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Revision of Loricaria Linnaeus, 1758 (Pisces, Siluriformes, Loricariidae).

Authors:
51
Beaufortia
INSTITUTE
OF
TAXONOMIC
ZOOLOGY
(ZOOLOGICAL
MUSEUM)
UNIVERSITY
OF
AMSTERDAM
Vol.
31
no.
3
October
30,
1981
Revision
of
Loricaria
Linnaeus,
1758
(Pisces,
Siluriformes,
Loricariidae)
I.J.H.
Isbrücker
Institute
of
Taxonomic
Zoology,
University
of
Amsterdam,
P.O.
Box
20125,
1000
HC
Amsterdam,
The
Netherlands
&
Musee
de
Zoologie
de
I'Universite
et
de
la
Ville
de
Nancy,
34,
rue
Sainte-Catherine,
54000
Nancy,
France
Abstract
The
South
American
genus
of
mailed
catfishes
Loricaria
Linnaeus,
1758
is
reviewed.
Almost
all
primary
type-material
and
additional
specimens
are
examined.
Descriptions,
tabulated
morphometric
and
meristic
data,
and
illustrations
are
presented
of
eleven
species:
Loricaria
cataphracta
Linnaeus,
1758
(syn.:
Plecostomus
flagellaris
Gronovius,
ed.
Gray,
1854,
and
Loricaria
carinata
de
Castelnau,
1855),
L.
lata
Eigenmann
&
Eigenmann,
1889,
L.
simillima
Regan,
1904,
L.
parnahybae
Steindachner,
1907
(syn.:
L.
piauhiae
Fowler
1941),
L.
piracicabae
Von
Ihering,
1907,
L.
clavipinna
Fowler,
1940,
L.
nickeriensis
Isbrücker,
1979,
L.
tucumanensis
Isbrücker,
1979,
L.
apeltogaster
Boulenger,
1895,
L.
prolixa
Isbrücker
&
Nijssen,
1978,
and
L.
lentiginosa
Isbrücker,
1979.
The
nomenclatural
history
of L.
cata-
phracta
is
revised:
contrary
to
the
author’s
previous
(1972)
assumption,
two
syntypes
of
this
species
are
still
in
existence.
Definitions
are
presented
of
the
tribe
Loricariini,
the
subtribe
Loricariina,
the
Loricaria-
and
Pseudohemio-
don-groups,
and
of
the
Loricaria
cataphracta
species
complex.
In
addition,
the
Pseudoloricariina
(new
subtribe),
the
Ricolina
(new
subtribe),
and
the
Planiloricariina
are
defined.
A
key
to
the
genera
of
Loricariinae
with
filamentous
lip
structures
is
given.
A
list
of
species
originally
proposed
as
representatives
of
the
genus
Loricaria,
now
excluded,
is
added
together
with
their
current
identification.
INTRODUCTION
Loricaria
Linnaeus,
1758,
is
a
genus
of
substrate
dwelling,
South
American
primary
freshwater
fishes.
It
is
the
earliest
described
taxon
in
the
family
of
armoured
or
mailed
catfishes,
the
Lori-
cariidae.
A
total
of
61%
of
all
Loricariinae
has
at
some
time
been
assigned
to
Loricaria.
However,
authors
of
important
works
on
this
group,
like
Eigenmann
&
Eigenmann
(1889,
1890,
1891),
Regan
(1904),
Eigenmann
(1910),
and
Gosline
(1945)
had
adopted
a
subdivision
of
Loricaria
into
several
subgenera,
each
of
which
is
considered
to
be
of
generic
rank
by
me,
resulting
in
a
considerable
reduction
of
the
number
of
species
in
Loricaria.
Compared
to
the
works
of
Eigenmann
&
Eigen-
mann,
Regan,
and
Gosline,
Bleeker's
(1862,
1863)
subdivision
of
the
Loricariidae
is
surprisingly
modern
or
the
recent
subdivision
by
me
(1980)
is
perhaps
conservative
in
terms
of
the
number
of
recognized
genera.
A
list
of
nominal
species
proposed
originally
as
members
of
Loricaria
sensu
52
Ricola
Isbrucker
&
Nijssen,
1978,
Paraloricaria
Isbrucker,
1979,
Brochiloricaria
Isbrucker
&
Nijs-
sen,
1979,
Crossoloricaria
Isbrucker,
1979,
Pseu-
dohemiodon
Bleeker,
1862,
Rhadinoloricaria
Is-
brucker
&
Nijssen,
1974,
and
Planiloricaria
Is-
briicker,
1971,
share
a
single
external
character
with
Loricaria,
absent
in
all
other
genera
of
Lori-
cariinae:
numerous
filamentous
labial
barbels
and
subbarbels.
A
key
to
these
genera
is
included.
In
previous
papers
(especially
1971b,
1972)
I
have
attempted
to
point
out
some
historical
dif-
ficulties
in
correctly
naming
specimens
belonging
to
the
type-species
of
the
genus,
Loricaria
cata-
phracta.
Recently,
Boeseman
(1976)
demon-
strated
some
errors
in
these
publications.
With
his
criticism
in
mind
I
have
revised
my
previous
interpretations
(see
Addendum).
ABBREVIATIONS
ANSP
:
Academy
of
Natural
Sciences
of
Philadel-
phia,
Philadelphia,
Perm.
BMNH
:
British
Museum
(Natural
History),
London.
MCZ
:
Museum
of
Comparative
Zoology,
Cam-
bridge,
Mass.
MNHN
:
Museum
National
d'Histoire
Naturelle,
Paris.
MZUN
:
Musee
de
Zoologie
de
l'Universite
et
de
la
Ville
Nancy,
Nancy.
MZUSP
:
Museu
de
Zoologia
da
Universidae
de
Sao
Paulo,
Sao
Paulo.
NMW
:
Naturhistorisches
Museum,
Vienna.
NRS
:
Naturhistoriska
Riksmuseet,
Stockholm.
USNM
:
National
Museum
of
Natural
History,
formerly
United
States
National
Museum,
Washington
D.C.
ZMA
:
Instituut
voor
Taxonomische
Zoologie
(Zoologisch
Museum),
Amsterdam.
ZMB
:
Museum
fur
Naturkunde
der
Humboldt-
Universitat,
Berlin.
hi
:head length
si
:
standard
length
METHODS
The
methods
of
taking
morphometric
and
meristic
data
were
defined
by
Isbriieker
&
Nijssen
(1978a:
180-182).
ACKNOWLEDGEMENTS
Many
persons
have
contributed
importantly
to
this
study
in
many
ways,
particularly
by
the
loan
or
gift
of
specimens,
providing
working
space
in
their
laboratories,
extensive
curatorial
assistance,
or
by
information.
I
am
much
obliged
to
all
these
individuals
for
their
kindness:
Mrs
J.
Abel
(MNHN),
Mrs
Dr
M.
L.
Bauchot
(MNHN),
Dr
J.
E.
Bohlke
(ANSP),
Dr
H.
A.
Britski
(MZUSP),
Mrs
Ph.
Choanier
(MNHN),
Mrs
M.
Clark-McLellan
(BMNH),
Dr
B.
Conde
(MZUN),
Mrs
M. M.
Dick
(MCZ),
Dr
W. L.
Fink
(MZC),
Dr
P.
H.
Greenwood,
Mr
G.
J.
Ho-
wes
(BMNH),
Dr
P.
Kahsbauer
(NMW),
Mrs
S.
Karnella
(USNM),
Mrs
Dr
C.
Karrer
(formerly
ZMB),
Dr
E.
Lachner
(USNM),
Dr
K.
Liem
(MCZ),
Mr
C.
Powell
(USNM),
Dr
W.
G.
Saul
(ANSP),
and
Mr
R.
Schoknecht
(MCZ).
Mr
Howes
and
Mr
A.
Wheeler
(both
BMNH)
criticized
parts
of
the
typescript
dealing
with
the
history
of
Loricaria
cataphracta.
I
am
very
grate-
ful
for
their
suggestions
for
improvement.
Mr
Wheeler
provided
me
with
information
on
the
still
existing
syntypes
of
Loricaria
cataphracta,
and
gave
permission
to
use one
of
his
photographs
(fig.
2
in
this
paper).
I
am
much
indebted
for
the
friendly
and
ex-
tensive
interest,
advice,
assistance,
critical
com-
ments
and
stimulating
discussions
while
working
(in
ZMA)
to
Dr
P.
J.
H.
van
Bree,
Mrs
C.
Lin-
denaar-Sparrius,
Dr
H.
Nijssen,
and
Mr
L.
van
Tuyl;
Mr
L.
A.
van
der
Laan
and
Mr
J.
Zaag-
man
(both
also
ZMA)
prepared
all
but
one
of
the
illustrations
in
this
paper.
Mrs
Drs
F. F.
J.
M.
Pieters
(Artis
Library,
Amsterdam)
provided
me
with
many
of
the
valuable
publications
in
her
care.
I
deeply
appreciate
the
stimulation
given
by
both
Dr
J.
Gery
(St.
Cyprien)
and
Professor
Dr
B.
Conde
(MZUN),
while
proposing
to
present
this
study
to
the
University
of
Nancy
I.
At
an
earlier
occassion
(1978)
I
had
already
dedicated
the
MZUN-scription
containing
the
basis
of
this
publication
to
Dr
H.
Nijssen,
who
strongly
supported
my
study
of
fishes
throughout
many
years:
without
his
support
none
of
my
publications
would
have
been
realized.
With
gratitude
I
acknowledge
the
financial
sup-
port
received
by
a
grant
from
the
K.
F.
Hein
Foundation
(Utrecht)
to
complete
the
present
study.
lato,
presently
assigned
to
other
genera
of
Lori-
cariidae,
is
given
at
the
end
of
the
systematic
part
of
this
paper.
53
THE
TRIBE
LORICARIINI
The
Loricariini
are
characterized
by
having:
a)
the
origin
of
the
dorsal
fin
about
opposite
to
the
origin
of
the
pelvic
fins;
b)
6
branched
dorsal
fin
rays,
the
last
one
split
to
its
base;
c)
10
branched
caudal
fin
rays;
d)
no
more,
and
usually
considerably
less
than
18
teeth
in
each
premaxilla
in
adults
(against
at
least
27
in
the
adults
of
the
Harttiini);
and
e)
great
differentiation
in
denti-
tions
and
lip
shapes
and
structures.
The
characteristic
dentition
present
in
almost
all
Hypostominae,
Ancistrinae,
Hypoptopomatinae,
all
Harttiina
and
in
the
Farlowellini
(viz.,
very
numerous,
slender,
filiform
teeth
with
a
strongly
curved
and
bilobate
crown),
never
occurs
in
the
Loricariini.
In
spite
of
the
diversity
in
lip
struc-
tures
and
shapes
in
the
Loricariini,
none
is
similar
to
those
present
in
other
Loricariid
groups.
The
outer
(naked)
surface
of
the
upper
lip
is
devoid
of
dermal
ossifications.
The
orbital
rim,
with
few
exceptions,
is
provid-
ed
with
a
more
or
less
conspicuous
posterior
notch.
The
fin
rays
are
usually
dichotomously
branch-
ed.
Often
the
upper
unbranched
caudal
fin
ray
extends
as a
fragile,
extremely
long
filament;
a
similarly
prolonged
lower
unbranched
caudal
fin
ray
is
rarely
encountered,
in
some
of
the
Rine-
loricariina
only.
Within
the
Loricariini,
several
different
types
of
secondary
sexual
dimorphism
have
evolved.
These
include
(a)
excessive
growth
of
odontodes
in
certain
areas
of
the
head,
on
dorsum
of
the
body
in
front
of
the
dorsal
fin
origin,
and/or
on
dorsum
of
the
pectoral
fin,
(b)
change
in
the
shape
of
certain
minute
odontodes
for
example,
those
on
the
spines
of
the
pelvic
and
anal
fins
(c)
change
in
the
shape
of
the
teeth,
and
(d)
the
development
of
a
very
long
and
broad
lower
lip.
All
these
changes
occur
in
the
male
of
the
various
representatives
of
the
Loricariini.
The
dimorphism
indicated
under
(a)
is
restricted
to
the
more
primitive
genera,
that
indicated
under
(d)
is
found
only
in
the
more
specialized
sub-
tribes.
In
addition,
secondary
sexual
dimorphism
is
unknown
in
various
species
and
may
prove
not
to
exist
at
all
in
various
instances.
The
Loricariini
are
the
sistergroup
of
the
Harttiini.
The
more
generalized
representatives
of
both
tribes
do
not
appear
to
be
very
distantly
related
to
each
other,
although
the
probably
most
primitive
genus
of
the
Loricariini
(Ixinandria)
is
more
advanced
than
the
probably
most
primitive
genus
of
the
Harttiini
(Harttiella).
There
appears
to
be
more
variability
among
the
Loricariini
than
among
the
other
tribes
of
the
Loricariinae
(Hart-
tiini,
Farlowellini,
and
Acestridiini).
The
Loricariini
are
subdivided
into
8
subtribes:
the
Rineloricariina,
Ricolina
(new),
Loricariina,
Planiloricariina,
Reganellina,
Pseudoloricariina
(new),
Loricariichthyina,
and
the
Hemiodontich-
thyina.
Various
species
of
Loricariini
reach
a
larger
size
than
most
other
Loricariinae,
many
reaching
over
300
mm
in
si
(e.g.,
some
Rineloricariina,
Loricariina,
Pseudoloricariina,
and
Loricariich-
thyina).
Anal
fin
with
1,4
rays,
last
one
split
to
its
base;
pectoral
fin
1,6;
pelvic
fin
1,5.
Thirteen
genera
of
Loricariini
differ
from
the
Rineloricariina
(consisting
of
Ixinandria,
Rine-
loricaria,
Dasyloricaria,
and
Spatuloricaria),
from
the
Harttiini
(
(Harttiella,
Harttia,
Cteniloricaria,
Lamontichthys,
Pterosturisoma,
Sturisomatich-
thys,
Sturisoma,
and
Metaloricaria)
and
from
the
Farlowellini
(
(Farlowella
only)
by
the
apparent
absence
of
prominent
odontode
development
in
mature
males:
Ricola,
Paraloricaria,
Loricaria,
Brochiloricaria,
Crossoloricaria,
Pseudohemiodon,
Rhadinoloricaria,
Planiloricaria,
Reganella,
Lima-
tulichthys,
Pseudoloricaria,
Loricariichthys,
and
Hemiodontichthys.
The
absence
of
this
sexual
dimorphism
needs
confirmation
in
several
of
these
13
genera
(not
all
species
are
known
from
both
sexes,
rendering
the
conclusion
preliminary).
PSEUDOLORICARIINA
new
subtribe
Type-genus:
Pseudoloricaria
Bleeker,
1862
The
subtribe
Pseudoloricariina
as
here
proposed
is
equivalent
to
the
genus
Pseudoloricaria
sensu
Isbriicker
&
Nijssen,
1976b.
At
that
time,
Pseudo-
loricaria
was
considered
to
contain
two
species,
but
since
then
(Isbriicker
&
Nijssen,
in
Isbriicker,
1979a)
we
established
Limatulichthys
to
accom-
modate
the
second
species.
Limatulichthys
(a
less
specialized
genus
than
54
the
next)
and
Pseudoloricaria
probably
are
de-
scendants
of
some
Rineloricaria/
Dasyloricaria-
like
lineage,
which
gradually
lost
the
ability
to
develop
masculine
odontode
enlargement
during
maturation.
This
dimorphism
became
replaced
by
a
tendency
towards
improvement
of
the
masculine
lower
lip
enlargement.
It
appears
that
the
Pseudo-
loricariina
are
intermediate
between
Rinelorica-
riina
and
Loricariichthyina.
The
latter
subtribe
shows
a
much
higher
specialization
in
dentition
and
lip
structure,
restricted
to
a
single
genus,
Loricariichthys.
As
a
definition
of
the
Pseudoloricariina,
our
previous
(1976b)
diagnosis
of
Pseudoloricaria
is
still
valid.
RICOLINA
new
subtribe
Type-genus:
Ricola
Isbriicker
&
Nijssen,
1978
Ricola
consists
of
a
single
species,
R.
macrops,
occurring
in
the
Rio
de
la
Plata
and
higher
up
the
Rio
Parana,
in
Uruguay
and
Argentina.
Ricola
macrops
may
reach
a
standard
length
of
219
mm.
It
is
characterized
by
three
features:
a)
it
is
strikingly
similar
in
appearance
(shape
of
head
and
body,
structure
of
the
odontodes,
arrangement
of
scutes
and
development
of
abdom-
inal
scutelets,
and
colour)
to
species
of
Loricaria,
particularly
of
the
Loricaria
cataphracta-g
roup;
b)
its
dentition
is
strikingly
reminiscent
(shape
and
number,
as
well
as
relative
size
of
the
teeth)
of
a
Rineloricaria,
particularly
of
the
Rinelori-
caria
platyura-g
roup;
c)
it
is
unique
among
Loricariidae
by
the
struc-
ture
of
its
lips
and
barbels.
These
barbels
are
not
only
very
numerous
(like
in
Loricaria
and
related
genera),
but
many
of
them
are
further
subdividing
into
minute
branches.
The
upper
lip
is
narrow;
a
series
of
about
5
barbels
at
either
side
along
the
posterior
edge
of
this
lip,
increasing
in
length
towards
the
maxillary
barbel.
Posterior
to
these
series
are
three
thick,
deeply
branched
barbels
present
along
the
outer
surface
of
the
premaxillae.
Outer
side
of
the
maxillary
barbels
with
a
series
of
long
barbels
(actually
being
a
continuation
of
the
series
ante-
rior
to
the
premaxillae),
each
barbel
being
provid-
ed
with
numerous
small
barblets
in
a
linear
series.
Inner
side
of
producing
part
of
the
maxillary
barbels
likewise
with
several
long
barbels
with
small
barblets.
Lower
lip
narrow,
the
anterior
half
consisting
of
a
thick,
semicircular
cushion-like
structure.
This
structure
bears
irregular,
low
papillae
on
the
surface.
The
posterior
part
of
the
lower
lip
has
numerous
slender,
simple
papillae
or
subbarbels
on
its
surface
(like
those
in
Loricaria
spp.).
Edge
of
the
lower
lip
with
numerous
long
barbels,
each
provided
ventrally
with
shorter,
slender
sub-
barbels.
A
short,
thick,
triangular
papilla
between
premaxilla
and
dentary.
Three
long
papillae
in
the
buccal
cavity
posterior
to
the
premaxillae,
one
in
the
middle
and
one
at
either
side.
Ricola
has
up
to
15
teeth
in
each
premaxilla
and
up
to
14
teeth
in
each
dentary.
Those
in
the
premaxillae
are
about
twice
as
long
as
those
in
the
dentary.
They
have
a
prominent
inner
lobe
and
a
smaller
outer
lobe.
Ricola
tends
to
have
many,
sometimes
slightly
more
lateral
body
scutes
(37-39)
than
the
Lori-
cariina
(32-38).
Secondary
sexual
dimorphism:
The
pectoral
fin
spine
is
thicker
in
the
male
than
in
the
female,
just
like
in
males
of
Loricaria.
Tooth
lobes
in
mature
males
become
broader
and
more
rounded
at
the
tip
than
in
females
and
juveniles,
which
have
acute
tips.
This
type
of
change
in
tooth
shape
also
occurs
in
males
of
various
other
sub-
tribes
of
Loricariini
(e.g.,
Rineloricariina,
Lori-
cariina,
Pseudoloricariina,
Loricariichthyina,
and
Hemiodontichthyina).
Considering
the
three
mentioned
characters
of
Ricola
macrops,
it
can
neither
satisfactorily
be
included
within
the
Rineloricariina,
nor
within
the
Loricariina.
I
propose
a new
subtribe
for
Ricola,
because
it
is
hard
to
imagine
that
this
genus
evolved
from
some
Loricaria-like
ancestor
which
secondarily
would
have
attained
the
primitive
dentition,
compared
to
the
Loricariina.
One
could
judge
Ricola
as
some
highly
specialized
derivation
of
the
Rineloricariina
stock
which
retained
its
ancestral
dentition
and
attained
its
unusual
lip
structures
plus
the
Loricaria-
like
appearance
in-
dependently
from
the
Loricariina.
Because
of
the
combination
of
characters
(general
appearance
of
Loricaria,
dentition
of
Rineloricaria
,
and
unique
barbel
structure)
I
think
it
is
justified
not
to
55
unite
the
Rineloricariina
with
the
Loricariina
on
account
of
Ricola.
PLANILORICARIINA
Planiloricaria
Isbriicker,
1971
Planiloricaria
is
known
only
from
the
holotype
of
P.
cryptodon,
214
mm
in
si,
originating
from
the
Rio
Ucayali
near
Pucallpa,
Peru.
Superficially,
it
is
reminiscent
of
a
Pseudo-
hemiodon
with
a
disk-like
head
shape
in
dorsal
and
ventral
view,
and
much
depressed
in
lateral
view.
It
differs
not
only
from
that
genus
but
also
from
all
other
species
of
the
tribe
Loricariini
by:
its
produced
dorsal
fin
spine,
2.4
in
si
(against
3.4
or
more);
its
small
eyes,
13.
1
in
hi
(against
9.6
or
less);
—•
its
long
maxillary
barbels,
1.0
in
hi,
each
with
about
20
subsidiary
barbels
(maxillary
barbel
is
1.1
in
Rhadinoloricaria
and
more
in
other
Loricariini);
its
24
long,
papillose
barbels
along
the
posterior
edge
of
the
lower
lip,
2.6
in
hi
(against
at
least
4.5);
its
numerous
lateral
body
scutes,
40
(against
27-39);
and
the
absence
of
premaxillary
teeth
and
its
reduced
number
of
mandibular
teeth,
3
(against
up
to
18
and
up
to
34,
respectively).
Planiloricaria
has
no
orbital
notch;
no
dorsal
flap
on
the
pupil;
its
lower
lip
is
very
narrow,
11.5
times
in
hi;
numerous
filaments
and
papil-
lose
extensions
about
the
(rudimentary?)
pre-
maxillae;
a
smooth
dorsum
of
head
and
snout,
except
for
a
feeble
coalescing
double
ridge
running
anterior
as
well
as
posterior
to
the
supraoccipital
process;
head
and
snout
are
devoid
of
prominent
odontodes;
a
very
long
upper
caudal
filament;
no
produced
pelvic
fin
spines;
abdomen
almost
com-
pletely
covered
with
minute
scutelets.
Planiloricaria
shares
the
absence
of
premaxil-
lary
teeth
with
Reganella
and
with
Hemiodon-
tichthys;
with
both
it
does
not
seem
to
have
a
close
relationship.
Secondary
sexual
dimorphism
is
not
known.
LORICARIINA
With
the
exception
of
the
Ricolina
and
the
Plani-
loricariina,
the
Loricariina
embrace
the
other
genera
of
the
Loricariinae
with
numerous
long,
fleshy
filaments
on
the
surface
and/or
along
the
outer
edges
of
both
the
upper
and
the
lower
lips.
Loricariina
have
up
to
9
premaxillary
and
up
to
11
mandibular
teeth
(Ricolina
have
up
to
15
and
14,
respectively;
Planiloricariina
have
no
premaxillary
teeth
and
3
in
each
dentary).
The
number
of
lateral
body
scutes
ranges
in
the
Lori-
cariina
from
31
to
38;
in
the
Ricolina
there
are
37-39
and
in
the
Planiloricariina
40.
The
Loricariina
occur
all
over
tropical
South
America.
The
largest
member
of
the
subfamily
belongs
here:
Paraloricaria
vetula,
which
reaches
437
mm
in
si.
All
members
of
the
subfamily
Loricariinae
(like
those
of
the
other
subfamilies
of
Loricarii-
dae)
have
replacement
teeth.
Specimens
of
species
now
assigned
to
other
genera
than
Loricaria
(e.g.,
Paraloricaria,
Crossoloricaria)
may
be
found
with
five
or
less
teeth
in
each
premaxilla.
The
teeth
are
embedded
between
axial,
fleshy
lamellae,
and
counting
these
lamellae
results
in
a
higher
number
of
(potential)
teeth.
A
difference
in
tooth
shape
and
relative
size
is
usually
asso-
ciated
with
differences
in
the
structure
of
barbels
and
other
soft
parts
of
lips
and
in
the
buccal
cavity.
I
am
of
the
opinion
that
these
differences
are
not
necessarily
related
to
feeding
habits,
but
to
the
different
trends
in
the
evolution.
The
subtribe
Loricariina
may
be
further
divided
into
two
genus-groups,
presumably
representing
distinct
phyletic
lineages,
which
I
here
informally
refer
to
as
the
Loricaria-group
(also
including
Paraloricaria
and
Brochiloricaria),
and
the
Pseu-
dohemiodon-group
(also
including
Crossoloricaria
and
Rhadinoloricaria).
The
Loricaria-g
roup
seems
to
be
rather
close
(perhaps
more
so
than
the
Pseudohemiodon-
group)
to
the
Rineloricariina.
The
ancestor
of
the
Loricaria-
group
probably
was
a
Rineloricariina
which
developed
a
tendency
towards
(a)
fewer
teeth;
(b)
slightly
larger
lips,
especially
the
lower
lip
becoming
longer;
(c)
more
numerous
and
more
prominent
papillae
which turned
into
fila-
ments;
(d)
more numerous
lateral
body
scutes,
tending
to
coalesce
more
posterior
than
in
the
Rine-
loricariina
stock;
and
(e)
a
gradual
loss
of
the
secondary
sexual
dimorphism
in
the
odontode
development
in
the
males.
56
The
LORICARIA-
and
PSEUDOHEMIODON-groups
The
genera
of
the
Loricaria-g
roup
have
larger
and
less
premaxillary
teeth,
and
the
teeth
in
the
dentary
are
likewise
more
robust
than
in
the
Pseu-
dohemiodon-group
genera.
The
teeth
of
the
Pseudohemiodon-group
are
rather
inconspicuous,
either
simple
or
bilobed,
and
more
or
less
spoon-
shaped.
The
filaments
on
the
surface
and
along
the
posterior
edge
of
the
lower
lip
are
longer
(some-
times
bifid
or
trifid)
and smooth
in
the
Loricaria-
group.
In
the
Pseudohemiodon-group
the
fila-
ments
may
be
almost
absent
on
the
surface
of
the
lower
lip,
sometimes
being
reduced
to
small,
elongate
papillae.
The
long
filaments
on
and
especially
along
the
lower
lip
are
not
smooth,
but
more
or
less
papillose;
frequently
these
papillae
are
elongate.
The
snout
of
the
Loricaria-
group
genera
is
not
produced;
in
the
Pseudohemiodon-group
the
snout
is
hardly
or
not
produced
in
some
species
and
quite
produced
in
others.
The
following
comparison
of
166
specimens
of
Paraloricaria,
Loricaria,
and
Brochiloricaria
with
46
specimens
of
Crossoloricaria,
Pseudohemiodon,
and
Rhadinoloricaria
was
made.
Those
of
the
Loricaria-
group
are
given
first,
followed
between
parentheses
by
those
of
the
Pseudohemiodon-
group:
standard
length
up
to
437
mm
(246.5
mm);
postdorsal
length
1.6-1.8
(1.6-2.1);
length
first
dorsal
fin
ray
3.6-6.6
(5.1-7.3);
anal
spine
length
4.6-7.2
(6.0-13.2);
pectoral
spine
length
2.2-6.6
(5.0-6.6);
pelvic
spine
length
3.9-7.3
(6.0-8.7);
upper
caudal
fin
spine
1.0-7.4
(0.4-6.9);
lower
caudal
fin
spine
4.9-8.7
(6.1-9.9);
ventrorostral
length
nihil
(4.2-25.8);
length
lower
lip
3.7-9.4
(3.9-10.6);
cleithral
width
1.0-1.5
(°-8-i-3);
supracleithral
width
r.
5-2.1
(r.3-1.9);
head
width
1.0-1.5
(0.8-1.3);
head
depth
1.9-3.1
(2.5-3.5);
body
depth
at
dorsal
fin
origin
1.6-3.2
(2.2-3.5);
length
maxillary
barbel
x.5-7.4
(1.1-2.4);
length
longest
barbel
of
lower
lip
5.6-26.0
(4.5-
12.1);
lateral
body
scutes
32-38
(31-34);
coalescing
lateral
body
scutes
17-26
(13-21);
thoracic
scutes
4-13
(5-11);
premaxillary
teeth
up
to
6
(up
to
9).
KEY
TO
GENERA
OF
LORICARIINAE
WITH
FILAMENTOUS
LIP
STRUCTURES,
TRIBE
LORICARIINI
ia
No
teeth
in
upper
jaws
.
(Planiloricariina)
Planiloricaria.
ib
Teeth
in
both
upper
and
lower
jaws
.
2.
2a
Upper
jaws
with
6
or
more
teeth
on
each
side
3.
2b
Upper
jaws
with
5
or
less
teeth
on
each
side
4.
3a
Upper
jaws
with
up
to
9
teeth
on
each
side;
maxillary
(=
rictal)
barbels
with
subbarbels
which
are
not
subdivided
into
minute
branches
(Loricariina)
5.
3b
Upper
jaws
with
up
to
15
teeth
on
each
side;
maxillary
barbels
with
subbarbels
which
are
subdivided
into
minute
branches
.
(Ricolina)
Ricola.
4a
Teeth
in
upper
jaws
long,
about
twice
the
length
of
teeth
in
lower
jaws
.
Loricaria.
4b
Teeth
in
upper
jaws
about
r/3
longer
than
those
in
lower
jaws;
teeth
in
lower
jaws
about
as
long
as
teeth
in
the
upper
jaws
of
Loricaria
........
Brochiloricaria.
5a
Abdomen
naked,
or
covered
with
different
pat-
terns
of
scutelets,
not
however,
arranged
into
a
single
median
strip
6.
5b
Abdomen
naked
except
for
a
single
median
strip
of
small,
roundish
scutelets
Crossoloricaria.
6a
A
ventrorostral
extension
present
.
.
7.
6b
No
ventrorostral
extension
.
Paraloricaria.
7a
Sides
of
head
and
snout
more
or
less
trian-
gular
in
dorsal
view;
cleithral
width
0.8-1.i
in
hi;
supracleithral
width
1.3-1.7
in
hi;
head
depth
2.5-3.3
i
n
hi;
maxillary
barbel
1.4-2.4
in
hi;
depth
caudal
peduncle
12.3-17.3
in
hi
Pseudohemiodon.
yh
Sides
of
head
tapering,
of
snout
narrow
and
somewhat
concave
in
dorsal
view;
cleithral
width
1.2
in
hi;
supracleithral
width
1.9
in
hi;
head
depth
3.5
in
hi;
maxillary
barbel
1.1
in
hi;
depth
caudal
peduncle
9.7
in hi
Rhadinoloricaria.
57
THE
GENUS
LORICARIA
LINNAEUS,
1758
Loricaria
Linnaeus,
1758:
307-308
(type-species,
by
mono-
typy,
Loricaria
cataphracta
Linnaeus,
1758).
Fusiloricaria
Fowler,
1940a:
247
(subgenus
of
Loricaria;
type-species,
by
original
designation
and
monotypy,
Loricaria
(Fusiloricaria)
clavipinna
Fowler,
1940).
Loricaria
consists
of
11
valid
species,
which
occur
in
a
wide
range
of
localities
in
Surinam,
French
Guiana,
Brazil
(States
of
Para,
Amazonas,
Mato
Grosso,
Acre?,
Roraima),
Paraguay,
Uruguay,
Argentina,
Bolivia,
Peru,
Ecuador,
Venezuela,
and
Guyana.
The
distributions
shown
in
figs.
14-15
include
examined
material
only.
Loricaria
is
unique
among
Loricariidae
in
its
dentition.
There
are
up
to
5,
usually
3
or
4,
teeth
in
each
section
of
the
premaxilla, distinctly
longer
(about
twice)
than
the
teeth
in
the
dentary.
In
juveniles
and
females,
these
teeth
have
a
large,
oblong
crown
with
a
more
or
less
rounded
tip.
Small
specimens
(e.g.,
one
of
29.1
mm
in
si)
already
have
a
completely
developed
number
of
teeth.
The
teeth
are
either
simple,
or
bilobate,
usually
with
a
much
smaller
outer
lobe.
In
each
section
of
the
dentary
there
may
be
up
to
11
(usually
less)
teeth.
Generally,
the
crowns
of
the
mandibular
teeth
have
a
much
shorter,
broader,
and
more
rounded
inner
lobe
and
a
more
con-
spicuous
outer
lobe
than
the
premaxillary
teeth.
The
upper
lip
is
short,
its
margin
with
numerous
slender,
simple,
bifurcate
and
rarely
trifurcate,
barbels,
subbarbels
and
long
papillae.
Cirrhi
are
present
on
its
ventral
surface,
around
the
base
of
the
teeth,
in
the
buccal
cavity,
and
on
the
surface
of
the
maxillary
barbel.
The
lower
lip
is
well-developed,
usually
with
a
more
or
less
deep
median
notch.
Posterior
to
the
dentary
the
surface
of
the
lip
usually
has
a
cushion
bearing
low
papillae.
Posterior
to
this
cushion,
the
lip
has
many
long,
simple
filaments
which continue
along
the
posterior
margin.
A
mostly
inconspicuous
posterior
orbital
notch
is
usually
present.
The
pelvic
fin
spine
is
almost
always
longer
than
the
adjacent
branched
ray.
The
upper
un-
branched
caudal
fin
ray
often
extends
as a
fragile
filament
as
long
as,
or
exceeding
the
standard
length.
The
abdomen
may
be
naked,
or
is
partially
or
entirely
covered
with
small
scutelets,
which
de-
velop
with
age.
There
are
specific
differences
in
the
speed
of
development
of
these
scutelets:
some
species
are
fully
covered
at
an
earlier
stage
than
others.
In
the
late
development
of
these
scutelets,
Loricaria
apeltogaster,
L.
prolixa,
and
L.
lentigi-
nosa
are
reminiscent
of
the
Paraloricaria
spp.
and
of
Brochiloricaria
macrodon.
The
morphometric
and
meristic
variation
within
155
specimens
representing
all
species
of
Lori-
caria
is
summarized:
standard
length
29.1-346
mm;
axial
length
31.7-377
mm;
total
length
35.0-515
mm;
smallest
mature
male
103.8
mm;
head
length
3.7-5.2;
predorsal
length
2.9-3.7;
postdorsal
length
1.6-1.8;
postanal
length
1.8-2.2;
dorsal
spine
length
3.4-6.5;
length
first
dorsal
fin
ray
3.6-6.6;
anal
spine
length
4.6-7.2;
pectoral
spine
length
2.2-6.6;
pelvic
spine
length
3.9-7.4;
length
upper
caudal
'spine'
1.0-7.4;
length
lower
caudal
'spine'
4.9-8.7;
snout
length
r
.7-2.1;
length
lower
lip
3.7-9.4;
thoracic
length
1.1-1.8;
abdominal
length
1.2-2.0;
maximum
orbital
diameter
4.2-9.2;
interorbital
width
4.3-6.4;
cleithral
width
1.0-1.5;
supracleithral
width
1.5-2.1;
head
width
1.0-1.5;
head
depth
1.9-3.1;
body
depth
at
dorsal
fin
origin
1.6-3.2;
body
width
at
dorsal
fin
origin
1.1-3.2;
body
width
at
anal
fin
origin
1.1-3.3;
depth
caudal
peduncle
12.0-21.1;
width
caudal
peduncle
3.6-9.9;
length
maxillary
barbel
2.0-7.4;
longest
barbel
along
the
lower
lip
5.6-26.0;
lateral
scutes
32-37;
coalescing
lateral
scutes
17-26;
thoracic
scutes
4-12;
premaxillary
teeth
up
to
5;
mandibular
teeth
up
to
TT.
58
Secondary
sexual
dimorphism.
Mature
males
of
Loricaria
(of
the
L.
cataphracta-group,
which
excludes
L.
apeltogaster,
L.
prolixa,
and
L.
lenti-
ginosa)
have
hypertrophied
pectoral
fin
spines,
blunter
odontodes
on
the
pelvic
and
anal
fin
spines,
and
shorter,
more
rounded
tooth
lobes
than
females.
Rarely,
males
have
long
and
broad
lips
(e.g.,
a
male
of
L.
clavipinna
in
USNM
124930,
from
the
Rio
Ampiyacu
in
Peru,
156.5
mm
in
si,
has
the
lower
lip
shape
suggestive
of
those
Lori-
cariidae
which
protect
their
eggs
by
the
enlarged
lower
lip:
Dasyloricaria,
Limatulichthys,
Pseudo-
loricaria,
Loricariichthys,
and
Hemiodontichthys).
A.
de
Miranda
Ribeiro
(1912:
10)
reported
a
specimen
of L.
cataphracta
from
S.
Manoel,
Tapa-
joz,
"...
com
uma
placa
de
ovos
sobre
a
lado
ab-
dominal."
Males
of
Loricaria
apeltogaster,
L.
pro-
lixa,
and
of
L.
lentiginosa
are
unknown;
maybe
it
is
more
appropriate
to
say
that
they