Indian Journal of Animal Research

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Feeding Aspects and Reproductive Biology of Smalltooth Emperor, Lethrinus microdon (Valenciennes, 1830) in the Gulf of Mannar along the Indian Southeast Coast

Sanjay Chandravanshi1,*, N. Jayakumar1, R. Durairaja1, S. Aanand2, E. Suresh3, Chandran Sudhan4, Narsingh Kashyap3, Swapnil Ananda Narsale5
1Department of Fisheries Biology and Resource Management, Fisheries College and Research Institute, Tamil Nadu Dr. J. Jayalalithaa Fisheries University, Thoothukudi-628 008, Tamil Nadu, India.
2Erode Centre for Sustainable Aquaculture Bhavanisagar, Erode-638 451, Tamil Nadu, India.
3Department of Fish Genetics and Breeding, Institute of Fisheries Postgraduate Studies, Tamil Nadu Dr. J. Jayalalithaa Fisheries University, Vaniyanchavadi-628 008, Tamil Nadu, India.
4Department of Fisheries Resource Management, Dr. M.G.R. Fisheries College and Research Institute, Tamil Nadu Dr. J. Jayalalithaa Fisheries University, Pooneri-601 204, Tamil Nadu, India.
5Department of Fish Pathology and Health Management, Fisheries College and Research Institute, Tamil Nadu Dr. J. Jayalalithaa Fisheries University, Thoothukudi-628 008, Tamil Nadu, India.

Background: The Gulf of Mannar, situated on the southeast coast of India, is a unique distinctive productive marine ecosystem and is rich in biodiversity. Its significance stems from the presence of a string of 21 islands spanning 140 km between Rameswaram and Tuticorin, each adorned with a wealth of coral reefs and hosting a variety of ornamental and marine fish species despite their small and uninhabited nature. It is also first Marine Biosphere Reserve in South and South-East Asia, covers approximately an area of 19,000 sq. km.

Methods: Fish samples were collected monthly from three major fishing harbours situated in the Gulf of Mannar along the southeast Coast: Thoothukudi Fishing Harbour, Pamban Fishing Harbour and Chinnamuttom Fishing Harbour from December 2022 to November 2023. These species are caught using trawl net, gill net and hook and line and collected specimens were identified using FAO sheets and standard literature.

Result: In this study, a total of 405 specimens were collected. The length weight relationship was estimated for male: W = 0.0522.57, r2 = 0.94, female; W = 0.0572.51, r2 = 0.94; and pooled; W = 0.0632.54, r2 = 0.90, respectively. Fulton condition factor (KF) value for male: 1.22, female: 1.33; and pooled: 1.19, respectively. Gonadosomatic index (GSI) of L. microdon was ranged between 1.21-2.21 for male and 1.85-2.31 for female and Gastrosomatic index (GaSI) varied from 1.45-2.37 for male whereas 1.89-4.43 was found for female. Hepatosomatic index (HSI) of L. microdon was ranged between 0.45-2.54 for male while for female was ranged between 1.11-3.21. The male and female (M:F) sex ratio was estimated 1:0.67. The peak spawning season was observed in June to August. The results provide the baseline biology information required to evaluate the fisheries and sustainable management of these underexploited fishes in the Gulf of Mannar ecosystem.

Lethrinids are one of the most commercially significant groups of fish along many tropical coasts and are regarded as an important source of animal protein for communities in the western Pacific and the Indian Ocean (Carpenter and Allen, 1989). Lethrinus microdon, are commercially important food fish and highly significant due to their abundance and high market value. It is widely distributed throughout the Indo-West Pacific, commonly found in sandy areas near coral reefs at depths of approximately 80 meters (Ebisawa, 2006). L. microdon is primarily caught using longline, gill nets, trawl nets and hook and line. Despite being a significant component of artisanal fisheries and widely distributed in the Gulf of Mannar (GoM), there is limited knowledge about the feeding and reproductive biology and stock status of lethrinid fishes on national and international levels.

The Gulf of Mannar was designated as the first Marine Biosphere Reserve in India as well as South East Asia. Covering an area of about 10,500 km2, this region hosts approximately 3600 species of flora and fauna (Magesh and Krishnakumar, 2019).  The Gulf of Mannar, located on the southeast coast of India, is referred to as a “Marine Biological Paradise” because of the remarkable richness and diversity of its fauna; hence its conservation is very important of this valuable flora and fauna thriving in this region (Retnamma et al., 2021).

While a comprehensive understanding of the principles of biology and stock is crucial for the development and execution of effective conservation programs (Mawa et al., 2021), there is a lack of comprehensive information regarding the feeding and reproductive characteristics of L. microdon despite the economic significance in the GoM. Hence, investigating feeding and reproductive biology facilitates understanding the basic feeding habits and resource utilization. The study of biology of emperor therefore becomes very important not only for using the data to develop improved exploitation strategies, but also to comprehend the substantial structural changes brought about in the ecosystem when they are removed by fishing. Therefore this study aimed to explore certain aspects of the feeding and reproductive biology of L. microdon, including gonadosomatic index, gastrosomatic index sex ratio, hepatosomatic index, sex ratio, length at first maturity stages and fecundity. Therefore, the current study aimed to gather information on the feeding and reproductive biology of the Lethrinus microdon in the Gulf of Mannar, India may potentially improve further stock status study of this species at GoM.
Study area

Fish samples were collected fortnightly from three major fishing harbours situated in the GoM along the southeast Coast: Thoothukudi Fishing Harbour, TFH (8°43' 26.7"N; 78° 56' 23.12" E), Pamban Fishing Harbour, PFH (9° 27' 84. 82" N; 79°15' 11. 99  E) and Chinnamuttom Fishing Harbour, CMFH (8° 23' 92. 65" N; 77° 16’ 83.17" E) (Fig 1).

Fig 1: Sampling grounds of the gulf of mannar.



Sample collection and identification

A total of 405 specimens were collected during the present study from December 2022 to November 2023. These species were caught using longline, trawl nets, gill netsand hook and line. Upon collection, the specimens were identified using standard literature (Carpenter and Allen, 1989) and transported to the laboratory under refrigerated conditions for further examination. The total length and total weight were measured to the nearest 0.1 cm and 0.01 g, respectively.
 
Length-weight relationship and condition factor

The relationship between length (L) and weight (W) were estimated according to Le Cren (1951) using the following formula:
                               
                                      W = aLb                                                        
Where:
W = Body weight (g).
L = Total body length (cm).
a = Intercept.
b = Slope value.

The parameters ‘a’ and ‘b’, along with their corresponding 95% confidence limits (CL) and coefficient of determination (r2), were calculated using linear regression analysis in Microsoft Office Excel. When the coefficient of allometry (b) is equal to 3, the growth rates of length and weight are equal (Ricker 1975); hypo-allometry, or negative allometry growth, occurs when length increases more than predicted weight; hyper-allometry, or positive allometry growth, occurs when length increases less than predicted weight (Shingleton, 2010). The b value for each species was estimated using Student’s t-test (Sokal and Rohlf 1987) with a 95% confidence interval to determine if it significantly differed from the isometric value (H0: b = 3).

The condition of fish species under study was determined using Fulton’s condition factor (KF) (Fulton, 1904) by using the following formula:
 
                                                                       
Where:
W = Observed body weight (g).
L = Observed length of fish (cm) and the factor 100 is utilized to normalize K to unity.
 
The fish is said to be in good condition when K=1, while average or poor growth condition is indicated if K<1.
 
Biological characteristics

The collected specimens were dissected to extract all internal organs. The gut, intestine, liverand gonads were weighed using a digital balance, which were then used to assess biological parameters such as the Hepatosomatic index (HSI), Gastrosomatic index (GaSI) and Gonadosomatic index (GSI). The weights of the fish, gut, liverand ovaries were measured with a digital electronic balance to the nearest 0.01 g.

Gut content analysis

The analysis of the gut contents was conducted through close examination under a microscope. Visual estimation of gut content was performed for all specimens to identify the diet composition present in the fish gut. The GaSI (Desai, 1970) and HSI (Welcomme, 2001) were calculated using the standard protocol:
 
Reproductive characteristics

Gonadosomatic Index

The female of ovaries was dissected using very fine forceps after that whole ovary was carefully removed and preserved in 5% formalin solution. The weight of the ovaries was measured in electronic weighing balance with accuracy 0.01 g. The Gonadosomatic index (GSI) was calculated by following formula (Bal and Rao, 1984):
 

Where:
GW = Gonad weight in (g).
TW = Total body weight in (g).

Sex ratio

The sex ratio was determined by considering the total number of male and female individuals collected during the sampling period. The significance differences in sex ratio were assessed using a Chi-square test (X2) at a 1% level of significance (P<0.05) (Snedecor and Cochran, 1989) to test if the observed ratio between males and females differs from the expected 1:1 (male: female) ratio.

Length at first maturity

The mean length at sexual maturity (L50) is the length at which 50% of individuals are matured (Stages III, IV and V). The length at first maturity was estimated by using the logistic curve formula (Udupay, 1986):
 
                                P = 1/[1+ e(-r(L-Lm))]
Where:
P = Proportion of mature individuals by length (L).
r = Slope of the curve.
Lm = Length corresponding to a proportion of 50% of mature individuals (L50).

Maturity stage

The maturity stages of female gonads were determined based on visualization of the morphometric characteristics of ovaries such as colour, shape, vascularization and size of the ova. The reproductive stages in female gonads were determined using the four-stage of the gonads maturity scale and categorized as Stage-I Immature; Stage II maturing; Stage III matured and Stage IV ripe (Brown-Peterson et al., 2011).

Fecundity

Fecundity, defined as the number of oocytes present in the ovary and weight, was estimated by using the gravimetric method. A subsample of 0.5 g eggs was taken from each portion of the ovary (anterior, middle and posterior) and preserved in Gilson fluid solution (Simpson, 1951) for easy release of the oocytes, which were then counted and measured with the help of a Sterozome microscope (20x) (Nikon DS-Fi1, Japan). Fecundity was calculated using the formula proposed by Garg et al. (2002):

Where:
F= Estimated fecundity of the individual.
Fs = Number of oocytes in a sample.
GW = Total weight of the ovary.
GWs = Weight of the subsample of the ovary.
Length-weight relationship and condition factor

In the present study, the b value of L. microdon was estimated 2.57 (male), 2.51 (female)and 2.54 (pooled) whereas coefficient of determination (r2) was 0.94 (male), 0.94 (female) and 0.90 (pooled) respectively (Table 1 and Fig 2).

Table 1: Length-weight relationship of Lethrinus microdon.


Fig 2: Length-weight relationship of Lethrinus microdon.



In the length-weight relationship of fishes, the b values were typically found between the estimated ranges of 2.5-3.5 (Froese, 2006). Ontomwa et al., (2018) and Matthews et al., (2019) have estimated growth co-efficient (b) estimates for L. microdon (2.6), which are in the same line of the present findings. Mehanna et al., (2017) and Al Kamel et al. (2020) observed b values as 2.9 and 3.1 for L. microdon which is higher than the recent findings, confirming the presence of a negative allometric pattern in this species, which is in line with the present study. Rajathy et al., (2021) estimated the slope value for L. lentjan (2.9) and L. microdon (2.8) which is higher than the current investigation. Grandcourt et al., (2010) and Daliri et al., (2012) registered a b value of 3 for L. microdon from the Southern Arabian Gulf and Northern Persian Gulf, which is higher than present observation. Thangaraj et al. (2022) reported b value for L. microdon (2.8) which is slightly higher than the present estimation. Vasantharajan et al., (2014) and Zaahkouk et al., (2017) estimated an almost identical b value of 3.0 for another species of lethrinid (L. lentjan) along the Indian coast and Red Sea, which are lower than the current findings. Currey et al. (2009) and Vasantharajan et al., (2014) reported a slope value of 2.9 for L. nebulosus from the Great Barrier Reef and Thoothukudi, Southeast coast, India, confirming the negative allometric as observed in the present study. Additionally, Daliri et al., (2012) reported a slope value of 2.6 for L. nebulosus from the Northern Persian Gulf, which is in conformity with the current observation. In comparison with the previous estimates, some variations in the b values were noted in the current study. These differences could be due to several factors, including fish physiology, growth phase, sex, sexual maturity, season, stomach fullness, length range and sample size, habitat, feeding rate, dietand health (Le Cren, 1951; Froese, 2006; Mondol et al., 2017). Additionally, sampling bias might also occur due to the use of different gears, or absence of an independent and uniform sampling technique or relatively small sample size or narrow size range which may also affect the accuracy of b values as mentioned by Roul et al., (2017c).

The fulton condition factor (KF) of Lethrinus microdon were estimated 1.22 for male, 1.33 for female and 1.19, for pooled, which reveals that these species are in good condition in the waters of the Gulf of Mannar (Table 1). Rajathy et al., (2021) recorded a KF value of L. nebulosus as 1.44 from the Thoothukudi coast, India, which is in conformity with the present study. Ravikumar et al., (2023) reported the Fulton condition factor values from 1.3357 to 2.5846 for demersal reef fishes from the Gulf of Mannar, India, which is higher than the present findings. The results indicate that the ecological conditions in the waters of the Gulf of Mannar, located off the southeast coast of India, are favourable for the growth of Lethrinus microdon species. Additionally, the condition factor values nearing to one and more than 1 could be attributed to the well-being of the fish in the ecosystem besides the optimal environmental factors which are very conducive for the better growth and reproduction of the species. The health of fish varies slightly from habitat to habitat, possibly due to factors such as population size, food intake, reproductive development, sex, time of year and geographic location. When the condition factor e” 1, it indicates healthy growth condition of fish, whereas <1 signifies poor growth condition (Jisr et al., 2018). It can be influenced by both living and non-living factors, such as water quality and the food availability and predators (Blackwell et al., 2000).

Food and feeding habits
 
The analysis of gut contents indicated that fish constitutes the majority of the diet (50-60%), followed by crustaceans (40-50%) and mollusca (20-30%) and echinoderms (10%). The primary and most significant prey items were teleosts, crustaceans (including brachyurans and carideans)and molluscs (both gastropods and bivalves). In addition to the prey items, coral rubble and sand particles were also found in the gut. After analysed the gut samples, L. microdon can be regarded as a piscivorous species. These fish have a low-bodied form with conical teeth, enabling them to feed on fast-moving prey like nekton (Carpenter, 1996). The present observations also found similar results like studies by Randall (1995) and Carpenter and Allen (1989) which indicate that their primary prey consists of fish, followed by crustaceans, mollusksand polychaetes. This result is confirmed by Ali et al., (2016), but does not correspond to Fischer and Bianchi (1984) who stated that L. microdon feeds on crustaceans, mollusks and small fish. L. microdon typically feed primarily on fish, crustaceans, molluscan, with a smaller portion of their diet consisting of benthic prey. Clupeidae was the main prey, making up the largest percentage of L. microdon diets. Smaller specimens were found to consume an abundance of benthic invertebrates, while larger individuals showed a dietary shift. This change is attributed to their solitary behavior, wider home rangeand body morphology, which allows for faster swimming and the ability to target more mobile prey compared to subadults or juveniles. In younger specimens, mollusks, particularly bivalves, were found in notable amounts, whereas in adult fish, nekton became the primary dietary component due to their higher nutritional needs. The results also indicate ontogenetic changes in diet composition, which vary based on growth patterns, seasonal factorsand prey availability. The Gastrosomatic index (GaSI) value of L. microdon were ranged from 1.45-2.37 for male and 1.89-4.34 for female, respectively (Table 2 and Fig 3).

Table 2: Gonadosomatic (GSI) index, Gastrosomatic (GaSI) and Hepatosomatic (HSI) of Lethrinus microdon.


Fig 3: Gastrosomatic Index of Lethrinus microdon.



The  Hepatosomatic index (HSI) value were ranged from 0.45-2.54 for male and 1.11-3.21 for female, respectively (Table 2).

Reproductive characteristics

Sex ratio
 
The overall sex ratio of male and female was observed to be 1:0.67 (Table 3).

Table 3: Month-wise sex ratio of Lethrinus microdon.



The observed sex ratio was tested against 1:1 using the chi-square test for (n-1) degrees of freedom at 5% level of significance. The chi-square test indicated that there was no significance difference (p>0.05) in the average sex ratio and month-wise sex ratio (Fig 4).

Fig 4: Month-wise percentage distribution of male and female of Lethrinus microdon.



Grandcourt et al., (2010) observed the overall male: female ratio of L. microdon was 1:1.6, which is higher than present findings. In his study, male was more dominant than female in the present study. According to Al-Areeki et al., (2007) reported a overall sex ratio for L. lentjan were estimates to be 1:1.1 (Male:Female) in the Red Sea coast of Yemen which is higher than the present findings. Other similar study were reported by  Damora (2018) 1:1.08 for L. lenjan  in Indonesia waters; Degoon and Ali (2016) estimated the overall sex ratio was 1:1.7 for  L. mahsena in the Sudanese Red Sea,. Kulmiye et al., (2002) estimated sex ratio was of L. lentjan were estimated to be 1.0:1.10 in Kenyan coastal waters, present findings. In the study males were more dominant than female. The sex ratio may fluctuate from the predicted 1:1 ratio from species to species, or even within the same population adaptability, reproductive behavior, food availability and environmental conditions (Asut et al., 2019).

The gonadosomatic index (GSI) value of Lethrinus microdon was ranged from 1.21-2.21 for male and 1.85-2.31 for female (Fig 5).

Fig 5: Gonadosomatic Index of Lethrinus microdon.



Al-Areeki et al., (2007) reported a maximum GSI value of 2.43 (March) for another Lethrinus species, (L. lentjan) which is lower than the present study. Similar results recorded by Currey et al., (2009) and Ebisawa (1990) revealed high GSI value in July-August in Australia and Okinawan waters. The highest GSI value was recorded in L. mahsena (3.23) in June 2023 and the lowest value (0.26) was recorded in September 2023. Similar results were recorded by Soondron et al., (1999) in Mauritius water. Interestingly, their stomachs are fullest between September to November, but less full during the spawning season from June to August.

Length at first maturity
 
The length at which 50% of all fish of a particular size reach maturity is referred to as the length at first maturity (L50). For both male and female L. microdon, the L50 attains maturity was 25.5 cm for male and 26.5 cm for female (Fig 6 and 7).

Fig 6: Length at first maturity of male Lethrinus microdon.


Fig 7: Length at first maturity of female Lethrinus microdon.



Toor (1964) documented an L50 of L. lentjan was found 30 cm for male and 28.7 cm for female in the Indian Ocean, whereas Grandcourt et al., (2010) observed an L50 of L. microdon was estimated 24.6 cm for male and 27.7 cm in the Arabian Gulf, which is similar with the present findings. The present study differs from  another species of Lethrinus (L. lentjan), attained length at first maturity of female at 26 cm in Red Sea Coast of Yemen by Al-Areeki et al., (2007), which is similar than the present investigation. Mrombo et al., (2019) observed that the L50 for male and female L. lentjan was 25.8 cm. and 26.2 cm, which are in agreement with the present findings. Toor (1964) reported an L50 of 30 cm for males and 28.7 cm for females (L. lentjan) in Indian Ocean waters, while Grandcourt et al., (2010) reported an L50 of 24.6 cm for males and 27.7 cm fork for females (L. lentjan) in the southern Arabian Gulf. Damora et al., (2018) reported L50 of L. lentjan as 25.1 cm for males and 25.3 cm for females, which is similar with the present findings. The gaps in first sexual maturity values could be changed due to variations in diet, water temperature and stock population (Reznick, 1993). These differences may be also linked to varying levels of fishing pressure in different regions, as Lappalainen et al., (2016) suggested, noting that L50 can be a potential indicator of fishing pressure in fish stocks.

Maturity stages

The occurrence of various maturity of the gonad in different months showed that the peak spawning of L. microdon was in June to August. The simultaneous occurrence of immature, maturing, mature and ripe individuals in a number of months and evidence obtained from the hepatosomatic index and gonadosomatic index support above inferences. The observations of gonad maturity stages across different months revealed that peak spawning of L. microdon typically occurs in June to August in whole year. Motlagh et al., (2010) reported peak spawning of L. nebulosus in March in the Persian Gulf and Oman Sea. Al-Areeki et al., (2007) reported peak spawning of L. lentjan in April-May in the Yemen coastal water. Mrombo et al., (2019) reported peak spawning of L. lentjan in January in Kenya waters, there was no reported study on the maturity stages in L. microdon, although in the same genus of different species were reported.  Toor (1964) also observed the peak spawning of Lethrinus species occurring June – August in Indian waters. The present study also indicated same peak spawning season. Spawning season can vary from region to region due to the environmental factors.

Fecundity

The total fecundity was ranged from 70,345 to 5,09,735 eggs. Al-Areek et al. (2007) reported 80295 to 837251 eggs of another Lethrinus species (L. lentjan) in Yemen coastal water, which is higher than the present findings.  Mrombo et al., (2019) reported 90 to 3,80,364 eggs of L. lentjan in Kenya waters, which is higher than the present investigations. Toor (1964) reported 12,146-77,922 eggs of L. lentjan in Indian water, which is lower than the current findings. Degoon and Ali (2016) estimated absolute fecundity of L. mahsena was 2,26,667-1,646,667 eggs per mature female, which is higher than the present investigation. According to Toor (1964) the fecundity of L. mahsena was 26,700-166200 eggs (Carpenter and Allen, 1989) which is lower than the present findings. The fluctuation may arise from various environmental factors like temperature, sunlightand weather conditions (Jonsson and Jonsson, 1999).
Our results provide partial information about the feeding habits and reproductive characteristics of Lethrinus microdon, of major prominence for fisheries management. The percentage occurrence of various food items in the gut content showed that the primary preferable food item was the fish. Thus, the present investigation revealed that L. microdon could be considered a piscivorous species, whose diet is mainly composed of teleostean fishes, crustaceans, molluscs and echinoderms.  The information got by this study would be useful as a first step for the transition from single-species management to ecosystem-based management. Reproductive features provide baseline information for understanding the population dynamics which would help in formulating strategies for sustainable exploitation and conservation of this species in the region. The present investigation calls for further research in identifying the Smalltooth emperor fishing and spawning aggregation grounds and initiate measures for reducing fishing efforts with input and output controls to sustain the Smalltooth emperor fishery in the Gulf of Mannar coastal waters.
The authors acknowledge TNJFU PG Merit Fellowship provided by the Tamil Nadu Dr. J. Jayalalithaa Fisheries University, Nagapattinam, during the study period. Author greatly acknowledges the support from Fisheries College and Research Institute, Thoothukudi, for providing all the necessary facilities to carry out this research work by extending all the laboratory facilities.

Ethical statement

No live specimens here used in the present study.
All authors declared that they have no conflict of interest.

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