ from the mainland in hurricane (Vandermeer et al. 1990;Brokaw & Walker 199 1 ) and treefall
gap frequency and dynamics (Brokaw 1985; Lawton &
Putz 1988), landslides may be the natural disturbance
which occurs in the most places. Consequently, I examine biotic responses after landslides because they may
be useful in illuminating insular/continental similarities
and differences, and, therefore, provide a good framework for comparison.
The little regeneration from saplings, seedlings or
even from the seed pool in landslides (Guariguata 1990;
Myster 1993b; Myster & Fernindez 1995; Walker et al.
1997)points to a seed-driven revegetation process, where
seed dispersal (Walker & Neris 1993), seed mortality
mechanisms and seed germination are filters (Myster
1993a). Therefore, this study focuses on seed-oriented
field experiments on landslides both in Puerto Rico and
56
Myster, R.W.
Costa Rica and addresses these questions:
1. What are the effects of predation (invertebrate and
vertebrate) and fungal disease on seeds collected directly from the tree (pre-dispersal) and on seeds placed
on landslides (post-dispersal)? Also, is germination affected by dispersal?
2. How do seed predation, seed disease, and seed germination vary among tree species, compared to within and
among landslides?
3. How do the levels of predation, disease and germination differ between Neotropic island and mainland landslides with similar life-zones (lower montane wet forest) and flora?
Methods
Stuclq. sites and spec,ies
The Neotropic island study site was the Luquillo
Experimental Forest (LEF), a Long-Term Ecological
Research site of the U.S. National Science Foundation,
in the northeastern corner of Puerto Rico, USA
(18" 20' N, 66" 00' W). Within the LEF, this study concentrated on the lower montane wet forest (Ewe1 &
Whitmore 1973) characterized by Due,,-yodes escelsa
(tabonuco), Cyr.illa I-ac,cmiflor-a(palo colorado) and
Prestoea motitanu (sierra palm) (Waide & Lugo 1992).
Common landslide vegetation includes the ferns
Cyathea ar.hor.ea and Glcic,hetlia hifida in the bare soil
areas and Cecropia sc,ht.chcriana, Mic,onia racemosa,
and Nej7ser.a acuatic,a in the lower debris areas (Myster
& Fernandez 1995). Five landslides were selected: ES2
and ES3 in the Rio Espiritu Santo watershed (both < 5
yr old), RB2 and RB9 in the Rio Blancos watershed
(15 and < 5 yr old, respectively), and one in the Bisley
watershed (< 5 yr old). All are close to 2000 m' in size
with the exception of RB2 which is 25 000 m' (Myster
& Walker in press). In addition all landslides are between 500 and 600 m in elevation and, except for Rio
Blancos landslides - which are on diorite intrusions have soils derived from volcanoclastic primary substrate
that has weathered into clay Ultisols (Walker 1994).
The seeds used in the LEF experiments came from
five bird-dispersed woody species (Cect.opia schreheriatia, Miconia t.acclmosa, Pulicourea riparia, Gotizalugutiia spicata and Ocotea lerlc,osylon; Parrotta 1995)
and one bird-dispersed herb, Phytolacca ri~itioicles.
Ph~tolaccaoccurs in the initial grass-herb stage of
LEF landslide succession (Myster et al. in press), which
stabilizes the soil and builds up soil nutrients needed
for the first tree invader Cect-opia, which may then
moderate environmental extremes for the other woody
test species which follow (Myster & Walker in press).
All species are abundant in both vegetation (Myster &
Walker in press) and seed rain on LEF landslides
(Walker & Neris 1993).
The Neotropic mainland study site was the Monteverde Cloud Forest (MCF) Reserve of Costa Rica
(10" 12' N, 84" 42' W). The MCF is dominated in the
overstory by Lauraceae, Moraceae and Araliaceae, and
in the understory by Ruhiaceae, Solanaceae, Acarzthaceae, Gesneriaceae, Piperaceae and Arecaceae
(Palmae) (Lawton & Dryer 1980). Landslide vegetation is dominated by Gunnet-a insigrzis in the bare soil
areas and Chl4squea pohlii, Cecropia pol~phlehia,and
Palicourea standleyana in the lower debris areas (Myster
1993b). MCF has well-weathered soils of volcanic origin.
The climates of both LEF and MCF are dominated
by trade winds that produce moisture-bearing clouds
year-round (Odum & Pigeon 1970; Nadkarni & Matelson
1991). Total water input is similar between the two
sites, although MCF receives more moisture deposition
from mist-bearing winds (MCF averages 3.3 m/yr with
2.5 m/yr as rain and 0.8 m/yr as deposition; LEF averages 3-5 m/yr, almost all in rainfall with small amounts
in deposition; Nadkarni & Matelson 1991: D. Schaefer
pers. comm.). In addition, LEF has a dry season between late spring and summer and MCF has a dry
season during the first few months of the year (Holdridge
1967; Lawton & Dryer 1980: Janzen 1983). Normal
wind speeds above the canopy also have overlapping
ranges (MCF 5 - 15 km/h; LEF 2 - I0 km/h; Brown et
al. 1983; Nadkarni & Matelson 199 1 ).
As a further indicator of the site similarity, the
majority of tree genera found in Puerto Rican landslides
also occur in Costa Rican landslides, and, therefore, the
LEF landslide flora appears to be a large subset of the
MCF landslide flora, at least at the generic level (Myster
1993b). Because of this floristic similarity and in order
to magnify comparisons, test species were chosen to
include a species of the genus C~c.r.opiaand a species of
the genus Gotizalag~rtiiaat each site. The seed species
used in the MCF experiments were Cec.t.opiupolyphlehia,
Gon:alag~4nia kallutikii, Uwra c.ar.ac.usutia and Wither.itigia coccolohoicles which are common species in
Costa Rican landslide seed rain and vegetation (Myster
1993b) and may fill similar ecological roles as LEF
landslide species of the same genera. C ~ c r o p i ais both
bird and mammal-dispersed; the other species are birddispersed.
The two landslides studied at MCF were in the Rio
Penas Blancos watershed at 1300 m, approximately
2000 m' in size and were 5-15 yr old (M. Marin pers.
comm.). Although MCF landslides are at a higher elevation than LEF landslides, the 'Massenerhebung' or
mass elevation effect (Grubb 197 1 ) should reduce any
- Seed predation, disease and germination in wet forest on landslides
vegetation differences due solely to elevation. Taken
together, the similarity in climate, soils and vegetation
between MCF and LEF makes this pair of sites appropriate for comparative islandlmainland studies.
Predation, disease and germination espel-iments
At both sites, I hand-collected ripe seeds from 25-30
trees of each test species and examined seed predation,
disease and germination both before and after dispersal
- which provided more detail into the regeneration biology of the individual species. The following were
considered evidence of pre-dispersal seed predation:
empty seedlfruit stalks obviously removed by birds or
mammals and non-abortive seeds with extensive insect
damage either seen with a hand lens or implied by the
ability of the seed to float in water (Myster & Pickett
1993). Intact (presumed viable) seeds were taken back
to the laboratory, placed on moist paper in plastic-wrap
covered petri dishes, and put under plant growth lights
(Sylvania 40 watt wide spectrum GRO-LUX) to create
a warm, humid environment. This was necessary to
determine which non-germinating seeds were lost to
fungus and which were non-viable. After one month
seeds were examined under a dissecting microscope for
germination and fungal growth, and assigned to one of
three categories: germinated, diseased or other. Seeds
that failed to germinate and had visible fungal damage
were classified as 'diseased' and seeds that failed to
germinate but had no microscopically visible fungal
damage were assigned to the 'other' category. This
categorization is reasonable, though limitations must be
kept in mind: germinated seeds with fungal damage
might have died later from that damage, fungi might
have attacked seeds that would not germinate anyway
or that died due to other agents (e.g. bacteria; Groth
1989), and ungerminated seeds might have germinated
given more time.
For both the Puerto Rican and the Costa Rican postdispersal field experiments, I hand-collected more seeds
from the study trees and examined them for predator
effects as before; only those seeds without damage were
used. The wide differences in seed mass and size necessitated using different numbers of each species in the
post-dispersal treatments to standardize the seed area
presented to predators in the treatments (Myster & Pickett
1993). At LEF, seed mass was measured and 20 Cecl-opia
(0.0003 g/seed) and 20 Miconia (0.002 glseed) seeds
were placed in each petri dish. Palieout-ea (0.2 glseed),
Gorzzalaglrnia (0.02 glseed), and Ocotea (0.1 glseed)
had four seeds per dish and Phytolacca (0.01 glseed)
had 10 seeds per dish. At MCF the same procedure was
followed with 20 Cecr.opia and 20 Urel-a (0.0003 g)
seeds placed per petri dish. Gon:alagl4nia (0.029) had
-
57
four seeds per dish and Withel-inga (0.01) had 10 seeds
per dish. Consequently, percentage of seeds remaining
in the dishes after placement in a landslide was the
dependent response variable instead of raw number of
seeds remaining or lost (Myster & Pickett 1993). Neither Puerto Rican nor Costa Rican seeds had passed
through a bird gut. Because gut passage did not affect
germination success for Phytolacca r.i~~irzoicies,
Wither.inga solarzac~aand W . c~occ~olohoides
in another MCF
experiment (Murray 1988), I assume that any effect of
bird gut passage on the results here would be minimal.
The effect of mammal and insect exclusion on postdispersal seed removal in all landslides was tested using
four treatments:
(1) cages of 1 cm' mesh hardware cloth enclosing
open plastic petri dishes of 9 cm diameter to exclude
mammals - including the roof rat Rattus rattlis and the
mongoose Herpestes nurrzgo in Puerto Rico (Weinbren
et al. 1970) and, in Costa Rica, the rodents Peron~ysclls
m~.ricanus,Heteromys desrnal-estianus and Olyzomys
alhigularis, the variegated squirrel, Scila-us \>at-iegatoides, and the agouti, Dasypl-octa pllnctata (Hayes &
Lava1 1989) and large invertebrates such as the Puerto
Rican land crab Epilohoc,era sirzuatifi-ons;
(2) tanglefoot (a sticky resin) spread on the inside of
petri dishes to exclude small non-flying invertebrates;
(3) both treatments on the same petri dish; and
(4) a control without any treatment.
For each of the four treatments, seeds of each of the
six species were placed on top of local soil spread in
separate dishes for a total of 24 dishes. Dishes were then
arranged randomly and glued to a wooden platform
(0.5 m x 0.5 m; surface area = 0.25 m2). Over the platform, a plastic roof at a height of 0.5 m was placed to
protect dishes from the weather and to reduce any seed
loss due to rain and wind. Platforms were anchored in
place with a 30 cm long plastic stake. Five colored
inedible plastic seed mimics of similar shape and mass
were placed in each control petri dish to see if any seed
losses occurred due solely to rain impact. In addition,
extra dishes with seeds but no soil were placed on the
Puerto Rican Rio Blancos landslides to determine if soil
contact was needed for fungal infection of seeds.
All replicates were placed in the microhabitats of
forest border (in the forest just outside the lip where soil
and vegetation slid), landslide border (just inside the
lip), and center of the landslide on three transects, for a
total of nine replicates per landslide. The transects were
20 m apart starting 20 m from the top of the slide. The
experiment design was six species (four in Costa Rica)
x four treatments x nine replicates. The Puerto Rico
experiment ran for 26 days from mid-March to midApril 1992, and the Costa Rica experiment ran for 24
days in May 1992. At the end of the field experiment
Myster, R.W.
PUERTO RICAN SPECIE
Re Post
Cecropia Gonzalagunia
schreberiana
spicata
Miconia
racemosa
Ocotea
leucoxylon
A
Palicourea
riparia
Phytolacca
rivinoides
Fig. 1. The fate of seeds of plant species that colonize landslides in Puerto Rico: eaten by predators (clear), killed by disease (darkcrossed), germinated (light-crossed) or unaffected by these three mechanisms ('other' in the Methods: black) both before dispersal
(pre) and after (post).
and on the same day, the number o f seeds remaining in
all dishes was recorded. Then all seeds left in the control
dishes were pooled for subsequent incubation, to detect
fungal infection and germination, for 60 days in the
laboratory either in Puerto Rico or Costa Rica (same
protocols as for pre-dispersal seed incubation).
Treatment effects were investigated using nested
analysis o f variance ( A N O V A ; Sokal & Rohlf 1981;
Anon. 1985) where the dependent variable was percentage o f seeds remaining. For species with four seeds per
dish. the range o f response values was too narrow to
approximate a continuous function and so logistic regression analysis was used (Anon. 1985; Myster &
Pickett 1993). In both cases, only significant results are
given. Nested A N O V A allowed me to partition the
variance in seed percentage remaining and identify significant treatment effects.Here, this means groups (sensu
Sokal & Rohlf 1981) o f dishes pooled in individual
microhabitats (e.g. landslide center dishes vs. landslide
border dishes vs. forest border dishes. referred to as the
within-landslide variance component in the results) vs.
groups o f dishes pooled in each landslide (e.g. ES2
dishes vs. RB2 dishes, referred to as the among-landslide variance component in the results). Finally, to
assess significant differencesamong seed process mechanisms, sites or pre/post dispersal levels, a simple t-test
was used (Sokal & Rohlf 198 I).
Results
Because fewer than 1 % o f the colored mimic seeds
were lost to the weather in either the Puerto Rican or
the Costa Rican experiments, significant treatment seed
losses were assumed to be due to active removal by
animals which leads to predation (Myster & Pickett
1993). In addition, seeds that were obviously eaten but
not removed were counted as predation. Both pre- and
post-dispersal predation losses were small for all six
Puerto Rico species, only once exceeding 10 % (Fig.
I). In addition, post-dispersal predation was greater
than pre-dispersal in four o f the six test species; Miconiu
had the only significant increase (3 C/o pre/l5 56 post; t
= 7.65, d f = 2, p = 0.05).There was a trend o f increased
predation for smaller-seeded species (e.g. Miconiu,
Cec,ropia, G O I I : U ~ U ~ M
M ~ U ) to larger-seeded
compared
but see
species (e.g. Or,oteu, Pulicoureu, Pli~toluc,c~u:
Uhl 1987).
The only significant treatment e f f e c t was for
Cec~.opiuschreheriu~luunder the non-volant insect exclusion treatment (Table 1 ) which reduced predation
from 9 % to 2 5%. Ants may have been the predators
because they were sampled in the five Puerto Rico
landslides (Myster in press) and fed on Cecl-opiu seeds
when offered seeds from all test species in feeding
trials. Most post-dispersal predation occurred in the
first week regardless o f species and, consequently, the
- Seed predation, disease and germination in wet forest on landslides -
Table 1. Complete ANOVA table for the dependent variable
of percentage of seeds remaining in the Puerto Rican predation
field experiment. Significant effects are highlighted in bold
type and only those species are given. ss = sum of squares: ms
= mean squares.
Source of variation
df
Cec i.oplu s<~h~.rhr~~.iariu
In\ects excluded
I
1
Mammal\ excluded
Insect\ and mammals excluded
1
Error
41
us
m\
F
1.82
0.23
0.39
31.16
1.82
0.23
0.39
0.76
3.45
0.50
0.67
P
0.05
0.55
0.43
rate of seed loss was approximately 1 - 2 C/c per day
(average of 10 % for all species divided over 7 days),
similar to temperate forest disturbance studies in New
Jersey, USA (Myster & Pickett 1993). Partitioning the
variance for the only species having significant losses
(Cecropia) showed 80%' variance among the five slides
(ANOVA variance component: VC = 0.435), leaving
20 8 variance within the five slides (VC = 0.061).
Here, as in all Puerto Rican results, among-slide
variances were not decreased by grouping landslides
into common watersheds even through local hydrology
is presumed to be important in recovery from disturbance (Waide & Lugo 1992).
Puerto Ricai? seed disease
Seeds in dishes that contained soil did not have a
greater infection percentage than those in dishes without
soil. and so contact with the soil may not be necessary
for infection (but see Augspurger & Kelly 1984). Levels
of seed loss to pathogens were similar to the small
predation losses for four of the six species (22 r/c loss
maximum) with post-dispersal losses greater than predispersal losses for those four species (Fig. 1) (t-values
with df = 2 and p = 0.01 for each species) were:
Cecropia
Micoi~ia
10.45
8.87
Gonzalagunia
Phytolacca
5.2 1
9.02
Alternatively Ocotea and Palicourea showed much
greater loss to pathogens (86 r/c maximum), more predispersal losses compared to post-dispersal; (t-values
with df = 2 and p = 0.05):
Ocotea
6.35
Palicourea
6.98
There were much greater seed losses to pathogens than
to predators (t-values with df = 2 and p = 0.01):
Ocotea
t = 15.45
Palicourea t = 20.02
In addition. Cecropia and Ocotea lost a relatively greater
59
proportion of seeds to pathogens at the forest edge
compared to the center of the landslide (21 % vs. 9 8;
similar to seedlings in gaps; Augspurger 1984).
When seeds were isolated and incubated, the fungal
genus could be identified under the microscope in many
cases. A species of Colletotrichuni (Melanc.o~liules)
grew from dead Pulicozlriu seeds and is known to inhibit germination and cause seed mortality in some
species (Tripathi 1974; Ellis et al. 1979). A young
Phytolucca individual that died after germination exhibited watersoaking and softening of the hypocotyl
tissues, which are typical symptoms of P~thiurndamping-off disease (Hering et al. 1987), and a P~thiurn
species was found growing in the affected hypocotyl.
Fusui-iuni species (Moniliales) grew from Phytolucca,
Pulicoui-ia and Ocoteu seeds, but it is not known if this
was the cause of mortality. However, a Fusurium spec.
probably contributed to the death of a Cec,ropia schreheriuna seedling which exhibited a pinkish lesion on
the hypocotyl and dead feeder-roots, symptoms typical
of Fzlsai-ium damping-off and root rot (Huang &
Kuhlman 1989).
Another fungus in the Moniliules, belonging to the
genus Arthrosl~oi-ium(Wang 1972). grew from Cecropia
schreheriunu seeds although this genus is generally
saprophytic (D. J. Lodge pers. comm.). Rhizoctoniu
spec. infected a Ceci-opiu sc.hi-ehei-ianu seedling and
possibly a Phytolacca seed and is a known seedling
pathogen (Sinclair & Backman 1989). In addition, two
unidentified fungi that produced rhizomorphs or rootlike structures apparently destroyed seeds of Palic,ouria.
The effects of fungi may have been greater than scored
because infected seedlings may be more susceptible to
other stresses such as drought, and this may thus contribute to seedling mortality later (D. J. Lodge pers.
comm.).
Puerto Rican seed gernzinatioi~
Except for the pre-dispersed seeds of Ocotea and
Palicourea, the majority of seeds of all six test species
germinated both pre-dispersal and post-dispersal (Fig.
I). Because species that were abundant in landslides
were used, this result is consistent with Walker & Neris
(1993) who also found a close positive correspondence
between abundance and germinability in Puerto Rican
landslides. With the exception of the small-seeded
Cecropia and Miconia, germination either increased or
stayed the same after dispersal for all species, suggesting that some seeds that were lost to predators and
pathogens before dispersal were viable and could have
germinated after dispersal. G o n ~ a l a g u ~and
~ i aPalicourea
showed the largest percentage of non-viable seeds (between 32 % and 45 %). Comparison between dishes
- Seed predation. disease and germination in wet forest on landslides -
TOTAL SEEDS
Pre
~ost
Pre
61
1988) with significantly more Cecropia seeds germi~ost
nated in Puerto Rico compared to Costa Rica ( t = 5.29,
df = 2, p = 0.05) and Gonzalugurzia had no significant
predation at either site, with similar levels of predation
and pathogens in Puerto Rico and Costa Rica.
Discussion
PUERTO RICO
COSTA RICA
Fig. 3. Combining the results of all plant species of Figs. 1 and
2 into mean percent loss of seeds to predation, disease, germination and 'other' of all Puerto Rican plant species and of all
Costa Rican plant species both pre- and post-dispersal using
the same shading indicators as Fig. 1.
minated at 14 R in a Puerto Rican landslide study;
Walker & Neris 1993) with the relatively large-seeded
Witheringia germination increasing dramatically after
dispersal (Murray 1988). The proportion of total seeds
that were non-viable showed large species variation,
e.g. Ur.er-awith small percentages and Withel-ingia with
large percentages. Perhaps some of these could have
been dormant and still viable, e.g. Ceci.opiu spp. and
Witheringiu (Murray 1988) can have a long dormancy
period.
Puerto Rican 1,s. Costa Ricun seedp~.ocesses
Comparing the results from Puerto Rico and from
Costa Rica (Fig. 3) reveals these similarities: (1) the
level of both pre-dispersal and post-dispersal seed predation is low in both sites, (2) germination levels are
similar when species are combined and increase after
dispersal at both sites, and (3) non-viable seeds are in
the 10 - 20 C/c range of total seeds and decrease in percentage after dispersal. By contrast, even though more
seeds are lost from pathogens than from predation at
both sites, this trend is only significant in Puerto Rico
( t = 6.33, df = 2, p = 0.05). For the two plant genera
used at both sites, insect predation reduced Cecr-opiu
seeds significantly at both sites (the loss of Azteca or
other ant species association in Puerto Rico did not
seem to affect insect predation levels; Putz & Holbrook
Plant species had individualistic (Myster & Pickett
1988) predation, disease and germination responses
(Myster & Pickett 1993) and this species variation dominated over seed process, site, and pre/post dispersal
differences. Because germination levels are a function
of the seeds which escape predation and pathogens,
these three regeneration processes are not mutually exclusive and, furthermore, may have probabilistic (e.g.
influenced by animal behavior) or stochastic (e.g. fungi
dispersal) interactions. In addition the use of plant characteristics to predict seed losses had mixed support.
For example there was weak positive correspondence
between seed mass and level of predation and pathogens (Louda & Zedler 1985; Uhl 1987), but there may
be more support for this hypothesis when the predators
are vertebrates rather than invertebrates (Myster &
Pickett 1993). Similar to seedlings (Augspurger & Kelly
1984), the vulnerability of seeds to pathogens varies
among neotropical species but seed mass is not a good
predictor of vulnerability to pathogens (Augspurger
1984) even though light-seeded early successional species may be more susceptible to seedling pathogens
than later successional species (Augspurger & Kelly
1984). In general, little is known about what seed characteristics permit or encourage seed infection.
Although higher levels of predation than found in
this study have been suggested for Cecr-opiu and
Palicour.eu in other LEF studies (Weinbren et al. 1970),
Walker & Neris (1993) found in landslides similar levels to mine and agree that Ph~tolaccuand Gonzalaguniu
are successful germinators and Palicour~euis relatively
unsuccessful. Microsite germination cues (light increasing rates but litter, high soil moisture and high temperatures decreasing rates) were shown in a LEF gap
study by Everham et al. (1996) suggesting that the
small number of moderate temperature microsites in
landslides (Myster & Fernhndez 1995) could be contributing to slow invasion and establishment rates on
landslides. Finally, combining the results with other
landslide studies shows relationships between seed processes and establishment patterns. Using permanent plot
data on LEF landslides which included those used in
this study (Myster & Walker in press), there is a positive correspondence between stem density and seed
process results seen here; those species with small losses
62
Myster, R.W
to predation and disease and large germination levels
were common in permanent plots. This pattern held
true for some species (Cecl-opiu, Gonrulugilniu) but
not for others ( U I - e m ) in MCF landslides (Myster
1993b).
Species interactions on islands - e.g. seed predation
(Janzen 197 1) - have been thought to be relatively weak
(Janzen 1973) compared to the mainland. However,
experimental studies on islands have shown that insect
seed predation can be intense (Louda & Zedler 1985).
Perhaps surprisingly, this study showed similar levels
of all three seed processes (predation, disease, germination) on a neotropical island site and neotropical
mainland site with similar life-zone, climate, and tlora.
The biggest difference between the Puerto Rican ecosystem and the Costa Rican ecosystem may be the lack
of native terrestrial mammals in Puerto Rico (Nilsson
et al. 1985), which could lead to a lack of mammaldispersed plant species and allow other animal species
to move up in the island food chain to replace the
mammals (as in Puerto Rico: Waide 1987). Taken together, the results suggest that islands and the mainland may not differ in much more than basic climate or
species composition (here mammal: Lugo 1987) and,
consequently, run counter to the model of reduced
strength of biotic processes (e.g. competition. predation, herbivory) on island ecosystems compared to mainland ecosystems (Janzen 1973).
Acknowledgements. I hish to thank D. Jean Lodge of the
Center for Forest Mycology Research, Forest Products Laboratory. USDA - Forest Service, for her help in identifying
fungal genera. I also wish to thank my field assistants in Puerto
Rico (Maria Aponte and Mildred Cruz) and in Costa Rica
(Marcos Marin and Gabriel Barbosa). Finally, I thank L.
Walker. C. Augspurger, J. Zimmerman, D. J. Lodge. D. Clark,
C. Peterson. A. Sabat and E. Schupp for conlnlenting on
earlier drafts of the manuscript. This research has performed
under grant BSR-8811902 from the National Science Foundation to the Institute for Tropical Ecosystem Studies, University of Puerto Rico, and the International Institute of Tropical
Forestry as part of the Long-Term Ecological Research Program in the Luquillo Experimental Forest. Additional support
was provided by the Forest Service (U.S. Department of
Agriculture) and the University of Puerto Rico.
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Accepted 3 June 1996.
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Seed Predation, Disease and Germination on Landslides in Neotropical Lower Montane Wet
Forest
Randall W. Myster
Journal of Vegetation Science, Vol. 8, No. 1. (Feb., 1997), pp. 55-64.
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