Age Determination of Microlepidotus brevipinnis (Steindachner, 1869) (Pisces: Haemulidae) in the Coast of Jalisco, Mexico, by Reading Otoliths and Scales
This study deals with age determination of the brassy grunt Microlepidotus brevipinnis by reading otoliths (sagittae) and scales which allowed the identification of 13 age groups. Growth of otoliths and scales is proportional to the growth of the fish. The time of formation of fast and slow growth bands in sagittae, as well as the time of ring formation in the scales is one year. The highest growth in length of this species takes place during the first year of life, in which the organism reaches 125.0 mm, this reduces natural mortality of the individuals by decreasing depredation. Growth in weight of this species is isometric. Sexual differentiation of the organisms is apparent after the age of two years and there were differences in the average length for each age in scales and otoliths for males and females. The age of other members of the Haemulidae family were compared with those obtained in the present study and M. brevipinnis reaches its oldest age in 13 years. It occupies a middle point (442.4 mm) in relation to the total lengths of other haemulids in the Mexican Pacific coast.
to cite this article:
K. Granados-Flores, M. Gallardo-Cabello, E. Espino-Barr and E.G. Cabral-Solis, 2010. Age Determination of Microlepidotus brevipinnis (Steindachner, 1869) (Pisces: Haemulidae) in the Coast of Jalisco, Mexico, by Reading Otoliths and Scales. International Journal of Zoological Research, 6: 1-12.
Microlepidotus brevipinnis (Steindachner, 1869) (Pisces: Haemulidae)
(Fig. 1) represents 10% of the catch of the artisanal fishery
in the coast of Jalisco, Mexico (Espino-Barr et al.,
2004a), 2 to 11 tons from 2000 to 2007. Locally it is very appreciated due
to its low price and great flavor.
This and other species of the Haemulidae family are mentioned in the taxonomic
lists of Castro-Aguirre (1978), López
and Bussing (1982), Van Der Heiden (1985), Chávez-Ramos
et al. (1994), Allen and Robertson (1994),
Béarez (1996), Madrid-Vera
et al. (1998), Espion-Barr et al. (2003),
Mariscal-Romero and van der Heiden (2006). Age and growth
analysis of five species of this family were performed by Cruz-Romero
et al. (1993) using an indirect method of length-frequency. Gallardo-Cabello
et al. (2003) and Espino-Barr et al. (2004a,
b) also analyzed age growth of Anisotremus interruptus. Taking in
account the scarce information available on these species, it is important to
analyze the parameters of their population dynamics and determine the age by
||Brassy grunt Microlepidotus brevipinnis
The direct methods to determine the age based on the interpretation of growth
rings in hard structures as: scales (Gallardo-Cabello et
al., 2005), otoliths (Gallardo-Cabello et al.,
2006; Espino-Barr et al., 2006), vertebrae
(Anislado-Tolentino and Robinson-Mendoza, 2001) and
spines (Alvarado-Castillo and Félix-Uraga, 1998),
among others. These direct methods allow determining the age groups with a higher
precision as compared to the indirect methods of length frequency analysis,
eliminating most of the biases produced by small samples taken with low periodicity
and allowing to identify the first age groups (Peñailillo
and Araya, 1996; Campana, 1999; Campana
and Thorrold, 2001; Mascareñas-Osorio et al.,
2003; Berg et al., 2005; Francis
et al., 2005).
The purpose of this study was
age groups of M. brevipinnis using otoliths and scales, to validate
||Evaluate the time of formation of growth bands
||Relationship between the growth of these structures and of
||Analyze differences between sexes
||Compare lengths at age with other species of the Haemulidae
These results will provide information on some aspects of the life history of this species that will help assess the fishery and suggest regulation measures.
MATERIALS AND METHODS
During five days of every month from January 2005 to June 2006, samplings of 1050 individuals from the commercial catch were taken in Melaque, coast of Jalisco (104°4030 to 104°4330W and 19°1200 to 19°1330N). Of each individual total and standard length, height, total and eviscerated weight was measured. Sex was determined for 480 individuals; the other 570 were eviscerated by fishers before landing. For age study two structures were obtained from 353 individuals: otoliths (sagittae) and scales.
To obtain the otoliths sagittae, a transverse cut was made in the ventral cranial
cavity and the brain was removed; the left and right semicircular canals were
extracted and otoliths separated, rinsed with water, dried and stored dry in
Eppendorf tubes labeled with number, date, total length and sex. Data on the
length and the width were regarded for each otolith through their observation
in a stereoscopic microscope with a graduated ocular lens (Fig.
2a, b). All measures were made on the right-sided sagitta, in order to eliminate
possible differences due to variations between the otoliths in the same organism.
The sample size was calculated with the formula described by Daniel
In the case of the sagittae, constants of the relationship were calculated
for Rostrum Length (RL), Antirostrum Length (AL) and Width (Wi). The rostrum
length was also related to the total length of the fish.
|| Left and right sagittae of M. brevipinnis, (a) external
and (b) internal aspect
|| Scale of M. brevipinnis
Regressions were done by least squares and a variance analysis (anova) was
carried out (Mendenhall, 1987; Zar, 1996).
This last test was also applied to analyze possible morphometric differences
between the otoliths of males and females. Identification of growth rings was
done observing the sagittae in a stereoscopic microscope with transmitted light.
The time of the growth ring formation was analyzed, observing whether the sagittae
borders had fast or slow growth rings.
Around 15 scales were taken from the area under the left pectoral fin, below
the lateral line (Ehrhardt, 1981; Holden
and Raitt, 1975; Ruiz-Durá et al., 1970)
and stored in dry labeled envelopes. Following the method described by Holden
and Raitt (1975) the scales were washed to clean them of any tissue stuck
to them. Later, 10 scales of each individual were placed between two slides,
sealing them with adhesive tape and labeled. Reading of the scales was done
with the help of a transparency projector Kodak Ektagraphic with a 127 mm lens
(which increases the size of the scale 13.4 times). Scales were read independently
by two different individuals and the results were compared (Fig.
Determination of the marginal increment was carried out monthly for each individual, in order to determine the date in which the mark is formed and to validate its periodicity.
Analysis of otoliths and scales showed that M. brevipinnis recruited by the fishing gear has four growth rings, therefore the sampling strategy was modified and a biological sampling was included to capture younger specimens, with a ½ inch mesh net.
Morphometric Analysis of the Species
Table 1 shows the relationship between total length vs.
standard length, height, total and eviscerated weight of M. brevipinnis.
The relationship between total length and standard length and total length and
height are expressed by the values of the exponents b = 1.000 and b = 1.042,
respectively for the species which correspond to an isometric growth (Table
2). Some differences of these values are shown for sexes and the undetermined
The relationship between total length and total and eviscerated weight are described by the values of the exponents b = 3.042 and b = 2.954, respectively, which represents an isometric growth, which means that the fish grows similarly in length and weight (Table 3).
|| Values of standard length, height, total and eviscerated
weight for total length classes of M. brevipinnis
||Values for total length vs. standard length and height for
the species, females, males and undetermined of M. brevipinnis
||Values for total length vs. total and eviscerated weight for
the species, females, males and undetermined individuals of M. brevipinnis
In the case of males and females, values show a tendency to a negative allometry. The highest values of these indexes are reached by undetermined specimens with values b = 3.085 and b = 3.027 for total and eviscerated weight, respectively.
Otoliths Description of the Sagittae
The anterior margin exhibits a small excisura, as a tiny notch, showing
an underdeveloped antirostrum (Fig. 2). The shape of the rostrum
varies between individuals and also between the right and the left sagittae
in the same specimen. The posterior margin presents a sharped postrostrum without
an excisura minor and a pararostrum. The dorsal margin is rectilinear from the
antirostrum to the center of the otolith, beyond which the dorsal margin descends
with a marked inclination towards the posterior margin. The ventral margin is
curved from the rostrum to the posterior margin. The anterior, posterior and
ventral margins show regular denticles that spread irregularly and disappear
in some sections of the otolith as the fish ages. The internal aspect of the
otolith is convex; this feature increases with age; its surface is smooth and
marked lengthwise by a deep sulcus which decreases in width from the center
of the otolith to the end of its posterior margin, showing a clear difference
between the ostium and the cauda. The external aspect is concave; its thickness
decreases abruptly in the longitudinal sense from the middle area to the anterior
margin. The average width of the sagitta is 2.1 times its length.
Growth of the Sagitta
Results of fish length classes and different measures of the otolith are
shown in Table 4. The relationship between length and width
of the sagitta is expressed by the value of exponent b = 0.64 which corresponds
to a negative allometric growth (Table 5). The determination
index of the relationship of the two series of data is R2 = 0.75,
with F = 997 of the anova, which indicates that the sagitta grows more in length
||Rostrum (RL), width (Wi) and antirostrum (AL) length in different
size classes of the sagitta of Microlepidotus brevipinnis
|| Relationships between the rostrum and other measures of the
The allometric relationship between the length of the rostrum and of the antirostrum
is expressed in the isometric growth index b = 1.02, with the highest values
of R2 and F obtained for this analysis, i.e., 0.96 and 7751, respectively.
This shows that among the sizes and age groups, the length of the antirostrum
and rostrum increase in a similar way.
The relationship between the fish total length and the length and width of the sagitta is shown in the Table 6. The highest value of the allometric index related to the total length of the fish corresponds to the rostrum and antirostrum length, b = 0.60; this means that there is a proportionality between the increase of the length of the sagitta and of the fish. This structure is suitable to describe the growth in length of the organism. The values of the determination index R2 and anova (F) show a high correlation between the analyzed structures in every case. The values of the allometric growth indexes are lower than 1 due to the difficulty to relate very small structures with the corporal length of the fish.
Identification of the Growth Rings
Analysis of the growth bands in the sagittae allowed the identification
of thirteen groups (Table 9). The percentage of the otoliths
which showed perfectly defined bands of growth was 93.8%. The growth bands can
be seen more clearly in the rostrum aspect of the otolith (Fig.
2), where the deposit of aragonite and otolin takes place with more intensity.
Formation Time of the Fast and Slow Growth Rings
In most of the sagittae the fast growth band (opaque) occurs in November
and the slow growth band (hyaline) in July, which proves that formation of growth
rings takes place once a year.
Scales Description of the Scale
Scales of M. brevipinnis are catenoid and rectangular-shaped; the
length is contained around 1.6 times the width. The anterior area is divided
in sectors by very marked radios that converge in the focus; their number varies
between 6 and 8. The anterior margin presents lobes and in some adults shows
serial fissures as distinct bifurcations. The focus is well defined and in an
eccentric position due to the faster growth of the anterior area than the posterior
area. The lateral borders are smooth. The growth rings are well defined and
were interpreted as the formation of a double dark and continuous line defined
by a clear and translucent space. The triangular-shaped ctenii are in the rear
area and project toward the posterior margin which is thinner and brittle compared
with the rest of the scale (Fig. 3).
Growth of the Scale
The relationship between length and width of the scale is expressed by the
value of exponent b = 0.885 which corresponds to a negative allometric growth.
The determination index of the relationship of the two series of data is R2
= 0.79, with F = 967 of the anova, which indicates that the scale grows more
in length than widthwise (Table 7).
|| Relationships between the total length of the fish (L) and
the measures of the sagittae
|| Relationship between length classes, length and width of
the scale of M. brevipinnis
|| Age-length distribution of M. brevipinnis with scales
The relationship between total fish length and length of the scale is b = 0.977, which is very close to isometry (R2 = 0.483 and anova F = 229.05). This means that there is a high proportionality between the increase of scale length and of fish length; for this reason the scale is suitable to describe the growth in length of the fish.
Identification of the Growth Rings
The analysis of the growth rings in the scales allowed the identification of thirteen groups (Table 9). The percentage of the scales which showed perfectly defined rings of growth was 85%; the rest (15%) were regenerated scales. Table 8 shows the age-length distribution.
|| Mean total length (mm) for each ring identified with scales
and sagittae of M. brevipinnis
|| Mean total length (mm) for each ring identified with scales
and sagittae for sexes of M. brevipinnis
Determination of the Marginal Increment
Maximum, average and minimum data of the margin between the last ring and
the scale border for every month. June was the month with the smallest value
of the margin which validates the use of scales to determine the age in this
species and its annual rhythm.
Differences Between Sexes
Table 10 shows the differences in the mean total length
for each ring for males and females observed by the study of the sagittae and
scales. Sexual differentiation is possible from ring 2 on, that is, 220 mm of
total length. Data for rings 12 and 13 correspond to 8 individuals that were
eviscerated by fishers before landing and their sex could not be determined.
Morphometric analysis of the species showed that M. brevipinnis is stouter
during the first stage of its life; once it reaches sexual maturity (approximately
at 220 mm, Table 10) it becomes slimmer because waste of
fatty acids is higher during the spawning season and the individuals loose
weight. Cruz-Romero et al. (1993) in Manzanillo
coast and Espino-Barr et al. (2004a, b)
in Jalisco coast, also found negative allometric indexes between standard length
and eviscerated weight for mature individuals of M. brevipinnis with
values of b = 2.45 and b = 2.53, respectively.
Other members of the Haemulidae family show a negative allometric index of
the relationship between standard length and eviscerated weight: Haemulon
steindachnerii (b = 2.13), H. flavigutattum (2. 35), Pomadasys
panamensis (2.49), Anisotremus caesius (2.65), A. taeniatus
(2.71) and Xenichthys xanti (2.76). Other species show an isometric
index: H. sexafasciatum (2.91), H. maculicauda (2.99), A. interruptus
(3.027, P. elongatus (3.21) and P. bayanus (3.26) (Espino-Barr
et al., 2004b). Microlepidotus brevipinnis is in a middle
position in the Haemulidae family.
Table 9 shows that the mean length of the organisms with rings 1 to 5 have similar sizes in otoliths and scales and greater differences occur in groups 6 to 13. Ring 9 has the highest difference between these structures, because from 6th ring on there is a decrease in the number of organisms (Table 8) and possibly a higher bias in the data. Both cases show the highest growth rate from 0 to 1 ring, with an increment of 125 mm. From the 2nd ring on, growth rate decreases abruptly. Reduction of organisms from age six on, is due to fishing pressure.
There is very little information on M. brevipinnis, but as a family,
Table 11 shows the average lengths for each age of different
Haemulidae species in different countries. According to this Table
11, M. brevipinnis reaches the higher age (13 years), followed by
H. plumieri (12 years) in North and South Carolina (Manooch,
1976) and A. interruptus in Manzanillo, Colima (Gallardo-Cabello
et al., 2003) and H. aerolineatum in the Southeastern USA
(Manooch and Barinas, 1979) both with 9 years. Other
reports show that Haemulids do not reach more than 7 years of age.
The smaller species are Pomadasys jubelini (3 years) (Alberdi,
1971), H. steindachnerii (3 years) (Eslava, 1991),
P. striatum (4 years) (Latif and Shenouda, 1972),
Brachydeuterus auritus (4 years) (Raitt and Sagua, 1969)
and P. hasta (from Western India, 5 years, Deshmukh,
1973). However P. hasta reaches the largest length of this family
(569 mm, Deshmukh, 1973).
Haemulids which reach larger lengths in Jalisco, according to Espino-Barr
et al. (2004a) are H. sexfasciatus (650.2 mm), A. interruptus
(621.8 mm), P. panamensis (557.5 mm), H. flaviguttatum (492.1
mm), A. caesius (445.6 mm) and M. brevipinnis (442.4 mm). Smaller
species of Haemulidae are: P. elongates (430.2 mm), P. bayanus
(395.7 mm), A. taeniatus (334.0 mm), H. scudderi (310.0 mm),
H. maculicauda (301.0 mm), A. dovii (283.0 mm), H. steindachnerii
(274.8 mm), A. pacifici (249.0 mm) and Xenichthys xanti (238.4
mm). According to these results, M. brevipinnis occupies a middle point
in the mid Mexican Pacific coast. Species that reach smaller ages as B.
auritus, H. steindachnerii, P. hasta, P. jubelini,
P. striatum, H. plumieri and H. aurolineatum these last two in
Campeche are found in lower latitudes, near the equator, where temperature is
higher, which coincides with Taylors (1958, 1960),
who reported that organisms living in high temperatures reach smaller ages.
Conversely Haemulids living in higher latitudes reach older ages, i.e., H.
aurilineatum (Southeastern USA., Manooch and Barans 1979),
H. plumieri (North and South Carolina, Manooch, 1976),
A. interruptus (Manzanillo, Colima, Gallardo-Cabello
et al., 2003) and M. brevipinnis (Jalisco, present study).
This study provided information on population structure in length, weight and age classes, necessary for the assessment plans of the fishery.
We want to thank the artisanal fishers from Jalisco, who were very helpful in giving us information and facilitating organisms. Arturo Garcia-Boa and Marcos Puente-Gómez helped in the field and lab., work; also MS Alejandro Gómez-Ponce and Biol. Claudio Padilla-Hernández for their help in the lab. Several institutions gave financial support: UNAM, CONACyT and INP.
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