Acta Zoologica (Stockholm) 90: 101– 109 ( April 2009) doi: 10.1111/j.1463-6395.2008.00336.

REVIEW
Blackwell Publishing Ltd

Morphology of the digestive tract and feeding habits of

Loricaria lentiginosa Isbrücker, 1979 in a Brazilian reservoir
L. F. Salvador-Jr, G. N. Salvador and G. B. Santos
PPG Zoologia de Vertebrados, PUCMinas, Av. D. José Gaspar 500, prédio 41, 30535-610 Belo Horizonte, MG, Brazil

Gilmar Bastos Santos, PPG Zoologia de
Vertebrados, PUCMinas, Av. D. José Gaspar,
500, prédio 41, 30535-610 Belo Horizonte,
MG, Brazil. E-mail: astyanax@pucminas.br
Abstract
Salvador-Jr, L.F., Salvador, G.N. and Santos, G.B. 2009. Morphology of the
digestive tract and feeding habits of Loricaria lentiginosa Isbrücker 1979
(Loricariidae) in a Brazilian reservoir — Acta Zoologica (Stockholm) 90: 101–
109.

Keywords
The present study describes some aspects of the morphology of the digestive
food items, Loricariidae, Rio Grande basin
tract and the feeding patterns of Loricaria lentiginosa. Sixty-three specimens
with gut contents were captured quarterly from November 2001 to August
2002 in Porto Colômbia reservoir, Rio Grande basin, southeastern Brazil. The
species has bicuspid teeth inserted in the lips, unicuspid teeth in the dentary
bone and pharyngeal teeth, besides having short, numerous and slightly
separated gill rakers. The stomach is rudimentary and the intestine is relatively
short (Intestinal Quotient = 1.32 ± 0.15). Molluscs, especially bivalves,
represented the most important item on the diet of this species, followed by
organic matter. The characteristics of the digestive tract as well as the dietary
pattern, suggest that L. lentiginosa is able to explore bottom food resources,
being classified as benthophagous with a tendency to malacophagy, and
utilizing organic matter as an associated or additional food resource.
Loricariidae species are able to use not only algae, sediment and detritus, but
also plant and invertebrate items.

characterization have been performed mostly for the genera

Introduction
Loricariid fish are characterized by having a body covered
with bony plates or scutes, and a ventral sucker-like mouth
(Montoya-Burgos et al. 1998). They constitute the most
diverse group of Siluriformes, with 673 recognized species
grouped in at least 100 genera (Isbrucker 2002; Reis et al.
2003).
Their natural distribution encompasses the Neotropical
region, reaching Panama and Costa Rica in the north
(Fowler 1954; Reis et al. 2003). Although widely distributed
on the South American continent, the majority of species in
this group are found in limited numbers in small geographical
areas (Burgess 1989).
Angelescu and Gneri (1949), Rapp Py-Daniel and Py-
Daniel (1984), Fugi et al. (1996, 2001) and Delariva and
Agostinho (2001) carried out studies on the functional
morphology of this family. In addition, works related to dietary
Hypostomus (Angelescu and Gneri 1949; Delariva and
Agostinho 2001; Casatti 2002; Alvim and Peret 2004;
among others) and Loricariichthys (Fugi et al. 1996, 2001;
Hahn et al. 1998; da Luz et al. 2001; among others).
The ‘Acari’ or ‘Cascudo-chinelo’, Loricaria lentiginosa, is
an endemic species of the tributaries of the upper Paraná
River basin, upstream of the Paranapanema River (Agostinho
and Júlio 1999). Little is known about the biology of this
Loricariinae species, with data in the literature restricted to
works by Agostinho and Júlio (1999), Andrade and Braga
(2005a,b) and Guimarães-Cruz et al. (2005).
In Porto Colômbia reservoir (Rio Grande, southeastern
Brazil), L. lentiginosa is restricted to a narrow stretch
(Pimenta 2003) comprising the ‘riverine’ region according to
the Thornton (1990) classification, with rocky bottom and
aquatic macrophytes. As a result of these peculiar features,
that characterize the species’ preferential biotope, its current

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Journal compilation © 2008 The Royal Swedish Academy of Sciences 101
Feeding of Loricaria lentiginosa • Salvador-Jr et al.
distribution along the other 11 reservoirs constructed
consecutively in the Rio Grande probably follows the same
pattern.
The present study aimed to describe some aspects of the
morphology of the digestive tract and the feeding habits of
L. lentiginosa specimens from Porto Colômbia reservoir as
well as to discuss the dietary patterns of Loricariidae fish.
Materials and Methods
Collection of fish
Specimens of L. lentiginosa were captured quarterly from
November 2001 to August 2002 in the Porto Colômbia
reservoir (20°01′69′′S 48°13′40′′W) (Fig. 1), upper Rio
Paraná drainage system.
Fish were caught using 20-m lengths of gill nets with 3, 4,
5, 6, 7, 8, 10, 12, 14, 16 and 18 cm between opposing knots,
exposed for 12 h. A ventral incision was made on every
fish captured; fish were then fixed in a solution of 10% for-
maldehyde and later measured in the laboratory.
Morphology of the digestive tract and feeding habits
Sixty-three specimens were dissected for analysis of food
items according to Hyslop (1980). Ten of them, with stand-
ard lengths ranging from 35.0 cm to 44.5 cm, were also used
for the characterization of mouth, branchial rakers, stomach,
intestine, and pharyngeal, dentary and upper lip teeth.
Intestine length was measured from pyloric sphincter to anus
using a millimeter-graduated rule. The analysed structures
of the digestive tract were sketched by hand with Nanking
ink, scanned at 600 d.p.i. resolution and had their layout
plotted using Corel Draw software version 12, keeping the
proportions.
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Acta Zoologica (Stockholm) 90: 101– 109 ( April 2009)
Fig. 1—Map of Porto Colômbia reservoir,
located on the Rio Grande, showing the
sampled area.
The first third of the digestive tracts had their contents
identified to the possible lowest taxonomic level. Because of
the small size, the volume of each ingested item was evaluated
using two 1 mm high aluminium plates covered by one glass
slide upon a Petri plate attached to a millimetre-graduated
paper. We converted the obtained values to millilitres
(1 mm3 = 0.001 ml) according to Hellawell and Abel
(1971).
In this study, we considered organic matter as animal and
vegetal fragments in advanced decomposition stages, which
could not be classified in any other group. Inorganic particles
from substrate, such as sand and clay, were classified as
sediment. Vegetal items (algae excluded) that had recently
entered the digestive process were classified as plant
fragments.
Frequency of occurrence of each item was correlated with
volumetric data using the Alimentary Importance Index
(AII); this was proposed by Kawakami and Vazzoler (1980)
and is described by the equation AII = (Fi × Vi )/ε(Fi × Vi ),
where Fi and Vi are the frequency of occurrence (%) and
volume (%) of each item, respectively. The results were
multiplied by 100 according to Hahn et al. (1997a).
From the same 10 specimens utilized for the digestive tract
description we estimated the Intestinal Quotient by dividing
the intestine length by the body standard length to obtain the
relationship between the intestine relative length and the
species diet.
Results
Digestive tract morphology
As shown in Fig. 2, the species has a ventral suckermouth
surrounded by expanded lips with well-developed barbels.
Bicuspid teeth are inserted on the upper lip and unicuspid
© 2008 The Authors
Journal compilation © 2008 The Royal Swedish Academy of Sciences
Acta Zoologica (Stockholm) 90: 101–109 (April 2009)
Fig. 2—Digestive tract morphology of
Loricaria lentiginosa. —A.Ventral view of the
mouth showing the well-developed barbells.
—B. Structure of branchial rakers. —C.
Stomach shape. —D. Intestine tract. —E.
Shape and distribution of pharyngeal teeth.
—F. Upper lip teeth shape. —G. Dentary
teeth shape. Bar length = 1 cm.
spatula-shaped teeth are located in the dentary bone. In
addition, well-developed teeth are present on both upper and
lower pharyngeal bones. It also has short, numerous and
slightly separated branchial rakers. The stomach is rudimen-
tary, tubular and has a thick wall. The intestine is relatively
short, such that it does not exceed the standard body length
of the examined specimens by much (mean Intestinal
Quotient = 1.32 ± 0.15; n = 10; CV = 11.1%).
Feeding habits
From the 92 captured specimens, 29 (31.5%) had com-
pletely empty guts, so only the contents of 63 specimens were
analysed. Frequency of occurrence (F ), volume (V ) and AII
© 2008 The Authors
Journal compilation © 2008 The Royal Swedish Academy of Sciences
Salvador-Jr et al. • Feeding of Loricaria lentiginosa
values for each item found in the digestive tracts are shown
in Table 1.
Molluscs, especially Bivalvia (F = 21.2%, V = 74.1%,
AII = 81.3%), represented the most important trophic
resource, followed by organic matter (F = 25.6%, V =
9.7%, AII = 12.8%) and Hydrobiidae gastropods (F =
10.2%, V = 3.2%, AII = 1.7%) Other items, such as Diptera,
Lepidoptera and Odonata larvae, Plecoptera nymphs,
other gastropods and plant fragments, were found in
small amounts. Sediment was observed in 27 (42.9%) of the
analysed tracts, always being associated with other items.
Like the organic matter, sediment showed high frequency
(19.7%) and low volume (3.2%) percentages in relation to
the total items found.
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Table 1 Percentages of frequency of occurrence (F ), volume (V) and

Discussion
alimentary importance index (AII) for the food items ingested by
Loricaria lentiginosa specimens captured in Porto Colômbia reservoir Suyehiro (1942), Al-Hussanni (1949) and Angelescu and
Gneri (1949) associated the digestive tract characteristics of
Items Sub-items F V AII
fish species with their diet and feeding behaviour. After them,
Algae 0.73 0.001 0.004
other authors such as Wooton (1996), Fugi et al. (1996,
Crustacea Ostracoda 1.46 0.01 0.001 2001), Hahn et al. (1998), Delariva and Agostinho (2001)
Insecta Diptera larvae 2.19 0.19 0.02 and Moyle and Cech Jr (2004) also commented on this
Hymenoptera 0.73 0.01 0.0002 association. Among the pioneer studies, the one by Ange-
Lepidoptera larvae 1.46 5.40 0.41 lescu and Gneri (1949) deserves special mention because
Odonata larvae 2.19 0.28 0.03
these authors compared the morphology of the digestive
Plecoptera nymph 1.46 0.46 0.03
tract of Loricaria vetula (= Paraloricaria vetula) and Loricaria
Trichoptera 0.73 0.05 0.002
Unidentified insects 5.84 0.54 0.16 anus (= Loricariichthys anus) with their respective diets. They
Mollusca Bivalvia 21.17 74.06 81.26 described well-developed pharyngeal teeth, a poorly developed
Gastropoda Hydrobiidae 10.22 3.15 1.67 stomach and an intestine that was 1.3 times the standard
Other Gastropoda 2.92 2.84 0.32 length in P. ventula and 1.7 times the standard length in
Organic matter 25.55 9.67 12.80 L. anus. These results are similar to those found in the
Plant fragments 2.92 0.12 0.01
present study. They also observed that the basic diet of these
Rotifera 0.73 0.01 0.0002
Sediments 19.71 3.21 3.28
species consisted of molluscs and crustaceans.
n 63 Delariva and Agostinho (2001) compared the morphology
Class of SL (cm) 21.2– 45.4 of the digestive tract and the diet of six Loricariidae species.
For three of them they relate the presence of larger and
SL = Standard length. Higher values are shown in bold type. strong spatulate teeth to the ability to scrape and ingest
coarse items associated with the substratum. Although the
species they studied fed on sponges, bryozoans, bryophytes,

Table 2 Trophic categories proposed for Loricariidae species according to the literature

Sub family/Species Locality Trophic category Source

Ancistrinae
Ancistrus spinosus
Ancistrus triradiatus
Chaetostoma fischeri
Chaetostoma milesi
Hemiancistrus vittatus
Hypancistrus contradens
Hypancistrus inspector
Megalancistrus aculeatus
Panaque albomaculatus
Panaque dentex
Panaque gnomus
Panaque maccus
Panaque nigrolineatus
Panaque nocturnes
Panaque purusiensis
Hypoptopomatinae
Hisonotus francirochai
Parotocinclus britskii
Schizolecis guntheri
Rio Frijoles, Panama Periphytivorous Power (1983, 1984)
Rio Las Marias, Orinoco basin, Venezuela Algae-eater Solomon et al. (2004)
Rio Frijoles, Panama Periphytivorous Power (1983)
Rio Apure, Orinoco basin, Venezuela Epibenthic Flecker (1992)
consumer
Rio Tocantins, Brazil Detritivorous Mérona et al. (2001)
Upper Rio Orinoco basin, Venezuela Omnivorous Armbruster et al. (2007)
Upper Rio Orinoco basin, Venezuela, Granivorous Armbruster (2002)
Upper Rio Negro and Rio Xingu, Brazil
Upper Rio Paraná basin, Brazil Detritivorous Agostinho et al. (1997);
Hahn et al. (1997b, 1998, 2004)
Upper Rio Amazonas basin, Ecuador and Peru Wood-eater Schaefer and Stewart (1993)
Upper Rio Amazonas basin, Brazil and Ecuador Wood-eater Schaefer and Stewart (1993)
Upper Rio Amazonas basin, Ecuador and Peru Wood-eater Schaefer and Stewart (1993)
Captivity experiments, RioOrinoco basin, Wood-eater/ Nelson et al. (1999); Nelson (2002);
Venezuela Herbivorous Schaefer and Stewart (1993)
Captivity experiments, Rio do Peixe, Brazil Herbivorous/ Nelson (2002); Nonogaki et al. (2007)
Xilophagous
Upper Rio Amazonas basin, Peru and Ecuador Wood-eater Schaefer and Stewart (1993)
Upper Rio Purus basin, Brazil Wood-eater Schaefer and Stewart (1993)
Upper Rio Paraná basin, Brazil Herbivorous Castro and Casatti (1997)1
Rio Araguaia basin, Brazil Detritivorous Melo et al. (2004)
Rio Ribeira do Iguape basin, Brazil Generalist Buck and Sazima (1995)
herbivorous

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104 Journal compilation © 2008 The Royal Swedish Academy of Sciences
Acta Zoologica (Stockholm) 90: 101–109 (April 2009)
Table 2 Continued
Sub family/Species
Hypostominae
Hypostomus ancistroides
Hypostomus commersonii
Hypostomus derbyi
Hypostomus emarginatus
Hypostomus garmani
Hypostomus microstomus
Hypostomus nigromaculatus
Hypostomus plecostomus
Hypostomus punctatus
Hypostomus regani
Glyptoperichthys
joselimaeanus
Pterygoplichthys multiradiatus
Liposarcus anisitsi
Liposarcus pardalis
Kronichthys heylandi
Rhinelepis aspera
Loricariinae
Harttia kronei
Loricaria lentiginosa
Loricaria prolixa
Loricariichthys acutus
Loricariichthys anus
Loricariichthys nudirostris
Loricariichthys platymetopon
Loricariichthys rostratus
Paraloricaria vetula
Rineloricaria latirostris
Rineloricaria uracantha
Spatuloricaria evansi
Sturisoma nigrirostrum
Named as: 1 Microlepidogaster francirochai; 2 Liposarcus multiradiatus; 3 Loricaria prolixa lentiginosa; 4 Loricaria anus; 5 Loricaria vetula.
© 2008 The Authors
Journal compilation © 2008 The Royal Swedish Academy of Sciences
Salvador-Jr et al. • Feeding of Loricaria lentiginosa
Locality Trophic category Source
Upper Rio Paraná basin, Brazil Herbivorous/ Castro and Casatti (1997);
Periphytivorous/ Uieda et al. (1997); Hahn et al. (2004);
Detritivorous Casatti (2002); Castro et al. (2005);
Ferreira (2007).
Rio de La Plata, Uruguai; Rio Tocantins Iliophagus/ Angelescu and Gneri (1949); Rapp
basin, Rio Ribeira do Iguape basin and Periphytivorous/ Py-Daniel and Py-Daniel (1984); Abelha
Rio Iguaçu, Brazil Detritivorous et al. (2005); Dourado et al. (2005)
Rio Paraná basin and Rio Ribeira Detritivorous Abelha et al. (2005);
do Iguape basin, Brazil Loureiro-Crippa and Hahn (2006)
Rio Araguaia basin and Rio Tocantins, Brazil Detritivorous Mérona et al. (2001); Melo et al. (2004)
Rio São Francisco basin, Brazil Algivorous Casatti and Castro (1998)
Upper Rio Paraná basin, Brazil Detritivorous Hahn et al. (2004)
Upper Rio Paraná basin, Brazil Periphytivorous Casatti (2002)
Rio Uruguai, Argentina; Rio Frijoles, Panama; Iliophagus/ Angelescu and Gneri (1949);
Rio Negro and Rio Amazonas, Brazil Periphytivorous/ Power (1983); Merona and
Detritivorous Rankin-de-Merona (2004)
Rio de La Plata, Argentina Iliophagus Angelescu and Gneri (1949)
Upper Rio Paraná basin, Brazil Detritivorous/ Hahn et al. (2004) Luz-Agostinho
Algivorous et al. (2006); Nonogaki et al. (2007)
Rio Negro, Rio Amazonas and Detritivorous Mérona et al. (2001); Mérona and
Rio Tocantins, Brazil Rankin-de-Merona 2004)
Rio Solimões basin, Brazil Iliophagus Lowe-McConnell (1999)2
Lower Rio Miranda and Upper Rio Detritivorous Resende (2000), Benedito-Cecilio
Paraná basin, Brazil et al. (2004b), Hahn et al. (2004)
Amazon basin, Brazil Specialized Yossa and Araújo-Lima (1998)
detritivorous
Rio Ribeira do Iguape basin, Brazil Generalist Buck and Sazima (1995)
herbivorous
Upper Rio Paraná basin, Brazil Detritivorous/ Agostinho et al. (1997); Hahn
Iliophagus et al. (1997b, 1998, 2004);
Delariva and Agostinho (2001)
Rio Ribeira do Iguape basin, Brazil Generalist Buck and Sazima (1995)
herbivorous
Upper Rio Paraná basin, Rio Grande, Brazil Specialized Andrade and Braga (2005b)3;
herbivorous/ Present study
Bentophagous
Upper Rio Paraná basin, Rio Paraná, Brazil Detritivorous Agostinho et al. (1997); Hahn
et al. (1997b, 1998); Fugi et al. (2005)
Rio Negro and Rio Amazon, Brazil Omnivorous Merona and Rankin-de-Merona (2004)
Rio Uruguai and Rio de La Plata, Argentina Omnivorous Angelescu and Gneri (1949)4
Rio Tocantins, Brazil Detritivorous Mérona et al. (2001)
Upper Rio Paraná basin and Detritivorous Fugi et al. (1996, 2001); Agostinho
Lower Rio Miranda, Brazil et al. (1997); Hahn et al. (1997b,
1998, 2004); Resende (2000);
Da Luz et al. (2001); Benedito-Cecilio
et al. (2004a,b), Dourado et al. (2005)
Upper Rio Paraná basin, Brazil Detritivorous Hahn et al. (2004)
Rio de La Plata, Argentina Omnivorous Angelescu and Gneri (1949)5
Upper Rio Paraná basin, Brazil Iliophagus Barbieri (1995)
Rio Frijoles, Panama Periphytivorous Power (1983)
Rio Tocantins basin, Brazil Omnivorous Rapp Py-Daniel and Py-Daniel (1984)
Rio Araguaia basin, Brazil Detritivorous Melo et al. (2004)
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Table 3 Trophic categories and items ingested according to literature for Loricariidae species

Trophic category Items ingested Source

Algivorous Algae and organic matter Casatti and Castro (1998)

Algivorous Green slurry, presumably comprised primarily of algae Nonogaki et al. (2007)
Detritivorous Vegetal organic matter, sediment and insect larvae Abelha et al. (2005)
Detritivorous Detritus and sediment and insect larvae Agostinho et al. (1997)
Detritivorous Detritus and sediments da Luz et al. 2001)
Detritivorous Detritus Mérona and Rankin-de-Mérona 2004)
Detritivorous Fine material from the bottom, mud, organic pellicle and sediment Mérona et al. (2001)
Detritivorous Non-particulated detritus, invertebrate excrements and rests Dourado et al. (2005)
Detritivorous Unicellular and filamentous algae, sand Ferreira (2007)
Detritivorous Moderate-degree dissociated vegetal matter and invertebrate fragments Hahn et al. (1998)
Detritivorous Organic and inorganic detritus Hahn et al. (2004)
Detritivorous Mainly detritus Loureiro-Crippa and Hahn (2006)
Detritivorous Detritus, algae, terrestrial plants, aquatic insects and aquatic plants Luz-Agostinho et al. (2006)
Detritivorous Detritus, algae, flowers, seeds and other materials Melo et al. (2004)
Detritivorous Organic detritus and its associated fungi and microorganisms Resende (2000)
Epibenthic consumers Algae and detritus Flecker (1992)
Generalist herbivorous Algae, insects larvae, crustaceans and macrophytes Buck and Sazima (1995)
Granivorous Seeds Armbruster (2002)
Herbivorous Algae Castro and Casatti (1997)
Herbivorous Wood Nelson (2002)
Iliophagous Sediment, vegetal organic matter, algae and sponges Angelescu and Gneri (1949)
Iliophagous Organic matter from mud on its premineralization phase Barbieri (1995)
Iliophagous Organic detritus, sediment and few periphytic organisms Delariva and Agostinho (2001)
Iliophagous Mud, high-degree dissociated organic matter and microorganisms Hahn et al. (1998)
Iliophagous Algae, vegetal material not identified and sand Rapp Py-Daniel and Py-Daniel (1984)
Omnivorous Fruits, seeds, insects, aquatic invertebrates and detritus Mérona and Rankin-de-Mérona (2004)
Omnivorous Insects, algae and sediment Rapp Py-Daniel and Py-Daniel (1984)
Periphytivorous Periphyton Power (1983, 1984); Casatti (2002)
Periphytivorous Organic matter, algae and Protozoa Uieda et al. (1997)
Specialized detritivorous Detritus in association with fungi and bacteria Yossa and Araújo-Lima (1998)
Specialized herbivorous Periphyton, Gastropoda, aquatic larvae and nymphs Andrade and Braga (2005b)
Xilophagous Wood Nonogaki et al. (2007)

sediment and detritus, all of them, except Rhinelepis aspera,
showed a defined stomach. The authors related this fact to
the diet of this species, composed mainly of organic matter.
They also agreed with Verigina (1990) who stated that
the intestine is generally more developed in species with
less-defined stomachs. However, a less-defined stomach
associated with a well-developed intestine was not observed
in L. lentiginosa or in some other Loricariidae species (Ange-
lescu and Gneri 1949; Rapp Py-Daniel and Py-Daniel
1984), probably because their diets were based essentially on
small animal items.
The digestive tract morphology of L. lentiginosa allows this
species to explore trophic resources associated with the
substratum. These features include the ventral position and
the anatomical shape of the mouth; the presence of teeth in
the dentary bone and upper lip that help it to scrape; and
well-developed pharyngeal teeth, responsible for triturating
and tearing food items apart before their ingestion (Ange-
lescu and Gneri 1949). This species also has numerous short
and slightly spaced gill rakers, structures associated with the
selection of the ingested food (Lammens and Hoogen-
boezem 1991; Delariva and Agostinho 2001).
The higher frequency and lower volume percentages of
organic matter ingested by L. lentiginosa compared to the
Bivalvia ingested may indicate an incidental consumption of
this item associated with other more valuable nutritional
resources, likewise the ingestion of sediment. Delariva and
Agostinho (2001) also report a similar pattern, involving
algae and other food resources for Loricariidae species.
Hahn et al. (1998) among others stated that in the upper
Paraná River basin, the congener L. prolixa feeds mostly on
detritus, but it also ingests aquatic invertebrates as a food
resource. Andrade and Braga (2005b), in a broad study on
the fish fauna in Porto Colômbia reservoir utilizing only
frequency data, categorized L. lentiginosa as specialized
herbivorous, ingesting periphyton as the main resource, and
gastropods and bivalves in smaller amounts. Our results,
however, point in another direction because the low Intestinal
Quotient values obtained would not be suitable to a diet
composed primarily of vegetal matter. According to

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106 Journal compilation © 2008 The Royal Swedish Academy of Sciences
Acta Zoologica (Stockholm) 90: 101–109 (April 2009)
Junger et al. (1989), short intestines indicate a tendency to
carnivory, as observed in the present study, as well as in the
studies of Angelescu and Gneri (1949), Rapp Py-Daniel
and Py-Daniel (1984) and, to some extent, Delariva and
Agostinho (2001).
The characteristics of the digestive tract as well as the
dietary pattern of L. lentiginosa, suggest that the species is
able to explore some bottom trophic resources, being classified
as benthophagous with a tendency to malacophagy, and
utilizing organic matter as an associated or additional food
resource.
Much of the literature concerning the feeding habits
of Loricariidae classifies these species as detritivorous/
iliophagous or herbivorous, usually able to exploit a particular
vegetal resource (Table 2). In fact, although detritivory/
iliophagy has been mentioned in all the Loricariidae sub-
families studied, this feeding habit is more related to Hypo-
stominae species. On the other hand, most of Ancistrinae
species are primarily herbivorous, feeding on items such as
algae or wood. In this group, only Hemiancistrus vittatus
(Merona et al. 2001) and Megalancistrus aculeatus (Delariva
and Agostinho 2001; among others) have been noticed to use
mostly detritus. Of the Hypoptopomatinae species found in
the literature, only Parotocinclus britski is reported as detritiv-
orous (Melo et al. 2004). In the Loricariinae, most species
are classified as detritivorous/iliophagous or omnivorous,
with some of them feeding mainly on small invertebrates
(Angelescu and Gneri 1949; Rapp Py-Daniel and Py-Daniel
1984; present study). For the two other subfamilies, Neople-
costominae and Lithogeneinae, no information was found
on their species feeding habits.
Some of the trophic categories into which Loricariidae
species have been inserted (Table 3) are subjective and
overlap each other to some degree. The detritivorous and
iliophagous categories, besides detritus, may encompass
other items such as algae (Rapp Py-Daniel and Py-Daniel
1984; Abelha et al. 2005); insect larvae (Agostinho et al.
1997; Luz-Agostinho et al. 2006); invertebrate excrement
(Dourado et al. 2005); sponges (Angelescu and Gneri 1949);
flowers and seeds (Melo et al. 2004) as well as sediment
(Angelescu and Gneri 1949; Rapp Py-Daniel and Py-Daniel
1984; Agostinho et al. 1997; da Luz et al. 2001; Delariva and
Agostinho 2001; Mérona et al. 2001; Abelha et al. 2005).
The herbivorous species are normally divided into more
specific categories related to one resource such as algivorous
(Casatti and Castro 1998; Nonogaki et al. 2007), peri-
phytivorous (Power 1983, 1984; Uieda et al. 1997; Casatti
2002), granivorous (Armbruster 2002) and wood eating/
xilophagous (Schaefer and Stewart 1993; Nelson et al. 1999;
Nonogaki et al. 2007). One specific food item is reported for
some species that are labelled as herbivorous in a broad
sense, as found in Castro and Casatti (1997) and Nelson
(2002).
On the other hand, the so-called omnivorous species
ingest several items, from algae to molluscs and crustaceans,
© 2008 The Authors
Journal compilation © 2008 The Royal Swedish Academy of Sciences
Salvador-Jr et al. • Feeding of Loricaria lentiginosa
using also detritus, sediment, plants and insects as trophic
resources (Angelescu and Gneri 1949; Rapp Py-Daniel and
Py-Daniel 1984; Merona and Rankin-de-Mérona 2004).
Our results associated with those found in the literature
about feeding habits and functional morphology of loricariids,
show that these species are able to use not only algae and
detritus, but also plant material and invertebrate items. This
ability to exploit a great variety of food resources is certainly
one of the main reasons for the evolutionary success of this
family in the colonization of Neotropical continental waters.
Acknowledgements
The authors would like to thank Furnas Centrais Elétricas
S.A, especially the staff of the Hatchery and Hydrobiological
Station for data collection and logistical support; CEMIG
for providing lodgings; R. J. Guimarães-Cruz (PUC-MG)
for collecting the digestive tracts; P. S. Pompeu (UFLA) for
comments and suggestions; I. Sazima (UNICAMP) and
C. T. Salomon (Cornell University, NY) for helping with the
references requested; and the staff of NUPELIA (UEM)
for their valuable support in the identification of doubtful
food items.
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