Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Lutjanus synagris (Acanthuriformes: Lutjanidae): reproductive biology

and synchronization with lunar phases in the Gulf of Salamanca,

Colombian Caribbean

Camilo B. García*1; https://orcid.org/0000-0003-0373-7916

Nelson Sandoval2

1. Departamento de Biología, Universidad Nacional de Colombia, Carrera 45 N° 26-85. Bogotá DC, Colombia;

cbgarciar@unal.edu.co (*Correspondence)

2. Facultad de Ingeniería-Ingeniería Pesquera, Universidad del Magdalena, Calle 29H3 N° 22-01, Santa Marta, Colombia;

nelsonenrique33@yahoo.es

Received 07-VI-2024. Corrected 03-II-2025. Accepted 30-V-2025.

ABSTRACT

Introduction: Lutjanus synagris is one of the most common demersal fishes in the Colombian Caribbean and is

of interest to artisanal fisheries, although in decline. The study of its reproductive patterns provides insight into

its biology relevant to the general study of tropical fishes.

Objective: To assess the condition factor and gonadosomatic index, to determine the monthly incidence of

mature fish, to estimate the mean length at sexual maturity (Lm), and to explore predictors affecting the condi-

tion factor and gonadosomatic index in the lane snapper L. synagris.

Methods: Extirpated gonads of males and females were classified according to their maturity status and related to

fish length. The condition factor and gonadosomatic index were estimated according to standard formulae. Their

dependency on variables related to climate, geographic space, and behavior were assessed using linear models.

Results: Condition factor was stable throughout the year, while the gonadosomatic index showed an increas-

ing pattern from the first months of the year (dry season), reaching a peak in the second semester of the year.

The incidence of mature fish was higher in about the same months as when the gonadosomatic index showed

high values. An Lm of 26 cm total length was estimated for both sexes, 27 cm for males and 25 cm for females.

Condition factor and gonadosomatic index were associated with several predictors, notably with moon phases.

Conclusion: A spatial and temporal signal for the reproductive patterns of L. synagris was found but decoupled

from climate seasonality. The reproductive rhythm of L. synagris was related to moon phases as the main envi-

ronmental cue.

Key words: biorhythms; condition factor; gonadosomatic index; tropical fishes.

RESUMEN

Lutjanus synagris (Acanthuriformes: Lutjanidae): biología reproductiva y sincronización con las fases

lunares en el Golfo de Salamanca, Caribe colombiano

Introducción: Lutjanus synagris es uno de los peces demersales más comunes en el Caribe colombiano y es de

interés para la pesca artesanal, que está en declive. El estudio de sus patrones reproductivos ayuda en la comp-

rensión de su biología, relevante para el estudio general de los peces tropicales.

Objetivo: Evaluar el factor de condición y el índice gonadosomático, determinar la incidencia mensual de peces

maduros, estimar la longitud promedio de madurez sexual (Lm) y explorar predictores que afectan el factor de

condición y el índice gonadosomático del pargo L. synagris.

https://doi.org/10.15517/rev.biol.trop..v73i1.60355

AQUATIC ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

INTRODUCTION

Condition factor, gonadosomatic index

and maturation patterns are functional features

of importance in the study of the biology of

tropical marine fishes with relevance in con-

servation and management. Condition factor

alludes to the physical health of individuals

and hence of the population (Gubiani et al.,

2020; Le Cren, 1951). It focuses on the question

whether an individual weights what it should

according to its measured length. An overall

measure of condition factor is indicative of

favorable or unfavorable environmental condi-

tions for the population. In turn, the gonadoso-

matic index focuses on the reproductive cycle

a fish goes through typically over a year. The

main purpose of such an index is to identify

the population time of spawning (Fontoura et

al., 2009; Lowerre-Barbieri et al., 2011). The

incidence of mature fish in time may shed light

onto the same question, but more importantly,

the study of the proportion of mature fish

in representative length samples provides an

estimation of one main life-history event in a

fish, the mean length at sexual maturity (Lm),

relevant for fishery management (Cope & Punt,

2009; Froese et al., 2008).

The lane snapper Lutjanus synagris Linnae-

us (1758) is a fish species of commercial interest

in the Caribbean coast of Colombia (García

& Duarte, 2024; Ministerio de Agricultura y

Desarrollo Rural, 2024), although in decline

globally (García, 2010; García & Duarte, 2024),

in our study area, the Golf of Salamanca (Mar-

tínez-Viloria et al., 2022; Salazar-Perez et al.,

2020), and in other areas of the Colombian

Caribbean (Martínez-Viloria, 2022; Ramírez

et al., 2017). It is a notorious member of soft

bottom fish assemblages (García et al., 1998;

Manjarres et al., 2001) occurring throughout

the extension of the coast from South to North

in soft bottoms (García & Armenteras, 2015).

Although Allen (1985) states that L. synagris

main habitat is around coral reefs and vegetated

sandy areas it turns out to be the most com-

mon fish in historical soft bottom trawls in the

Colombian Caribbean (García, 2018; García &

Armenteras, 2015).

Few studies have been conducted in

Colombian Caribbean waters on reproductive

aspects of L. synagris. Based on the gonadoso-

matic index different peaks in spawning activity

have been suggested for this species. Arteaga et

al. (2004) found two peaks in spawning, one

from April to August with a maximum in May

and a second peak in October in the Northern

Colombian Caribbean Sea (Guajira Peninsula

and Tayrona National Natural Park). Posada-

Peláez et al. (2012) reports a peak in spawning

from August to November in an area including

the Eastern corner of the Gulf of Salamanca, the

Tayrona National Natural Park, and some local-

ities to the East but not reaching the Guajira

Métodos: Las gónadas de machos y hembras extirpadas fueron clasificadas según su estado de madurez y asocia-

das a la longitud del pez. El factor de condición y el índice gonadosomático se calcularon con fórmulas estándar.

Su dependencia de variables asociadas al clima, espacio geográfico y comportamiento fue evaluada mediante

modelos lineales.

Resultados: El factor de condición fue estable en el año mientras que el índice gonadosomático mostró un patrón

de incremento en los primeros meses (época seca) para alcanzar un máximo el segundo semestre del año. La

incidencia de peces maduros fue mayor en los mismos meses en que el índice gonadosomático mostró valores

altos. Un Lm de 26 cm longitud total fue estimado para ambos sexos, 27 cm para machos y 25 cm para hembras.

El factor de condición y el índice gonadosomático se asociaron con varios predictores, notablemente, con las

fases lunares.

Conclusiones: Se encontró una señal espacial y temporal en los patrones reproductivos de L. synagris pero desa-

coplada de la estacionalidad climática. El ritmo reproductivo de L. synagris está vinculado a las fases lunares como

principal señal ambiental.

Palabras clave: biorritmos; factor de condición; índice gonadosomatico; peces tropicales.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Peninsula. In both cases it is suggested that

increased spawning is associated with the rainy

season environmental and climatic conditions,

that is, higher sea surface temperatures and

lower sea surface salinity in contrast to the dry

season (December to April) when an upwelling

phenomenon takes place in the area conductive

to lower sea surface temperature and higher

salinity (Andrade & Barton, 2005).

Regarding condition factor no anteced-

ent could be found in the primary literature

focusing on Colombian populations of L. syn-

agris. Estimations of Lm, on the other hand,

are numerous and disparate (see García &

Ramírez, 2016, their Table 1) for the Northern

Colombian Caribbean coast including the Gulf

of Salamanca with values ranging from 25.1

cm (Altamar et al., 2015) to 34.0 cm (Arteaga

et al., 2004) total length for both sexes. Geo-

graphically the closest estimate of Lm is that

of Posada-Peláez et al. (2012) whose study area

partly overlap ours, with 28 cm total length for

both sexes.

In this study, estimations of condition fac-

tor, gonadosomatic index and maturation pat-

terns in time are given for L. synagris in the Gulf

of Salamanca as well as an estimate of Lm. Via

linear models the dependency of these indices

on several climatic, environmental, and opera-

tive variables was explored.

MATERIALS AND METHODS

Fig. 1 shows the study area and the sam-

pling stations. Sampling was conducted month-

ly from April 1997 to March 1998, excepting

January and June. Stations were selected in

three clusters according to oceanographic con-

ditions in the Gulf of Salamanca which at the

center shows an intrusion of oceanic water

(García et al., 2013), plus some interspersed

stations. Trawling operations at a station lasted

an average 30 minutes at an average speed of

3.5 knots. On board L. synagris individuals were

measured (standard length) and weighted. A

representative sample of fishes were brought

to the lab (in total 166 males and 123 females)

and the stage of maturation was assessed by

the appearance of gonads according to the

scale of Holden & Raitt (1975). Afterwards the

gonads of males and females were extirpated

and weighted. Fishes classified in stage III, IV

Fig. 1. Study area Gulf of Salamanca, Colombian Caribbean, and sampling stations. The arrow shows the position of the Gulf

of Salamanca in the Colombian Caribbean coast.

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and V (Holden & Raitt, 1975) of gonadal devel-

opment were labeled as mature. At each station

measurements of bottom salinity and tempera-

ture were made with the aid of a CTD probe.

Condition factor was found with the for-

mula (Le Cren, 1951): CF = Wex / Wob, where

Wex is the expected weight of an individual

and Wob is the observed weight of the same

individual. Expected weights were estimated

from the length weight relationship derived

by García et al. (1998) that originated from

cruises in the study area in the period 1995 to

1998. The length weight relationship is based

on 2015 individuals, size ranging from 8 cm to

33.5 cm (García et al., 1998). The formula for

Wex is: Wex = 0.0898 × (SL / 10)2.63, where SL

is standard length (cm). The gonadosomatic

index was found with the formula: GDS = 100

× (Wgo / Wob), where Wgo is the weight of the

gonad and Wob is the observed weight of the

same individual.

Temporal trends of both indices and of the

incidence in percentage of mature versus unma-

ture fish were explored by means of boxplots

and barplots. To estimate Lm for males, females

and both sexes a logistic regression model was

adjusted to the raw mature-immature data with

individual lengths as the predictor variable. The

Lm estimated and their bootstrap confidence

intervals were transformed from standard

length to total length (TL) with the formula

TL = 0.0 + 1.203 × SL, found in FishBase for L.

synagris (Froese & Pauly, 2023).

The dependency of condition factor (CF)

and gonadosomatic index (GDS) on variables

related to climate (bottom salinity and tem-

perature), space (latitude and longitude decimal

transformed and depth), and fish behavior

(time of the day when captured decimal trans-

formed and moon phase of the day of capture)

were explored. Latitude, longitude, depth, and

time of the day correspond to the middle posi-

tion and time of the research ship during a

given tow. Correlations among the quantitative

variables were found to be less than 0.5 (Pearson

correlation test) in all cases. The linear model is

CF / GDS ~ Latitude + Longitude + Depth +

Time + Temperature + Salinity + Moon Phase.

Regarding the results of the model, the depen-

dency of CF and GDS on the moon phases was

further explored with the model: CF / GSD ~

Moon Phase.

Multiple comparisons of means were con-

ducted with the Tukey HSD test. Numerical

analysis and graphics were done with R base

(R Core Team, 2021) and R packages par-

zer (Chamberlain & Sagouis, 2021), car (Fox

& Weisberg, 2019), ggplot2 (Wickham, 2016),

cowplot (Wilke, 2020), Rmisc (Hope, 2022) and

multcomp (Hothorn et al., 2008).

RESULTS

Fig. 2 and Fig. 3 show the monthly esti-

mates of CF and GDS index, respectively.

Except for March samples, no peak is noticeable

in CF with ups and downs around one across

the year and this is true for males, females and

both sexes (Fig. 2A, Fig. 2B, Fig. 2C). The peak

in March median is probably due to a small

sample size (n = 8). CF medians grouped by

season do not differ, but CF is more dispersed

in the dry season (Fig. 2D). GDS index (Fig.

3) behaves in a richer fashion than CF. For

males, females and both sexes there appear to

be a building up of gonad material as the year

progresses to May in the case of males and

to September in the case of females, but with

great dispersion (Fig. 3A, Fig. 3B). When plot-

ted together the GDS index shows maximum

values in March and April (Fig. 3C). Female

gonads are noticeable heavier than male gonads

as evidenced by the scale of the plots (Fig.

3). When grouped by season no difference in

median GDS index is noticeable (Fig. 3D).

The incidence in percentage of mature

individuals in the year is shown in Fig. 4. Mature

males and females were found in all monthly

samples. Proportionally mature females showed

higher incidence than males (Fig. 4A, Fig. 4B).

In general, the incidence of mature individuals

is higher in the first months of the year with a

decreasing trend from February to August (Fig.

4C). However, when grouped by season no dif-

ference in percentage incidence of mature fish

is noticeable (Fig. 4D).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

The logistic model provides an estimate of

Lm of 27 cm 26-28 CI (confidence interval) for

males, 25 cm 24-26 CI for females and 26 cm

25-27 CI for both sexes (Fig. 5) which are with-

in the same order of magnitude of other Lm

estimates (see García & Ramirez, 2016, their

Table 1) in the Northern Colombian Caribbean.

Results of the linear models adjusted to

CF and GDS indices are shown in Table 1 and

Table 2, respectively. An interesting finding is

that moon phase has a significant effect in all

instances (p < 0.05) affecting both indices and

sexes (Table 1, Table 2). These results prompted

the formulation of an additional linear model

to closely explore the effect of moon phases,

which is referred further below.

Predictors affected differentially CF and

GDS indices of males and females. CF of males

is affected positively by latitude while it is

negatively affected by bottom temperature. CF

of females is affected positively by longitude

and negatively by time of the day when the

Fig. 2. Monthly series of condition factor of Lutjanus synagris in the Gulf of Salamanca, Colombian Caribbean from April

1997 to March 1998. A. Males. B. Females. C. Both. D. Seasonal comparison for both sexes.

Table 1

Association between predictor variables and condition

factor for Lutjanus synagris in the Gulf of Salamanca,

Colombian Caribbean.

Variables/Sex Males Females Both

Latitude < 0.05 + N.S. < 0.05 +

Longitude N.S. < 0.05 + N.S.

Depth N.S. N.S. N.S.

Time N.S. < 0.05 - N.S.

Temperature < 0.05 - N.S. < 0.05 -

Salinity N.S. N.S. < 0.05 +

Moon phase < 0.05 < 0.05 < 0.05

The sign positive or negative indicates the direction of the

association. Number in the table is the p-value of the test of

hypothesis of no association. N.S., not significant.

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individual was captured (Table 1). GDS index

of males, in turn, is affected positively by longi-

tude and negatively by bottom salinity while for

females GDS index was positively affected by

depth and bottom temperature and negatively

by bottom salinity (Table 2). When pooling

males and females the response of CF and GDS

index was contrasting. CF responded negatively

to the bottom temperature and positively to

bottom salinity while GDS index responded

positively to bottom temperature and negative-

ly to bottom salinity. CF was affected by latitude

while GDS was not. GDS index was affected by

depth while CF was not (Table 1, Table 2).

The linear model with moon phase as pre-

dictor confirmed the results of the full model.

For males and females and when pooled CF

and GDS indices were significantly affected by

the moon phase (p < 0.05), except for CF in

males. Table 3 shows the results of Tukey HSD

multiple comparisons test. Noticeably, while for

CF in females and both sexes the moon phase

that is different from the others is waxing moon

(Fig. 6), for GDS index the moon phase that

is different from the others is waning moon

(Table 3, Fig. 7).

Fig. 3. Monthly series of gonadosomatic index of Lutjanus synagris in the Gulf of Salamanca, Colombian Caribbean from

April 1997 to March 1998. A. Males. B. Females. C. Both. D. Seasonal comparison for both sexes.

Table 2

Association between predictor variables and gonadosomatic

index for Lutjanus synagris in the Gulf of Salamanca,

Colombian Caribbean.

Variable/Sex Males Females Both

Latitude N.S. N.S. N.S.

Longitude < 0.05 + N.S. N.S.

Depth N.S. < 0.05 + < 0.05 +

Time N.S. N.S. N.S.

Temperature N.S. < 0.05 + < 0.05 +

Salinity < 0.05 - < 0.05 - < 0.05 -

Moom Phase < 0.05 < 0.05 < 0.05

The sign positive or negative indicates the direction of the

association. Number in the table is the p-value of the test of

hypothesis of no association. N.S., not significant.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

DISCUSSION

The first thing to notice is that the health

condition of L. synagris was stable over the

year and that no time period be it monthly or

seasonal has bearing on health status at least in

what the condition factor can tell us. Food offer

in the year appears to be stable regardless of the

seasonal upwelling. Upwelling phenomena have

been traditionally associated with increased

Fig. 4. Monthly percentage incidence series of mature individuals of Lutjanus synagris in the Gulf of Salamanca, Colombian

Caribbean from April 1997 to March 1998. A. Males. B. Females. C. Both. D. Seasonal comparison for both sexes.

Table 3

Multiple comparisons of mean values of condition factor and gonadosomatic index of Lutjanus synagris by moon phase in

the Gulf of Salamanca, Colombian Caribbean.

Group/Index Condition Factor Gonadosomatic Index

Males p > 0.05 for all pairwise comparisons Waning Moon ≠ New Moon, Full Moon

Females Waxing Moon ≠ New Moon, Full Moon,

Waning Moon Waning Moon ≠ Full Moon

Both Waxing Moon ≠ New Moon, Full Moon,

Waning Moon Waning Moon ≠ New Moon, Waxing Moon, Full Moon

Statistical significance was set at < 0.05. Tukey HSD test.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Fig. 5. Sigmoid curves representing the relation between

standard length and the probability of being mature for

Lutjanus synagris in the Gulf of Salamanca, Colombian

Caribbean. Length at which there is 50 % probability

of being mature is signaled. Model line plus standard

deviation. A. Males. B. Females. C. Both sexes.

Fig. 6. Mean condition factor plus 95 % confidence

intervals by moon phases for Lutjanus synagris, Colombian

Caribbean. A. Males. B. Females. C. Both sexes.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

primary productivity and via the food web to

better food offer in the water column. However,

for the Gulf of Salamanca it has been found that

the effect of the upwelling is more reflected on

temperature and salinity changes rather than

on increased primary production (Ramírez,

1990). For instance, García et al. (2013) found

that concentrations of chlorophyll did not differ

in the dry (upwelling) and rainy seasons. On

the other hand, seasonal change is not without

consequences. Duarte & García (1999) found

that the importance of crustaceans in the diet

of L. synagris diminishes in favor of teleost in

the dry season.

Males and females of L. synagris associ-

ated differentially to the predictors explored

here in terms of CF. Males respond to the

latitudinal gradient while females respond to

the longitudinal gradient. Thus, the distribu-

tion of individuals in the geographic space is

not homogeneous. Further research should

be conducted focused on characterizing the

environmental gradients offered to L. synagris

in the Gulf of Salamanca to find a mechanistic

explanation of these patterns.

The effect of movement of individuals dur-

ing the day was detected only for females. The

later the capture, the lower the CF. A hypothesis

to be explored is whether this pattern is related

to a feeding cycle with more incidence of empty

stomachs at night. In turn, CF of males had a

negative association with bottom temperature

while for both sexes a positive association with

bottom salinity emerges. These conditions, low

temperature and high salinity are reminiscent

of the conditions in the dry season (upwelling)

thus this pattern enters in contradiction with

the finding that no monthly or seasonal trend

was found in the year. García et al. (2013) found

three water masses in the Gulf of Salamanca

with characteristic temperature and salinity

and moving limits as the season change. One to

the East under the influence of brackish water

of the Ciénaga Grande de Santa Marta, one to

Fig. 7. Gonadosomatic index plus 95 % confidence

intervals by moon phases for Lutjanus synagris, Colombian

Caribbean. A. Males. B. Females. C. Both sexes.

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the center sector of the Gulf of Salamanca cor-

responding to a salty oceanic intrusion, and a

third to the West under the influence of dis-

charge of the Magdalena River. Thus, the Golf

of Salamanca is not a homogeneous water body.

The location of the stations may explain this

pattern that is in line with the associations with

latitude and longitude discussed earlier.

Unlike CF that showed no monthly ten-

dency, the GDS index shows a tendency of

increasing median values as the year progresses

including and from the dry season (upwell-

ing) and extending beyond. Median monthly

values, however, showed substantial variability.

On the other hand, the percentage incidence of

mature individuals indicates predominance of

mature fish approximately in the same months

when high GDS index values were seen. Thus,

with caveats, it appears that the main spawning

activity of L. synagris occurred the first part of

the year from February to a peak in September

for females but continued the year round. These

months partly overlap with the range given by

Arteaga et al. (2004) and less so with the range

given by Posada-Peláez et al. (2012). The main

difference is that the range given here fully

includes the dry season months. Given that

bottom water temperatures and salinity are

associated significantly with the GDS index,

positively for temperature and negatively for

salinity, and temperature increases, and salinity

decreases as the year progresses thus it appears

that spawning augmented activity is triggered

by the gradients of these environmental vari-

ables until temperature and salinity reach a

maximum and a minimum, respectively.

The same case as with CF, no seasonal dif-

ference in median GDS index was found. Con-

sidering these findings, the idea that the climate

season per se correlates with the spawning cycle

in L. synagris should be abandoned in favor of a

scenario in which it is the gradient of increasing

temperature and decreasing salinity that trig-

gers augmented spawning activity.

The estimated length at first maturity, Lm,

is well in the range of Lm values calculated for

L. synagris in the Colombian Caribbean coast.

However, these measures have been criticized

because many of them come from sampling

the fishery instead of sampling the popula-

tion. García and Ramírez (2016) demonstrated

that estimates of Lm significantly correlated to

mean size of captured individuals that in turn

depended on the selectivity patterns of the fish-

ing gears. Thus, Lm estimates obtained from

sampling the fishery do not necessarily reflect

what is happening in the population and man-

agement measures based on Lm as indicator

obtained that way may not be optimal. It would

be interesting to conduct research on the cur-

rent status of this trait in the Gulf of Salamanca

for comparison.

The most interesting finding in this study

is the significant association between moon

phases and both CF and GDS indices values.

Many marine species, including fishes, use

moon phases as cue to synchronize relevant

events in their life cycle (Andreatta & Tessmar-

Raible, 2020; Tessmar-Raible et al., 2011) hence

it might not be a surprise that L. synagris does

so. However, to our knowledge in the Colombi-

an Caribbean, apart from some mentions based

on fishermen local knowledge (Altamar et al.,

2010), the phenomenon of synchronization of

biological activity with moon phases has not

been documented, excepting Lopez and García

(2001) in reference to postlarvae and juveniles

of shrimps. In the context of fisheries Altamar

et al. (2010) related CPUE with moon phases

for the bigeye scad (Selar crumenophthalmus).

It is suggestive that while CF is signifi-

cantly larger in the waxing moon, GDS index is

significantly larger in the waning moon. Fishes

appear to be preparing for the spawning events

along the lunar cycle. This hypothesis should

be tested with new research and field sampling

designed to test it. At any rate it is clear that the

lunar cycle superimposes on the yearly patterns

for CF and GDS indices in L. synagris in the

Gulf of Salamanca.

Shortcomings of the studies referenced

here, including this, are the time period of

observations that rarely extends more than

one year and also that many are dependent on

fishery landings instead of unbiased samples of

the population. This renders the observations

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

to criticisms as just anecdotical. However, as

reproduction is an event so central in the life

history of species it can be viewed as a conser-

vative set of traits. The challenge is to learn how

reproduction characteristics that emerged in

evolution are modulated by biological, climatic,

environmental factors and impacted by human

activities like fishing through clear cut ques-

tions and well-planned sampling.

Condition factor of the lane snaper L.

synagris is stable throughout the year when

followed at monthly time steps. The gonadoso-

matic index builds up the first months of the

year to a maximum in September. The gradi-

ent of increasing temperature and decreasing

salinity appears to trigger augmented spawning

activity rather than a strict conformance with

the dry and rainy seasons. Of the predictive

variable tested the moon phases were the most

determinant with the waxing moon signifi-

cantly associated to the highest values of condi-

tion factor and the waning moon significantly

associated to the highest values of the gonad-

osomatic index.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

This work was supported by COLCIEN-

CIAS Grant No 2105-09-176-94 for the project

Estudio ecológico pesquero de los recursos

demersales del Golfo de Salamanca, Caribe

colombiano. Estimación de la variabilidad de

los componentes biológicos del Sistema. Com-

ments by two anonymous reviewers helped to

improve the manuscript.

REFERENCES

Allen, G. R. (1985). Snappers of the world. An annotated

and illustrated catalogue of lutjanid species known to

date. FAO Fishery Synopsis, 125(6), 1–208.

Altamar, J., Manjarrés-Martínez, L., Duarte, L. O., Cuello

F., & Escobar-Toledo, F. (2015). Qué tamaños debería-

mos pescar? [Technical Report]. Autoridad Nacional

de Acuacultura y Pesca (AUNAP), Universidad del

Magdalena.

Altamar, J., Parraga, D., & Duarte, L. O. (2010). Influencia

de la fase lunar en las tasas de captura de Selar crume-

nophthalmus (Carangidae) en el caribe de Colombia.

Una aproximación por modelos lineales generali-

zados. Acta Biológica Colombiana, 15(3), 121–130.

https://doi.org/10.15446/abc

Andrade, C. A., & Barton, E. D. (2005). The Guajira upwe-

lling system. Continental Shelf Research, 25, 1003–

1022. http://dx.doi.org/10.1016/j.csr.2004.12.012

Andreatta, G., & Tessmar-Raible, K. (2020). The still dark

side of the moon: Molecular mechanisms of lunar-

controlled rhythms and clocks. Journal of Molecular

Biology, 432, 3525–3546. https://doi.org/10.1016/j.

jmb.2020.03.009

Arteaga, E., Criales-Hernandez, M. L., Rodriguez, D. J.,

Gomez-Canchong, P., & Manjarres, L. (2004). Diná-

mica reproductiva de los stocks de pargo rayado

Lutjanus synagris (Linnaeus, 1758) en el área norte

del Caribe Colombiano. In L. Manjarres (Ed.), Pes-

querías demersales del área norte del mar Caribe de

Colombia y parámetros biológico-pesqueros y pobla-

cionales del recurso pargo (pp. 203–215). Universidad

del Magdalena.

Chamberlain, S., & Sagouis, A. (2021). parzer: Parse messy

geographic coordinates. R package (Version 0.4.1.)

[Software]. CRAN. https://CRAN.R-project.org/

package=parzer

Cope, J. M., & Punt, A. E. (2009). Length-based reference

points for data-limited situations: Applications and

restrictions. Marine and Coastal Fisheries: Dynamics,

Management, and Ecosystem Science, 1, 169–186.

https://doi.org/10.1577/C08-025.1

Duarte, L. O., & García, C. B. (1999). Diet of the lane

snapper, Lutjanus synagris (Lutjanidae), in the Gulf of

Salamanca, Colombia. Caribbean Journal of Science,

35(1-2), 54–63.

Fontoura, N. F., Braun, A. S., & Milani, P. C. (2009).

Estimating size at first maturity (L50) from gona-

dosomatic index (GSI) data. Neotropical Ichthyo-

logy, 7(2), 217–222. http://dx.doi.org/10.1590/

S1679-62252009000200013

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Fox, J., & Weisberg, S. (2019). An R Companion to Applied

Regression (3rd ed.). SAGE Publications.

Froese, R., & Pauly, D. (Eds.). (2023). FishBase. World Wide

Web electronic publication. https://www.fishbase.org/

Froese, R., Stern-Pirlot, A., Winker, H., & Gascuel, D.

(2008). Size matters: How single-species management

can contribute to ecosystem-based fisheries manage-

ment. Fisheries Research, 92, 231–241. http://dx.doi.

org/10.1016/j.fishres.2008.01.005

García, C. B. (2010). Conocimiento tradicional: Lo que los

pescadores artesanales del Caribe colombiano tie-

nen para decirnos. Pan-American Journal of Aquatic

Science, 5(1), 78–90.

García, C. B. (2018). Spatial richness patterns of soft bottom

fish in the Colombian Caribbean shelf and slope. Acta

Biológica Colombiana, 23(1), 59–65. http://dx.doi.

org/10.15446/abc.v23n1.66530

García, C. B., & Armenteras, D. (2015). Atlas de la ictiofau-

na demersal de fondos blandos del Caribe Continental

Colombiano: Aproximación a su biodiversidad. Uni-

versidad Nacional de Colombia, Facultad de Ciencias.

García, C. B., & Duarte, L. O. (2024). A diagnosis of the arti-

sanal fishery landings in the Colombian Caribbean

coast by means of indicators. Marine and Fishery

Sciences, 37(2), 277–293. https://doi.org/10.47193/

mafis.3722024010504

García, C. B., & Ramirez, J. (2016). Perceived length at

first maturity in the lane snapper, Lutjanus synagris

(Linnaeus, 1758) (Perciformes: Lutjanidae), along the

Caribbean coast of Colombia. Pan-American Journal

of Aquatic Science, 11(1), 60–69.

García, C. B., Duarte, L. O., & Ramírez, G. (2013). Fisiogra-

fía y oceanografía del Golfo de Salamanca (mar Cari-

be, Colombia). In L. M. Melgarejo, C. B. García (Eds.),

Investigación en Ciencias del Mar: Aportes de la Uni-

versidad Nacional de Colombia (pp. 111–140). Uni-

versidad Nacional de Colombia. Facultad de Ciencias.

Red de Estudios del Mundo Marino, REMAR.

García, C. B., Duarte, L. O., & von Schieller, D. (1998).

Demersal fish assemblages of the Gulf of Salamanca,

Colombia (southern Caribbean Sea). Marine Ecology

Progress Series, 174, 13–25. https://doi.org/10.3354/

meps174013

Gubiani, E. A., Ruaro, R., Ribeiro, V. R., & Gomez de Santa

Fe, U. M. (2020). Relative condition factor: Le Cren’s

legacy for fisheries science. Acta Limnologica Brasile-

ra, 32, e3. https://doi.org/10.1590/S2179-975X13017

Holden, M. J., & Raitt, D. F. (1975). Manual de ciencia pes-

quera. parte 2. Métodos para investigar los recursos y su

aplicación. Documento Técnico de Pesca Nº 115. FAO.

https://openknowledge.fao.org/handle/20.500.14283/

f0752s

Hope, R. M. (2022). Rmisc: Ryan miscellaneous. R package

(Version 1.5.1.) [Software]. CRAN. https://CRAN.R-

project.org/package=Rmisc

Hothorn, T., Bretz, F., & Westfall, P. (2008). Simulta-

neous inference in general parametric models. Bio-

metrika, 50(3), 346–363. https://doi.org/10.1002/

bimj.200810425

Le Cren, E. D. (1951). The length-weight relationship and

seasonal cycle in gonad weight and condition in the

perch (Perca fluviatilis). Journal of Animal Ecology,

20(2), 201–219. http://dx.doi.org/10.2307/1540

Lopez, A. C., & García, C. B. (2001). Postlarvas y juveniles

de camarones Farfantepeneus spp y Xiphopenaeus kro-

yeri en la Boca de la Barra (Ciénaga Grande de Santa

Marta), Caribe Colombiano. Boletín de Investigaciones

Marinas y Costeras, 30, 177–198.

Lowerre-Barbieri, S. K., Ganias, K., Saborido-Rey, F.,

Murua, H., & Hunter, J. R. (2011). Reproductive

timing in marine fishes: Variability, temporal scales,

and methods. Marine and Coastal Fisheries, 3, 71–91.

http://dx.doi.org/10.1080/19425120.2011.556932

Manjarres, L., García, C. B., & Acero, A. (2001). Carac-

terización ecológica de las asociaciones de peces

demersales del Caribe Colombiano norte con énfasis

en los pargos (Lutjanidae). Boletín de Investigaciones

Marinas y Costeras, 30, 77–107.

Martínez-Viloria, H. M., Franke-Ante, R., Narváez-Baran-

dica, J. P., & De La Hoz-Maestre, J. (2022). Esta-

do de Lutjanus synagris en tres áreas protegidas

del Caribe colombiano: Evaluación basada en datos

limitados. Boletín de Investigaciones Marinas y Cos-

teras, 51(2), 83–106. https://doi.org/10.25268/bimc.

invemar.2022.51.2.1104

Ministerio de Agricultura y Desarrollo Rural. (2024). Series

Históricas. Servicio Estadístico Pesquero Colombia-

no (SEPEC). http://sepec.aunap.gov.co/Estadisticas/

SeriesHistoricas

Posada-Peláez, C., Santafé-Muñoz, A., Grijalba-Bendeck,

M., Bustos-Montes, D., González-Sarmiento, E., de

la Hoz-Maestre, J., Castro-Martinez, L., Gómez-

Cubillos, C., Guerrero-Berna, P., Olaya-Reyes, C.,

Palacio-Barros, C. M., & Restrepo-Gómez, D. (2012).

Aspectos biológicos de las principales especies comer-

ciales de la zona costera del departamento del Mag-

dalena, Caribe colombiano. In M. Grijalba-Bendeck,

D. Bustos-Montes, C. Posada-Peláez, & A. Santafé-

Muñoz (Eds), La pesca artesanal marítima del depar-

tamento del Magdalena (Colombia). Una visión desde

cuatro componentes (pp. 115–415). Fundación Uni-

versidad de Bogotá Jorge Tadeo Lozano, Ministerio de

Agricultura y Desarrollo Rural.

R Core Team (2021). R: A language and environment for

statistical computing. R Foundation for Statistical

Computing. https://www.R-project.org/

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e60355, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Ramírez, G. (1990). Evaluación de parámetros hidrográfi-

cos y su relación con la surgencia en las aguas cos-

teras. In J. M. Díaz (Ed.), Estudio ecológico integrado

de la zona costera de Santa Marta y Parque Nacional

Natural Tayrona (pp. 19–54). INVEMAR.

Ramírez, J. G., Lleonart, J., Coll, M., Reyes, F., & Puentes, G.

(2017). Improving stock assessment and management

advice for data-poor small-scale fisheries through

participatory monitoring. Fisheries Research, 190,

71–83. http://dx.doi.org/10.1016/j.fishres.2017.01.015

Salazar-Perez, C., Choles-Rodriguez, E., & Manjarres-

Martinez, L. (2020). Short-term changes in demersal

fish assemblages exploited by an artisanal set gill net

fishery in the Caribbean Sea (Colombia). Ciencias

Marinas, 46(1), 39–56. http://dx.doi.org/10.7773/

cm.v46i1.3041

Tessmar-Raible, K., Raible, F., & Arboleda, E. (2011).

Another place, another timer: Marine species and the

rhythms of life. BioEssays, 33, 165–172. https://doi.

org/10.1002/bies.201000096

Wickham, H. (2016). ggplot2: Elegant Graphics for Data

Analysis. Springer.

Wilke, C. O. (2020). cowplot: Streamlined Plot Theme and

Plot Annotations for ‘ggplot2’. R package (Version

1.1.1.) [Software]. CRAN. https://CRAN.R-project.

org/package=cowplot