Natural Feeding Of Lutjanus Novemfasciatus Juveniles in the Cuyutlan Lagoon, Colima, Mexico

Trophic dynamic offers knowledge on the function and structure of a population, which is necessary for the management of commercially exploited resources [1,2]. Studies focused on the feeding of organisms allow to understand the biology of species, since the diet depends on organisms fulfilling their life cycle [3]. However, the feeding of the species can vary according to the site, the season of the year and physiological requirements during growth and reproduction, known as trophic adaptability [4]. Fish from the Lutjanidae family are active demersal predators, which feed regularly during the night, sunrise or sunset and their principal food source are demersal organisms as invertebrates and fish. Due to the quality of their meat, many of these species are of commercial importance in tropical and subtropical regions [5-7]. One the ten species of snappers in the Pacific, from Northern Meico to Peru, Lutjanus novemfasciatus is the largest with a maximum length of 1.7 m [6] and a weight of 35.7 kg [8]. Adults live in rocky zones and coral reefs, and during their juvenile stages they can be found in estuarine systems and river mouths [9]. According to Duncan et al. [10], the length of first sexual maturation is from 55 to 61 cm in males and 60 to 68 cm in females.  

The Cuyutlan Lagoon is the largest lagoon in the state of Colima, Mexico, and one of its vessels was declared a RAMSAR site in 2011 [11,12], and despite the fact that multiple studies have been carried out on some organisms [13-16], till now there is no study related to this species even though it is an important resource in the study area.

The objective of this study is to determine the feeding composition, as to its ontogenic and seasonal variations of the juveniles of this species.

The study area is the Cuyutlan Lagoon, Colima, Mexico, located on the Pacific Ocean littoral (18° 57' - 19° 03' N y 104° 00' - 104° 19' W), it measures 37 km long by 6 m in width. It is divided by natural and artificial physical barriers into four water bodies called vessels. The communication with the sea is in the middle portion of the lagoon body and the main supply of fresh water comes from the Armeria river [12,17-19]. 

The analyzed organisms come from biological samples taken from the commercial catches of coastal fishermen. Data of each organism is total length (TL cm). the stomach was conserved in a solution of formalin 10% till the analysis time. Once in the laboratory, the food entities were identified down to the lowest possible taxon based on the degree of digestion of the prey, using guides and specialized keys for each category [9,20-26]. 

The food entities of the stomach content were counted and weighed, by the numerical method (%N), the gravimetric method (%P) according to Rosecchi & Nouaze [27] and the frequency of occurrence (%FO) of Pinkas et al. [28]. The relative importance of each species of prey of L. novemfasciatus was established by the Relative Specific Importance Index for preys (%PSIRI) according to Brown et al. [29], expressed as: 

%PSIRI= [%Fi x (%PNi + %PPi)] / 2 

Where: %PSIRIi is the Relative Specific Importance Index for the prey i, %PNi is the specific number in percentage for the prey i, and %PPi is the specific weight percentage for the prey i.

To evaluate if the size of the sample (number of stomachs) is enough to describe the diet of L. novemfasciatus, trophic diversity accumulation curves of were made from the total number of prey identified, with the use of EstimateS 9.1.0. software [30], per size class and annual season; with a 100 randomizations without replacement, to assure that the curves reach an asymptotic value. Shannon index value [31] was represented in function of the accumulated number of examined stomachs [32]. The curve was considered asymptotic if at least two values prior to the trophic diversity (H´TOT) were in the interval of H´TOT ±0.05 H´TOT [33]. 

Organisms were grouped by size intervals (<28.0, 28.1-34.0, 34.1-40.’, y >40.1cm TL) to analyze the diet related to the increase of the length of the fish. To determine the length values of the species of prey, total length was used. To determine the number of length classes Sturges formula was used: 

K = 1+3.322 (Log10 n) 

Where: K = number of class intervals and n is the number of total values. 

Once the number of class intervals was obtained, the width of this interval was determined, which was calculated with the following formula [34]:

W = R/k

Where: W = interval width, R = maximum value minus the minimum value and k = number of intervals. 

In addition, an evaluation of the amplitude of the trophic spectrum (niche amplitude) was carried out for the dry and rainy seasons in order to determine the degree of specialization in the diet of L. novemfasciatus. For this purpose, the Levins measure [35], standarized by Hespenheiden method [36] was used, with the scale of 0 to 1, considered as a specialist when the value is close to zero and generalist when the value is close to one [37].

General composition of the diet 

From 2012 to 2013 148 stomachs of L. novemfasciatus juveniles were analyzed, of which 99 67%) presented stomach contents and 49 (33%) were empty. Maximum total length of the sampled organisms was 49 cm and the minimum 21.30 cm. the most representative prey in numeric percentage was Portunus xantusii (%N = 20.88), while the percentage in weight was P. vannamei (%P = 34.75). The remains of fish presented the most occurrence frequency in percentage (%FO = 21.21), however, the most important prey for its contribution to the PSIRI was P. vannamei (%PSIRI = 20.05) and P. xantusii (%PSIRI = 15.18), (Table 1). 

In the stomach contents 91 food entities were found, which were grouped in seven superfamilies corresponding to the order or decapoda, of the subphylum crustacean: Galathoidea, Xanthoidea, Goneplacoidea, Cancroidea, Portunoidea, Penaeoidea and Cymothoidae, and other two more categories were added: remains of fish and crustaceans. The main food component for its contribution to PSIRI with 81.1% were the Decapods, of which the Peneids were the preferred food (%PSIRI=41.55) (Table 1). 

Table 1: List of species in the composition of the trophic content of juveniles of L. novemfasciatus %FO (frequency of occurrence), %N (numerical percentaje). %W (weight porcentaje), %PSIRI (Relative Specific Importance Index for preys). Total %PSIRI for each functional group is marked in bold leters. 

In this study fish species nwere not identified because of their advanced stage of digestión. The Decapod that obtained the highest percentage was P. vannamei (%PSIRI= 27.27). Total number of stomachs were considered enough to describe the throphic spectrum of L. novemfasciatus, since the accumulation curve presented a tendency toward asymptote (Figure 1).

Figure 1: Trophic diversity accumulation curve for juveniles of L. novemfasciatus. The solid line corresponds to the Shannon diversity (H’) and the dashed horizontal lines indicate the index H’ ± 0.05 H´.

Diet composition by size

In the composition of the diet by size, the number of stomachs examined was adequate for its description ( < 28.0=17, 28.0-34.0=34, 34.1-40.0=33 and >40.1=14) (Figure 2). The prey with higher contribution to the gravimetric index in organisms with smaller sizes than 28.0 cm LT was P. xantusii (%P =28.19). Organisms from 28.1 to 3.0 cm LT and from 34.1 to 40.0 cm the most important prey was P. vannamei (%P =44.52 and %W=29.09), respectively. For those with lengths higher than 40.1 cm LT, Portunus asper (%P =47.34) was the most important. The preys with highest numeric percentage in organisms with lengths smaller than 28.9 cm LT was the Xanthidae family (%N=20.00). From 28.1 to 34.0 cm LT Eury panopeous (%N=18.75) and from 34.1 to 40.0 cm LT P. vannamei (%N=23.33). Finally, in bigger lengths than 40.1 cm LT P. xantusii was the most important with %N=46.15. (Figure 3). The most frequent prey in organisms of sizes smaller than 28.0 cm LT was P. vannamei (%FO=23.53). The most frequemt preys in sizes from 28.1 to 34.0 cm LT were cataloged as fish remains (%FO=26.47). From 34.1 to 40.0 cm LT, also fish remains and also Penaeus sp. (%FO=15.15). In sizes bigger than 40.1 cm LT fish remains were found with higher frequency %FO=42.86 (Figure 4).

Figure 2: Cumulative curves of trophic diversity (H’) by length class of L. novemfasciatus: a)  < 28 cm, b) 28.1- 34 cm, c) 34.1 – 40 cm y d) >40 cm total length.

Figure 3: Gravimetric composition (W%), in number (N%), occurrence frequency (FO%) and Relative Specific Importance for preys (PSIRI%), by length class in the diet of L. novemfasciatus  juveniles.

 Figure 4: Frequency of Occurrence (%FO) of preys in the diet of L. novemfasciatus juveniles, by size clases.

P. xantusii (%PSIRI=22.09) and P. vannamei (% PSIRI=20.46) were the most important preys for their contribution to PSIRI% in sizes smaller than 28.0 cm LT, from 28.1 to 34.0 cm LT and 34.1 to 40.0 cm LT, the most important prey was P. vannamei (%PSIRI=30.07, %PSIRI=26.21), and finally for the range of lengths bigger than 40.1 cm LT, the preys with the highest percentage were P. xantusii (%PSIRI=30.97) and P. asper (% PSIRI=31.36) (Figure 3).

Diet composition by time of year

According to the composition of the diet by time of year in both seasons (rainy and dry) the predominant prey in terms of weight percentage was P. vannamei with %P=38.07 and %P=32.65, respectively. Also as according to the relative specific importance for preys with %PSIRI=29.33 and %PSIRI=25.34, respectively. Regarding the numeric index, the preys that presented a higher percentage during rainy and dry seasons were P. xantusii (%N=23.53 and %N=18.03) and P. vannamei (%N=20.59 and %N=10.03), respectively. During both seasons fish remains were the most frequent ítem (rainy season: %FO=21.05, and dry season: %FO=20.97) (Figures 5&6).

Figure 5: Gravimetric composition (W%), in number (N%), occurrence frequency (FO%) and Relative Specific Importance for preys (PSIRI%), by time of year (dry and rainy) in the diet of L. novemfasciatus juveniles. 

Figure 6: Occurrence frequency (%FO) of preys by time of year (dry and rainy) in the diet of L. novemfasciatus  juveniles.

Regarding the amplitude of the trophic niche, a higher value was observed during dry season. Despite the differences found in observed data between times of the year, there was no statistic significant differe ce between rainy and dry seasons. During all the months a value less tan 1 sas obtained (number: 0.3, weight: 0.3), therefore it can be considered a diet with low trophic diversity. It is considered that the total number of stomachs is enough to be able to describe the trophic spectrum by time of year, since the accumulation curve presented a tendency towards the asymptote (Figure 7).

Figure 7: Cumulative curves of trophic diversity (H’) by time of year: a) rainy and b) dry, of L. novemfasciatus juveniles.

In Barra de Navidad lagoon, Aguilar- Betancourt et al. [38] report that the diet of L. novemfasciatus was mainly composed of decapods (49.9 %) and fish (43.7%) for their contribution to PSIRI and in less proportion stomatopods and mollusks, however, in the present study decapods predominated (81.1%) for their contribution to PSIRI, and in less proportion by stomatopods and mollusks. The variation between the two studies may be due to the characteristics of each estuarine system and the availability of food. The Cuyutlán Lagoon has a wider extension and depth range than the Barra de Navidad lagoon, which could influence the distribution of species and a mainly muddy bottom, with a large amount of detritus [13], which provides optimal conditions for the development of invertebrates. Cruz-Romero & Espino-Barr [19] noted that shrimp and crabs are some of the most abundant invertebrates in the lagoon. Other authors have documented species of the Lutjanidae family base their diet on crustaceans and fish, such as L. peru, L. guttatus, L. campechanus, L. griseus, L. argentiventris, L. analis and L. colorado [2,38-43]. According to Ruíz-Nieto [44], the most predominant prey of L. novemfasciatus in estuarine habitats of Sinaloa were shrimp (Penaeidae), crabs (Portunus spp. and Callinectes spp.), and fish (not identified), which is similar to the present work. Studies in some species of the Lutjanidae family (juveniles and adults) have shown a preference for crustaceans in their diets, such as in L. jordani, L. synagrisy and L. argentiventris [38,45,46]. 

In this study, variations in the weight and number of food entities were observed between small and large sizes. This may be because as the size of the predator increases, it looks for heavier prey to maximize the energy gain with respect to the capture effort, possibly due to the approach of the reproductive stage, as mentioned by Rojas [47] and Rojas et al. [48]. Szedlmayer & Lee [40] found that the diet of juvenile L. campechanus changes with increasing fish size: before 30.0 cm, their diet consists of shrimp, quetognathids, squid and copepods, after this size, the organisms feed on fish, larger quantities of squid and crabs, and continue to feed on shrimp. These variations in diet during fish growth could be due to morphological changes due to the increase in size, making the capture of prey more efficient [49].  Aguilar-Betancourt et al. [38] found that L. novemfasciatus has more piscivorous habits than L. colorado and L. argentiventris. Due to their morphological characteristics such as a more slender body and flatter head, they are more hydrodynamic and allow for easier displacement, the width of their mouth is larger and they have longer teeth, which makes them more efficient in their capture of prey. Some authors have documented the transition from invertebrate to fish feeding in species of the family Lutjanidae [41,48,50-52].

In the dry season, 14 food components were found and 13 in the rainy season. On the feeding of Syacium ovale and Achirus mazatlanus in Barra de Navidad lagoon, Padilla-Gutiérrez [49] mentions that a greater diversity was found in dry season, in terms of number and frequency, despite the fact that it is not about the species and lagoon body, the results are similar in terms of diversity in both study periods. Seasonal changes and variation in diet may be due to the availability and abundance of food during the rainy or dry season. Diet variation may be due to changes in the ecosystem to which organisms must adapt [53]. In the study of the feeding habits of L. argentiventris and L. colorado, Santamaría-Miranda et al. [54] found a low niche breadth, and mentioned that the diet of these species is cyclical in nature because the prey species are always present. This coincides with the present study, since in general the trophic niche breadth was low showing a specialist tendency; L. novemfasciatus consumed P. vannamei, P. xantusii and fish throughout the year. In tropical regions there is a relative stability with respect to climatic changes throughout an annual cycle, which allows stability in the food base of fish, however, there may be variations in the hydro-meteorological cycles that can influence the feeding habits of fish in different regions [55].

This work was made possible by the invaluable support of the coastal fishermen of Laguna Cuyutlán. As well as †Arturo Garcia-Boa and Marcos Puente-Gómez from the Regional Fisheries Research Center (CRIP-Manzanillo) of INAPESCA who were in charge of collecting the samples. Likewise, the Department of Sustainable Development of Coastal Zones of the University of Guadalajara, where the data were analyzed and processed.

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