The Lagenar Otoliths of Teleosts: Their Morphology and Its Application in Species Identification, Phylogeny and Systematics

Journal of Fish Biology (2003) 62, 1268–1295
doi:10.1046/j.1095-8649.2003.00106.x, available online at http://www.blackwell-synergy.com
The lagenar otoliths of teleosts: their morphology and

its application in species identification, phylogeny and
systematics
C. A. A S S I S
Instituto de Oceanografia and Departamento de Zoologia e Antropologia, Faculdade
de Cieˆncias da Universidade de Lisboa, Campo Grande, 1749-016 Lisboa, Portugal
(Received 29 May 2001, Accepted 28 March 2003)
The general morphology of the asterisci from recent European teleosts is described, terminology
for their parts is proposed, their three major morphological types are identified and the
relevance of the morphology of these otoliths in species identification, and as a source of
information for studies on fish phylogeny and systematics is discussed.
# 2003 The Fisheries Society of the British Isles
Key words: asteriscus; lagenar otoliths; morphology; phylogeny; systematics.
INTRODUCTION
According to Chaine & Duvergier (1934), the existence of a ‘stone’, much later
called the asteriscus, in the lagena of fishes was first reported by Athenoeus
Naucratita in 1597.
Higgins (1868) stated that the ‘posterior’ otoliths, as he then referred to the
asterisci, were completely devoid of value for fish species identification and in
studies of fish systematics, phylogeny and palaeontology. According to him, the
asterisci, although varying considerably in shape, are extremely small, rather
fragile and very difficult to extract from the fishes’ heads.
In the first half of the 20th century many studies on fish otoliths were
undertaken. In a few of them the asterisci were represented, alone, or together
with the sagittae and lapilli (Shepherd, 1910a, b; Frost, 1925a, b, c, 1926a, b, c,
1927; Sanz Echeverrı́a, 1928, 1929, 1930, 1932, 1935, 1936, 1949; Bauzá Rullán,
1956, 1958). Apart from not being constant in the inclusion of the asterisci,
most authors were very disproportional in the relevance attributed to the
sagittae and to the other two pairs of otoliths.
Three main reasons seem to have discouraged the study of fish asterisci:
(1) the fact that they are generally small and fragile; (2) the idea that they are
seldom found in geological layers and in stomach contents or predator food
remains; (3) the perpetuated belief that their morphology is not species specific,
Tel.: þ351 21 7500000; fax: þ351 21 7500009; email: carlos.assis@fc.ul.pt
1268
# 2003 The Fisheries Society of the British Isles
THE ASTERISCI OF TELEOSTS 1269
and that they are useless as a source of taxonomic and phylogenetic information
(Assis, 2000). As a result, the morphology of these anatomical parts of fishes is
still almost unknown (Frizzell & Koenig, 1973; Nolf, 1985).
Only in the Otophysi are the asterisci rather large and robust, when compared
with the respective sagittae, and have been used for species identification when
occasionally found in the geological record (Frizzell & Koenig, 1973), or in
stomach contents and in other predator food remains (Adams, 1940; Berinkey,
1956; Tilak, 1963; Mollo, 1981; Martı́nez & Monasterio de Gonzo, 1991).
The authors who have described fossil (Frizzell & Koenig, 1973) or recent
(Adams, 1940; Berinkey, 1956, 1957; Tilak, 1963) asterisci from cyprinids or
silurids, or those who prepared keys for their identification (Mollo, 1981;
Martı́nez & Monasterio de Gonzo, 1991) have arbitrarily assigned provisional
terminologies for their parts. Their terminologies, however, are based on a very
narrow range of taxa and are too incomplete to be useful (Assis, 2000). Besides,
most terms proposed are coincident with those already in use for features of the
sagittae.
In fact, no author, to date, has found the asterisci of fishes worthy of a
detailed comparative morphologic study (Assis, 2000).
It is in this context that this account of the asterisci of fish species aims to:
(1) present the general morphology of the asterisci of teleosts; (2) propose a
terminology for their parts; (3) describe the three types of asterisci found in
fishes; (4) discuss their relevance as a source of additional information in species
identification and in studies on fish phylogeny and systematics.
MATERIALS AND METHODS

This study is based on the analysis of a collection of lagenar otoliths from 4735
positively identified teleosts, including 183 species, and representing 62 families and
23 orders. All samples were collected in Portuguese coastal waters, estuaries and rivers.
Before dissection, the total length (LT) of each specimen was recorded. The otoliths
were then extracted through ventral dissection of the neurocranium, with the help of a
dissecting stereo microscope, cleaned in 90% ethanol with a small brush, and stored dry.
All drawings were made with a camera lucida, using the most convenient magnification
in each case.
RESULTS
THE MORPHOLOGY OF THE ASTERISCI

The asterisci are laterally compressed otoliths and, with the exception of those
of the Otophysi, are normally very small and fragile. Their flattened appear-
ance, and the presence of a furrow on their medial face makes it possible to
establish a parallel between the morphology of the asterisci and the sagittae.
This fact has been recognized by some authors who have used the same terms to
designate parts of both otolith pairs (Adams, 1940; Berinkey, 1956, 1957; Tilak,
1963; Frizzell & Koenig, 1973; Mollo, 1981; Martı́nez & Monasterio de Gonzo,
1991).
As the features of the two pairs of otoliths are not homologues, the coin-
cidence in names is not appropriate and may lead to confusion. On the other
# 2003 The Fisheries Society of the British Isles, Journal of Fish Biology 2003, 62, 1268–1295
1270 C. A. ASSIS
hand, in order not to create too much disruption in the terminology already
used by some authors, it is proposed that the names that are common to
features of the sagittae and asterisci be differentiated by adding the name of
the respective otolith in the genitive single. In the particular case of the word
asteriscus, the genitive single is similar to the nominative plural: asterisci.
Whenever there is no possibility of confusion between the parts of different
otoliths, the names may be abbreviated through the elimination of the unneces-
sary particles.
In terms of outline, the asterisci of teleosts may assume many different
shapes, as shown by the examples presented in Figs 1 and 2. Morphologically,
they are composed of two lobes of various relative sizes (Fig. 3), forming an
angle between themselves to the inside of the fishes’ heads. One of the lobes is
always larger than the other and the terms lobus major (larger lobe) and lobus
minor (smaller lobe) are proposed to designate each of them. The terms campus
major (larger field) and campus minor (smaller field) may be used to indicate
their respective surfaces (Fig. 4). Although in some taxa the two lobes are
almost similar in size [Fig. 3(a)], in most fish groups the lobus minor is reduced
[Fig. 3(b), (c), (d)], and in some it is almost inconspicuous, or even undiffer-
entiated [Fig. 3(e), (f)].
Viewed from the medial side, in a position equivalent to the one occupied in
the fish skull, several features of the asterisci can be used in the diagnosis of
taxa (Fig. 5).
When the lobus minor is evident, the campus minor is normally convex and
bent towards the medial face, forming a conspicuous concavity on its lateral
face, the concavitate lobi minor (depression of the smaller lobe), which assumes
an anterior position.
On the medial face, the two lobes are separated by a furrow of variable shape,
width and depth, the fossa acustica (acoustic pit). Although other authors have
used the designation of sulcus to describe this furrow (Adams, 1940; Berinkey,
1956, 1957; Tilak, 1963; Frizzell & Koenig, 1973; Mollo, 1981), in the large
majority of the asterisci it does not look like a groove at all. It is for this reason
that the retrieval of the designation of fossa acustica, used by Berinkey (1956,
1957) to name a furrow like depression which exteriorly surrounds the fossa in
the cyprinids, is proposed.
The fossa acustica may be open on one or both ends, in association with one
or two more or less deep indentations in the outline, the excisura major asterisci
(larger indentation of the asteriscus) (always anterior) and the excisura minor
asterisci (smaller indentation of the asteriscus) (either anterior or posterior).
The excisurae define protuberances similar to those found in the sagittae,
which are proposed to be termed rostrum asterisci, antirostrum asterisci (those
associated with the larger indentation), pseudorostrum asterisci and pseudo-
antirostrum asterisci (those associated with the smaller indentation).
When the fossa acustica is closed in the ventral end, and there is no excisura
minor, one point of the lobus major, which would correspond to the pseudo-
antirostrum asterisci, or to the fusion between the pseudorostrum and pseudoanti-
rostrum asterisci, forms one more or less pointed angle in the ventral region of
the otolith, the extremum ventralis [Fig. 6(a)]. On the other hand, when the fossa
is horizontal and closed in its posterior end, and an excisura minor is lacking,
FIG. 1. Medial (proximal) face of the right asterisci of several teleost fish species. All specimens have their
dorsal region to the top of the page and the anterior margin to the left. The shaded area
corresponds to the fossa acustica.
the homologue of the extremum ventralis is located in the posterior region

of the otolith and assumes the designation of extremum posterior asterisci
[Fig. 6(b)].
1272 C. A. ASSIS
FIG. 2. Medial (proximal) face of the right asterisci of several teleost fish species. All specimens have their
dorsal region to the top of the page and the anterior margin to the left. The shaded area
corresponds to the fossa acustica.
The fossa acustica, as the sulcus acusticus in the sagittae, is normally delimited
by two walls. While the wall in the lobus minor is coincident with the campus
minor, the other wall does not normally cover the entire campus major, and is
FIG. 3. Medial face (proximal side) of the right asterisci of several teleost species showing different
relative sizes of the two lobes (&, lobus major and &, lobus minor). Specimen (a) has its dorsal
margin to the right and the anterior margin to the top of the page, to allow direct comparison with
the other specimens, which have their dorsal margins to the top of the page and anterior margins to
the left.
often raised over the surface of the otolith, its border forming a conspicuous
crest, the crista medial (medial crest) (Fig. 5).
The crista medial is usually well developed and, in some species, it even
exceeds the outline of the lobus major. It may be raised or flattened over the
campus major, and defines a slit, crevice, furrow or step-like depression, the
fissura periferica asterisci, which surrounds it.
The floor of the fossa acustica, as well as its walls, is normally paved with a
substance similar to the colliculum found in the sagittae, the colliculum asterisci,
which, in these otoliths, is always undifferentiated and frequently has a very
irregular outline. In some species, however, the deepest region of the fossa is not
covered by colliculum. In such cases, the collicular substance forms a type of
ring which only covers the walls.
In the asterisci of some Otophysi, a large indentation is frequently formed in
the middle of the anterior margin of the otolith which, due to the extreme
1274 C. A. ASSIS
FIG. 4. Proposed terminology and orientation of the lobes of the otoliths asteriscus. Right otoliths are
represented. The shaded area corresponds to the fossa acustica. D, V, A and P: dorsal, ventral,
anterior and posterior directions, respectively.
reduction of the lobus minor, is limited by the antirostrum and pseudoantiros-

trum, or by the dorsal and ventral ends of the medial crest. Such indentation
does not correspond to any of the excisurae and is termed pseudoexcisura
asterisci [Fig. 5(c)].
THE THREE BASIC MORPHOLOGICAL TYPES OF

ASTERISCI
In terms of lobe shape and orientation, three morphological types of asterisci
may be defined: the vertical type [Fig. 5(a)], found among the Elopomorpha,
FIG. 5. Proposed terminology for the features displayed by teleost asterisci. Three types, (a) vertical,
(b) horizontal and (c) gyro, of right side otoliths are represented. The shaded area corresponds to
the fossa acustica. D, V, A, P, L and M: the dorsal, ventral, anterior, posterior, lateral (distal) and
medial (proximal) directions, respectively.
Clupeomorpha, Scopelomorpha, Protacanthopterygii and Acanthopterygii; the

horizontal type [Fig. 5(b)], found among the Paracanthopterygii; and the gyro
type [Fig. 5(c)], found among the Otophysi.
The vertical type is the most widespread type of asterisci among fish taxa. It
is characterized by having both lobes with their major axes vertical; a lobus
minor reduced, considerably less developed than the lobus major, and occupying
an anterior position in the otolith; a fossa acustica vertical or strongly bent to
the ventral direction close to its dorsal opening; and an excisura minor, a
pseudorostrum and a pseudoantirostrum that, when present, are always anterior
or ventral [Figs 5(a) and 7].
1276 C. A. ASSIS
FIG. 6. Medial face of right side asterisci, illustrating the extremum ventralis and the extremum posterior.
(a) Vertical type asterisci of (from left to right) Plectorinchus mediterraneus (Guichenot) and
Argentina sphyraena L. (b) Horizontal type asterisci of (from top to bottom) Nezumia scleror-
hynchus (Valenciennes) and Merluccius merluccius (L.). The shaded area corresponds to the fossa
acustica. All otoliths have their dorsal region to the top of the page and the anterior margin to the left.
The horizontal type is characterized by having both lobes with their major
axes horizontal; a lobus minor little less developed than the lobus major, and in a
ventral position; a fossa acustica horizontal; and an excisura minor, a pseudo-
rostrum and a pseudoantirostrum asterisci that, when present, are always
posterior [Figs 5(b) and 8].
The gyro type is characterized by having a well-developed lobus major,
surrounding the central part of the otolith and describing a circumference of
almost 360 around the fossa acustica; a lobus minor extremely reduced or
absent that, when identifiable, is very narrow and fused to one of the margins
(normally the dorsal margin) of the pseudoexcisura; a curved fossa acustica; a
more or less conspicuous anterior pseudoexcisura; and a pseudoantirostrum
asterisci that, when present, is always anterior [Figs 5(c) and 9]. In some species,
the fossa acustica dives into the middle part of the lobus major and, as it is bent
downwards, may not be completely exposed to the surface.
THE ASTERISCI OF FISHES AS A SOURCE OF

I N F O R M A T I O N F O R S P E C I E S ID E N T I F I C A T I O N ,
PHYLOGENY AND SYSTEMATICS
The usefulness of the knowledge on the morphology of the asterisci for
species identification, and in studies of fish phylogeny and systematics is largely
dependent on three conditions: (1) the characteristics presented by the otoliths
must be subjected to a considerable diversity; (2) they must possess a sufficiently
diagnostic power to allow a precise identification to a low taxonomic
level, ideally to the species level; (3) the degree of similarity displayed by the
characteristics of otoliths from different species must reflect, with considerable
precision, the affinity resultant from the distance to a common ancestor.
FIG. 7. Medial face of right side, vertical type asterisci. The shaded area corresponds to the fossa acustica.
All otoliths have their dorsal region to the top of the page and the anterior margin to the left.
In spite of representing a very small fraction of the total diversity of forms

displayed by the asterisci, Figs 1 and 2 clearly illustrate the fact that these
otoliths are far from being morphologically homogeneous among the fish
taxa, and suggest that the first condition may be considered as fulfilled.
The other two conditions, related to the diagnostic power of the morpho-
logical characteristics and to their reflection of the phylogenetic relationships,
were analysed.
The diagnostic power of the asterisci

The major difficulty of using the asterisci in species identification is related to:
(1) the differences in the shape of their parts are in most cases very subtle, not
quantifiable and difficult to describe by words; (2) frequently the same struc-
tures are present, in spite of being different in the degree of expression, which
gives them a relative value; (3) the number of usable descriptors is rather small.
All these limitations lead to the situation, unfortunately very common, that it is
rather hard to describe the characteristics that allow a precise discrimination
FIG. 8. Medial face of right side, horizontal type asterisci. The shaded area corresponds to the fossa acustica.
1278 C. A. ASSIS
FIG. 9. Medial face of right side, gyro type asterisci. The shaded area corresponds to the fossa acustica.
among otoliths from close species. Notwithstanding, when in presence of repre-

sentatives of all the forms in question, it is frequently easy to visually confirm
that they are considerably different.
The otoliths asteriscus of some species of the family Sparidae will be used to
illustrate this point.
The right asterisci of nine species belonging to various genera of the family
Sparidae are shown in Fig. 10. Although it is obvious that the otoliths are rather
different in shape, the existence of a common morphology among them is
evident. All asterisci are of the vertical type, approximately semi-elliptical; the
lobus minor is reduced and considerably smaller than the lobus major; the
campus major is moderately concave, while the campus minor is strongly convex;
the rostrum asterisci is well differentiated, variable in length, and always curved
to the lateral side; the excisura major is always present, although variable in
extension; the fossa acustica is rather deep; the crista medial is conspicuous and
elevated over the campus major, forming a concentric fissura periferica along the
posterior region; the extremum ventralis slightly tapers and is orientated in the
antero-ventral direction; and the concavitate lobi minor is very strong and
antero-lateral.
On the other hand, the outline of the otolith and that of its lobes, the shape
and depth of the excisura major, the presence and development of the excisura
minor, the shapes and relative sizes of the rostrum and antirostrum asterisci, the
dorsal origin of the crista medial, the extension of the fissura periferica, and the
shape and orientation of the extremum ventralis, seem to be the most important
characteristics for the discrimination of taxa within this family.
At a lower taxonomic level the differences among species are not so obvious,
but some of the above characteristics can still be used to identify them with an
acceptable precision.
The right and left asterisci of several specimens belonging to four species of
genus Diplodus are represented in Fig. 11. A detailed look at the images shows
that the otoliths of specimens <7 cm in LT do not allow a precise discrimination
of the species, as the main diagnostic characters are not yet developed in them.
From that length on, the following key can be used to discriminate among the
four species:
FIG. 10. Medial face of the right side asterisci of several species of the family Sparidae. The shaded area
represents the part of the fossa acustica covered by colliculum. All otoliths have their dorsal region
to the top of the page and the anterior margin to the left.
1 – Ventral limit of the lobus minor conspicuous in the outline. Excisura

major markedly indented with variable depth and borders forming an
acute angle. Ventral region of the anterior margin of the otolith rather
sinuous. ...................................................................................................... 2
1280 C. A. ASSIS
FIG. 11. Medial face of the right and left side asterisci of several species of genus Diplodus (Sparidae). The
shaded area represents the part of the fossa acustica covered by colliculum, and the heavy line parallel
to the posterior outline represents the medial crest. All otoliths have their dorsal region to the top of the
page and the anterior margin to the middle of each pair of columns (L, left otoliths; R, right otoliths).
– Ventral limit of the lobus minor inconspicuous in the outline. Excisura major
shallowly indented, with borders forming an obtuse angle. Ventral region of
the anterior margin regularly concave, with a strong curvature. .............. 3
2 – Rostrum asterisci clearly supramedian, long and tapering to an

almost pointed tip. Antirostrum asterisci triangular and prominent in the
outline. Excisura major deep and angular. Ventral region of the otolith
tapered. .................................................................................. D. sargus (L.)
– Rostrum asterisci almost median, short and tapered, but most frequently
round. Antirostrum asterisci round and not very prominent in the outline.
Excisura major shallow and round. Ventral region of the otolith broadly round
in the large majority of the specimens. ...... D. vulgaris (Geoffroy St. Hilaire)
3 – Excisura major clearly indented, although not very deep. Antirostrum
asterisci with an apex almost horizontal and, commonly, little behind the
level of the rostrum. Postero-dorsal region of the lobus major dividable in
two sections by a pronounced bend. ................................ D. annularis (L.)
– Excisura major little or not indented. Antirostrum asterisci with an antero-
dorsal apex, considerably behind the level of the rostrum. Curvature of the
postero-dorsal region of the lobus major regular. ... D. bellottii (Steindachner)
This may serve as evidence of the fact that, once the characteristic features of
the asterisci are fully developed, and at least in some taxa, these otoliths have a
distinct morphology that can be used in the identification of fish taxa, in certain
cases to the species level.
The reflection of phylogenetic relationships

As stated before, three considerably different basic types of asterisci may be
defined. Among them, the vertical type is the most generalized form of
asteriscus, found along the evolutionary sequence of the Teleostei, from the
Elopomorpha to the Percomorpha, with the exception of the Paracanthopterygii
and Ostariophysi. It is thus legitimate to assume that the vertical type of
asteriscus corresponds to the plesiomorphic condition, and that the other
more specific types of asterisci would have evolved from it. In fact, the assumed
changes suffered by a vertical type asteriscus to originate a horizontal or a gyro
type may be explained through simple rotations of the lobes and the fossa
acustica (Fig. 12).
As it will be shown, the horizontal and the gyro types are probably specific
and universal among most paracanthopterygian and ostariophysan taxa, respect-
ively. In that case, their emergence from the vertical type may be assumed to
have occurred very close to the origins of those taxa. The type of asteriscus may
thus be considered as a diagnostic character, and as an additional argument to
support each group’s holophyly.
The vertical type of asteriscus

The otoliths of the Elopomorpha and Clupeomorpha (although rather defi-
cient in number in the samples) seem to be characterized by possessing a well-
developed lobus minor, wide and almost as high as the lobus major; a deep and
almost vertical fossa acustica open at both ends; and by the association of both
extremes of the fossa with a clear excisura major and with an excisura minor
which, though variable in size and width, is always present (Fig. 13).
In the direction of the Percomorpha, although some of these plesiomorphic
characteristics are retained in several groups, it seems that there is a tendency
1282 C. A. ASSIS
FIG. 12. Distribution of the three types of asterisci among teleostean superorders, and assumed origin of
the horizontal and gyro types from a vertical type otolith. The lobus major is represented in white,
the lobus minor is shaded, and the dashed line corresponds to the crista medial. All otoliths
correspond to right side asterisci, and have their dorsal region to the top of the page and their
anterior margin to the left. The following otoliths are represented: Alosa alosa (L.) (ancestral
vertical type); Barbus bocagei Steindachner (Ostariophysi); Coelorhynchus coelorhynchus (Risso)
(Paracanthopterygii); Polymetme corythaeola (Alcock) (Stenopterygii); Notoscopelus bolini
Nafpaktitis (Scopelomorpha); Argentina sphyraena L. (Protacanthopterygii); Dicentrarchus punctatus
(Bloch) (Acanthopterygii, Percomorpha); Atherina boyeri Risso (Acanthopterygii, Atherinomorpha);
Liza ramada (Risso) (Acanthopterygii, Mugilomorpha).
for a reduction in the size of the lobus minor, which becomes lower and
narrower than the lobus major. Along with this general tendency, other correl-
ated changes can be noticed. Among them, (1) the ventral closure of the fossa
acustica, with the consequent reduction of the excisura minor, which becomes
vestigial or absent in the majority of the groups; (2) the fusion between the
ventral regions of both lobes; and (3) the rotation of the dorsal opening of the
fossa in the anterior direction, are evident (Figs 13 and 14).
FIG. 13. Medial face of the right asterisci of several fish species, representing some teleost superorders.
The lobus major is represented in white and the lobus minor is shaded. All specimens have their
dorsal region to the top of the page and the anterior margin to the left.
As each of these tendencies is displayed by the different taxa in a different

degree, remaining unnoticed in some of them, and as the diversity covered in the
collections of asterisci is still rather small, specially in what concerns some taxa
considered as basal in the Euteleostei, any attempt to deepen this analysis to a
lower taxonomic level seems to be rather premature.
1284 C. A. ASSIS
FIG. 14. Medial face of the right asterisci of several fish species, representing some acanthopterygian
orders. The lobus major is represented in white and the lobus minor is shaded. All specimens have
their dorsal region to the top of the page and the anterior margin to the left.
The horizontal type of asteriscus

The asterisci of the horizontal type are assumed to have evolved from those
of the vertical type through a simultaneous or, which is more likely, sequential
rotation of their dorsal and ventral regions, around the centre of the otolith, in
the anterior and posterior directions, respectively (Fig. 12).
In any of the cases, the parts that occupied the antero-dorsal position in the
vertical type otolith became the anterior region in the horizontal, while those
that occupied the antero-ventral region became postero-ventral or posterior.
This hypothesis is supported by two facts. Firstly, the two types of otoliths
differ almost exclusively in the orientation of their parts and structures, which
remain identifiable and keep the same associations. Secondly, otoliths with an
arrangement apparently intermediate between the vertical and horizontal types
exist among some of the paracanthopterygian orders (Fig. 15).
Concerning the distribution and possible tendencies displayed by the hori-
zontal type asteriscus among the Paracanthopterygii, the limited number of
sampled species belonging to this group, specially in the assumed basal region
of the lineage (Percopsiformes and Ophidiiformes), prevents a detailed analysis
to be carried out. Based on the characteristics of the asterisci of some Batra-
choidiformes, Lophiiformes and Gadiformes, however, some statements can be
made.
Assuming that the lagenar otoliths of the Batrachoidiformes, Lophiiformes
and Gadiformes sampled are representative of the respective orders, their pos-
session of typically horizontal asterisci may be taken as one additional synapo-
morphy defining the group of these orders, and as one more argument to justify
its holophyly. This supports the phylogenetic arrangement of Patterson &
Rosen (1989), according to which Batrachoidiformes and Lophiiformes
FIG. 15. Medial face of the right asterisci of several fish species, representing some paracanthopterygian
orders, and illustrating intermediate forms between the vertical and the horizontal type otoliths.
The shaded area corresponds to the fossa acustica. Homologous parts or regions are indicated,
even if undifferentiated: LMa, lobus major; LMi, lobus minor; R, rostrum; PR, pseudorostrum;
AR, antirostrum; PAR, pseudoantirostrum; EMa, excisura major; EMi, excisura minor; CM, crista
medial. All specimens have their dorsal region to the top of the page and the anterior margin to
the left.
1286 C. A. ASSIS
constitute sister groups, the Pediculati, which, in turn, are the sister group of the
Gadiformes.
In fact, a superficial observation of the morphology of the asterisci of the
taxa involved, makes intuitive their association in such a way (Fig. 15).
The total ignorance concerning the morphology of the asterisci of the
potential sister groups (Percopsiformes and Ophidiiformes) for the above set of
orders is a serious drawback to any attempt to deduce the tendencies displayed
by the features of the horizontal asterisci or the plesiomorphic state of the
characteristics.
Nevertheless, if it is taken into consideration that the main uncertainties
concerning the polarity of the characters are almost exclusively related with
the characteristics of the excisura minor and its associated structures (the
pseudorostrum and the pseudoantirostrum asterisci), and with the moment
when their complete reduction occurred (before or after the origin of the
Paracanthoperygii), some hypotheses can be advanced.
Three facts indicate that the non-existence of excisura minor, pseudorostrum
and pseudoantirostrum should be considered as the plesiomorphic state among
the Paracanthopterygii: (1) the Batrachoidiformes and Lophiiformes do not
possess excisura minor, pseudorostrum or pseudoantirostrum and, among the
Gadiformes, there are several species in which these structures are very much
reduced or even undifferentiated; (2) the asterisci of Benthocometes robustus
(Goode & Bean), an ophidiiform, show characteristics intermediate between the
vertical and horizontal types, and lack excisura minor, pseudorostrum and
pseudoantirostrum (Fig. 15); and (3), among the Paracantopterygii, the excisura
minor and associated structures only occur in a posterior position and asso-
ciated with an horizontal fossa acustica (i.e. after the otolith has completed the
rotation and the fossa is reoriented).
Based on these facts, it seems legitimate to assume that the otoliths of the
Batrachoidiformes and Lophiiformes are closer to the plesiomorphic condition,
while the ones of the Gadiformes are farther from it.
Consequently, it will have to be assumed that: (1) the tendency to the reduc-
tion and closure of the excisura minor has resulted in its disappearance and in
the undifferentiation of the associated structures, at least in the basal group of
the Paracanthopterygii; and that (2) the lengthening of the fossa acustica, the
association of its terminal region with the posterior position and its opening to
an excisura minor correspond to a secondary condition.
Either of these assumptions seems acceptable, not only because none of the
above structures is found in taxa such as the Protacanthopterygii, Stenopterygii
or Scopelomorpha, considered as more primitive than the Paracanthopterygii,
but also because the formation of an indentation in the ventral union between
the lobi seems to have independently occurred in several rather unrelated
groups; probably as an homoplasy, in many of them.
Taking this into account, it is possible to identify seven synapomorphies,
exclusively related with the asterisci, which justify the phylogenetic arrangement
proposed by Patterson & Rosen (1989) of the Anacanthini (Fig. 16).
Despite the little information available on the morphology of the lagenar
otoliths for some of the more diverse gadiform families, such as the Moridae,
Macrouridae and Gadidae, the observation of Fig. 17 allows some more
FIG. 16. Phylogenetic relationships among the orders of Anacanthini (Patterson & Rosen, 1989). The
Ophidiiformes should be considered sensu lato, according to Nelson (1994). Each of the orders in
the cladogram is represented by the right side otolith of one of its members, with the dorsal region
to the top of the page and the anterior margin to the left. The numbers 1–7 represent synapo-
morphies which may be utilized to define the groups: (1) unfolding of the lobus minor, with the
consequent increase in the angle formed between the lobi; (2) rotation of the antero-dorsal region
of the otolith in the anterior direction; (3) fossa acustica approaching an horizontal tendency;
(4) increase in the dimensions of the lobus minor, which forms all the ventral region of the otolith;
(5) development of the crista medial along the dorsal margin of the fossa; (6) rotation of the
excisura major in the ventral direction; (7) fossa acustica horizontal or slightly ascending.
considerations to be made, concerning some possible tendencies of the horizontal

type of asteriscus among the Gadiformes.
Assuming that the figured otoliths are representative of the respective taxa,
and taking into consideration that the conditions: absence of excisura minor,
medial crest undifferentiated, lobus minor little developed, and fossa acustica
straight or ventrally concave, are considered as plesiomorphic, the asterisci of
the Merlucciidae are closer to the plesiomorphic situation, while those of the
Gadidae seem to correspond to the apomorphic condition for those characters.
If these tendencies are true, and assuming that the characteristics related to
the differentiation of the excisura minor and associated structures suffer a
reversion to the original undifferentiated state, the Moridae and Phycidae may
be considered as sister groups and form the sister group of the Gadidae. All
these three families, in turn, may be the sister group of the Macrouridae and,
finally, of the Merlucciidae (Fig. 18).
It is a fact that the information on the asterisci of many gadids is lacking,
which makes it impossible to evaluate to what extent the otoliths of the sampled
taxa are representative of the characteristics of the groups involved in the
analysis. The cladogram obtained (Fig. 18), built exclusively with the information
1288 C. A. ASSIS
FIG. 17. Medial face of the right asterisci of several paracanthopterygian species, grouped according to
orders and families. The shaded area corresponds to the fossa acustica. All specimens have their
dorsal region to the top of the page and the anterior margin to the left.
FIG. 18. Tentative phylogenetic relationships among the asterisci of some gadiform families. Each family in
the cladogram is represented by the right side otolith of one of its members, with the dorsal region to
the top of the page and the anterior margin to the left. The numbers 1–12 represent the synapo-
morphies utilized to define the groups: (1) increase in the relative dimensions of the lobus minor; (2)
deepening of the excisura major and development of a rostral projection; (3) lengthening of the fossa
acustica in the posterior direction; (4) differentiation of an excisura minor in the posterior region of the
otolith; (5) development of the medial crest; (6) occurrence of a bend in the fossa acustica with a dorsal
concavity; (7) medial crest composed of two segments forming between themselves an angulous bend
with a dorsal concavity; (8) medial crest becomes higher; (9) increase in the curvature of the fossa
acustica and loss of its posterior opening (reversal); (10) closing of the excisura minor (reversal);
(11) lengthening of the crista medial, along the posterior region of the fossa acustica; (12) curvature
of the dorsal and ventral margins of the otolith becoming parallel to the curvature of the fossa acustica.
provided by the otoliths asteriscus, however, is not significantly different, in

terms of the arrangement of the groups that are common, from the one
proposed by Nolf & Steurbaut (1989) for the Gadiformes, the last one based
on the information provided by the otoliths sagitta. Between the two, the main
differences reside in the position attributed to the Merlucciidae and to the
Moridae, which were considered by Nolf & Steurbaut (1989) as closer to the
Phycidae and Gadidae, and to the Macrouridae, respectively.
Although it can be argued that the above presentation involves a lot of untested
assumptions and has a rather speculative character, the fact that it was possible to
use data on the morphology of the asterisci without falling into serious contra-
diction with other sources of information demonstrates the potential of these
otoliths as a valuable source of additional information in this type of study.
The gyro type of asteriscus

The asterisci of the gyro type probably evolved from those of the vertical type
through the rotation of the ventral region in the anterior direction, bringing the
1290 C. A. ASSIS
dorsal (antirostral) and ventral (pseudoantirostral) regions of the lobus major to

close proximity (Fig. 12). In this process, the lobus minor would have initially
been compressed between the dorsal and ventral regions of the anterior margin
of the lobus major, eventually becoming inconspicuous (either through its com-
plete fusion to the larger lobe or its loss).
Considering that, in some cyprinid species, (1) the limits of the lobes are
apparently possible to identify (although with some doubts); (2) the crista
medial surrounds almost completely the fossa acustica; (3) the fossa has com-
pletely lost the opening in its ventral region; and (4) the fossa possesses a round
and curved terminal end, away from the margin of the otolith which, in some
species, dives deep under the wall of the lobus major to the interior of the
otolith, the above hypothesis is perfectly acceptable (Fig. 19).
As a consequence of the assumed morphological transformations and of the
extreme reduction of the lobus minor, the pseudoexcisura, very frequent in the
gyro type asterisci, should not be confused with any of the excisurae. Other
indentations that, in some species, occur above or below the pseudoexcisura
correspond to a simple marginal sculpture of the otolith (Fig. 19).
Concerning the possible tendencies displayed by the gyro type of asteriscus
among the Ostariophysi, the limited material available (only eight cypriniform
species) does not allow a detailed analysis. As some images of the asterisci of
several ostariophysine taxa have been published (Frost, 1925b; Adams, 1940;
Berinkey, 1956; Tilak, 1963; Mollo, 1981; Martı́nez & Monasterio de Gonzo,
1991) some considerations, although speculative, can be advanced.
The Ostariophysi include the series Anotophysi (Gonorhynchiformes) and
Otophysi (Cypriniformes, Characiformes, Siluriformes and Gymnotiformes)
(Nelson, 1994; Fink & Fink, 1996). Among them, the Gonorhynchiformes are
considered as being in the proximity of the origin of the remaining orders of
FIG. 19. Medial face of the right side gyro type asteriscus of Barbus bocagei (Steindachner), showing the
organization of the otolith parts. The shaded area corresponds to the fossa acustica. The specimen
has its dorsal region to the top of the page and the anterior margin to the left.
Ostariophysi (Nelson, 1994; Fink & Fink, 1996) and may represent an inter-
mediate link between them and the Clupeomorpha (Nelson, 1994).
Some knowledge of the morphology of the gonorhynchiform asterisci could
provide valuable data to locate the origin of the gyro type asterisci and to
confirm or refute the assumed morphological conversion of one type into the
other but, unfortunately, those otoliths are not known.
Nevertheless, if the following three facts are taken into consideration, it
seems admissible to assume that the anatomical and morphological changes
that characterize the otophysine ear are not yet fully developed among the
gonorhynchiforms. First, the otoliths sagitta of Gonorhynchus greyi (Richardson)
(Frost, 1925a; Nolf, 1985), Chanos chanos (Forsskål) and Kneria sp. (Nolf,
1985) are morphologically similar to the otoliths of non-otophysine fishes (to
those of the clupeiforms in particular), and very different from the highly
specialized otoliths that characterize the Otophysi. Second, the cephalic ribs
associated to the first three vertebrae may be considered, in the Gonorhynchi-
formes, as a Weber system in its primordial state (Nelson, 1994). Third, an
otophysic connection between the gas bladder and the inner ear does not exist in
these fishes (Fink & Fink, 1996).
Under these circumstances, it seems likely that the gyro type asterisci have
found their origin among the Ostariophysi and not before.
It is also worth noticing that the asterisci of all otophysine orders clearly
correspond to the gyro type, which indicates that the origin of this type would
have occurred near the origin of the group.
The asterisci of all otophysine orders have the following common character-
istics: (1) lobus major forming the whole otolith, around the fossa acustica;
(2) posterior region of the fossa acustica with a more or less drastic bend in
the ventral direction; (3) dorsal and ventral ends of the crista medial very close
to each other, converging, in the middle anterior region of the otolith; and
(4) fissura periferica surrounding the fossa acustica, forming an open depression,
or sunk under the medial crest. These characteristics, due to their specificity
and universality within the Otophysi, can also be added to the list of synapo-
morphies that define this taxon and confirm its holophyly.
In view of the present knowledge on the morphology of the gyro type
asterisci, a more detailed analysis is not possible. Not only there is a complete
lack of information concerning the asterisci of the Gonorhynchiformes, which
makes it difficult to have a precise idea about the plesiomorphic state of the
characteristics displayed by the gyro otoliths, but also, the poor quality of the
image of a Gymnotiform (Gymnotus carapo L.) asteriscus, from Frost (1925b)
invalidates its use. On the other hand, it has been impossible to directly observe
the otoliths of taxa other than the Cyprinids, and the number of taxa with
known asterisci, in this extremely diverse group, is still very reduced.
DISCUSSION
Despite of Higgins’ (1868) statements, and those of other authors who
subscribed some of his arguments (Rado, 1968; Stinton, 1968; Jonet, 1972/73;
Popper & Coombs, 1982), this contribution shows that the lagenar otoliths can
1292 C. A. ASSIS
be used in species identification and may provide valuable additional data for
studies on fish phylogeny and systematics.
It is true that the asterisci are normally rather small and sometimes very
fragile when compared with the corresponding sagittae, but some species have
very small and fragile sagittae when compared with the asterisci of other species.
Besides, if the fish heads are carefully dissected and the relative position of the
three pairs of otoliths is known, their retrieval does not present any major
difficulties. In this respect, the ventral dissection of the otic region of the
neurocranium is the best method to expose the membranous labyrinth and
extract the otoliths.
It is also true that the presence of asterisci in the fossil record and in the
stomach contents and food remains of piscivorous predators has seldom been
reported. What remains to be known is the extent to which they have been
neglected or considered as unidentified material. The little knowledge that
presently exists on the morphological characteristics of the asterisci and the
great difficulty of their association with a taxon, justify this situation.
In terms of shape, the asterisci seem to be as diverse as the sagittae, a fact that
has already been referred by Shepherd (1910a). Their fossa acustica, however,
does not present the degree of differentiation typical of the sulcus acusticus, and
the number of taxonomically usable features in the asterisci seems to be fewer
than in the sagittae.
As shown before, the asterisci may be used with as much precision as the
sagittae in the identification of at least some taxa (Adams, 1940; Berinkey, 1956,
1957; Tilak, 1963; Mollo, 1981; Martı́nez & Monasterio de Gonzo, 1991), and
although this study only focuses on some examples, it seems clear that these
otoliths can be used in the identification of taxa, in some cases to the species
level. This has been verified by Assis (2000) after the study of 183 species.
It is known that the morphology of the saccular otoliths, the sagittae, is not
only related to common ancestry, but also to the habitat where a particular
species of fish lives (Schwarzhans, 1978; Nolf, 1985, 1993) and to the anatomical
specialization connected with sound perception (Nolf, 1985, 1993; Popper &
Platt, 1993). There are therefore different degrees of morphologic convergence
among evolutionary distant taxa and divergence among evolutionary close ones
(Nolf, 1985, 1993). The same probably happens with the asterisci once that both
the sacculus and the lagena are specially engaged in sound detection (Popper,
1980; Rogers et al., 1988) and seem to have evolved in close association with the
specialization of the sense of hearing (Popper & Coombs, 1982).
For this reason, the otoliths alone should not be used to define taxonomic or
phylogenetic affinities (Schwarzhans, 1978; Nolf, 1985, 1993, 1995; Smale et al.,
1995). Nevertheless, it seems clear that the characteristics presented by them can
be of considerable relevance, in conjunction with other characters of the fish
body, to increase the information available in studies of fish systematics and
phylogeny, as already proposed by Adams (1940).
Due to the limited diversity for which the morphology of the asterisci is
known, most inferences concerned with phylogenetic relationships and with
the origin of the types of asterisci, as well as the value of the synapomorphies
utilized to define the groups, are endowed with a considerable margin of
uncertainty. The fact that the information obtained using the asterisci does
not contradict significantly the results reached by other authors, using other
information, is rather encouraging.
Future research on the subject should be directed to: (1) widen the range of
taxa with known asterisci; (2) ascertain the amount of intra- and interspecific
variation to which the features of these otoliths are subjected; (3) evaluate the
degree of convergence or divergence that can be imposed on the morphology of
these otoliths in relation to the differences in habitat and hearing capabilities of
fishes; (4) determine the features of this pair of otoliths that, in each taxon, are
taxonomically more relevant; and (5) determine, for each taxon, the polarity of
the various character states presented by each relevant feature.
Only when some progress in these areas is accomplished can the real potential
of the knowledge about the morphology of the otoliths asteriscus and its
usefulness in studies of fish phylogeny and systematics be fully evaluated.
I thank J. Vasconcelos for his assistance in the construction of the Latin names utilized
in this paper, and D. Nolf and the anonymous referees for their suggestions to improve
previous versions of this manuscript. This work was partially funded by the ‘Fundação
para a Ciência e a Tecnologia’ in the framework of the Pluriannual Program.
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The Lagenar Otoliths of Teleosts: Their Morphology and Its Application in Species Identification, Phylogeny and Systematics

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The Lagenar Otoliths of Teleosts: Their Morphology and Its Application in Species Identification, Phylogeny and Systematics

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Journal of Fish Biology (2003) 62, 1268–1295

doi:10.1046/j.1095-8649.2003.00106.x, available online at http://www.blackwell-synergy.com

The lagenar otoliths of teleosts: their morphology and

(Received 29 May 2001, Accepted 28 March 2003)

Key words: asteriscus; lagenar otoliths; morphology; phylogeny; systematics.

Tel.: þ351 21 7500000; fax: þ351 21 7500009; email: carlos.assis@fc.ul.pt

MATERIALS AND METHODS

THE MORPHOLOGY OF THE ASTERISCI

the homologue of the extremum ventralis is located in the posterior region

reduction of the lobus minor, is limited by the antirostrum and pseudoantiros-

THE THREE BASIC MORPHOLOGICAL TYPES OF

Clupeomorpha, Scopelomorpha, Protacanthopterygii and Acanthopterygii; the

THE ASTERISCI OF FISHES AS A SOURCE OF

In spite of representing a very small fraction of the total diversity of forms

The diagnostic power of the asterisci

among otoliths from close species. Notwithstanding, when in presence of repre-

1 – Ventral limit of the lobus minor conspicuous in the outline. Excisura

2 – Rostrum asterisci clearly supramedian, long and tapering to an

The reflection of phylogenetic relationships

The vertical type of asteriscus

As each of these tendencies is displayed by the different taxa in a different

The horizontal type of asteriscus

considerations to be made, concerning some possible tendencies of the horizontal

provided by the otoliths asteriscus, however, is not significantly different, in

The gyro type of asteriscus

dorsal (antirostral) and ventral (pseudoantirostral) regions of the lobus major to

Frost, G. A. (1926c). A comparative study of the otoliths of the neopterygian fishes.

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