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Aquatic fungi associated with decomposing Ficus sp. leaf litter in a neotropical
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DOI: 10.1899/07-084.1

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J. N. Am. Benthol. Soc., 2009, 28(2):416–425
Ó 2009 by The North American Benthological Society
DOI: 10.1899/07–084.1
Published online: 7 April 2009

Aquatic fungi associated with decomposing Ficus sp. leaf litter

in a neotropical stream

José Rincón1 AND Roberto Santelloco2

Departamento de Biologı́a, Facultad Experimental de Ciencias, Bloque A-2, Apartado 526,
Universidad del Zulia, Maracaibo, Venezuela

Abstract. Aquatic fungi are important decomposers of plant litter in temperate streams. However, these
microorganisms have been poorly studied in tropical streams. We assessed the dynamics of aquatic
hyphomycetes during the decomposition of Ficus sp. (Moraceae) leaves in an intermittent lowland stream of
northwestern Venezuela. Our goals were to: 1) determine the composition of aquatic hyphomycetes during
leaf decomposition, 2) assess the effect of different mesh sizes (i.e., coarse vs fine mesh) on assemblage
composition and activity, and 3) assess the role of habitat (i.e., pools vs riffles) on fungal assemblages.
Coarse- and fine-mesh litter bags were incubated in a pool and a riffle for 31 d. For each habitat and
treatment, we determined physicochemical variables, sporulation activity, and the structure and
composition of fungal assemblages. Leaf litter decomposition rates ranged from 0.058/d in fine-mesh bags
in the riffle to 0.031/d in coarse-mesh bags in the pool. Decomposition rates were significantly influenced by
habitat type, with the highest decomposition rates found in riffle habitats. Sporulation rates were
significantly different between habitats but not between mesh sizes. The highest sporulation rate (399 spores
mg1 leaf dry mass d1) and fungal diversity (16 spp.) was found in coarse-mesh bags in the riffle. Twenty-
one fungal species were identified during the study. Anguillospora longissima and Lunuluspora curvula
contributed up to 75% of the conidia produced in the coarse- and fine-mesh bags in the riffle, whereas
Filosporella aquatica and Lemonniera terrestris accounted for 65 to 75% of the conidia produced in the coarse-
and fine-mesh bags in the pool. In ordination space based on species composition, fungal assemblages
formed 2 clear groups that corresponded to habitat type. A temporal change in fungal species composition
was observed in riffle samples. Early succession assemblages (2–14 d) were characterized by the presence of
A. longissima, L. curvula, and F. aquatica, whereas late assemblages (24–31 d) were dominated by
Pyramidospora fluminea, Pyramidospora casuarinae, and unidentified sp 4. Overall, habitat type (pool or riffle)
influenced sporulation rates and assemblage composition of aquatic hyphomycetes. Our study suggests that
environmental factors associated with hydrology (e.g., stream velocity, width, depth) regulate fungal
composition and activity and, hence, leaf litter decomposition in this tropical stream.

Key words: tropical, decomposition, aquatic hyphomycetes, sporulation, intermittent stream.

Aspects of fungal dynamics associated with leaf
litter decomposition are well studied in temperate
streams (e.g., Bärlocher and Kendrick 1974, Suber-
kropp and Klug 1976, Griffith and Perry 1994, Baldy et
al. 1995, Gessner 1997, Grattan and Suberkropp 2001,
Happala et al. 2001, Crenshaw and Vallet 2002, Hieber
and Gessner 2002, Laitung et al. 2002, Gessner and
Robinson 2003, Robinson and Gessner 2003), but less is
known about the role of fungi during leaf litter
decomposition in tropical streams (Mathuriau and
Chauvet 2002). A few studies in tropical Asia and
1
E-mail addresses: jerincon04@gmail.com
2
rsantelloco@hotmail.com
416
subtropical Africa have addressed colonization and
diversity patterns of aquatic hyphomycetes during leaf
decomposition in streams (Raviraja et al. 1996, 1998,
Rajashekhar and Kaveriappa 2003, Abdel-Raheem
1997), and a few taxonomic studies are available for
neotropical streams (Hudson and Ingold 1960, Betan-
court and Caballero 1983, Betancourt et al. 1987,
Justiniano and Betancourt 1989, Schoenlein-Crusius
and Piccolo 2003). In Venezuela, initial taxonomic
studies were published by Nilsson (1962), and more
recent studies have been published by Smits et al.
(2007) and Cressa and Smits (2007). To our knowledge,
few studies have addressed patterns of aquatic
2009] AQUATIC FUNGI
hyphomycetes associated with leaf litter decomposi-
tion in neotropical streams (Mathuriau and Chauvet
2002, Rincón et al. 2005).
Litter quality (i.e., lignin, secondary compounds,
and nutrient content) (Abdel-Raheem 1997, Mathuriau
and Chauvet 2002) and habitat type (Rincón et al.
2005) influence biomass and colonization patterns of
aquatic hyphomycetes and leaf litter decomposition in
tropical streams. High fungal activity associated with
leaf decomposition appears to be characteristic of leaf
processing in tropical streams (Mathuriau and Chau-
vet 2002). Moreover, fungal colonization of decompos-
ing leaves appears to affect shredder feeding
preferences in tropical streams, as it does in temperate
streams (Graça et al. 2001, Rincón and Martinez 2006).
Abiotic factors, such as water temperature, nutrient
concentrations, pH, and alkalinity, seem to affect
fungal activity during leaf litter decomposition (Cha-
mier 1992, Suberkropp and Chauvet 1995). The effects
of other factors, such as habitat type (riffles vs pools),
on aquatic hyphomycete activity have been less
studied. The roles of macroinvertebrates and fungi
appear to change with habitat in temperate regions,
and such changes have the potential to affect leaf litter
decomposition (Baldy et al. 2002, Langhans et al.
2008). Pool–riffle sequences are particularly important
in intermittent streams where seasonal hydrological
changes can produce dramatic changes in some
physical and chemical variables affecting the stream
biota (Rincón and Cressa 2000). In a temporary
Moroccan river, fungi were most active in spring
when water was constantly flowing and decomposi-
tion was fastest (Maamri et al. 1998). Our understand-
ing of the effects of stream habitat on fungal activity in
tropical streams is still very limited.
Little is known about how macroinvertebrate
species and aquatic fungal species interact to affect
leaf litter processing rates. Differences in the feeding
mode of macroinvertebrate species (e.g., scraping,
shredding) might influence fungal colonization and
leaf decomposition rates. Leaf decomposition rates can
be accelerated in the presence of macroinvertebrates
because aquatic fungi can penetrate damaged leaf
tissue rapidly. We studied leaf decomposition in fine-
and coarse-mesh leaf bags to assess the dynamics of
aquatic hyphomycetes in an intermittent neotropical
stream in northwestern Venezuela. Our goals were to:
1) determine the composition of aquatic hyphomycetes
during leaf decomposition, 2) assess the effect of
different mesh sizes (i.e., coarse vs fine mesh) on
assemblage composition and activity, and 3) assess the
effect of habitat (i.e., pools vs riffles) on fungal
assemblages.
IN NEOTROPICAL STREAMS 417
Methods
Study site
We studied leaf litter decomposition in Carichuano
Creek, an intermittent 2nd-order stream in northwest-
ern Venezuela (lat 10842 0 –11808 0 N, long 72842 0 –
72822 0 W). The stream is in the northern foothills of
Sierra de Perijá (Estado Zulia, Venezuela), which is a
major structural component of the Venezuelan Andean
Cordillera system. Carichuano Creek is a tributary of
the Guasare River, which flows into Lago de Mara-
caibo.
The stream drains a diverse tropical semideciduous
dry forest, with ;17 to 20 plant species present in the
riparian zone. Tabebuia rosea and Calliandra sp. are
dominant in this area, and Anacardium excelsum, Cartan
sp., Luehea sp., Lecithis ollaria, Hura crepitans, Centro-
lobium parense, Cecropia peltata, Guapira sp., Euphorbia
cassearea, Ficus sp., and Inga spurea also are present
(Cifuentes 2002). Catchment geology is dominated by
limestone, and streams have a characteristic water
chemistry classified as the bicarbonate–calcium type
(152–320 mg/L CaCO3) (Cressa et al. 1993, Rincón and
Cressa 2000).
The study area has a mean annual rainfall of 1102 6
285 mm, and the annual average water temperature is
27.78C. Two periods of rainfall, interspersed with dryer
intervals of variable duration, occur each year (Espi-
noza 1987). Typically, the 1st rainy period begins in
April and continues until June. A short dry period
follows for 1 to 2 mo. The 2nd rainy period starts in
September and lasts until November or December, and
is followed by another longer dry period (2 or 3 mo).
Litterfall is strongly seasonal, with peaks at the end of
the 2 dry periods (February and July) (JR, unpublished
data). Water flow is not continuous during the dry
seasons and isolated pools are common.
Leaf-litter decomposition experiment
Freshly fallen Ficus sp. (Moraceae) leaves with no
indication of partial decomposition or fungal coloni-
zation were collected from the riparian forest floor in
January 2004. Air-dried leaves were weighed into 3.0
6 0.1 g groups and enclosed in 15- 3 10-cm litter bags
(240-lm [fine] and 5-mm [coarse] mesh). Litter bags
were anchored to the stream bottom with steel bars.
Leaf litter decomposition experiments were con-
ducted from June to August 2004 (incubation periods
of 2, 7, 14, 24, and 31 d) in a riffle and a pool. Four leaf
litter treatments were used: coarse mesh in riffle,
coarse mesh in pool, fine mesh in riffle, and fine mesh
in pool. Five bags of each mesh size (coarse or fine)
were retrieved from each habitat (riffle or pool) on
418 J. RINCÓN AND R. SANTELLOCO [Volume 28

each sampling date. An additional 20 bags were used TABLE 1. Values of physicochemical variables in riffle and
to evaluate handling losses. Leaf bags were transferred pool habitats in Carichuano Creek, Venezuela, between July
to plastic zip-lock bags and transported to the and August 2004.
laboratory in ice chests. In the laboratory, the bags
Riffle Pool
were disassembled, and leaves were washed gently
with running tap water over a sieve (250 lm) to Variable Mean Range Mean Range
remove debris and invertebrates. On each sampling
Temperature (8C) 28.3 27.6–29.6 28.4 27.6–29.0
date, the contents of 3 randomly selected bags (n ¼ 3) pH 8.0 7.95–8.02 8.0 7.99–8.06
from each treatment were used to estimate leaf mass Dissolved O2 (mg/L) 9.5 8.9–10.5 9.8 8.7–10.4
loss; the contents of another 2 bags (n ¼ 2) were used to Conductivity (lS/cm) 580 562–590 555 447–607
determine fungal spore production. Leaf mass loss was Depth (m) 0.23 0.20–0.30 0.42 0.38–0.46
Width (m) 1.99 1.43–2.25 11.39 10.6–13.2
estimated after drying the leaf litter at 608C for 48 h. Current velocity (m/s) 0.46 0.31–0.57 0.11 0.10–0.13

Spore production
Eight leaf discs were cut from the leaves in each bag
with a cork borer (1-cm diameter). Discs were
incubated in a 250-mL flask containing 100 mL of
autoclaved and filtered (0.45-lm pore size membrane
filter) stream water and agitated in a shaker at 258C for
72 h (slightly modified from Bärlocher 2005). The
entire volume with suspended conidia was filtered
through an 8-lm pore size membrane filter. The filter
was stained with cotton blue in lactophenol for 30 min
at 408C, and the conidia trapped on the filter were
counted and identified at 4003 and 10003 magnifica-
tions under a light microscope. Generally, the entire
surface of the filter was scanned and all conidia were
counted. If conidial densities were very high, 25
randomly chosen microscope fields were scanned
and conidia in these fields were counted. Aquatic
hyphomycetes species were identified with available
keys for tropical areas (Marvanová 1997, Santos-Flores
and Betancourt-López 1997). Leaf discs were dried and
weighed to measure the amount of leaf material and
results were expressed as conidia mg1 leaf dry mass
(DM) d1.
Physicochemical variables
On each sampling date, temperature, conductivity,
and pH were measured with field instruments (Check
Mate 90; Corning, New York). Dissolved O2 (Winkler
method; Hauer and Hill 1996) was measured on 2
water samples collected at ;60% depth from each
habitat type. In addition, stream width, depth, and
flow (Digimeter Model 9000; Scientific Instruments,
Milwaukee, Wisconsin) were measured.
Data analysis
Percentage mass remaining through time was fit to
an exponential model mt/m0 ¼ ek t; where mt is the
mass remaining at time t, m0 is the initial mass, and k is
the decomposition rate coefficient (Boulton and Boon
1991). Differences in decomposition coefficients (de-
termined from linear regression of ln[x]-transformed
data) between habitat types (pool and riffle) and mesh
sizes (coarse and fine) were tested using analysis of
covariance (ANCOVA) in Prism 5.01 (GraphPad
Software, San Diego, California). Slope estimates were
used to represent decomposition coefficients for each
treatment and were compared with the Student–
Newman–Keuls (SNK) multiple comparison method
(a ¼ 0.05).
The effects of mesh size and habitat type on
sporulation rates were determined with 2-way analy-
sis of variance (ANOVA) (Sokal and Rohlf 1981). Data
were log(x þ 1)-transformed before analysis, and the
analysis was done with Statgraphics centurion XV
(Statpoint Technologies Inc, Warrenton, Virginia).
Fungal assemblage composition in pools and riffles
were analyzed with nonmetric multidimensional
scaling (NMDS) in PRIMER (version 5.2.9; PRIMER-
E, Plymouth, UK). NMDS is a procedure for plotting a
set of assemblages in a space such that the distances
between assemblages correspond as closely as possible
to a given set of dissimilarities/similarities among the
assemblages (Clark and Warwick 2001). Similarities
between 2 assemblages were calculated with the Bray–
Curtis similarity index. The abundance of each fungal
species was 4th-root(x)-transformed before the similar-
ities were calculated (Clark and Warwick 2001). In
addition, a SIMPER routine (PRIMER-E) was done to
inspect the contribution of individual species to
average dissimilarity between groups.
Results
Physicochemical variables
Physicochemical variables varied slightly during the
study period (Table 1). In the riffle, current velocity
increased steadily from day 2 to day 24, then
decreased sharply until day 31. In the pool, the current
2009] AQUATIC FUNGI IN NEOTROPICAL STREAMS 419

TABLE 2. Mean (SE) leaf-litter decomposition rates (k) for

Ficus sp. leaves in coarse- (5 mm) and fine- (0.24 mm) mesh
bags incubated in the riffle and the pool of Carichuano
Creek, Venezuela. Treatments with the same letter were not
significantly different on the basis of Student–Newman–
Keuls (SNK) multiple comparison method (a ¼ 0.05).

Habitat Mesh k SNK grouping

Riffle Coarse 0.04693 (0.0045) ac
Fine 0.05847 (0.0036) a
Pool Coarse 0.03132 (0.0037) b
Fine 0.03819 (0.0047) bc

faster in fine- and coarse-mesh bags in the riffle than in

coarse-mesh bags in the pool.

FIG. 1. Mean (61 SE; n ¼ 3) percentage dry mass (DM)
remaining of Ficus sp. leaves during decomposition in fine-
and coarse-mesh litter bags in the riffle and pool of
Carichuano Creek, Venezuela.
velocity decreased slightly from day 2 to day 14, then
peaked at day 24, and decreased by day 31. Stream
width and depth were ;63 and 23, respectively,
higher in the pool than in the riffle, but current velocity
was 43 higher in the riffle than in the pool (Table 1).
Riffle and pool habitat did not differ in dissolved O2
concentration, pH, temperature, or conductivity.
Leaf litter decomposition
Spore production
Spore production peaked after 14 d in the riffle (399
and 383 conidia mg1 DM d1 in coarse- and fine-mesh
bags, respectively; Fig. 3A) and after 7 d in the pool
(339 and 166 conidia mg1 leaf DM d1 in coarse- and
fine-mesh bags, respectively; Fig. 3B). Spore produc-
tion declined by the end of incubation in all treatments
and habitats.
Sporulation rates were significantly higher in the
riffle than in the pool (F ¼ 13.09, p ¼ 0.0010; Table 4).
However, sporulation rates did not differ between
coarse- and fine-mesh bags (F ¼ 4.12, p ¼ 0.0508), nor
was the interaction term (mesh-size 3 habitat) statis-
tically significant (F ¼ 0.44, p ¼ 0.5126; Table 4).

Loss of leaf litter mass was rapid during the first 2 d Fungal assemblages
of immersion (40 and 17% for coarse- and fine-mesh
bags, respectively, in the riffle; 31 and 30% for coarse- Twenty-one taxa of aquatic hyphomycetes were
and fine-mesh bags, respectively, in the pool; Fig. 1). identified from sporulating Ficus leaves in both riffle
After 7 and 14 d of immersion, leaf litter mass loss in and pool habitat (Table 5). Anguillospora longissima and
coarse- and fine-mesh bags was higher in the pool than Lunuluspora curvula contributed ;75% of conidia
in the riffle. After 24 and 31 d of immersion, leaf litter produced in the coarse- and fine-mesh bags in the
mass loss was higher in the riffle than in the pool. riffle, whereas Filosporella aquatica and Lemonniera
Decomposition rates (k) ranged between 0.0585 terrestris contributed ;65 and ;75%, respectively, of
(fine mesh in the riffle) and 0.0313 (coarse mesh in the the conidia produced in the coarse- and fine-mesh
pool; Table 2), and differed significantly among
TABLE 3. Results of 2-way analysis of variance for the
treatments (ANCOVA; p , 0.001). Decomposition rate
effects of habitat (riffle vs pool) and litter bag mesh size
was significantly affected by habitat type and mesh (coarse vs fine) on Ficus sp. leaf-litter decomposition rate (k).
size, but not by their interaction (ANOVA; Table 3).
Habitat type accounted for 20.9% and mesh size Sum of Mean
accounted for 2.8% of the total variance. Decomposi- Source df squares square F-ratio p-value
tion rates did not differ between coarse- and fine-mesh Main effects
bags in the pool or between coarse- and fine-mesh Habitat 1 0.0057 0.0057 19.06 ,0.0001
bags in the riffle (SNK; Table 2, Fig. 2). Decomposition Mesh size 1 0.0015 0.0015 5.01 0.0285
rates did not differ significantly between coarse-mesh Mesh size 3 habitat 1 0.0001 0.0001 0.32 0.5719
Residual 67 0.0201 0.0003
bags in the riffle and fine-mesh bags in the pool (SNK; Total (corrected) 70 0.0274
Table 2, Fig. 2). Decomposition rates were significantly
420 J. RINCÓN AND R. SANTELLOCO [Volume 28

FIG. 2. Mean (61 SE; n ¼ 3) leaf decomposition rates (k)

of Ficus sp. leaves during decomposition in fine- and coarse-
mesh litter bags in the riffle and pool of Carichuano Creek,
Venezuela. Treatments with the same letter were not
significantly different on the basis of Student–Newman–
Keuls (SNK) multiple comparison method (a ¼ 0.05).

bags in the pool (Table 5). Species richness was higher

in coarse- (16) and fine-mesh (14) bags in the riffle than
in coarse- (9) and fine-mesh (5) bags in the pool.
Conidia of Anguillospora crassa, Anguillospora gigantea,
Heliscus tentaculus, Triscelophorus acuminatus, Campylo-
spora filicladia, Flagellospora penicillioides, Flagellospora
FIG. 3. Mean (range) rate of conidia production on Ficus
curvula, Helicomyces sp., and 3 unidentified species sp. leaves during decomposition in fine- and coarse-mesh
were restricted to riffle samples, whereas L. terrestris litter bags in the riffle (A) and pool (B) of Carichuano Creek,
and 1 unidentified species were present only in pool Venezuela.
samples.
Differences in conidial composition between coarse- right and above samples from fine-mesh bags. The
and fine-mesh bags were small in both the riffle and species that contributed most to the separation of pool
the pool (Table 5). In the riffle, A. longissima and
and riffle samples were A. longissima (22%), Lunulu-
Lunuluspora curvula dominated both coarse- and fine-
spora curvula (16.5%), F. aquatica (12.2%), and Pyramido-
mesh bags, although some additional less-abundant
spora fluminea (11.9%).
species (,1% of total conidial production) were
Assemblage composition did not appear to vary
present (Pyramidospora casuarinae, T. acuminatus, F.
temporally in the pool samples, but assemblage
curvula, and 1 unidentified species). In the pool,
Filosporella aquatica and Lemonniera terrestris dominated composition in riffle samples showed a temporal trend
both coarse- and fine-mesh bags. Three species on axis 2 (Fig. 4). Two assemblages were defined for
(Campylospora chaetocladia, Lunuluspora cymbiformis,
and P. casuarinae) were found in coarse-mesh bags in TABLE 4. Results of 2-way analysis of variance for the
effects of habitat (riffle vs pool) and litter bag mesh size
the pool.
(coarse vs fine) on conidia production during decomposition
of Ficus sp. in Carichuano Creek.
Spatial variation of aquatic fungal communities
Sum of Mean
NMDS ordination of fungal samples indicated
Source df squares square F-ratio p-value
separation of pool and riffle assemblages on axis 1
(Fig. 4). Only 4 samples (2 replicates from day 2 coarse- Main effects
mesh bags, 1 replicate from day 24 coarse-mesh bags, Habitat 1 2.05 2.05 13.09 0.0010
Mesh size 1 0.64 0.64 4.12 0.0508
and 1 replicate from day 7 fine-mesh bags) from the
Mesh size 3 habitat 1 0.07 0.07 0.44 0.5126
pool grouped with samples from the riffle. No riffle Residual 32 5.01 0.16
samples grouped with pool samples. In the riffle, Total (corrected) 35 7.56
samples from coarse-mesh bags tended to group to the
2009] AQUATIC FUNGI IN NEOTROPICAL STREAMS 421

TABLE 5. Relative abundance (%) of identified and unidentified taxa of aquatic hyphomycetes sporulating on Ficus leaves in
coarse- and fine-mesh bags incubated in the riffle and the pool of Carichuano Creek, Venezuela.

Riffle Pool

Taxon Coarse mesh Fine mesh Coarse mesh Fine mesh

Anguillospora crassa Ingold 2.0 1.9 — —
Anguillospora gigantea Ranzoni 1.0 0.9 — —
Anguillospora longissima Ingold 47.1 54.8 7.7 2.1
Campylospora chaetocladia Ranzoni 3.3 1.4 0.6 —
Campylospora filicladia Nawawi 1.6 — — —
Clavariopsis azlanii Nawawi 1.3 1.4 — —
Filosporella aquatica Nawawi 6.5 1.4 36.9 37.5
Flagellospora curvula Ingold 1.0 — — —
Flagellospora penicillioides Ingold 1.3 2.9 — —
Heliscus tentaculus Umphlett 0.3 0.5 — —
Helicomyces sp. — 0.5 — —
Lemonniera terrestris Tubaki — — 26.2 37.5
Lunulospora curvula Ingold 28.4 22.4 12.5 8.3
Lunulospora cymbiformis Miura 2.0 1.4 3.6 —
Pyramidospora casuarinae Nilsson 0.3 — 1.8 —
Pyramidospora fluminea Miura and Kudo 3.3 9.1 7.7 14.6
Triscelophorus acuminatus Nawawi 0.3 — — —
Unidentified sp.1 — — 1.8 —
Unidentified sp. 2 0.3 — —
Unidentified sp. 3 — 0.5 — —
Unidentified sp. 4 — 0.5 — —
Total number of species 16 14 9 5

riffle samples: early (days 2–14) and late (days 24–31).
Early assemblages often were dominated by A. longis-
sima, L. curvula, and F. aquatica, whereas late assem-
blages often were dominated by P. casuarinae, P.
fluminea, and unidentified sp. 4.
FIG. 4. Nonmetric multidimensional scaling ordination
plot for fungal assemblage composition on Ficus sp. leaves
during decomposition in coarse- and fine-mesh bags placed
in a pool and riffle in Carichuano Creek, Venezuela. Samples
were collected on days 2, 7, 14, 24, and 31 after deployment
of the bags. Points within ellipses labeled Pool or Riffle
represent samples from those respective habitats unless
otherwise labeled.
Discussion
Leaf decomposition
Ficus leaves decomposed rapidly in both riffle and
pool habitats of this intermittent stream. Decomposi-
tion rates were comparable with those reported for
Ficus benghalensis in southern India (Raviraja et al.
1998), Ficus insipida in Costa Rica (Rosemond et al.
1998), Croton in the Colombian Andes (Mathuriau and
Chauvet 2002), and Hura crepitans in mid-north
Venezuela (Abelho et al. 2005). In contrast, Protium
brasiliense leaves in a tropical Cerrado stream in Brazil
(Gonçalves et al. 2007) and 3 species (Myrsine
guianensis, Cupania latifolia, and Nectandra lineatifolia)
in a headwater stream of Colombian Andes (Chara et
al. 2007) had slower decomposition rates than were
observed for Ficus in our study. Ficus decomposition
rates in our study were considerably higher than those
of Anacardium excelsum in the same stream (Rincón et
al. 2005). This difference might be a consequence of
differences in the chemical composition of Ficus and
Anacardium leaves (Rincón and Martinez 2006). Ana-
cardium leaves have high lignin and total polyphenols,
whereas Ficus leaves have low lignin and polyphenols.
Content of lignin and total polyphenols are good
predictors of leaf litter decomposition rates (Campbell
and Fuchshuber 1995, Ostrofsky 1997, Gessner and
Chauvet 1994).
422 J. RINCÓN
Decomposition rates were not affected by mesh size
within a habitat. This result indicates that macroin-
vertebrates, particularly shredders, played a minor
role in decomposition, as has been reported for some
tropical streams (Mathuriau and Chauvet 2002, Want-
zen and Wagner 2006, Gonçalves et al. 2006, 2007). In
contrast, shredders have been associated with differ-
ences in decomposition rates of leaves in bags with
different mesh sizes in other tropical streams (Cheshire
et al. 2005, Chara et al. 2007). Scarcity of shredders
might be a biogeographical rather than a latitudinal
issue (Cheshire et al. 2005). The high abundance of
shredders in Colombian Andean streams (Chara et al.
2007) contrasts with the low abundance of shredders
in lowland tropical streams (Dobson et al. 2002,
Mathuriau and Chauvet 2002, Wantzen and Wagner
2006).
Differences in current velocity between pool and
riffle habitats might partially explain the differences in
decomposition rates in our study. Current velocity in
riffles can accelerate decomposition by increasing
physical fragmentation, improving the transport and
colonization of fungal species, and supplying dis-
solved O2 and nutrients to microbial communities.
Differences in sporulation activity between habitats
appear to support this idea. Rincón et al. (2005)
attributed differences in decomposition rates of A.
excelsum between pools and riffles in Carichuano
Creek to current velocity. Langhans et al. (2008)
suggested that current velocity promoted faster leach-
ing and fragmentation and, hence, faster leaf litter
decomposition in river channels than in ponds in the
floodplain of the Tagliamento River (Italy). In contrast,
decomposition rates did not differ significantly be-
tween main stem and floodplain ponds of the Garonne
River system in France, and fungal production was
similar between the 2 sites (Baldy et al. 2002). Baldy et
al. (2002) suggested that fungi might be important
agents of litter decomposition in standing water bodies
of fluvial systems when the physicochemical condi-
tions are favorable to aquatic hyphomycetes.
Sporulation rates and activity of aquatic hyphomycetes
Habitat type influenced fungal activity, assemblage
composition, and decomposition rates of Ficus sp. in
Carichuano Creek. Sporulation rates in our study were
within the range reported in other tropical and
temperate studies (Mathuriau and Chauvet 2002, Gulis
and Suberkropp 2003, Rincón et al. 2005, Ferreira and
Graça 2006). Early sporulation rates similar to those in
our study have been reported for other fast-decom-
posing leaf species in tropical (Mathuriau and Chauvet
2002) and temperate streams (Gessner et al. 1993,
AND R. SANTELLOCO [Volume 28
Gessner 1997). This result is consistent with the strong
relationship between the maximum sporulation rate
and the decay coefficients for 7 deciduous leaf species
(Gessner and Chauvet 1994).
In our study, current velocity and turbulent condi-
tions in riffles provided favorable conditions for
aquatic hyphomycetes activity. Sporulation rate on
Ficus leaves was greater in the riffle than in the pool
habitat, a result similar to that reported for sporulation
rates on A. excelsum leaves in Carichuano Creek
(Rincón et al. 2005). In a fluvial system in France,
sporulation activity was highest in the main channel
and very low in the floodplain pond (Baldy et al. 2002).
Fungal biomass was highest in the river channel and
lowest in ponds of a riverine floodplain in Italy
(Langhans et al. 2008). In addition, Maamri et al.
(1999) reported a lower fungal biomass in a pool than a
riffle during leaf decay in an intermittent stream in
North Africa. Fungal activity and biomass might be
limited by low turbulence and oxygenation in pools
(Bärlocher 1985, Chamier 1987, Suberkropp 1992,
Abdel-Raheem 1997).
Fungal assemblages
The fungal assemblage on Ficus leaves in Carichua-
no Creek was similar to that found in other tropical
studies (Santos-Flores and Betancourt-López 1997,
Mathuriau and Chauvet 2002, Rincón et al. 2005).
Species richness in our study (21 species) was nearly
23 greater than previous reports of aquatic hyphomy-
cetes for Venezuela (11 species; Nilsson 1962), but only
;½ of the number (50 species) reported by Smits et al.
(2007) from 7 streams in Venezuela. All species
identified in our study were included in a recent
compilation made for South America (Schoenlein-
Crusius and Piccolo 2003). However, the fact that we
were unable to identify 4 species with available keys
indicates the need for more surveys and taxonomical
studies to improve our knowledge of neotropical
fungi.
Sporulating fungal assemblages differed between
riffle and pool habitat. Higher species richness of
conidia in the riffle than in the pool suggests that
conditions for reproduction of aquatic hyphomycetes
are better in riffles than in pools. Fungal assemblages
in the riffle were dominated by A. longissima and L.
curvula. Lunuluspora curvula is a widespread species
commonly found during the summer in temperate
streams (Webster and Descals 1981). This species also
is commonly reported as a dominant species of
tropical (Justiniano and Betancourt 1989, Mathuriau
and Chauvet 2002) and subtropical streams (Abdel-
Raheem 1997). Anguillospora longissima is a widespread
2009] AQUATIC FUNGI
species commonly found in tropical (Schoenlein-
Crusius and Piccolo 2003) and temperate (Webster
and Descals 1981) streams. Different species (Filospo-
rella aquatica and Lemonniera terrestris) were dominant
in the pool, a result that might indicate habitat
preferences by the aquatic fungi. Filosporella aquatica
and L. terrestris have a typical tropical distribution but
also can be found in fungal assemblages in temperate
lowland rivers (Marvanová 1997, Santos-Flores and
Betancourt-López 1997).
Successional patterns in fungal development, such
as the pattern observed in the riffle in our study, have
been reported previously, but to our knowledge, ours
is the first study to assess temporal dynamic patterns
in fungal assemblage composition in a neotropical
stream. Fungal succession moved through 3 stages and
was coupled with the dynamics of leaf mass loss in a
temperate stream in France (Gessner et al. 1993). In our
study, Lunuluspora curvula, A. longissima, and F.
aquatica were associated with the early stage of
colonization, whereas P. casuarinae, P. fluminea, and
unidentified sp. 4 were associated with the advanced
stages of decay. Changes in nutritional and chemical
values of leaves might promote changes in species
assemblages during leaf decomposition (Arsuffi and
Suberkropp 1984).
Changes in water temperature can influence tempo-
ral changes in fungal assemblages in subtropical
streams (Abdel-Raheem 1997). However, water tem-
perature was constant during our study, and thus, was
unlikely to have influenced fungal assemblage com-
position in Carichuano Creek. In our study, stream
flow varied considerably during the experiment, and
environmental factors associated with changes in
hydrological conditions (e.g., stream velocity, width,
and depth) could have affected fungal assemblages.
Gonçalves et al. (2006) attributed low microbial
colonization in Mediterranean streams to flow-related
conditions. In addition, Maamri et al. (1998, 1999,
2001) showed that fungal assemblages changed during
the flow transition from wet to dry period in an
intermittent stream in Morocco (North Africa). The
intermittent character of Carichuano Creek makes
hydrology a critical driver of physical, chemical, and
biological processes (Rincón and Cressa 2000). A better
understanding is needed of the responses of fungal
communities to hydrological changes and their impli-
cations for decomposition rates.
Acknowledgements
Our work was funded by the Fondo Nacional de
Ciencia, Tecnologı́a e Innovación de la República
Bolivariana de Venezuela (FONACIT) (grant no. S1-
IN NEOTROPICAL STREAMS 423
2002000392). We thank Jeimmy Uzcátegui and Mario
Nava for field assistance. We also thank Luz Boyero
and Alonso Ramı́rez for their kind invitation to the
Tropical Session at the annual meeting of the North
American Benthological Society in Anchorage, Alaska,
USA. We thank Felix Bärlocher for his comments on
earlier versions of the manuscript. Several anonymous
referees provided useful comments on earlier versions
of the manuscript, which are much appreciated.
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Received: 23 July 2007
Accepted: 17 November 2008