http://www.icn.unal.edu.co/
Caldasia 36(2):xx-xx.
204
Román-P. et
al.
TROPHIC AND REPRODUCTIVE ECOLOGY OF A
NEOTROPICAL CHARACID FISH HEMIBRYCON
BREVISPINI (TELEOSTEI: CHARACIFORMES)
Ecología tróica y reproductiva del pez carácido neotropical
Hemibrycon brevispini (Teleostei: Characiformes)
CRISTIAN ROMáN-P.
Universidad del Valle, Facultad de Ciencias Exactas y Naturales, Departamento de Biología,
Cali, Colombia. cromanpa94@gmail.com
CéSAR ROMáN-VALENCIA
Universidad del Quindío, Laboratorio de Ictiología, Apartado 2639, Armenia, Quindío,
Colombia. ceroman@uniquindio.edu.co
DONALD C. TAPHORN
1822 North Charles Street, Belleville, Illinois, 62221, USA. taphorn@gmail.com
ABSTRACT
Hemibrycon brevispini is a Neotropical characid ish endemic in La Venada Creek,
a headwater tributary of the Quindío River of Colombia (Cauca River drainage). It is
mainly a diurnal insectivore with a diet dominated by benthic dipterans (Chironomidae,
Simuliidae, Psychodidae, Culicidae, Calliphoridae, Dixidae and Muscidae),
hymenopterans (Formicidae and Vespidae) and ephemeropterans (Baetidae), as
well as allochthonous prey and items eaten accidentally. Microhabitats of mountain
streams with lower water velocity tend to have more riparian vegetation and the
associated terrestrial arthropods that are consumed by H. brevispini. It has three peaks
in reproduction: December, April and August. Average fecundity was 776 mature
oocytes per female.
Key words. Diet, reproduction, conservation, Neotropical ishes, natural history.
RESUMEN
Hemibrycon brevispini es un pez carácido neotropical endémico de la quebrada
La Venada, un aluente del río Quindío en Colombia (cuenca del río Cauca). Esta
especie es predominantemente insectívora diurna, con una dieta dominada por
dípteros (Chironomidae, Simuliidae, Psychodidae, Culicidae, Calliphoridae, Dixidae
y Muscidae), himenópteros (Formicidae y Vespidae) y efemerópteros (Baetidae),
además de presas de origen alóctono y otras deinidas como consumo accidental. Los
hábitats con baja velocidad de agua sustentan mayor vegetación ribereña asociada a
artrópodos terrestres, consumidos por H. brevispini. Su reproducción tiene tres picos:
diciembre, abril y agosto. Su fecundidad promedio es de 776 oocitos por hembra.
Palabras clave. Dieta, reproducción, conservación, peces neotropicales, historia
natural.
Trophic and reproductive ecology of Hemibrycon brevispini
INTRODUCTION
The genus Hemibrycon consists of fishes
characterized by the presence of more
than four teeth on the maxilla (in adults)
(Eigenmann 1927, Román-Valencia et al.
2013). A phylogenetic analysis of Hemibrycon
determined its monophyly based on four
synapomorphies: ectopterygoids with widened
ventral anterior projection, four to six times
wider than posterior part; a red spot present
in life on ventral margin of caudal peduncle;
a postero-ventral projection on the pterotic
and irst infraorbitals gradually decreasing
in width from posterior tip and located near
posterior part of antorbital (Arcila-Mesa 2008).
Hemibrycon brevispini Román-Valencia &
Arcila-Mesa was described from La Venada
and Quebrada Negra Creeks tributaries of the
Santo Domingo River, Quindío River basin,
upper Cauca, Andes of Colombia (RománValencia & Arcila-Mesa 2009).
Fourteen species of Hemibrycon have been
described from the Cauca-Magdalena River
Basin in Colombia, but there are few studies of
their ecology that provide baseline information
to determine their conservation status or
provide guidelines for the management of
many species that have relatively small
geographical ranges and populations. In fact,
habitat, extensive diet and reproductive data
are only available for two Hemibrycon species
from the Magdalena-Cauca River Basin: H.
boquiae (Román-Valencia et al. 2008) and H.
quindos (Román-Valencia & Botero 2006),
but short notes have been published about
of H. brevispini (Román-Valencia & ArcilaMesa 2009), H. antioquiae, H. fasciatus and
H. cardalensis (Román-Valencia et al. 2013),
H. cairoense (Román-Valencia & Arcila-Mesa
2009), H. paez, H. raqueliae, H. virolinica and
H. yacopiae (Román-Valencia & Arcila-Mesa
2010), H. palomae (Román-Valencia et al.
200) and H rafaelense (Román-Valencia &
Arcila-Mesa 2008). H. brevispini is endemic
to La Venada and La Negra Creeks that are
2
both tributaries of the Quindío River, in the
upper Cauca River drainage (Román-Valencia
& Arcila-Mesa 2009). Aspects of H. brevispini
diet, reproduction and habitat were analyzed
order to provide baseline information useful
for conservation and management efforts
of this endemic species and its habitat,
especially considering the increasing impacts
of hydropower and mining development in the
Colombian Andes.
MATERIALS AND METHODS
Data collection and study area description.
Sampling sites are distributed along the entire
length of La Venada Creek, from its origin to
its mouth where it discharges in to Quebrada
Negra Creek. Fishes were collected in the
middle and lower reaches of La Venada Creek
(4o 26’ 47.4” N & 75 o 40’ 44.3” W, 1661
m.a.s.l. and 4o 26’ 54.9” N & 75o 40’ 48.8”
W, 1307 m.a.s.l.), a tributary of Quebrada
La Negra Creek, which in turn is a tributary
of the Santo Domingo/Quindío/upper Cauca
River system in the Andes of Colombia. Thus,
La Venada Creek is a primary or secondary
stream, in the hierarchical classification
system of Allan (1995). Fish were captured
on two days of each month from July 20 to
November 2012, using a 2 x 0.5 m seine net,
with 5 mm mesh and a 2 m cast net with a 10
mm mesh between 0900 and 1300 hr, sampling
much of the creek. This period included dry
seasons (June-August and January-February)
and wet seasons (March-May and SeptemberDecember) (Fig. 1).
A total of 122 specimens of H. brevispini were
collected and placed on ice to decrease the rate
of enzymatic digestion of stomach contents,
as recommended by Bowen (1996). Samples
were dissected the same day as collection at
the Ichthyology Laboratory of the Universidad
of Quindío, Armenia, Colombia (IUQ). An
incision was made along the ventrum, and the
stomach, intestine and gonads were extracted.
After dissection, specimens were ixed in
Román-P. et al.
formalin (10%) for 15 days and then in 70%
ethanol and deposited in the ish collection
(IUQ). Gonads were weighed using an
analytical balance (Adventurer-Ohaus H226)
with 0.000 g precision and subsequently
preserved in 70% ethanol. Mitutoyo digital
calipers with 0.0 mm precision were used to
measure standard and total length of the ish,
stomach length, stomach width and intestine
length.
contents were analyzed using numeric and
frequency methods (Hyslop 1980, Hynes
1950) and a volumetric method (Capitoli,
1992, Pedley & Jones 1978). The Index of
Relative Importance (IRI) (Oda & Parrish,
1981), proposed by Pinkas et al. (1971) was
used to determine the importance of each
food item.
Habitat measurements. A pH meter (Hanna
HI 921ON) was used to measure pH, air
temperature and water temperature. A
digital oxygen meter (OX1196) was used to
measure dissolved oxygen and saturation.
Geographical coordinates were recorded from
a GPS unit (Garmin eTrex 10).
Where, %V= percent volume, % Fo= observed
frequency percent and %N= proportion of
food type.
Diet. Prey items in ish stomach contents
were identified to the lowest taxonomic
resolution possible (order, family, genus)
(Borror et al. 1992, Roldàn 1996). Stomach
IRI = %Fo (%N + %V)
Prey item identification was done for
individuals in different states of digestion
or based on diagnostic structures, however
unidentiied remains or parts of organisms
were not treated as items; these are listed, but
not included in the statistical diet analyses. The
emptiness coeficient (V) (Hyslop 1980) was
also calculated to reveal the months included
Figure 1. Multi-year monthly rainfall 1998-2005, station La Bella, Calarcá, Quindio, Colombia.
Data of Cenicafé.
3
Trophic and reproductive ecology of Hemibrycon brevispini
on the feeding period of the species,
V=
n x 00
N
Where, n= number of empty stomachs, N=
total number of stomachs examined.
A centered principal component analysis was
made using stomach contents abundances
(%N) and the proportion of the total volume
of each prey item (called %PCA after Billy et
al. [2000]). In this method, total row (of diet
items encountered in an individual stomach)
is equal to 1. For the analysis, the families
were grouped within orders to enable a better
explanation of the diet. This analysis reveals
the pattern of dispersion of prey found in
stomachs. Representation on two axes (Gabriel
1981) was done with the irst factorial plane;
the position of each item is equivalent to the
position of a stomach containing 00% of the
prey species. Each stomach is at the centroid
of the prey items, with each prey species
being given a weight equal to its proportion
in the stomach. Computations and graphical
displays were performed with the ADE-4
package (Thioulouse et al. 1997) running in Rsoftware (R Development Core Team 2013).
Correlation analysis between the %PCA
scores and standard length was performed to
examine patterns of prey consumption among
ishes of different sizes.
Abundance distribution normality of food
items was evaluated using the KolmogorovSmirnov test with a 5% of signiicance (α=
0.05). Based on this a Kruskal-Wallis was
done to evaluate differences in prey abundance
(numbers of individuals) among seasons, sex
and maturation stage. A correlation analysis
based on log-transformed data was applied to
examine relationships among the variables:
standard length (SL), total length (TL),
intestine length (IL) it is in mm, stomach
width (STW) in g, stomach length (STL) in
mm, stomach weight (STWE) in g, gonad
4
weight (GW) in g, fecundity (FE) in number of
oocytes and total weight (TWE) in g. The Past
2.11 (Hammer et al. 200) and R-software
(R Development Core Team 2013) programs
were used for statistical analyses.
ANOSIM was used to test dietary preferences
among food items. The similarity matrix
was generated with the transformed data
(log[x+]) for consumed prey using the
Bray-Curtis similarity and the observed
relationships were compared based on 9999
permutations. The value of R lies between -
and 1, where 0 indicates that low and high
similarities are perfectly mixed, thus there is
no preference in dietary items.
Condition factor (K). A condition factor
(K) was calculated and used to evaluate
the population’s condition (Wootton 1992,
Vazzoler 1996, Bagenal & Tesch 1978),
K = Wt x 00
Lsb
Where, Wt (g) = total weight, Ls (mm) = standard length and b, la relation length weight.
Reproduction. To determine the reproductive
season, temporal variation in the gonadosomatic index (GSI) was evaluated. GSI was
equal to
GSI = Wo x 00
Wc
where Wc= Wt-Wo, and Wo (g) = gonad
weight, Wt (g) = total weight, and Wc (g)
= body weight (Vazzoler 1996). Spawning
seasons were identiied as peaks in mean GSI.
Size at sexual maturity was determined using
the graphic method of Sokal & Rohlf (1995)
that identiies the maturation size as that for
which 50% of the population is reproducing.
Sex ratio was evaluated using chi-squared (X2)
and proportion of males and females.
Román-P. et al.
Fecundity. Fecundity was determined using
the dry subsample method (Ricker, 1971),
and absolute fecundity (Fa) was calculated
using only mature females according to the
formula
Fa =
∑ no
No
Where no= number of oocytes per female, and
No= total number of females. Oocyte diameter
was measured using millimetric graph paper
by counting the number of oocytes itting into
0 mm of the line on the paper and dividing by
10, and later calculating the average number
of oocytes on a one-dimensional space for
each ovary with oocytes.
RESULTS
Habitat. La Venada Creek is a primary
stream in both its highest, and lower sections,
with a width of 2-3 m, and a depth 0.5-1
m during both the rainy and dry seasons.
Substrate is mostly rocky, with some sand
and decomposing vegetation. For most of the
length studied shore vegetation is not natural,
consisting of white ginger (Hedychium
coronarium), bamboo (Guadua angustifolia),
coffee trees (Coffea arabica) and banana
plantations (Musa spp.).
Ambient and surface water temperature
ranged on average from 18˚C to 20.0°C
during low-water season, and from 16.2°C
to 20.5°C during the rainy season. Oxygen
saturation was around 84% and dissolved
oxygen was 6.0mg/L during the dry season,
but values were in average higher during the
rainy season: 92% and 7.9mg/L respectively.
In the dry season pH was near 7.6 and during
the wet season 7.1 (Table 1).
Hemibrycon brevispini was found in the middle
and lower reaches of La Venada Creek along
with Carlastyanax aurocaudatus (Eigenmann
1913), Astroblepus cf. cyclopus Humboldt
1805, Brycon henni Eigenmann 1913,
Bryconamericus caucanus Eigenmann 1913,
Cetopsorhamdia boquillae Eigenmann 1922,
Chaetostoma cf. ischeri Steindachner 1879,
Parodon caliensis Boulenger 1895, Poecilia
caucana Steindachner 1880, Trichomycterus
caliense Eigenmann 1918 and T. chapmani
Eigenmann 1918.
Digestive tract morphology. The stomach of
H. brevispini is longer (mean=12.7 mm, S.D.
= 2.79) than wide (mean 7 mm, S.D. = 1.95)
and is located in the anterior portion of the
coelomic cavity, sometimes thickly covered
with fat. Two pyloric caecae are present on
the anterior part of the stomach. A signiicant,
Table 1. Physico-chemical variables during wet and dry season for La Venada Creek, Quindío
River drainage, upper Cauca River, Colombia. July 2011 - November 2012. AT: Ambient
temperature, WT: Water temperature, DO: Dissolved oxygen, PS: Percent saturation, CO:
Conductivity.
AT (°C)
WT (°C)
DO (mg/L)
PS (%)
CO (us)
pH
Range
6.00-20.00
18.50-21.56
5.00-7.03
83.00-85.00
51.00-62.00
7.01-8.2
Dry season
Mean
18.00
20.03
6.0
84.00
56.50
7.60
V (%)
15.71
10.80
23.86
1.68
13.76
.06
Range
14.00-18.46
18.00-23.00
7.62-8.20
90.00-94.00
67.00-79.00
5.85-8.40
Wet season
Mean
6.23
20.50
7.91
92.00
73.00
7.125
V (%)
19.43
17.24
5.18
3.07
.62
25.30
5
Trophic and reproductive ecology of Hemibrycon brevispini
positive correlation was found between the
intestine length and total length (r=0.67;
p=0.017) as well as the intestine length and
standard length (r=0.67; p=0,017; Fig. 6).
Diet. The feeding activity of this species
is constant throughout the year (emptiness
coeficient [V] = 0.82%). Prey abundance
did not show normal distribution (p>0.05).
The Kruskal-Wallis analysis revealed no
difference in the abundance of the diet items
between males vs. females or immature vs.
adults (KW P= 0.25; df= 1; KW P= 0.31, df=
respectively). This was also the case when
diet abundances were compared between wet
and dry seasons (KW P= 0.98; df= 1). Thus
season, sex and maturity are not determinant
factors of the prey abundance in the diet of H.
brevispini, and hence it appears that the items
found in its diet are in constant supply all year
round. Stones, feathers and nematodes found
in the digestive tract of some individuals were
considered occasional and accidental.
Stomach contents analysis revealed 4 total
prey categories consumed by H. brevispini
(Table 2, Fig. 2), with Diptera being the most
frequently consumed (%N= 11.44; %FO=
16.96; %V= 15.01; IRI= 448.39), followed
by Hymenoptera: Formicidae (%N= 15.84;
%FO= 13.84; %V= 13.59; IRI= 407.21)
and Ephemeroptera: Baetidae (%N= 16.09;
%FO= 9.86; %V= 13.44; IRI= 291.19).
Occasionally ingested organisms (classiied
as such based on their low consumption
frequency) included Coleoptera, Araneae,
Dyctioptera, and other allochthonous
material. The degree of digestion of items
found in stomach contents and the hour
of capture allow us to infer that feeding
is diurnal in H. brevispini, when they take
mostly benthic organisms, some arthropods
from the water column or that have fallen
into the water from shoreline vegetation.
If this species were a nocturnal feeder, the
samples made during the day would not
6
have found identiiable stomach contents
that for the most part showed little effects
of digestion (soft tissue still present) and
indication that prey ingestion had occurred
shortly before capture.
In the normalized principal component
analysis (%PCA) (Fig. 2) of prey item
abundances components one (17.18%),
two (13.62%) and three (9.33%) explain
only 40.4% of total variance. This is a
consequence of the elevated heterogeneity
of volume of prey consumed and large
number of different prey items consumed
by different individuals of H. brevispini.
The principal component analysis recovered
only a small percentage of this variability,
leaving 59.06% of the variance unexplained.
Thus, only a preliminary approximation of
the trophic characteristics of this species
is possible based on PCA, but it is evident
that widely different amounts of a wide
variety of prey ítems are eaten. The items
Coleoptera, bird feathers, gravel, Diptera,
Nematoda and Hemiptera accounted for
much of the variation in individual diets,
whereas Ephemeroptera, Hymenoptera and
others items accounted for little variation and
were distributed near the sample centroid.
Diptera were very abundant in only a few
stomachs and, overall, were not as common
as the other items located near the centroid.
Most of the individual stomachs in Figure
grouped near the origin are indicating
that individuals employ a foraging strategy
that exploits both dominant and rare prey
items. The analysis indicates a broad trophic
niche and relatively low between-individual
variation in diets. Correlation analysis did
not reveal an association between the irst
axis scores (% PCA1) and ish size (r=0.12,
p=0.615), indicating no prey preferences in
relation to ish length. ANOSIM revealed
non-signiicant differences between the items
found among individuals (R= -0.45, P=1).
Román-P. et al.
Table 2. Diet of Hemibrycon brevispini in La Venada Creek, upper Cauca, Colombia. %N=
numerical percent, %FO= observed frequency percent, %V= percent volume %I.A. = index
of alimentary importance, All: allochthonous, Auto: autochthonous. Only the main items are
identiied in the table.
Item
Hymenoptera
Formicidae
Vespidae
Diptera
Chironomidae
Simuliidae
Psychodidae
Ceratopogonidae
Calliphoridae
Culicidae
Dixidae
Muscidae
Hemiptera
Heteroptera
Auchenorrhyncha
Coleoptera
Hydrophilidae
Ptilodactylidae
Lampyridae
Gyrinidae
Chrysomelidae
Ephemeroptera: Baetidae
Odonata
Anisoptera
Zygoptera
Trichoptera
Hydropsychidae
Helicopsychidae
Hydrobiosidae
Lepidoptera
Miriapoda: Diplopoda
Crustacea: Isopoda
Arachnida: Araneae
Nematoda (Parasite)
Gastropoda
Vegetal Material
Seeds
Vegetative tissue
Pteridophyta
Cyanophyceae (Oscillatoria sp.)
Feather
Oocytes
Scales
Dyctioptera: Ootheca
Rocks
Stage
Adult
Adult
Larvae
Larvae
Larvae
Larvae
Larvae
Larvae
Larvae
Larvae
Adult
Adult
Adult
Adult
Adult
Adult
Larvae
Adult
Adult
Nymph
Nymph
Nymph
Larvae
Larvae
Larvae
Adult
Adult
Adult
Adult
-
Origin
All
All
Auto
Auto
Auto
Auto
Auto
Auto
Auto
All
All
All
Auto
Auto
All
Auto
Auto
All
Auto
Auto
Auto
Auto
Auto
Auto
All
All
Auto
All
Auto
Auto
All
All
All
Auto
All
Auto
Auto
All
Auto
%N
15.83
0.
11.43
0.05
0.28
0.
1.29
0.02
.0
0.59
0.08
0.50
0.08
2.47
0.
0.22
0.05
0.02
0.05
16.08
0.42
0.
1.97
0.05
0.
0.19
0.19
0.4
0.05
0.33
0.02
2.08
0.6
0.02
0.4
0.25
0.02
0.02
0.87
%V
13.58
0.24
15.00
0.
0.44
0.32
1.76
0.03
1.29
.02
0.22
0.66
0.4
3.06
0.26
0.55
0.06
0.04
0.39
13.43
1.07
0.53
6.3
0.3
0.52
0.58
0.63
0.39
0.09
1.45
0.24
.30
0.88
0.04
.66
0.29
0.6
0.06
0.06
0.57
%FO
13.84
0.51
16.95
0.17
.03
0.69
1.73
0.17
2.42
.2
0.51
1.73
0.34
6.05
0.69
.03
0.17
0.17
0.17
9.86
1.73
0.69
6.74
0.17
0.69
.03
0.86
0.86
0.34
1.38
0.17
.03
0.86
0.17
.03
0.69
0.34
0.178
0.17
2.42
IRI
407.21
0.18
448.39
0.029
0.75
0.30
5.29
0.0
5.58
1.96
0.6
2.03
0.08
33.60
0.25
0.81
0.02
0.0
0.07
291.19
2.593
0.45
54.73
0.032
0.43
0.81
0.72
0.46
0.05
2.47
0.04
3.52
0.90
0.0
1.72
0.30
0.4
0.0
0.0
3.50
7
Trophic and reproductive ecology of Hemibrycon brevispini
Figure 2. Biplot of prey and stomachs obtained from a Centered Principal Component
Analysis (irst factorial plane) for Hemibrycon brevispini, in La Venada Creek, Quindío River.
Dots represent stomachs. Where, Hym: Hymenoptera, Dip: Diptera, Hemi: Hemiptera, Col:
Coleoptera, Eph: Ephemeroptera, Odo: Odonata, Trich: Trichoptera, Lepi: Lepidoptera, Miri:
Miriapoda, Crus: Crustacea, Arac: Aracnae, Nem: Nematoda, Gas: Gastropoda. Plan: Plantae,
Plu: Feather, Ooc: Oocyte, Esc: Escale, Dyc: Dyctioptera, Ro: Rocks.
Condition factor (K). In adults (female),
the lowest K values were obtained from July
2011 (dry season), November-December 2011
and October 2012, both of which coincide
with the wet season when fewer prey items
were found in stomachs; low K values were
obtained for adult males in September 20
(wet season), February 2012 and July 2012
both of which coincide with the dry season.
8
Maximum K values were obtained for females
in October 2011, February and June 2012,
during the wet and dry seasons, respectively;
maximum K values for males were observed
in July 2011 (dry season), October 2011
and 2012, both during the wet season. The
lowest and maximum values in condition
factor K contrast with the gonadosomatic
index value GSI (Fig. 4) and cannot be
Román-P. et al.
interpreted as related to gonadal development.
In immature specimens, variation was
observed in condition factor values with the
notable increase occurring between May and
November of 2012 and July 2011, which
corresponds to the wet and dry season; the
variation in K values is considerable in
both immature and adults. Comparison of
maximum and minimum values by sex for
both adults and immatures does not reveal
great differences, but lowest values are from
April 2012 and highest just one month later
in May of 2012.
Reproduction. The gonosomatic index (GSI)
(Fig. 4) showed high variation, up to one order
of magnitude, among months of this study.
Hemibrycon brevispini reproduction has three
peaks during the year, with two large peaks
in GSI in December 2011 and August 2012,
and another smaller peak in April 2012.The
two larger peaks coincide with the transition
from wet to dry season (December) and from
dry to wet season (August), and the smaller
peak (April) is during the rainy season (Fig.
1). Females had higher GSI values than males
throughout the year.
Figure 3. Condition factor (K) of Hemibrycon brevispini, in La Venada Creek,Quindío River
drainage, upper Cauca, Colombia. July 2011 - November 2012.
Figure 4. Mean gonosomatic index (GSI) for male and female Hemibrycon brevispini, in La
Venada Creek,Quindío River drainage, upper Cauca, Colombia. July 2011 - November 2012.
9
Trophic and reproductive ecology of Hemibrycon brevispini
The high number of males present in the
population is remarkable: 65.2% were males
and 34.7% of the population individuals
were females, giving a sex ratio of 1.9 with a
predominance of males during the entire study
period; signiicant differences exist as a result
(X2= 9.26, df= 1, p= 0.05). For females, the
size at sexual maturity is 76.3 mm SL and for
males 68.7 mm SL (Fig. 5). Moreover, the size
difference between the sexes is statistically
signiicant (KW, P< 0.05, df= 1).
Fecundity. Average fecundity was 776
oocytes, and the mean diameter of mature
oocytes was 0.85 mm (S.D. = 0.26). A nonsigniicant low correlation value was found
between fecundity and SL (r= 0.1, P> 0.05,
Fig. 6). The mean weight of an oocyte was
3.4 x 10-4 g (S.D. =1,37 x 10-6). Total body
weight was significantly and positively
correlated with gonad weight (r= 0.66, p=
0.019).
DISCUSSION
In streams of the upper Cauca River drainage,
two species of Hemibrycon feed heavily
on benthic insects such as Ephemeroptera,
Odonata, and Trichoptera (Román-Valencia
& Botero 2006, Román-Valencia et al. 2008),
and this was also found for H. brevispini.
Similar diets have been reported for other
characid genera of the upper Cauca River
drainage such as Creagrutus brevipinnis
(Román-Valencia 1998), Roeboides dayi
(Román-Valencia et al. 2003), Argopleura
magdalenensis (Román-Valencia & Perdomo
2004), Carlastyanax aurocaudatus (RománValencia & Ruiz 2005), and Bryconamericus
caucanus (Román-Valencia & Muñoz 2001a,
Román-Valencia et al. 2008). It is commonly
accepted that immature aquatic stages of
insects are an abundant alimentary resource
in Neotropical montane streams; however H.
brevispini also consumed large amounts of
ants (terrestrial Hymenoptera).
Figure 5. Size at sexual maturity distribution for male and female Hemibrycon brevispini, in
La Venada Creek,Quindío River drainage, upper Cauca, Colombia. July 2011 - November
202.
0
Román-P. et al.
Figure 6. Scatter plot matrix between morphometric variables. The lower panel corresponds
to the scatter plot. Upper panel indicates the correlation coeficient and the signiicance.
Abbreviations: standard length (SL), total length (TL), intestine length (IL), stomach width
(STW), stomach length (STL) and stomach weight (STWE), gonads weight (GW) and fecundity
(FE), stomach width (STW) and total weight (TWE).
Pools and other habitats with low current
velocities (0.23 to 0.67 m/s, mean= 0.35 m/s)
in La Venada Creek inhabited by H. brevispini
tended to have more riparian vegetation that
probably supports ants and other terrestrial
arthropods. Field observations of riparian
vegetation in La Venada Creek and other
similar Andean streams indicate that areas
with lower water velocity are often wider
than swift-water reaches and support denser
riparian vegetation that in turn offers refuge
and food for ishes.
Among allochthonous items found in their
diet (14 of 42 categories), ants (Hymenoptera:
Formicidae) were much more important than
other categories (e.g., Vespidae, Diptera:
Muscidae, Heteroptera, Auchenorrhynca,
Trophic and reproductive ecology of Hemibrycon brevispini
Chrysomelidae, Ptilodactylidae, Lepidoptera,
Miriapoda: Diplopoda, Arachnida: Araneae,
Seeds, Vegetative tissue, Feather, Pteridophyta,
Dyctioptera).
Similar to the observed diet of H. brevispini,
Román-Valencia et al. (2008) reported that
Hemibrycon boquiae consumed a large
proportion of Diptera larvae (Chironomidae,
Simuliidae, Tipulidae, Ceratopogonidae and
Muscidae): to evaluate why some items but
not others are found in stomach contents
it would be necessary to conduct prey
preference and relative abundance studies,
not stomach content occurrence as we present
here. However, the ecological characteristics
of Neotropical dipterans coincides with the
preponderance of exploitation of these prey
by H. brevispini. Diptera have pupae and
larvae with aquatic or semi-aquatic habitats,
in both running and quiet waters (Foote 1987,
Brown 2001, Merritt et al. 2003, Courtney
& Merritt 2008, Courtney et al. 2008).
Among Diptera, Nematoceran families
(especially Tipuloidea and Chironomoidea)
are a preponderant component of aquatic
communities, frequently eaten by primary
consumers. Armitage et al. (1995) mention
that Chironomidae are one of the most
abundant families present in freshwater
habitats, making them prone to capture.
We also found nematodes in H. brevispini,
and interpret from their intact state that they
are parasites and not prey; which coincides
with indings reported for Bryconamericus
caucanus (Román-Valencia & Muñoz
200a). The relationship of intestine and body
length is a general diet indicator. Herbivores
have relatively long intestines compared to
carnivores and omnivores (Wootton 1992,
Kramer & Bryant 1995). The relatively short
intestine length of H. brevispini indicates
carnivory. Hemibrycon brevispini consumed
mostly benthic arthropods (35 of the 45 food
categories), and plant material was rare in
stomachs.
2
The spawning period for H. brevispini differs
greatly from patterns reported for other
species in the genus. Hemibrycon boquiae
spawns from July to September, the transition
from dry to wet season (Román-Valencia
et al. 2008). Hemibrycon quindos spawns
from March to September in both wet and
dry seasons (Román-Valencia & Botero
2006) (Fig. 1). Variation observed in the
gonosomatic index of H. brevispini; could be
caused by a seasonal life history (Winemiller
1989, 1992) inluenced by a seasonal increase
in food availability.
The low abundance of females during the
sampling period could be due to bias imposed
by males in sex ratios, since according some
authors (Daiber 1977, Morse 1981, Petrie
1983, Clutton-Brock 1988, Goto et al. 1999,
Goto et al. 2000, Trivers 1972) this could be
a consequence of protandria, in which later
maturation of females and dominance by males
during juvenile life phases causes differential
early mortality. Temperature affects are often
cited as a principal cause of sex ration bias,
but, the differential reproductive success
between sexes or sampling bias have also been
indicated as possible explanations. So our
results may be a consequence of protandria
and later maturation in females. Size at sexual
maturity was larger for females than that of
males. This may be a strategy of higher initial
investment in somatic body weight which
allows an increased investment in gonads at
a later time. This trade-off strategy implies
an extension of the time transcurred before
entering the reproductive life phase, which
in turn may permit higher mortality before
reproduction. Although itness in one hand is
augmented by morphological characteristics,
it is at the same time reduced by the inverse
relationship between development time and
the possibility of death.
Winemiller (1992) noted that medium-sized
characids usually have a periodic life strategy
characterized by late maturity, large numbers
Román-P. et al.
of eggs and low survival. Mean fecundity
(776 oocytes) for H. brevispini is high when
compared to other species of the genus: H.
boquiae (376 oocytes) (Román-Valencia et al.
2008) and H. quindos (445 oocytes) (RománValencia & Botero 2006). When compared
to other characids in the area, H. brevispini
has lower fecundity than Bryconamericus
caucanus (3759) (Román-Valencia & Muñoz
2001a), B. galvisi (1391 oocytes) (RománValencia & Muñoz 2001b), and Creagrutus
brevipinnis (613 oocytes) (Román-Valencia
1998), but it has higher fecundity than
Carlastyanax aurocaudatus (181 oocytes)
(Román-Valencia & Ruiz-C. 2005).
The periodic life strategy is also evident in the
larger average size of females with respect to
males, a tendency shared with other species of
characids common in the area. A larger size at
sexual maturity was found for H. brevispini
(76.3 mm SL for females and 68.7 mm SL
for males) when compared to other species of
the genus: H. boquiae (65 mm SL for females
and 45 mm SL for males) (Román-Valencia
et al. 2008) and H. quindos (53 mm SL for
females and 50 mm SL for males) (RománValencia 1998); and when compared to other
characids in the area: C. brevispinnis (40 mm
SL) (Román-Valencia 1998), Bryconamericus
caucanus (50 mm SL for female and 40 mm
SL for male) (Román-Valencia et al. 2008),
Carlastyanax aurocaudatus (35 mm SL for
female and 40 mm SL for male) (RománValencia & Ruiz-C. 2005), except for B.
galvisi (57.5-89.9 mm SL for female and 61.381.1 mm SL for males) (Román-Valencia &
Muñoz 2001b).
Although we found no statistically signiicant
seasonal differences in water quality
parameters measured (pH, temperature,
dissolved oxygen) between the wet and dry
seasons (see also other data and parameters
measured in Román-Valencia et al. 2005), this
does not mean that changes in these parameters
do not affect Hemibrycon brevispini. Physico-
chemical parameters for habitats of H.
boquiae (Román-Valencia et al. 2008), H.
quindos (Román-Valencia & Botero 2006)
and Hyphessobrycon poecilioides (GarcíaAlzate & Román-Valencia 2008) also showed
few signiicant seasonal differences.
Although physico-chemical parameters
measured do not yet indicate decreased water
quality in La Venada Creek (see also other
data in Román-Valencia et al. 2005), threats
not evaluated here, occurring near the study
site from mining and current cultivated areas
make this endemic species’ future uncertain.
In this study area, other endemic species
of Hemibrycon are present: H. boquiae, H.
palomae and H. quindos, and are threatened
in similar ways as H. brevispini. Studies
of trophic and reproductive ecology of this
species will provide a useful baseline for
future impact studies.
ACKNOWLEDGMENTS
We thank K. Winemiller and three anonymous
reviewers for valuable criticism and
suggestions on the manuscript. We thank
Biology students of the University of Quindío,
Armenia (IUQ) for help with sampling.
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Recibido: 07/10/2013
Aceptado: 29/10/2014