JOURNAL OF MORPHOLOGY 00:00–00 (2013)
Functional Morphology of the Andean Climbing
Catfishes (Astroblepidae, Siluriformes): Alternative
Ways of Respiration, Adhesion, and Locomotion
using the Mouth
Wannes De Crop,1* Elin Pauwels,2 Luc Van Hoorebeke,2 and Tom Geerinckx1
1
Evolutionary Morphology of Vertebrates, Ghent University–UGent, K.L. Ledeganckstraat 35, 9000 Ghent, Belgium
2
UGCT, Department of Physics and Astronomy, Ghent University–UGent, Proeftuinstraat 86, 9000 Ghent, Belgium
ABSTRACT Astroblepidae or “climbing catfishes”
encompass a single genus of species living in high
altitude rivers in the Andes of South America. They
are characterized by a specialized head morphology
closely resembling their better known, widely radiated
sister family Loricariidae, or armored suckermouth
catfishes. Existent data show that even though both
families share important traits, there are some striking
differences as well. Albeit poorly known, Astroblepus
species possess a duplicated gill opening, and have the
ability to climb vertical rocks or waterfalls. In this
study, morphological and kinematic data are combined
to yield insights into the functions of the mobile
elements of the astroblepid head, and to compare head
morphology and biomechanics with those of Loricarii-
dae. We found that, even though there is substantial
similarity in head structure of both families, there are
major differences in functionally important structures.
These include a different lower lip muscle configura-
tion, an alternative oral valve system, and an incurrent
gill opening only found in astroblepids. Kinematic anal-
yses confirm that the astroblepid suckermouth, freed
from its inhalatory function, offers advantages for
climbing in the high-altitude environment, and is used
alternately with the extremely mobile pelvic girdle, in
a crawling, nonundulatory motion. J. Morphol.
000:000–000, 2013. VC 2013 Wiley Periodicals, Inc.
KEY WORDS: respiration; locomotion; head morphology;
kinematics
INTRODUCTION
Various taxa of benthic rheophilic fishes have
acquired anatomical modifications allowing them
to maintain position in fast flowing riverine envi-
ronments. Examples of these structures include a
dorsoventrally flattened body, ventral odontodes
directed backward (Hora, 1930), and attachment
structures formed by the fins or the mouth (Nelson,
2006; Gerstner, 2007). Among catfishes, a ventrally
placed suckermouth evolved independently within
the paleotropical family Mochokidae and in the
neotropical families Astroblepidae and Loricariidae.
The outcome of evolutionary transformations of
the ancestral morphology toward an adhering
C 2013 WILEY PERIODICALS, INC.
V
structure is heavily influenced by the presence
and configuration of the available anatomical ele-
ments. This becomes especially interesting when
comparing the attachment mechanisms in closely
related families with common ancestral morpholo-
gies. Loricariidae and Astroblepidae are sister
families in the basal siluriform clade Loricarioidea
(Sullivan et al., 2006). At first sight, their
morphology is similar: bodies are dorsoventrally
flattened, the paired fins are positioned ventrally,
and the ventral mouth with expanded lips forms a
suction disk. Both are benthic feeders, with lori-
cariids feeding mainly on plant material whereas
astroblepids are omnivorous (Pouilly et al., 2006).
These sister families inhabit fast flowing rivers in
South American basins, with some overlap in
spatial niche occupation. Astroblepids, however,
are restricted to the higher altitude Andean
waters (Nelson, 2006).
In contrast to Loricariidae, the largest of the
siluriform families and quite diverse in body
shapes and sizes, sister group Astroblepidae is a
more modest family, including about 54 species in
the single genus Astroblepus (Schaefer et al.,
2011). They are known for their amazing ability to
climb along vertical rock faces of waterfalls using
their mouth and pelvic fins in an alternating
pattern (Johnson, 1912). Although astroblepid
ecology, especially in relation to conservation, has
been studied in the past (Rom an-Valencia, 2001;
Contract grant sponsor: Ghent University; Grant number: GOA
01G01008.
*Correspondence to: Evolutionary Morphology of Vertebrates,
Ghent University–UGent, K.L. Ledeganckstraat 35, Ghent 9000,
Belgium. E-mail: wannes.decrop@ugent.be
Received 7 December 2012; Revised 8 April 2013;
Accepted 26 April 2013.
Published online in
Wiley Online Library (wileyonlinelibrary.com).
DOI 10.1002/jmor.20169
2 W. DE CROP ET AL.

Fig. 1. The head and pectoral fins of Astroblepus sp. “Peru” (MUSM 46925 nr. 1, 106-mm SL) during the inhalation phase while
being attached. A: Dorsal view. Note that the incurrent gill openings (indicated by arrows) are opened and the excurrent gill open-
ings are closed; B: Ventral view. Scale bars 5 1 cm. Points of interest: a, medioventral point representing hyoid bar; b, suspensorium;
c, first pectoral fin ray; d, caudal tip of opercular; e, valve closing the dorsal incurrent gill slit; f, caudalmost point of branchiostegal
membrane; g, upper jaw; h, lower jaw; i, lower lip; j, upper oral valve.

Velez-Espino, 2003a,b,2004, 2006; Ortega et al.,
2007; Schaefer and Arroyave, 2010), many aspects
of their biology remain unknown. A handful of
articles comprise most of what is known about the
cranial morphology of astroblepid catfishes, even
though their main focus lies elsewhere (Howes,
1983; Schaefer and Lauder, 1986, 1996; Schaefer,
1988, 1990; Arratia, 1990).
Many characters of the astroblepid-loricariid
clade evolved before the two families separated
(e.g., Schaefer and Lauder, 1986; Schaefer, 1988).
Astroblepids lack some muscular specializations
found in Loricariidae (Howes, 1983); they also
lack the bony armour of loricariids. They do, how-
ever, possess a duplicated gill opening (Fig. 1A;
Johnson, 1912), which in turn does not occur in
loricariids. In this study, we investigate how the
presence of the novel incurrent gill opening in
astroblepids relates to a new functional mecha-
nism of respiration during adhesion, which might
facilitate climbing. To achieve this, a thorough
examination of the astroblepid morphology is
essential to fill a gap in siluriform comparative
knowledge and deliver a solid comparison between
both families. Because no electromyography or cin-
eradiography was performed, inferences on muscle
functions remain hypothetical. Still, the performed
kinematic analysis allows us to propose functional
theories regarding astroblepid respiration and
attachment mechanisms.
MATERIAL AND METHODS
Astroblepus sp. specimens were collected at night using a
cast net in the Rıo Acuzaz
u, a tributary of Rıo Chorobamba in
the community of Santa Clara (10 310 4.658400 S, 75 240 56.030400
W, ca. 1800 m altitude), Pasco, Peru. Attempts of identification
indicate that these specimens most probably belong to morpho-
species F as described by Schaefer et al. (2011). In total, 29-
numbered specimens were collected and will be referred to as
Astroblepus sp. “Peru.” Twenty-specimens are deposited in the
Museo de Historia Natural of the Universidad Nacional Mayor
de San Marcos, Lima, Peru (lot MUSM 46925), while nine are
kept at the research group Evolutionary Morphology of Verte-
brates at Ghent University (further abbreviated as EVM).
Apart from the main study on these specimens from Peru, a
limited comparative osteological study was performed on one
specimen from Bolivia (Astroblepus sp. “Bolivia”; Zoological
Museum Amsterdam (ZMA) 119.552; Rio Espıritu Santo near
Cochabamba) and one from Colombia (Astroblepus cf. longifilis;
ZMA 111.43; Est. Caldas, Cecilia).
High-speed recordings (at 120 frames per second) of Astroble-
pus sp. “Peru” (nr. 1, 106 mm-standard length (SL); nr. 3, 147-
mm SL; nr. 5, 123-mm SL of MUSM 46925; and nr. 13, 136-mm
SL at EVM) were performed using an aquarium setup and a
Casio Exilim EX-FH100 high-speed camera at the Instituto del
Bien Com un, Oxapampa, Peru. Individuals were recorded from
different angles during two main modes of respiration. The first
mode involved respiration with suckermouth attachment on the
aquarium floor or wall. The second mode involved unattached
respiration, with the specimens resting on the aquarium floor
without attachment of the suckermouth. Water inflow and out-
flow was visualized by the motion of minute suspended par-
ticles in the water. Additional recordings allowed analysis of
crawling locomotion.
The main anatomical points of interest (Fig. 1) are a medio-
ventral point representing the hyoid bar, the lateral and dorsal
side of the suspensorium, the first pectoral fin ray, the caudal
tip of the opercular, the medial edge of the valve closing the
dorsal incurrent gill slit, the caudalmost point of the branchios-
tegal membrane, upper and lower jaws, the lower lip and the
midline of the upper oral valve, with two additional reference
points on the neurocranium. These points were digitized in the
first of every two frames using a marker tracking and direct
linear transformation program (Hedrick, 2008) in Matlab
R2011b (The Mathworks). Data were filtered using a fourth
order zero-phase shift butterworth low-pass data noise filter
with a cut-off frequency of 7 Hz (add-in for Excel written by
Sam Van Wassenbergh; procedure in Erer, 2007). Statistical dif-
ferences between inhalation and expiration phase durations
were tested using PopTools 3.2 (Monte-Carlo Analysis, 10,000

Journal of Morphology
 MORPHOLOGY OF ASTROBLEPID CATFISHES
replicates, Greg Hood, Australia) on Excel 2010 (Microsoft).
This approach uses a randomized permutation procedure with
the null hypothesis stating that the distance (difference)
between two groups of variables (phase durations) does not
differ from the distance between any pair of groups with
random inclusion of the variables. The P-value (the relative
number of times the distance is in the upper 0.975 percentile of
replicates; Collyer et al., 2007) can then be calculated from the
result. Since it was not possible to follow the exact same point
on a particular anatomical element in different film fragments
(due to recording angle, shade or specimen coloration), absolute
measurements of excursion amplitudes of these structures were
not compared between different movie sequences.
For the study of anatomy, Astroblepus sp. “Peru” specimens
were first anesthetized in a MS 222 solution (ethyl 3-
aminobenzoic acid methanesulfonate salt, Sigma Chemical),
and then euthanized by MS 222 overdose. Specimens were fixed
using a 4% buffered formalin solution (at neutral pH), followed
by 70% ethanol for later storage. Dissections were performed
on Astroblepus sp. “Peru” (nr. 8, 89-mm SL; nr. 16, 83-mm SL;
nr. 17, 69-mm SL; nr. 19, 105-mm SL; nr. 29, 100-mm SL, at
EVM), using an Olympus SZX 9 stereo microscope. Serial sec-
tions (5 mm) of the head of one Astroblepus sp. “Peru” specimen
(nr. 14, 57-mm SL at EVM) were made with a Leica SM2500
sliding microtome, and examined with a Reichert-Jung Polyvar
light microscope. One Astroblepus sp. “Peru” specimen (nr. 19,
105-mm SL at EVM) was used for in toto clearing and staining
(protocol based on Taylor and Van Dyke, 1985). The modular
micro-CT setup at the “Centre for X-ray Tomography” of the
Ghent University (UGCT; Masschaele et al., 2007) was used for
a CT scan of the head of a specimen of Astroblepus sp. “Peru”
(nr. 15, 89-mm SL at EVM). The specimen was scanned using a
dual-head X-ray tube from Feinfocus (FXE 160) with a direc-
tional high-power head, at 100-kV tube voltage. The detector
was an aSi flat panel (PerkinElmer XRD 1620 CN3 CS) with a
CsI scintillator. A number of 1,441 projection images of 1,648
by 2008 pixels were recorded, covering 360 , with a total expo-
sure time of 2 s per projection. The isotropic voxel size of the
sample was 183 mm3. The raw data were processed and recon-
structed using the CT software Octopus (Vlassenbroeck et al.,
2007) and rendered using VGStudio Max 2.0 (Volume Graphics,
Germany). Osteological data were extracted from these three-
dimensional scans. Both Astroblepus sp. “Bolivia” (ZMA
119.552, 66-mm SL) and Astroblepus cf. longifilis (ZMA 111.43,
159-mm SL) were scanned with similar settings.
RESULTS
Anatomical elements described herein are paired
unless mentioned otherwise. Only those structures
relevant to the functioning of the feeding, respira-
tion, or climbing apparatus will be described in
detail. All results, tables, and figures are based on
Peruvian specimens.
Osteology
Neurocranium. The neurocranium (Figs. 2
and 3) is horizontally flattened, resulting in a
broad skull. Anteriorly, the single mesethmoid
bears a ventral protuberance, to which the upper
jaws connect. The frontal forms a large part of the
skull roof, flanked by the lateral ethmoid, which
provides a prominently broad articulation facet for
the autopalatine, and forms part of the skull floor
and wall, along with the orbitosphenoid and
pterosphenoid. The medial portion of the skull
floor is formed by the unpaired, toothless pre-
vomer and parasphenoid, both exhibiting a ventral
3
mediosagittal ridge. More posteriorly the skull is
broader and is composed out of the sphenotic and
parieto-supraoccipital dorsally and the prootic,
parasphenoid, basioccipital, and exoccipital ven-
trally. In addition the compound pterotic (see
Geerinckx et al., 2007) makes up a large part of
the caudolateral skull.
Splanchnocranium. The medial process of
the upper jaw or premaxilla (Figs. 2 and 3)
articulates with the rostral tip of the mesethmoid
via the mesethmoid-premaxillary cartilage, a
meniscus-like structure. Additional ligaments con-
tribute to the connection between the premaxilla
and the mesethmoid. Left and right premaxillae
are connected medially by a small meniscus-like
cartilage. The premaxilla is located ventrally to
the autopalatine head. This cartilaginous autopa-
latine head articulates with the basis of the elon-
gated maxilla (Figs. 2 and 3), which supports the
maxillary cartilages. Caudally, the autopalatine
(Figs. 2 and 3B) bears two processes ventral to the
articulation facet with the lateral ethmoid.
In the lower jaw (Figs. 2 and 3A), the proximal
anguloarticular and distal dento-mentomeckelium
are connected by the remnant of Meckel’s cartilage
caudally, and suture and partly overlap more
rostrally. Both bones contribute to a plate-like
coronoid process. No symphysis is present; left and
right lower jaws are separated by a cartilaginous
plug. Laterally, the anguloarticular of the
lower jaw articulates with the quadrate of the
suspensorium.
On the upper jaw, a first row of large, predomi-
nantly unicuspid teeth (Fig. 3A) is followed by
smaller bicuspid teeth. In the lower jaw, the large
teeth and posterior smaller teeth are all bicuspid,
the large ones asymmetrically so. The jaws
bear incompletely mineralized replacement teeth
of each tooth type. Additional unicuspid teeth are
present on the pharyngeal jaws, formed by the
fifth branchial arch.
The suspensorium (Figs. 2 and 3B) is formed by
the quadrate, hyomandibular, and metapterygoid.
These bones are centrally connected by narrow
strips of symplectic cartilage. The autopalatine-
metapterygoid ligament connects the metaptery-
goid and quadrate to the autopalatine and the
lateral ethmoid. Laterally, the preopercular firmly
attaches to the hyomandibular and the quadrate,
adding rigidity to the suspensorium. The hyoman-
dibular articulates with the neurocranium at the
junction of pterosphenoid, sphenotic, and prootic.
The hyoid (Figs. 2B and 3A) is composed of
three paired elements, being the hypohyal, ante-
rior ceratohyal and posterior ceratohyal, synchon-
drally contacting each other. On their ventral side
the latter two bones share a cartilaginous process
articulating with four branchiostegal rays support-
ing the branchiostegal membrane. Dorsally the
posterior ceratohyal is also connected with the
Journal of Morphology
4 W. DE CROP ET AL.

Fig. 2. The skull of Astroblepus sp. “Peru” (nr. 8, 89-mm SL at EVM). A: Dorsal view. B: Lateral view. Scale bars 5 1 mm. comp-pt,
compound pterotic bone; o-aa, os anguloarticulare; o-apal, os autopalatinum; o-cb, os ceratobranchiale; o-ch-a, os ceratohyale anterior;
o-ch-p, os ceratohyale posterior; o-cl, os cleithrum; o-den-m, os dentomentomeckelium; o-fr, os frontale; o-hh, os hypohyale; o-hm, os
hyomandibulare; o-int-h, os interhyale; o-int-op, os interoperculare; o-leth, os latero-ethmoideum; o-mes, os mesethmoideum; o-mpt,
os metapterygoideum; o-mx, os maxillare; o-nas, os nasale; o-op, os operculare; o-osph, os orbitosphenoideum; o-para, os parasphenoi-
deum; o-par-soc, os parieto-supraoccipitale; o-pmx, os praemaxillare; o-pop, os praeoperculare; o-psph, os pterosphenoideum; o-pvm,
os praevomerale; o-q, os quadratum; o-sph, os sphenoticum; r-br-I–IV, radius branchiostegus I–IV; tr-pr-c-v, transverse process of
complex vertebra.

ventral side of the preopercular and the quadrate
through a ligament (laterally) and a cartilaginous
articulation (medially). Medially, both hypohyals
are connected by their ligamentous contact to the
unpaired parurohyal. This sesamoid bone bears
two rostral processes, each fitting in a circular
cavity of a hypohyal.
The opercular series (Fig. 2B) comprises the
opercular, preopercular, and two subopercular
elements. Of these, the dorsal one is probably
homologous with the interopercular; the ventral
one probably is homologous with the interhyal
(Schaefer, 1988). The latter bone articulates
with the posterior ceratohyal. A medioventral
articulation facet of the opercular connects to the
hyomandibular. No relevant osteological differen-
ces were observed between Astroblepus sp. “Peru”
and the examined specimens of Astroblepus cf.
longifilis and Astroblepus sp. “Bolivia.”
Myology
Jaw and maxillary barbel musculature.
The adductor mandibulae (Figs. 4 and 5A) originates
on the dorsal face of the hyomandibular and preoper-
cular. A relatively small medial bundle stretches
forward and inserts on the dorsocaudal face of the
anguloarticular and dento-mentomeckelium. It is
covered by a larger, dorsolateral bundle inserting on

Journal of Morphology
 MORPHOLOGY OF ASTROBLEPID CATFISHES 5

Fig. 3. The skull of Astroblepus sp. “Peru” (nr. 8, 89-mm SL at EVM). A: Ventral view. B: Ventral view of the neurocranium, the
suspensorium, the opercular, the autopalatine and the maxilla. Scale bars 5 1 mm. comp-pt, compound pterotic bone; o-aa, os angu-
loarticulare; o-apal, os autopalatinum; o-boc, os basioccipitale; o-cb, os ceratobranchiale; o-ch-a, os ceratohyale anterior; o-ch-p, os
ceratohyale posterior; o-cl, os cleithrum; o-den-m, os dentomentomeckelium; o-exoc, os exoccipitale; o-hh, os hypohyale; o-hm, os
hyomandibulare; o-int-h, os interhyale; o-int-op, os interoperculare; o-leth, os latero-ethmoideum; o-mes, os mesethmoideum; o-mpt,
os metapterygoideum; o-mx, os maxillare; o-op, os operculare; o-osph, os orbitosphenoideum; o-para, os parasphenoideum; o-pmx, os
praemaxillare; o-pop, os praeoperculare; o-prot, os prooticum; o-puh, os parurohyale; o-pvm, os praevomerale; o-q, os quadratum;
r-br-I–IV, radius branchiostegus I–IV; tr-pr-c-v, transverse process of complex vertebra.

the coronoid process. Manipulation of the adductor
mandibulae, by experimentally pulling it during
dissection (before specimen fixation), results in an
upward and rotating motion of the lower jaw, pivot-
ing the tooth-bearing part anteriorly, in the direction
of the mouth cavity.
More medially the retractor premaxillae (Figs. 4
and 5C) originates on the hyomandibular and
metapterygoid and inserts upon the dorsolateral
premaxillary process with a tendon that continues
as a ventromedial aponeurosis in the anterior
portion of the muscle. When manipulated, the

Journal of Morphology
6 W. DE CROP ET AL.

Fig. 4. Dorsal view of the cranial musculature of Astroblepus sp. “Peru” (based on dissections) drawn upon CT images of Astroble-
pus sp. “Peru” (nr. 8, 89-mm SL at EVM). A: left: all superficial muscles; right: m-add-md and m-lev-tt are removed. B: left: m-add-
md-e and m-lev-tt are removed; right: m-add-md, m-lev-tt and m-rt-pmx are removed. Red 5 muscle tissue; Green 5 tendon. Not
shown: m-add-hh. Dashed circles indicate the position of the eyes. Scale bars 5 1mm. m-add-apal-d, musculus adductor arcus palatini
pars dorsalis; m-add-md, musculus adductor mandibulae; m-add-md-e, external part of musculus adductor mandibulae; m-add-md-i,
internal part of musculus adductor mandibulae; m-add-op, musculus adductor operculi; m-dil-op, musculus dilatator operculi; m-ext-
tt-l, musculus extensor tentaculi pars lateralis; m-intm-p-d, musculus intermandibularis posterior pars dentalis; m-lev-apal, musculus
levator arcus palatini; m-lev-op, musculus levator operculi; m-lev-tt, musculus levator tentaculi; m-rt-pmx, musculus retractor
premaxillae.

premaxilla is rotated posteriorly, pivoting the teeth
inwards.
From the lateral ethmoid, the levator tentaculi
(Figs. 4A and 5A) stretches toward the proximal
part of the maxilla. It bifurcates near its anterior
end. The medial bundle inserts dorsally upon the
base of the maxilla, whereas the lateral bundle
inserts on the more ventrally placed basal process.
Manipulating this muscle pulls the maxillary bone
in a dorsocaudal direction. Given the absence of
the levator tentaculi in most related families
(except Loricariidae), its innervation has been
examined. The branchlet innervating this muscle
fuses with that of the retractor premaxillae and
then with the other branchlets innervating the
adductor mandibulae. The whole branch then joins
the complex formed by the buccomaxillary and
mandibular nerve branches.
The extensor tentaculi is divided in two separate
elements. Laterally, the pennate extensor tentaculi

Journal of Morphology
 MORPHOLOGY OF ASTROBLEPID CATFISHES 7

Fig. 5. Lateral view of the left cranial musculature of Astroblepus sp. “Peru” (based on dissections) drawn upon CT images
of Astroblepus sp. “Peru” (nr. 8, 89-mm SL at EVM). A: All superficial muscles. B: m-add-md, m-lev-tt and m-rt-pmx are removed.
C: m-intm-a-l and m-hh-add-br are shown (not in A and B), m-add-md, m-lev-tt and intm-p-d are removed; Red 5 muscle tissue;
Green 5 tendon. Not shown: m-hh-abd, m-hh-inf and all muscles on the right. Dashed circles indicate the position of the eyes. Scale
bars 5 1mm. m-add-apal-d, musculus adductor arcus palatini pars dorsalis; m-add-md, musculus adductor mandibulae; m-add-md-e,
external part of musculus adductor mandibulae; m-add-op, musculus adductor operculi; m-dil-op, musculus dilatator operculi;
m-ext-tt-l, musculus extensor tentaculi pars lateralis; m-ext-tt-m, musculus extensor tentaculi pars medialis; m-hh-add, musculus
hyohyoidei adductores (attached to branchiostegal rays and opercular); m-hh-add-br, part of musculus hyohyoidei adductores in bran-
chiostegal membrane edge; m-intm-a-d, musculus intermandibularis anterior pars dentalis; m-intm-a-l, musculus intermandibularis
anterior pars labialis; m-intm-p-d, musculus intermandibularis posterior pars dentalis; m-intm-p-l, musculus intermandibularis
posterior pars labialis; m-lev-apal, musculus levator arcus palatini; m-lev-op, musculus levator operculi; m-lev-tt, musculus levator
tentaculi; m-rt-pmx, musculus retractor premaxillae.

pars lateralis (Figs. 4, 5B, and 6C) originates on Manipulation forces the anterior tip of the autopa-
the lateral side of the lateral ethmoid and inserts latine in a lateroventral direction. The extensor
on the ventrolateral process of the autopalatine. tentaculi pars medialis (Fig. 6C) originates on the

Journal of Morphology
8 W. DE CROP ET AL.

Fig. 6. Ventral view of the cranial musculature (and m-hypaxialis and m-pr-ischii) of Astroblepus sp. “Peru” (based on dissections)
drawn upon CT images of Astroblepus sp. “Peru” (nr. 8, 89-mm SL at EVM). A: m-ext-tt-m not shown; left: m-intm-a-d is removed;
right: Mouth disk and m-intm-a-d are shown. B: m-ext-tt-m not shown; left: m-intm-a-d, m-hh-inf, m-hh-abd and m-hypaxialis are
removed; right: m-hh-inf is removed, mouth disk is shown. C: Only m-ext-tt-l, m-ext-tt-m, m-add-apal-d, m-add-apal-v and m-add-op
are shown with their insertions on the neurocranium, suspensorium, operculum and autopalatine; left: m-add-apal-v is removed;
right: m-add-apal-v is shown. Red 5 muscle tissue; Green 5 tendon; Blue 5 cartilage plug. Not shown: m-intm-a-l, m-hh-add and
m-hh-add-br. Dashed lines indicate position underneath bone or mouth disk. Scale bars 5 1mm. crt-p, cartilage plug; m-add-apal-d,
musculus adductor arcus palatini pars dorsalis; m-add-apal-v, musculus adductor arcus palatini pars ventralis; m-add-md-e, external
part of musculus adductor mandibulae; m-add-op, musculus adductor operculi; m-ext-tt-l, musculus extensor tentaculi pars lateralis;
m-ext-tt-m, musculus extensor tentaculi pars medialis; m-hh-abd, musculus hyohyoideus abductor; m-hh-inf, musculus hyohyoideus
inferior; m-hypaxialis, musculus hypaxialis; m-intm-a-d, musculus intermandibularis anterior pars dentalis; m-intm-p-d, musculus
intermandibularis posterior pars dentalis; m-intm-p-l, musculus intermandibularis posterior pars labialis; m-lev-tt, musculus levator
tentaculi; m-pr-ischii, musculus protractor ischii; m-rt-pmx, musculus retractor premaxillae; m-stern, musculus sternohyoideus.

Journal of Morphology
 MORPHOLOGY OF ASTROBLEPID CATFISHES
ventral side of the lateral ethmoid, the orbitosphe-
noid and the parasphenoid ridge and anteriorly
attaches onto the ventromedial process of the
autopalatine, with a broad, ventral tendon. This
muscle is hard to manipulate, but its orientation
strongly suggests that contraction would swing
the anterior tip of the autopalatine medioventrally.
Suspensorial musculature. The adductor
arcus palatini (Figs. 5B and 6C) is subdivided in
two parts. The upper part originates on the lateral
ethmoid and orbitosphenoid, while the lower part
originates ventrally on the parasphenoid. Both
parts insert upon the suspensorium; however, the
upper part inserts on the medial edge of both the
metapterygoid and the hyomandibular, whereas
the lower part inserts on the ventral side of the
hyomandibular. While neither of the two can be
manipulated, the only probable outcome of con-
traction is adduction of the suspensorium to a
more medial position.
Dorsally on the head, the levator arcus palatini
(Fig. 4) reaches from the frontal, pterosphenoid
and sphenotic toward the mediocaudal process on
the hyomandibular. Manipulation of this muscle
has no visible effect.
Opercular musculature. The dilatator oper-
culi (Fig. 4) originates on the sphenotic. Its narrow
tendon inserts on the opercular, anterior to the
articulation with the hyomandibular. Due to its
small size, this muscle is hard to manipulate.
The adductor operculi (Figs. 4 and 6C) connects
the medioventral face of the opercular to the com-
pound pterotic of the neurocranium. Its only possi-
ble action thus is a medial motion of the opercular.
Interestingly, some of its (posterior) fibers are
somewhat curved around the dorsal incurrent gill
opening. A small group of dorsal fibers, inserting
marginally more dorsally, probably corresponds to
a (remnant of) the levator operculi.
Ventral musculature. The ventrally placed
intermandibularis muscles are divided in an ante-
rior and posterior group, each containing two parts.
In the anterior group the intermandibularis ante-
rior pars labialis (Fig. 5C) is located within the dor-
sal side of the posteriorly expanded lower lip, just
underneath the dermis. The muscle has a fan-like
appearance, with fibers spreading out centrifugally
to the peripheral rim of the lower lip disk.
Medially, the contralateral muscles are completely
separated from each other. The intermandibularis
anterior pars dentalis (Figs. 6A,B) connects the
dorsal side of the lower lip to the cartilage plug
between both lower jaws. Just posterior to the
plug, both contralateral muscles also interconnect
in a raphe. The muscle also partly connects to the
intermandibularis anterior pars labialis.
More posteriorly, both the intermandibularis
posterior pars dentalis (Figs. 5A,B, 6A, and B)
and the intermandibularis posterior pars labialis
(Figs. 5C, 6A, and B) originate on the ventral side
9
of the posterior ceratohyal. The pars dentalis runs
forward toward the lower jaw, attaching on the
dento-mentomeckelium. Manipulation results in
rotation and caudal movement of the lower jaw,
consequently directing the teeth outwards. The
intermandibularis posterior pars labialis firmly
inserts into the lower lip tissue. When the muscle
is pulled on, the base of the lower lip is drawn in
a caudolateral direction.
The ventral faces of left and right anterior and
posterior ceratohyals are interconnected by the
broad hyohyoideus inferior (Fig. 6A). Medially
both halves of this muscle meet and form a raphe.
Manipulation of this muscle is hard and has little
visible effect.
The sternohyoideus (Fig. 6B) connects the dorsal
side of the cleithrum of the pectoral girdle with
the parurohyal of the hyoid. Manipulation is hard,
and has no clear effect.
The hyohyoidei adductores (Fig. 5) form the
musculature of the branchiostegal membrane and
its supporting structures. Only the lateralmost
two rays are interconnected by three small muscle
bundles. From the lateral ray, two muscles run
upwards to the opercular. Embedded in the soft
tissue of the branchiostegal membrane itself, an
additional muscle runs along the entire edge of
the membrane. It is probably derived from the
hyohyoidei adductores. Dorsally it embeds into the
basal tissue of the valve closing the incurrent gill
opening. Manipulation of any of these muscle bun-
dles is not possible.
From the medialmost branchiostegal ray, the
hyohyoideus abductor (Fig. 6B) runs in a medio-
rostral direction and contacts the ventral face of
the sternohyoideus where it is continuous with the
hypaxial muscles. When the hyohyoideus abductor
is manipulated, the branchiostegal membrane is
closed.
The protractor pectoralis connects the compound
pterotic of the neurocranium to the cleithrum of
the pectoral girdle, caudolateral to the insertion of
the sternohyoideus.
Two pelvic muscles are prominently visible
(Fig. 7) through the ventral skin. The protractor
ischii (Figs. 6B and 7A) is round in cross-section,
with fibers in a spiraling orientation. It connects
the lateral process of the basipterygium to the
cleithrum of the pectoral girdle. When contracted
it pulls the pelvic girdle anteriorly. The retractor
ischii (Fig. 7B) connects the external process of the
basipterygium to the basal elements of the anal fin
skeleton and the surrounding body musculature.
Its fibers are not spiraled. Contraction results in a
caudal displacement of the pelvic girdle.
Oral Valves and Gill Slit
Two oral valves (Fig. 8) are present, directly
posterior to both upper and lower jaws. They are
Journal of Morphology
10 W. DE CROP ET AL.

Fig. 7. Ventral view of freshly euthanized Astroblepus sp. “Peru” (nr. 29, 100-mm SL at EVM).
A: The pelvic girdle (dashed line) is in its caudalmost position. Arrows indicate the musculus
protractor ischii. B: The pelvic girdle (dashed line) is in its rostralmost position. Arrows indicate
the musculus retractor ischii. Scale bars 5 1 cm.

positioned in such a way that when both the upper
and the lower jaw are adducted, the upper oral
valve overlaps the lower jaw and (to a lesser
extent) the lower oral valve, separating a preva-
lvular cavity from a postvalvular cavity. The small
prevalvular cavity is bordered by the tissue of the
lips and jaws, the oral valves and (when attached)
the substrate. The larger postvalvular cavity forms
the main buccal cavity and is continuous with the
gill chamber or opercular cavity. Dorsal to the
excurrent gill slit an incurrent gill opening (Fig. 1)
is found, covered by a dorsal skin fold of the
opercular.
Kinematics
Attached respiration. During attached respi-
ration individuals are attached to the aquarium
wall or floor using their suckermouth, thereby
disconnecting the external environment from the
prevalvular cavity, which is now a separate
compartment. The average duration (mean 6 s.d.)
of one respiration cycle varies among individuals

Fig. 8. Mediosagittal view of the transected head of Astroblepus sp. “Peru” (nr. 17, 69-mm SL
at EVM). Note the presence of the upper and lower oral valves. Scale bar 5 1 mm. pov-c, post-
valvular cavity; prv-c, prevalvular cavity.

Journal of Morphology
 MORPHOLOGY OF ASTROBLEPID CATFISHES 11
TABLE 1. Average duration 6 standard deviation (in ms) of one respiration cycle, the inhalation and the expiration phase (during
attached and unattached respiration) of each Astroblepus sp. “Peru” specimen used in the digital kinematic analysis.
Specimen nr., SL (mm) Respiration mode Full cycle Inhalation phase Expiration phase
1 (MUSM 46925), 106 attached 317 6 28 (15) 193 6 56 (17) 134 6 15 (16)
5 (MUSM 46925), 106 attached 394 6 34 (11) 243 6 24 (12) 144 6 23 (13)
13 (EVM), 136 attached 500 6 26 (10) 308 6 39 (10) 186 6 34 (13)
3 (MUSM 46925), 147 attached 378 6 9 (3) 222 6 19 (3) 156 6 10 (3)
3 (MUSM 46925), 147 unattached 350 6 51 (20) 189 6 38 (22) 153 6 13 (25)

Sample size is shown in parentheses.

(383 6 76 ms), but also within individuals (Table 1).
During attached respiration the inhalation phase
(222 6 52 ms) lasts significantly longer (P < 0.005)
than the expiration phase (167 6 38 ms). Elements
inducing the volume changes during these phases
are the suspensorium forming the buccal cavity
wall, and the hyoid and pectoral girdle of the
buccal cavity floor.
During inhalation the hyoid depresses caudo-
ventrally, whereas it elevates rostrodorsally during
expiration. The lower jaw appears to be stationary:
no substantial movement could be detected.
The hyoid movements are synchronous to those of
the pectoral girdle, with both caudoventral and
rostrodorsal peaks of the pectoral girdle occurring
3 6 17 ms after the depression and elevation maxi-
mum of the hyoid, respectively (Table 2). The sus-
pensorium (Fig. 9) moves near-synchronously with
the hyoid (21 6 16 ms and 3 6 23 ms after the
maximum depression and elevation of the hyoid,
respectively). The opercular moves more or less
together with the suspensorium, with its abduc-
tion and adduction maximum 14 6 30 ms and
29 6 38 ms before those of the suspensorium,
respectively. The dorsal incurrent gill opening
(Fig. 9) is closed 76 6 23 ms and completely opened
32 6 19 ms after the abduction and adduction max-
imum of the suspensorium, respectively. The lat-
eral, excurrent gill slit (Fig. 9) opens and closes
5 6 19 ms and 13 6 10 ms after the suspensorium
is most abducted and adducted, respectively.
During attached respiration, no inflow of water
through the mouth is observed. The only inflow is
through the incurrent gill opening. In only one
movie sequence of one specimen the upper oral
valve was sufficiently visible, as most specimens

TABLE 2. Relative timing averages 6 standard deviation (in ms) of some important movements during both respiration modes in
Astroblepus sp. “Peru.”
Attached respiration Unattached respiration
1 (MUSM 5 (MUSM 13 (EVM) 3 (MUSM 3 (MUSM
Specimen nr., SL (mm) 46925), 106 46925), 123 46925), 136 46925), 147 mean 46925), 147
Suspensorium abduction max. 18 6 11 (8) 24 6 9 (7) 21 6 25 (7) – 21 6 16 (22) 248 6 16 (13)
after hyoid depression max.
Suspensorium adduction max. 12 6 16 (7) 22 6 12 (7) 0 6 38 (6) – 3 6 23 (20) 243 6 14 (13)
after hyoid elevation max.
First pectoral fin ray caudal max. 2 6 14 (8) 5 6 13 (7) 22 6 19 (3) – 3 6 17 (18) 8 6 9 (10)
after hyoid depression max.
First pectoral fin ray rostral max. 2 6 6 (7) 29 6 9 (7) 33 6 1 (3) – 3 6 17 (17) 22 6 10 (9)
ray after hyoid elevation max.
Opercular abduction max. after 0 6 23 (4) 29 6 12 (2) 267 6 23 (2) 24 6 16 (4) 214 6 30 (12) 20 6 21 (11)
suspensorium abduction max.
Opercular adduction max. after 0 6 13 (4) 239 6 27 (6) 264 6 16 (6) 22 6 19 (3) 229 6 38 (19) 33 6 22 (14)
suspensorium adduction max.
Excurrent gill slit opened after 13 6 21 (4) 6 6 23 (6) 23 6 14 (5) – 5 6 19 (15) 2 6 19 (8)
suspensorium abduction max.
Expiratory gill slit closed after 0 6 0 (4) 17 6 1 (6) 17 6 10 (6) – 13 6 10 (16) 18 6 9 (10)
suspensorium adduction max.
Incurrent gill slit closed after 77 6 26 (11) 67 6 13 (4) – – 76 6 23 (15) 80 6 9 (4)
suspensorium abduction max.
Inhalatory gill slit opened after 32 6 21 (10) 34 6 1 (4) – – 32 6 19 (14) 22 6 10 (3)
suspensorium adduction max.
Upper oral valve closed after – – – – – 60 6 11 (13)
hyoid depression max.
Upper oral valve open after – – – – – 27 6 24 (13)
hyoid elevation max.
Sample size is shown in parentheses. For each of the four specimens included in the analysis of attached respiration separate tim-
ings are shown, as well as the average (mean) of these specimens together. Only MUSM 46925 nr. 3 (147-mm SL) was included in
the analysis of unattached respiration.

Journal of Morphology
12
Fig. 9. Graph showing movements of anatomical points of
interest in a 1,200 ms recording of Astroblepus sp. “Peru”
(MUSM 46925 nr. 1, 106-mm SL) in dorsal view during attached
respiration. Timing of the excurrent gill slit and the incurrent
gill slit being open are indicated by horizontal bars.
tended to keep upper and lower jaw close together
during attached respiration. No movement of it
was observed.
Unattached respiration. In the aquarium,
individuals tended to stay attached to the wall or
floor and showed little initiative to rest on the
aquarium floor unattached, swim substantial dis-
tances, or climb. Only one specimen (MUSM
46925 nr. 3, 147-mm SL) could actually be
recorded exhibiting both attached and unattached
respiration, and in this specific film fragment, not
all points of interest are visible. Moreover, digital
analysis of unattached respiration was only possi-
ble on movie sequences of this individual. Due to
this limited amount of data, it is not possible to
compare temporal data between attached and
unattached respiration, as variations could just as
well be the result of variation between specimens
rather than actual differences between respiration
modes.
During unattached respiration, the suckermouth
does not adhere to the substrate. The prevalvular
cavity is connected to the external environment as
the mouth cannot be closed using the lips or jaws.
For the filmed and analyzed specimen, the average
duration of one respiration cycle is 338 6 35 ms
(Table 1). As in attached respiration, the inhala-
tory phase (198 6 41 ms) takes longer (P < 0.005)
than the expiration phase (158 6 9 ms).
The pectoral girdle also moves synchronously
with the hyoid, being at its most caudoventral
point 8 6 9 ms after the hyoids depression maxi-
mum and at its most rostrodorsal point 2 6 10 ms
before the hyoid reaches its most elevated position
(Table 2). The lower jaw appears to be moved by
the hyoid motion, but these movements are very
small in comparison. The suspensorium moves
slightly before the hyoid (abduction and adduction
maximum 48 6 16 ms and 43 6 14 ms before the
depression and elevation maximum of the hyoid,
respectively; Fig. 10). The opercular moves slightly
after the suspensorium with its abduction and
adduction maximum 20 6 21 ms and 33 6 32 ms
Journal of Morphology
W. DE CROP ET AL.
Fig. 10. Graph showing movements of anatomical points of
interest in a 1,200 ms recording of Astroblepus sp. “Peru”
(MUSM 46925 nr. 3, 147-mm SL) in ventral view during unat-
tached respiration. Timing of the excurrent gill slit and the
upper oral valve being open are indicated by horizontal bars.
after abduction and adduction maximum of the
suspensorium, respectively. The dorsal incurrent
opening (Fig. 10) is maximally appressed 80 6 9
ms and maximally opened 22 6 10 ms after the
abduction and adduction maximum of the suspen-
sorium, respectively. The excurrent gill slit
(Fig. 10) opens and closes 2 6 19 ms and 18 6 9 ms
after the suspensorium is most abducted and
adducted, respectively. During unattached respira-
tion water is inhaled via the dorsal gill opening as
well as via the mouth. In the expiration phase,
there seems to be little or no backflow through the
mouth. When the upper oral valve was visible
inside the mouth, it could be observed moving ros-
trally during hyoid depression and caudally during
hyoid elevation (Table 2). In addition to its caudal
movement, the valve appeared to bulge in a
ventral direction.
Mobility of the suckermouth, and locomo-
tion. The expanded lower lip is a highly mobile
structure: it can flatten out completely or move its
peripheral rim ventrally, forming a cup-like struc-
ture. Left and right lower jaws are able to move
back and forth quite substantially, and, appa-
rently, independent from each other. They are not
connected by a symphysis, and move slightly apart
when they are retracted. The upper jaws appear
less mobile than the lower jaws, with both pre-
maxillas staying close together as well. Mobility of
the upper lip is limited and it seems to be pas-
sively following movements of the upper jaws and
maxillary barbels. When the barbels move down-
ward, as is the case when attachment to a sub-
strate is initiated, the upper lip contributes to the
cup-like structure formed by the lower lip by mov-
ing slightly downward as well. During elevation of
the barbels the upper lip moves back dorsally.
Crawling locomotion with aid of the sucker-
mouth has been observed in all specimens. An
 MORPHOLOGY OF ASTROBLEPID CATFISHES
individual is able to move forward on the sub-
strate in a crawling way by the alternating use of
its sucker-like mouth and pelvic girdle, without
undulatory body movements. The pelvic girdle
moves in a caudal direction while the mouth is
unattached, resulting in the forward movement of
the fish. Then the mouth attaches to the substrate
and the pelvic girdle is pulled to a rostral position.
This rostrocaudal mobility of the pelvic girdle is
extreme and must be caused by the protractor
ischii and retractor ischii (Fig. 7). The ventral sur-
face of each principal pelvic fin ray is broadened
and has a rough skin functioning as a backward
antislip mat.
DISCUSSION
The Buccal Pump
In Astroblepus, buccal cavity volume is mediated
by the lateral suspensorium, opercular and bran-
chiostegal membrane, and the ventral lower jaws,
hyoid and pectoral girdle. Using both lateral and
ventral buccal walls for constriction and expansion
can be linked to a moderately dorsoventrally flat-
tened skull (Alexander, 1970). An important con-
tribution of the lower jaws, and mainly the hyoid
and pectoral girdle, is common in catfishes, includ-
ing the related Callichthyidae (e.g., Van Wassen-
bergh et al., 2009). Astroblepidae do not share the
loss of the involvement of the pectoral girdle in
respiration, as has been observed in sister family
Loricariidae, where the pectoral girdle is virtually
immobile (Geerinckx et al., 2007). Compared to
the hyoid and pectoral girdle, the contribution of
the lower jaws is low in astroblepids, and higher
in loricariids (Geerinckx et al., 2011). Apart from
the pectoral girdle, the opercular of Loricariidae
also has been decoupled from respiration, leaving
them with a prominent role for the lower jaws and
hyoid. The loricariid suspensorium has limited
mobility, and is often firmly attached to the bony
plates covering the skull (Howes, 1983; Geerinckx
and Adriaens, 2006; Geerinckx et al., 2007, 2011).
The astroblepid suspensorium, being very
mobile, lacks the adductor hyomandibulae, which
is found (but is small) in both the related Callich-
thyidae and Loricariidae (Huysentruyt et al.,
2007; Geerinckx et al., 2007). Adductive power
thus can only come from the large adductor arcus
palatini. Given the fact that the levator arcus
palatini is minute, the resting position of the sus-
pensorium presumably is the abducted position,
analogous to the loricariid situation (Geerinckx
et al., 2007).
In Astroblepidae, the opercular and large
branchiostegal membrane contributes to the
volume changes of the branchial cavity. Opposed
to the majority of the operculo-branchiostegal
wall, the loose posterior edge operates as a valve
preventing water inflow during inhalation. An
13
active contribution of adductor operculi and antag-
onistic dilatator operculi is possible, especially
during observations of extra wide opercular abduc-
tion from time to time. Direct observations of
adductor operculi contraction through the thin
skin imply a default action of this muscle during
respiration. No separate levator operculi was
found in this study, although Howes (1983) found
it in Astroblepus grixalvii and A. cf. praeliorum.
The levator operculi, being rudimentary in Astro-
blepidae, has been found to be larger in represen-
tatives of Callichthyidae (Huysentruyt et al., 2007)
and Loricariidae (Howes, 1983; Geerinckx et al.,
2007), where it is assumed to be a synergist of the
dilatator operculi. Its near-absence might well be
related to the incurrent gill opening situated at
exactly that location.
The Paradox of Simultaneous Attachment
and Respiration—A System of Valves
Using the mouth as a sucker device raises some
important constraints related to respiration. One
obvious problem is the fact that the sucker
capacity of the mouth is expected to be reduced
dramatically by water inflow through the mouth
during inhalation. In Astroblepus, the solution to
this problem is unique within the order of cat-
fishes. An incurrent gill opening replaces the
mouth as an inhalation device in an efficient way,
allowing astroblepids to sustain adhesion, without
any need for inhalation through the mouth. The
valve closing this opening seems to be largely pas-
sively controlled by the pressure differences
between the gill chamber and the surrounding
environment. During expiration, when the pres-
sure inside the gill chamber is higher than out-
side, the valve is pressed in dorsal direction,
closing off the opening and preventing water com-
ing from the buccal cavity to bypass the gills when
exiting. When the pressure difference is reversed
at the onset of the inhalation phase, the valve
moves back in ventral direction, allowing the
inflow of water through the incurrent opening.
During that phase, the valve rests on the underly-
ing gill arches, preventing water to flow directly
into the opercular cavity and probably forcing it to
flow in a medial direction toward the middle of the
buccal cavity instead, then back laterally through
the gills, into the opercular cavity. As such, the
possibility of a continuous ventilation of the gills
exists (but could not be proven). This is quite dif-
ferent from the mechanism found in Loricariidae.
During respiration with suckermouth attachment,
loricariids sustain adhesion via the mouth while
allowing water inflow through a furrow created in
the lateral lip tissue by slightly elevating the
maxillary barbel (Alexander, 1965; Geerinckx
et al., 2011). This action has been ascribed to the
Journal of Morphology
14 W. DE CROP ET AL.
levator tentaculi muscle (Geerinckx et al., 2007,
2009, 2011).
A second problem is that any water flowing
anteriorly into the suction cup during compression
of the postvalvular (or buccal) cavity would
strongly impair the sucker capacity of the mouth.
Moreover, as jaws nor lips can be closed, backflow
during unattached respiration would be problem-
atic as this backflowing water can not be used to
oxygenate the gills. In Astroblepus, this appears to
be solved by the presence of two oral valves. Their
position (directly caudal to the tooth-bearing parts
of the upper and lower jaws) suggests that, when
both jaws are adducted, and thus closer to each
other, the valves will close off the prevalvular cav-
ity. During compression of the postvalvular cavity,
the increasing pressure might help push both
valves even harder onto each other. The upper and
lower jaws are kept close together during attached
respiration, enabling the valves to completely
close, relative to the situation in unattached respi-
ration, supporting this hypothesis. During unat-
tached breathing, upper and lower jaws are
generally held further away from each other, but
nevertheless they approach each other during
expiration. This astroblepid mechanism is differ-
ent from the one found in loricariids: these possess
only one oral valve, located on the roof of the
mouth. However, this valve is actively pulled
dorsocaudally by the retractor veli muscle, thereby
closing the entrance to the postvalvular cavity
during expiration (Gradwell, 1971; Geerinckx
et al., 2007, 2009, 2011).
Function of the Lips and Suckermouth
Evidently, the lips of the suckermouth are of
great importance for attachment, especially the
mobile lower lip, moved by the labial parts of the
intermandibular muscles. Even though the inter-
mandibularis anterior muscles differ significantly
between astroblepids and loricariids, they are sub-
divided in two parts in both families and appear to
be involved in the ventral pursing of the lower lip.
The morphological difference is most obvious in
the labial part, in which left and right contralat-
eral muscles do not touch medially and have a fan-
ning appearance in Astroblepus. In Loricariidae,
the labial part is attached to both sides of the
inner lower lip tissue and where both contralateral
muscles meet medially, some fibers cross the mid-
line (Howes, 1983; Geerinckx and Adriaens, 2007;
Geerinckx et al., 2007). In Astroblepus, contralat-
eral parts of the pars dentalis originate laterally
in the lip tissue caudal to the lower jaws. They
each insert medially, on the cartilage plug anteri-
orly, and on a shared raphe posteriorly. In loricar-
iids, the dental part connects the posterior faces of
the lower jaws and both halves meet medially in a
slim raphe, connected to the medial tissue of the
Journal of Morphology
lower lip (Geerinckx and Adriaens, 2007;
Geerinckx et al., 2007). The intermandibularis
posterior muscles are very similar in astroblepids
and loricariids and the intermandibularis posterior
pars labialis serves as the antagonist of the inter-
mandibularis anterior muscles, stretching (flatten-
ing) the lower lip, in both astroblepids and
loricariids (Geerinckx et al., 2007).
Since no muscles insert directly into the tissue
of the upper lip it is less mobile and is mainly con-
trolled by movements of the upper jaw and the
maxillary barbel. The retractor premaxillae
retracts the upper jaw, in turn pulling the upper
lip along. Howes (1983) stated that the lower por-
tion of the retractor premaxillae is not completely
separated from the outer adductor mandibulae
complex in Astroblepidae. Such a muscular con-
nection was not observed in the present study. The
antagonist of the retractor premaxillae is the
extensor tentaculi pars medialis, which, based on
its orientation, appears to displace the autopala-
tine in a ventromedial direction, causing its carti-
laginous head to push down against the
dorsomedial process of the premaxilla, forcing the
upper jaw to rotate rostrally. This in turn pushes
the upper lip forward. The extensor tentaculi pars
lateralis is involved in movements of the maxillary
barbel, as its action swings the autopalatine in a
ventrolateral direction, which results in a similar
displacement of the maxillary, pushing the barbel
and lateral parts of the lip towards the substrate.
The idea that the extensor tentaculi controls the
ventral movements of the autopalatine is sup-
ported by the fact that this mechanism occurs in
other, related taxa as well and would have origi-
nated from a modified siluriform maxillary mecha-
nism (Gosline, 1975; Geerinckx et al., 2007). These
two parts of the extensor tentaculi are completely
separated from one another, as they are in loricar-
iids (Howes, 1983; Geerinckx et al., 2007). This
contradicts the findings of Howes (1983), Schaefer
and Lauder (1986), and Schaefer (1990) suggesting
that the extensor tentaculi is a single element in
Astroblepidae and that its subdivision is a unique
trait of Loricariidae. Diogo (2005) also noted the
subdivision of this muscle in Astroblepidae. The
medial part of the extensor tentaculi might previ-
ously have been overlooked because of its position,
covered laterally by the extensor tentaculi
pars lateralis and ventrolaterally by the adductor
arcus palatini, making it less apparent during
dissections.
The antagonist of the lateral part of the exten-
sor tentaculi is the levator tentaculi, which causes
a caudodorsal elevation of the maxillary barbel
and the upper lip when contracted. This muscle
was described by Howes (1983) and Schaefer and
Lauder (1986) as being the homologue of the
retractor tentaculi found in Callichthyidae, which
has its origin shifted from the suspensorium to the
 MORPHOLOGY OF ASTROBLEPID CATFISHES
lateral ethmoid in loricariids and astroblepids. A
more recent study by Geerinckx et al. (2009),
observing the ontogenetic origin of this muscle
alongside the extensor tentaculi bundles, well
away from the adductor mandibulae complex, in a
loricariid, however, proposed homology between
the dorsolateral part of the callichthyid extensor
tentaculi and the loricariid “retractor tentaculi,”
renaming the latter “levator tentaculi.” Even
though myological data from this study are largely
in line with this hypothesis, the innervation pat-
tern of the levator tentaculi does not support it. In
loricariids, the muscle is innervated by a branchlet
of the maxillary branch of the trigeminal nerve,
entering it via its ventral surface (Howes, 1983).
This ventral innervation is also observed in
Ancistrus cf. triradiatus (Geerinckx et al., 2009)
and Astroblepus sp. “Peru.” Geerinckx et al. (2009)
state that in the loricariid Ancistrus the nerve to
the levator tentaculi originate from the anasto-
mosed buccomaxillary branch and can only be
traced back to the point where the mandibular
branch forms part of the nerve. In Astroblepus,
however, the branchlet to the levator tentaculi
first fuses with the branchlet that innervates the
retractor premaxillae before joining the buccomax-
illary and mandibular branches. This contrasts
with the supposed homology between at least the
astroblepid levator tentaculi and the dorsolateral
part of the callichthyid extensor tentaculi. The
fact that the branchlets innervating the levator
tentaculi and retractor premaxillae fuse, rather
supports the idea of Howes (1983) and Schaefer
and Lauder (1986) about homology between the
astroblepid (and loricariid) levator tentaculi and
the callichthyid retractor tentaculi. Thus, because
of these contrasting observations, the evolutionary
origin of the levator tentaculi muscle remains
enigmatic.
Locomotion using Suckermouth
and Pelvic Fins
Actual near-vertical climbing out of the main
water body was not recorded, but it is likely to be
similar to crawling locomotion that was observed
in all specimens on flat surfaces out of the water,
and on the aquarium walls in the water, during
which the suckermouth and pelvic girdle are used
in an alternating fashion in order to move forward.
This unique form of locomotion is made possible in
Astroblepidae by two well-developed muscles, the
retractor ischii and the protractor ischii. Although
these muscles occur in the direct relatives of
astroblepids, they are much less pronounced in
Loricariidae and Callichthyidae. In these two
families, the protractor ischii is not completely sep-
arated from the hypaxial muscles (Shelden, 1937).
Apart from this pelvic locomotory apparatus,
one might hypothesize that the dorsal incurrent
15
gill opening has played a major role in the astro-
blepid evolution toward climbing, supposing that
stronger suction is required to withstand the
forces imposed on a climbing fish by falling water
and gravity than by a more horizontally flowing
river. The extra gill opening is a nice example of
convergent evolution, as it is also present in the
unrelated Gyrinocheilidae (Smith, 1945). Being a
popular fish in aquarium trade, Gyrinocheilus
aymonieri is the most well-known representative
of this Asian cypriniform family. It occurs in rela-
tively large rivers and flooded plains, attaching
itself on flat surfaces using its specialized mouth
while an incurrent gill opening allows the fish to
continue breathing (Nelson, 2006). Loricariidae
(and some African Mochikidae) have the remark-
able ability to use the mouth as an attachment
structure while inhaling through the mouth. The
lack of an incurrent gill opening might explain
why most loricariids are unable to climb, and con-
sequently occur more downstream than astroble-
pids. Representatives of one loricariid genus,
Lithogenes, are known to climb in a similar way
as Astroblepidae (Schaefer, 2003; Schaefer and
Provenzano, 2008). Whether there is a similar
incurrent gill opening as in astroblepids is not
mentioned in literature; further research on this
topic may well yield interesting results.
ACKNOWLEDGMENTS
The authors are grateful to Hern an Ortega
Torres (Universidad Nacional Mayor de San
Marcos, Lima), for essential support in organizing
the sampling. Authors also thank Denıs Orinhuela
and Yajaira Preising for making the collection a
success, and Edgardo Castro and the Instituto
del Bien Com un in Oxapampa, for kindly provid-
ing logistics for the aquarium and recording
setup. The authors wish to thank Scott Schaefer
(American Museum of Natural History, New York)
for aid in the identification of the specimens.
Ronald Vonk and Hielke Praagman of the ZMA
are acknowledged for the loan of specimens.
Finally we would like to thank Gert Van Goethem
for his contribution to the quality of the myological
images. T.G. is a postdoctoral researcher of the
fund for scientific research Flanders, Belgium
(FWO-Vl).
LITERATURE CITED
Alexander RMcN. 1965. Structure and function in the catfish.
J Zool (Lond) 148:88–152.
Alexander RMcN. 1970. Mechanics of the feeding of various
teleost fishes. J Zool (Lond) 162:145–156.
Arratia G. 1990. Development and diversity of the suspenso-
rium of trichomycterids and comparison with loricarioids
(Teleostei: Siluriformes). J Morphol 205:193–218.
Collyer ML, Stockwell CA, Adams DC, Reiser MH. 2007.
Phenotypic plasticity and contemporary evolution in
Journal of Morphology
16 W. DE CROP ET AL.
ecological invasions: Evidence from translocated populations
of White Sands pupfish. Ecol Res 22:902–910.
Diogo R. 2005. Morphological Evolution, Aptations, Homopla-
sies, Constraints, and Evolutionary Trends: Catfishes as a
Case Study on General Phylogeny and Macroevolution. New
Hampshire: Enfield: Science Publishers. 491 p.
Erer KS. 2007. Adaptive usage of the Butterworth digital filter.
J Biomech 40:2934–2943.
Geerinckx T, Adriaens D. 2006. The erectile cheek-spine
apparatus in the bristlenose catfish Ancistrus (Loricariidae,
Siluriformes), and its relation to the formation of a secondary
skull roof. Zoology 109:287–299.
Geerinckx T, Adriaens D. 2007. Ontogeny of the intermandibu-
lar and hyoid musculature in the suckermouth armoured
catfish Ancistrus cf. triradiatus (Loricariidae, Siluriformes).
Anim Biol 57:339–357.
Geerinckx T, Brunain M, Herrel A, Aerts P, Adriaens D. 2007.
A head with a suckermouth: a functional-morphological study
of the head of the suckermouth armoured catfish Ancistrus cf.
triradiatus (Loricariidae, Siluriformes). Belg J Zool 137:47–
66.
Geerinckx T, Huysentruyt F, Adriaens D. 2009. Ontogeny of the
jaw and maxillary barbel musculature in the armoured cat-
fish families Loricariidae and Callichthyidae (Loricarioidea,
Siluriformes), with a discussion on muscle homologies. Zool J
Linn Soc 155:76–96.
Geerinckx T, Herrel A, Adriaens D. 2011. Suckermouth arm-
ored catfish resolve the paradox of simultaneous respiration
and suction attachment: a kinematic study of Pterygoplich-
thys disjunctivus. J Exp Zool 315:121–131.
Gerstner CL. 2007. Effect of oral suction and other friction-
enhancing behaviors on the station-holding performance of
suckermouth catfish (Hypostomus spp.). Can J Zool 85:
133–140.
Gosline WA. 1975. The palatine-maxillary mechanism in cat-
fishes, with comment on the evolution and zoogeography of
modern Siluroids. Occas Pap Calif Acad Sci 120:1–31.
Gradwell N. 1971. A muscular oral valve unique in fishes. Can
J Zool 49:837–839.
Hedrick TL. 2008. Software techniques for two- and three-
dimensional kinematic measurements of biological and
biomimetic systems. Bioinspir Biomim 3:034001.
Hora SL. 1930. Ecology, bionomics and evolution of the torren-
tial fauna, with special reference to the organs of attachment.
Philos Trans R Soc London (Biol) 218:171–282.
Howes GJ. 1983. The cranial muscles of loricarioid catfishes,
their homologies and value as taxonomic characters. Bull Br
Mus Nat Hist (Zool) 45:309–345.
Huysentruyt F, Geerinckx T, Adriaens D. 2007. A descriptive
myology of Corydoras aeneus (Siluriformes: Callichthyidae),
with a brief discussion on adductor mandibulae homologies.
Anim Biol 57:433–452.
Johnson RDO. 1912. Notes on the habits of a climbing catfish
(Arges marmoratus) from the Republic of Colombia. Ann NY
Acad Sci 22:327–333.
Masschaele BC, Cnudde V, Dierick M, Jacobs P, Van Hoorebeke
L, Vlassenbroeck J. 2007. UGCT: New X-ray radiography and
tomography facility. Nucl Instr Meth Phys Res A 580:
266–269.
Nelson JS. 2006. Fishes of the World. New Jersey: Wiley. 601 p.
Ortega H, Guerra H, Ramirez R. 2007. The introduction of non-
native fishes into freshwater systems of Peru. In: Bert MT,
editor. Ecological and Genetic Implications of Aquaculture
Activities. Dordrecht: Springer. pp 247–278.
Pouilly M, Barrera S, Rosales C. 2006. Changes of taxonomic
and trophic structure of fish assemblages along an environ-
mental gradient in the Upper Beni watershed (Bolivia).
J Fish Biol 68:137–156.
Journal of Morphology
Rom an-Valencia U. 2001. Trophic and reproductive ecology of
Trichomycterus caliense and Astroblepus cyclopus (Pisces:
Siluriformes) in Quindio River, Colombia. Rev Biol Trop 49:
657–666.
Schaefer SA. 1988. Homology and evolution of the opercular
series in the loricarioid catfishes (Pisces: Siluroidei). J Zool
Lond 214:81–93.
Schaefer SA. 1990. Anatomy and relationships of the scolopla-
cid catfishes. Proc Acad Nat Sci Phila 142:167–210.
Schaefer SA. 2003. Relationships of Lithogenes villosus
Eigenmann, 1909 (Siluriformes, Loricariidae): Evidence from
high-resolution computed microtomography. Am Mus Novit
3401:1–55.
Schaefer SA, Arroyave J. 2010. Rivers as islands: Determinants
of the distribution of Andean astroblepid catfishes. J Fish
Biol 77:2373–2390.
Schaefer SA, Lauder GV. 1986. Historical transformation of
functional design: Evolutionary morphology of feeding mecha-
nisms in loricarioid catfishes. Syst Zool 35:489–508.
Schaefer SA, Lauder GV. 1996. Testing historical hypotheses of
morphological change: Biomechanical decoupling in loricar-
ioid catfishes. Evolution 50:1661–1675.
Schaefer SA, Provenzano F. 2008. The Lithogeninae (Siluri-
formes, Loricariidae): Anatomy, interrelationships, and
description of a new species. Am Mus Novit 3637:1–49.
Schaefer SA, Chakrabarty P, Geneva AJ, Sabaj P erez MH.
2011. Nucleotide sequence data confirm diagnosis and local
endemism of variable morphospecies of Andean astroblepid
catfishes (Siluriformes: Astroblepidae). Zool J Linn Soc 162:
90–102.
Shelden FF. 1937. Osteology, myology, and probable
evolution of the nematognath pelvic girdle. Ann NY Acad Sci
37:1–96.
Smith HM. 1945. The Fresh-Water Fishes of Siam, or Thailand.
Washington: Smithsonian Institution, US National Museum.
621 p.
Sullivan JP, Lundberg JG, Hardman M. 2006. A phylogenetic
analysis of the major groups of catfishes (Teleostei: Siluri-
formes) using rag1 and rag2 nuclear gene sequences. Mol
Phylogenet Evol 41:636–662.
Taylor WR, Van Dyke GC. 1985. Revised procedures for stain-
ing and clearing small fishes and other vertebrates for bone
and cartilage study. Cybium 9:107–119.
Van Wassenbergh S, Lieben T, Herrel A, Huysentruyt F,
Geerinckx T, Adriaens D, Aerts P. 2009. Kinematics of
benthic suction feeding in Callichthyidae and Mochokidae,
with functional implications for the evolution of food scraping
in catfishes. J Exp Biol 212:116–125.
V
elez-Espino LA. 2003a. Conservation aquaculture of the
Andean catfish Astroblepus ubidiai: Effect of light intensity
on growth rate and number of reproductive allocations.
J Aquacult Trop 18:327–342.
V
elez-Espino LA. 2003b. Taxonomic revision, ecology and
endangerment categorization of the Andean catfish Astroble-
pus ubidiai (Teleostei: Astroblepidae). Rev Fish Biol Fish 13:
367–378.
V
elez-Espino LA. 2004. Threatened fishes of the world:
Astroblepus ubidiai (Pellegrin, 1931) (Astroblepidae). Environ
Biol Fishes 71:296.
V
elez-Espino LA. 2006. Distribution and habitat suitability
index model for the Andean catfish Astroblepus ubidiai
(Pisces: Siluriformes) in Ecuador. Rev Biol Trop 54:623–
638.
Vlassenbroeck J, Dierick M, Masschaele M, Cnudde V, Van
Hoorebeke L, Jacobs P. 2007. Software tools for quantification
of x-ray microtomography at the UGCT. Nucl Instr Meth
Phys Res A 580:442–445.