You are on page 1of 9

Journal of Fish Biology (2006) 69, 1770–1778

doi:10.1111/j.1095-8649.2006.01245.x, available online at

Ontogenetic allometric coefficient changes: implications

of diet shift and morphometric traits in Hoplias
malabaricus (Bloch) (Characiforme, Erythrinidae)
*Aquatic Systems Research and Recovery Group, Ecology Department, Facultad de
Ciencias, Universidad de la República, Iguá 4225, CP. 11400 Montevideo,
Uruguay,†Basic and Applied Ecology Research Group, NGO Investigación y desarrollo
(IþD), Iguá 4225, CP. 11400 Montevideo, Uruguay and ‡Ecotoxicology and
Environmental Chemistry Group, Facultad de Ciencias, Universidad de la República,
Iguá 4225, CP. 11400 Montevideo, Uruguay

(Received 11 November 2005, Accepted 21 July 2006)

This study evaluated the relationship between body size and digestive tract characteristics of the
important predatory freshwater fish Hoplias malabaricus, which is widely distributed in South
America. The allometric coefficients were calculated for the mass and standard length (LS)
relationships for two different LS groups: (1) between 20 and 100 mm (characterized as
insectivores) and (2) >100 mm (characterized as piscivores). Differential growth measured from
the allometric coefficient, b, between the insectivore (b < 3) and the piscivore (b > 3) groups was
detected. Anterior intestine length and pyloric caeca zone length showed significant differences
between groups. Two complementary hypotheses were developed to explain the differential
growth: (1) H. malabaricus has a digestive tract adapted to a piscivorous diet, which is
independent of its ontogenetic stage of development, and (2) the negative allometry observed in
group 1 individuals agrees with a general behavioural strategy, allowing individuals to grow in
LS during a shorter period of time. # 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles

Key words: allometric coefficient; diet shift; Hoplias malabaricus; ontogenetic changes; South

The lack of basic biological knowledge about neotropical fish communities and
the implications for the development of management and conservation meas-
ures have been deplored by many researchers (Menni et al., 1984; Buti, 1995;
Buti & Miquelarena, 1995; Vari & Malabarba, 1998; Lizama & Ambrósio,
2002). Analyses of mass–length relationships contribute to reducing this lack
of knowledge.
The allometric coefficient of the mass–length relationships is influenced by,
among other factors, the quantity and quality of the ingested food. Many

§Author to whom correspondence should be addressed. Tel.: þ598 2 5258618 (ext. 162); fax: þ598 2
5258617; email:

# 2006 The Authors
Journal compilation # 2006 The Fisheries Society of the British Isles

studies of fishes have described the relationship between dietary habits and
variations in morphological attributes, such as the intestinal tract (Kramer &
Bryant, 1995a; Elliott & Bellwood, 2003) and body size (Mérigoux & Ponton,
1998). By using relative intestinal length as an index of fish diets, Al-Hussaini
(1947) revealed that planktivores had the shortest relative intestinal length, fol-
lowed by carnivores, omnivores and finally herbivores. Relative intestinal
length is a crude indication of diet quality and the most commonly used
descriptor of alimentary tract morphology in fishes (Horn, 1989).
Shorter digestive tracts, as observed in carnivores, may present a greater
number of pyloric caeca, which increase the surface area and compensate for
a relatively short intestine (Buddington et al., 1997; Kubitza, 1999). Function-
ally, these projections (pyloric caeca), which form part of the anterior-most
region of the intestine, increase the surface area for absorption (Buddington
& Diamond, 1986), but also appear to be sites of digestion with pancreatic en-
zymes (Bergot, 1981; Dimes et al., 1994).
Tararira Hoplias malabaricus (Bloch) is one of the most widely distributed
freshwater piscivores of South America and occurs in almost all hydrographic
basins, except west of the Andes and in rivers in the Patagonia (11° N–35° S;
85° W–35° W) (Fowler, 1950; Reis et al., 2003), where it is the top predator. It
is the species most frequently captured for human consumption in Argentinean
(Grosman et al., 2004), Colombian (Bentancur-Vásquez et al., 2004) and Uru-
guayan (Crossa, 1994; Amestoy, 2001) freshwater lakes and is also an attractive
game fish for anglers.
Hoplias malabaricus has three ontogenetic phases in its diet composition: (1)
a planktivore phase for sizes under 20 mm standard length (LS), (2) an insec-
tivore phase for sizes between 20 and 100–120 mm LS and (3) a piscivore phase
for specimens >160 mm LS. Individuals within the 100–120 mm and 160 mm
ranges have a mixed diet (Azevedo & Gomes, 1942; Oliveros & Rossi, 1991;
Galvis et al., 1997; Meschiatti & Arcifa, 2002).
A similar body size-related diet shift has been reported for other native spe-
cies of South America, i.e. Serrasalmus brandtii (Lutken) (Oliveira et al., 2004)
and Oligosarcus jenynsii (Günther) (Barros, 2004).
The ontogenetic change in diet relates to morphological limitations of each
developmental phase, such as mouth size (Wootton, 1990; Houde, 1997) and
locomotion capability (Wootton, 1990). Increasing size promotes not only
the development of foraging skills and the resultant progressively widening diet
but also reduces predation risk that is crucial for the survival of the individual
(Cowan et al., 1997; Fedatto-Abelha et al., 2001).
Although there are examples of allometric changes in the digestive traits
accompanying ontogenetic diet shifts in many species of vertebrates (Stevens
& Hume, 1995) and particularly in fishes, insectivory to frugivory (Drewe
et al., 2004), omnivory to herbivory (Benavides et al., 1994), no reports about
allometric changes in the digestive traits for insectivory to piscivory have been
found in the literature.
The aim of this study was to evaluate the relationship between the digestive
tract characteristics and the allometric coefficients of two different LS groups of
H. malabaricus (insectivores, <100 mm and piscivores, >100 mm). The follow-
ing questions were posed: (1) Do the two predefined dietary groups exhibit

# 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778

different growth types (shape)? (2) Are the differences related to modification
of the digestive tracts?


Hoplias malabaricus were collected (n ¼ 203) in lentic systems of the lower Santa
Lucia basin (South of Uruguay, between 33°429189 S; 34°5092499 S; 54°5992499 W–
57°0791199) during several sampling events between October 2004 and April 2006. Fish
were captured using a 30  15 m, 15  15 cm mesh beach seine, lines with fishhooks
and electrofishing (type FEG 1000). The LS and total mass (MT) were measured to the
nearest 1 mm and 001 g.
The coefficients (a and b) of the potential relationship MT ¼ a LSb (Ricker, 1973)
were estimated by a piecewise linear regression after log10 transformation of the varia-
bles (Bauchot & Bauchot, 1978). Given the diet shift of H. malabaricus reported by
Azevedo & Gomes (1942), Oliveros & Rossi (1991) and Meschiatti & Arcifa (2002),
the break point in the piecewise linear regression was fixed to the value 100 mm for
the LS axis, i.e. group 1 (G1) to all values <100 mm and group 2 (G2) to values
>100 mm (Table I). Because of possible overlapping in the diet of the two groups
due to a gradual transition from insectivory to piscivory and in order to obtain a better
regression and to avoid border interactions between the two segments of the regression,
LS data between 90 and 110 mm were omitted. To test for both size classes whether fish
growth was statistically different from isometric growth, a t-test (H0: b ¼ 3) with a ¼
005 was performed (Mayrat, 1970; Ricker, 1973, 1975).
For gut sampling, 54 individuals (30 belonging to G1 and 24 to G2) were sacrificed
with an overdose of anaesthesia (solution of 2-phenoxy-ethanol, 1 ml l1). They were
subsequently weighed and used to measure (1) total digestive tract length (from cardiac
sphincter to anus) (LTD), (2) stomach length (from the cardiac to the pyloric sphincter)
(LTS), (3) total intestine length (from the pyloric sphincter to the anus) (LTI), (4) ante-
rior intestine length (from pyloric sphincter to pyloric caeca zone) (LIA), (5) pyloric cae-
ca zone length (LIPC), (6) posterior intestine length (from pyloric caeca zone to the
anus) (LIPI) and (7) pyloric caeca maximum length (LPC) (Table II). Linear regressions
after log10 transformation of all the measured variables and the LS were conducted, and
t-test was employed to determine differences between the two dietary groups regression
Following this, some fish specimens were fixed in formalin (10%) and deposited in
the Vertebrates Collection of Facultad de Ciencias (institutional code ZVC-P).

From a total of 203 H. malabaricus individuals, 106 were classified as belong-
ing to G1 (LS  90 mm), 62 as belonging to G2 (LS  110 mm) and 35 were

TABLE I. Number of individuals (n), mean  S.E., minimum and maximum standard
length (LS) and total mass (MT) for insectivores (G1) and piscivores (G2) individuals of
Hoplias malabaricus used in the log10 transformed MT and LS regression

Ls (mm) MT (g)

Group n Mean  S.E. Minimum Maximum Mean  S.E. Minimum Maximum

G1 102 702  15 320 900 66  03 06 150

G2 62 2559  122 1100 5300 4461  629 99 36100

# 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778
TABLE II. Morphometric traits mesured for the two trophic groups of Hoplias malabaricus (G1 and G2). Number of individuals (n), mean 
S.E., minimum and maximum standard length (LS), total digestive tract length (LTD), stomach length (LTS), total intestine length (LTI), anterior
intestine length (LIA), pyloric caeca zone length (LIPC), posterior intestine length (LIPI) and pyloric caeca maximum length (LPC). Regression
equations of log10 transformed digestive traits and log10 LS are also given (S.E. of coefficients are given within parentheses)

G1 (n ¼ 30) G2 (n ¼ 24) Regressions fitted for digestive traits

# 2006 The Authors

LS (mm) Mean  S.E. 320  31 2443  166
Minimum 320 1100
Maximum 900 3450
LTD (mm) Mean  S.E. 754  41 3128  214 G1: log10 LTD ¼ 020 (010) þ 095 (006) log10 LS;
Minimum 428 1278 G2: log10 LTD ¼ 024 (013) þ 094 (005) log10 LS
Maximum 1302 4845
LTS (mm) Mean  S.E. 151  10 681  54 G1: log10 LTS ¼ 055 (018) þ 098 (010) log10 LS;
Minimum 87 279 G2: log10 LTS ¼ 046 (021) þ 096 (009) log10 LS
Maximum 300 1400
LTI (mm) Mean  S.E. 603  33 2448  166 G1: log10 LTI ¼ 012 (011) þ 094 (006) log10 LS;
Minimum 328 1000 G2: log10 LTI ¼ 015 (013) þ 094 (006) log10 LS
Maximum 10735 3445
LIA (mm) Mean  S.E. 40  02 165  14 G1: log10 LIA ¼ 057 (019) þ 067 (011) log10 LS*;
Minimum 22 64 G2: log10 LIA ¼ 150 (020) þ 113 (008) log10 LS*
Maximum 64 267
LIPC (mm) Mean  S.E. 129  06 488  31 G1: log10 LIPC ¼ 010 (020) þ 057 (011) log10 LS*;
Minimum 73 243 G2: log10 LIPC ¼ 035 (015) þ 085 (006) log10 LS*
Maximum 204 694
LIPI (mm) Mean  S.E. 433  28 1795  124 G1: log10 LIPI ¼ 031 (013) þ 110 (007) log10 LS;
Minimum 213 693 G2: log10 LIPI ¼ 001 (016) þ 095 (007) log10 LS

Maximum 865 2644

LPC (mm) Mean  S.E. 30  02 153  10 G1: log10 LPC ¼ 109 (023) þ 088 (013) log10 LS;
Minimum 14 75 G2: log10 LPC ¼ 085 (014) þ 085 (006) log10 LS
Maximum 57 221

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778
*Statistically significant differences between trophic groups, P < 005.

omitted (90 mm < LS < 110 mm). According to the literature, visual inspection
of gut contents revealed that the G1 individuals were insectivores and that ones
belonging to G2 were piscivores (the genera found in G2 gut contents were
Astyanax, Cheirodon, Charax and Characidium).
The relationships between MT and LS two dietary groups were G1: log10 MT ¼
437 (011) þ 279(006) log10 LS and G2: log10 MT ¼ 550 (015) þ
331(007) log10 LS. The  S.E. of the coefficients are given in parentheses.
The t-test showed significantly different allometric coefficient values (P <
005) for the two dietary groups compared to isometric growth (b ¼ 3). For
G1, the b value was significantly lower, corresponding to negative allometry,
while for the G2, b was significantly higher, corresponding to positive allometry.
Significant differences between the two dietary groups were found for LIA
(G1 < G2, t-test, d.f. ¼ 50, P < 001) and LIPC (G1 < G2, t-test, d.f. ¼ 50,
P < 005) (Fig. 1).

Allometric growth, measured as b, of the two size groups was reported. In
both cases, the b values fall within the range given by Le Cren (1951) for most
fish species (25–4). G1 individuals showed negative allometry (b < 3), while G2
showed positive allometry (b > 3). Lizama et al. (1999) found that several spe-
cies of Characiformes from the upper Paraná River floodplain tend towards
isometric growth; however, Villacorta-Correa & Saint-Paul (1999) found nega-
tive allometry in the characiform species of Central Amazon (Brazil) Colossoma
macropomum (Cuvier). Dománico et al. (1993) found positive allometry in H.
malabaricus in a lentic system in Argentina. This study is the first record of dif-
ferential ontogenetic allometry in response to a diet shift of H. malabaricus.
The present study did not reveal differences in the total digestive tract length,
stomach length, total intestine length, posterior intestine length and pyloric
caeca maximum length between the two groups. The expectation was to find
a larger intestine for the G1 individuals, in accordance with Zihler (1982)
who noted that insectivorous cichlids tend to have longer intestines that are
more adapted to degrade the chitinous and calcareous exoskeleton of
insects. The present results, however, are consistent with those of Kramer &
Bryant (1995a) who working with quotients found similar intestine and LS size
relationships between different size classes of Hoplias microlepis (Günther) to
those of this study. Kramer & Bryant (1995b) discovered that for species with-
out ontogenetic changes in intestine and LS relationship, ontogenetic modifica-
tions in the pyloric caeca zone and LS relationship may be found. This agrees
well with the present results, where the total digestive length and the total intes-
tine length did not differ but when the different segments of the intestine were
analysed a relative enlargement of pyloric caeca zone was observed. Such mod-
ification can be interpreted as a mechanism increasing the total absorption
surface, which allows maintenance of the absorption with enlarging body size
(Kramer & Bryant, 1995b).
Two complementary hypotheses are proposed to explain the differential
growth among the two groups. Hypothesis one claims that H. malabaricus

# 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778

FIG. 1. Significant differences between the two dietary groups (d, G1 < 90 mm and , G2 > 110 mm
standard length, LS) were found for (a) anterior intestine length (from pyloric sphincter to pyloric
caeca zone, LIA) (G1 < G2, P < 001) and (b) pyloric caeca zone length (LIPC) (G1 < G2, P < 005)
(see Table II).

has a digestive tract adapted to a piscivorous diet independently of its ontoge-

netic stage of development. This explains why the small G1 individuals showed
negative allometry, as their digestive tracts are not well adapted to efficiently
exploit the food ingested (insects). Hypothesis two proposes that the negative
allometry observed in G1 individuals agrees with a general behavioural strat-
egy, allowing individuals to grow longer in size during a shorter period of time
(Munch & Conover, 2003). This would imply allocation of more energy to
axial growth rather than biomass to reach the piscivorous phase as quickly
as possible, while simultaneously reducing the predation risk (Wootton,
1990). During the early life stages, the survival of organisms strongly depends
on body size (Sogard, 1997), and among mature individuals, the reproductive
success typically increases with body size (Roff, 1992; Stearns, 1992).

# 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778

Further field and experimental studies are needed to confirm or reject the
hypotheses, and careful analyses of the growth of fish species in South
American lakes would contribute to the understanding of the different growth
strategies associated with ontogeny.

We wish to thank Unidad de Ciencias de Epigenesis (UNCIEP) Facultad de

Ciencias-UDELAR for the use of their laboratory, M. Arim for invaluable statistical
advice and A.-M. Poulsen and M. Zuluaga for editorial and language assistance. Com-
ments from R. Wootton and two anonymous referees greatly improved the manuscript.
This work was supported by the Dirección Nacional de Ciencia y Tecnologı́a (Progra-
ma de Desarrollo Tecnologico, 07/16).

Al-Hussaini, A. H. (1947). The feeding habits and the morphology of the alimentary tract of
some teleosts living in the neighbourhood of the Marine Biological Station, Ghardaga,
Red Sea. Publications of the Marine Biology Station, Ghardaga, Red Sea 5, 1–61.
Amestoy, F. (2001). Hacia una cuantificación de estrés ecológico en el embalse de Rincón
del Bonete (Uruguay): distribución, biomasa y estructura de la comunidad ictı́cola.
PhD Thesis, Universidad de la República (PEDECIBA), Montevideo, Uruguay.
Azevedo, P. & Gomes, A. L. (1942). Contribuicxão ao estudo da biologia da traı́ra Hoplias
malabaricus (Bloch, 1794). Boletim de Indústria Animal 5, 15–64.
Barros, S. E. (2004). Alimentación de Oligosarcus jenynsii (Characiformes: Characidae)
en dos embalses sobre el rı́o Juramento, Salta, Subtrópico de Argentina. Revista
AquaTIC 20, 44–50.
Bauchot, R. & Bauchot, M. L. (1978). Coefficient de condiction et indice pondéral chez
les Téléostéens. Cybium (3e Se´rie) 4, 3–16.
Benavides, A. G., Cancino, J. M. & Ojeda, F. P. (1994). Ontogenetic changes in gut
dimensions and macroalgal digestibility in the marine herbivorous fish, Aplodacty-
lus punctatus. Functional Ecology 8, 46–51.
Bergot, P. (1981). Structure de l’appareil digestif. II. Structure et fonctions des caeca
pyloriques. In Nutrition des Poissons (Fontaine, M., ed.), pp. 45–53. Paris, France:
Editions du CNRS.
Buddington, R. K. & Diamond, J. M. (1986). Aristotle revisited: the function of pyloric
caeca in fish. Proceedings of the National Academy of Sciences 83, 8012–8014.
Buddington, R. K., Krogdhal, Å. & Bakke-McKellep, A. M. (1997). The intestines of
carnivorous fish: structure and functions and the relations with diet. Acta
Physiologica Scandinavica 161 (Suppl. 638), 67–80.
Buti, C. L. (1995). Ictiofauna del Embalse C. Gelsi (El Cadillal) y sus tributarios
provincia de Tucumán, República Argentina. Acta Zoologica Lilloana 43, 207–213.
Buti, C. L. & Miquelarena, A. M. (1995). Ictiofauna del Rı́o Sali superior, departamento
de Trancas, Tucumán, República Argetina. 1995. Acta Zoologica Lilloana 43, 21–44.
Cowan, J. H. Jr, Rose, K. A. & Houde, E. D. (1997). Size-based foraging success and
vulnerability to predation: selection of survivors in individual-based models of
larval fish populations. In Early Life History and Recruitment in Fish Populations
(Chambers, R. C. & Trippel, E. A., eds), pp. 357–386. London: Chapman & Hall.
Crossa, N. (1994). Aspectos relacionados a producão pesqueira no reservatório Rincón
del Bonete no rio Negro (Durazno-Tacuarembó, Uruguay). MSc Thesis, Uni-
versidade Estadual Paulista, Rio Claro, Brazil.
Dimes, L. E., Garcia-Carreño, F. L. & Haard, N. F. (1994). Estimation of protein
digestibility. III. Studies on the digestive enzymes from the pyloric ceca of rainbow
trout and salmon. Comparative Biochemistry and Physiology 109, 349–360.
Dománico, A. A., Delfino, R. & Freyre, L. R. (1993). Edad y crecimiento de Hoplias
malabaricus malabaricus (Bloch, 1794) en la laguna de Lobos (Argentina).
Iheringia, Se`rie Zoologica 74, 141–149.

# 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778

Drewe, K. E., Horn, M. H., Dickson, K. A. & Gawlicka, A. (2004). Insectivore to

frugivore: ontogenetic changes in gut morphology and digestive enzyme activity in
the characid fish Brycon guatemalensis from Costa Rican rain forest streams.
Journal of Fish Biology 64, 890–902. doi: 10.1111/j.1095-8649.2004.00357.x
Elliott, J. P. & Bellwood, D. R. (2003). Alimentary tract morphology and diet in three
coral fish families. Journal of Fish Biology 63, 1598–1609. doi: 10.1111/j.1095-
Fedatto-Abelha, M. C., Agostinho, A. A. & Goulart, E. (2001). Plasticidade trófica em
peixes de água doce. Maringá 23, 425–434.
Fowler, H. W. (1950). Os peixes de água doce do Brasil. Arquivos de Zoologia do Estado de
São Paulo 6, 205–404.
Galvis, G., Mojica, J. I. & Camargo, M. (1997). Peces del Catatumbo. Santafé de Bogotá
DC: Asociación Cravo Norte.
Horn, M. H. (1989). Biology of marine herbivorous fishes. Oceanography and Marine
Biology Annual Review 27, 167–272.
Houde, E. D. (1997). Patterns and consequences of selective processes in teleost early life
histories. In Early Life History and Recruitment in Fish Populations (Chambers, C. &
Trippel, E. A., eds), pp. 173–196. London: Chapman & Hall.
Kramer, D. L. & Bryant, M. J. (1995a). Intestine length in the fishes of a tropical stream:
2. Relationships to diet—the long and short of a convoluted issue. Environmental
Biology of Fishes 42, 129–141.
Kramer, D. L. & Bryant, M. J. (1995b). Intestine length in the fishes of a tropical stream:
1. Ontogenetic allometry. Environmental Biology of Fishes 42, 115–127.
Kubitza, F. (1999). Nutricxão e alimentacxão dos peixes cultivados, 3rd edn. Jundiaı́/SP
Piracicaba: Degaspari Ltda.
Le Cren, E. D. (1951). The length-weight relationship and seasonal cycle in gonad
weight and condition in the perch (Perca fluviatilis). Journal of Animal Ecology 20,
Lizama M. de los, A. P. & Ambrósio, A. M. (2002). Condition factor in nine species of
fish of the Characidae family in the upper Paraná river floodplain, Brazil. Brazilian
Journal of Biology 62, 113–124.
Lizama M. de los, A. P., Ambrósio, A. M. & Vazzoler, A. E. A. de M. (1999). Relacxão
peso-comprimento e estrutura da populacxão de nove espécies da famı́lia Char-
acidae na planı́cie de inundacxão do alto rio Paraná, Brasil. Revista Brasileira de
Zoologia 16, 779–788.
Mayrat, A. (1970). Allometrie et taxinomie. Revue de Statistique Applique´e 18, 47–58.
Menni, R. C., López, H. L., Casciota, J. R. & Miquelarena, A. M. (1984). Ictiologı́a de
las áreas serranas de Córdoba y San Luis (Argentina). Biologı´a Acuática 5, 1–63.
Mérigoux, S. & Ponton, D. (1998). Body shape, diet and ontogenetic diet shifts in young
fish of the Sinnamary River, French Guiana, South America. Journal of Fish
Biology 52, 556–569. doi: 10.1111/j.1095-8649.1998.tb02017.x
Meschiatti, A. J. & Arcifa, M. S. (2002). Early life stages of fish and the relationships with
zooplankton in a tropical Brazilian reservoir: lake Monte Alegre. Brazilian Journal
of Biology 62, 41–50.
Munch, S. B. & Conover, D. O. (2003). Rapid growth results in increased susceptibility to
predation in Menidia menidia. Evolution 57, 2119–2127.
Oliveira, A. K., Alvim, M. C. C., Peret, A. C. & Alves, C. B. M. (2004). Diet shifts related
to body size of the Pirambeba Serrasalmus brandtii Lütken, 1875 (Osteichthyes,
Serrasalminae) in the Cajuru reservoir, Sao Francisco river basin, Brazil. Brazilian
Journal of Biology 64, 117–124.
Oliveros, O. B. & Rossi, L. M. (1991). Ecologı́a trófica de Hoplias malabaricus
malabaricus (Pisces, Erythrinidae). Revista de la Asociación de Ciencias Naturales
del Litoral 22, 55–68.
Reis, R. E., Kullander, S. O. & Ferraris, C. J. (2003). The Check List of the Freshwater
Fishes of South and Central America. Porto Alegre: EDIPUCRS.
Ricker, W. E. (1973). Linear regressions in fishery research. Journal of the Fisheries
Research Board of Canada 30, 409–434.

# 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778

Ricker, W. E. (1975). Computation and interpretation of biological statistics of fish

populations. Bulletin of the Fisheries Research Board of Canada 191, 1–382.
Roff, D. A. (1992). The Evolution of Life Histories: Theory and Analysis. New York:
Chapman & Hall.
Sogard, S. M. (1997). Size-selective mortality in the juvenile stage of teleost fishes:
a review. Bulletin of Marine Science 60, 1129–1157.
Stearns, S. (1992). The Evolution of Life Histories. New York: Oxford University Press.
Stevens, E. D. & Hume, I. D. (1995). Comparative Physiology of the Vertebrate Digestive
System, 2nd edn. New York: Cambridge University Press.
Vari, R. P. & Malabarba, L. R. (1998). Neotropical ichthyology: an overview. In
Phylogeny and Classification of Neotropical Fishes (Malabarba, L. R., Reis, R. E.,
Vari, R. P., Lucena, Z. M. S & Lucena, C. A. S., eds), pp. 1–11. Porto Alegre:
Villacorta-Correa, M. A. & Saint-Paul, U. (1999). Structural indexes and sexual maturity
of tambaqui Colossoma macropomun (Cuvier, 1818) (Characiformes: Characidae)
in Central Amazon, Brasil. Revista Brasileira de Biologia 59, 637–652.
Wootton, R. J. (1990). Ecology of Teleost Fishes. New York: Chapman & Hall.
Zihler, F. (1982). Gross morphology and configuration of digestive tracts of Cichlidae
(Teleostei, Perciformes): phylogenetic and functional significance. Netherlands
Journal of Zoology 32, 544–571.

Electronic References
Bentancur-Vásquez, B., Humanez, J. C., Olaya-Nieto, C. W., Tordecilla-Petro, G.,
Sánchez-Banda, S., Segura-Guevara, F. F. & Brú-Cordero, S. B. (2004). Talla de
madurez sexual del Moncholo (Hoplias malabaricus Bloch, 1794) en la Ciénaga
Grande de Lorica (Colombia). Comunicación Cientı´fica CIVA, pp. 290–296. Avail-
able at
Grosman, F., Birnstil, V., Cabral, J., March, L., Merlos, C., Rudzik, G., Vavrin, L.
(2004). ‘El uso actual justifica acciones de gestión en la Tararira Hoplias
malabaricus’ Una visión desde los aspectos socioeconómicos y biológicos. Comu-
nicación Cientı´fica CIVA, pp. 362–374. Available at

# 2006 The Authors

Journal compilation # 2006 The Fisheries Society of the British Isles, Journal of Fish Biology 2006, 69, 1770–1778