# New insights into the reproductive physiology and culture approaches of Paratilapia sp., a Malagasy cichlid with potential interest for sustainable aquaculture

Marta Caballero-Huertas, Marc Canonne, Domoina Rakotomanana, Rovaniaina Jospin Césaire Tianandriatsiferana, Tsiriniaina Andrianjafisoa, Jean Robertin Rasoloariniaina, Jean-michel Mortillaro

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Marta Caballero-Huertas, Marc Canonne, Domoina Rakotomanana, Rovaniaina Jospin Césaire Tianandriatsiferana, Tsiriniaina Andrianjafisoa, et al.. New insights into the reproductive physiology and culture approaches of Paratilapia sp., a Malagasy cichlid with potential interest for sustainable aquaculture. Journal of the World Aquaculture Society, 2024, 56 (1), pp.e13103. 10.1111/jwas.13103. hal-04892655

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# New insights into the reproductive physiology and culture approaches of *Paratilapia* sp., a Malagasy cichlid with potential interest for sustainable aquaculture

Marta Caballero-Huertas<fnref n="1" />,<fnref n="2" /> |

Marc Canonne<fnref n="1" />,<fnref n="2" /> |

Domoina Rakotomanana<fnref n="3" /> |

Rovaniaina Jospin Césaire Tianandriatsiferana<fnref n="1" />,<fnref n="2" />,<fnref n="3" /> |

Tsiriniaina Andrianjafisoa<fnref n="1" />,<fnref n="2" />,<fnref n="3" /> |

Jean Robertin Rasoloariniaina<fnref n="4" /> |

Jean-Michel Mortillaro<fnref n="1" />,<fnref n="2" /> ID

## Correspondence

Jean-Michel Mortillaro, CIRAD, UMR ISEM, 34398 Montpellier, France.

Email: jean-michel.mortillaro@cirad.fr

## Funding information

European Commission; French Facility for Global Environment; French Development Agency

## Abstract

One of the primary objectives in freshwater aquaculture is the search for sustainable systems based on agroecological intensification. In this context, the cultivation of local species is decisive, although requiring the identification of new candidates. Malagasy aquaculture is an example of the development of such an approach. We aimed to deepen the knowledge of the biology of the *Paratilapia* sp., specifically, their reproductive traits and zootechnical aspects, with special emphasis on the juvenile stages. The information provided has interest for its commercial production and the species conservation, as populations are subjected to overexploitation and habitat degradation. This study is a consolidated report on morphological characteristics, sexual

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

© 2024 The Author(s). *Journal of the World Aquaculture Society* published by Wiley Periodicals LLC on behalf of World Aquaculture Society.

<footnote n="1">¹ISEM, Univ Montpellier, CNRS, IRD, CIRAD, Montpellier, France</footnote>

<footnote n="2">²CIRAD, UMR ISEM, Montpellier, France</footnote>

<footnote n="3">³Department of Zootechnical, Veterinary and Fish Research, FOFIFA, Antananarivo, Madagascar</footnote>

<footnote n="4">⁴Department of Environment, University of Antananarivo, Institut d'Enseignement Supérieur d'Antsirabe-Vakinankaratra, Antsirabe, Madagascar</footnote>
dimorphism, reproductive biology, and early developmental ontogeny. The main results suggested a marked sexual dimorphism, as well as greater growth of males compared with females. Aggressive behavior between males was observed before pair formation. Generally, female and males provided protection until fry are independent. Females produced 2756 ± 598 eggs per spawn on average (gonadosomatic index = 4.6%), with a reproductive frequency of one to two spawns. Despite the need to improve breeding protocols, this work represents a breakthrough in producing a local freshwater species of potential interest.

KEYWORDS

breeding, broodstock, embryogenesis, freshwater, fry, Madagascar

# 1 | INTRODUCTION

Freshwater fish are an important source of food, nutrition, income, and livelihood for hundreds of millions of people around the world (FAO, 2016). The total global inland water catches, although largely considered an underestimate, were 11.9 million tons in 2014, continuing a positive trend that resulted in a 37% increase over the last decade (Breuil & Grima, 2014). Freshwater fish are defined as those species that spend all or a critical part of their life cycle in fresh waters, such as rivers and lakes. Of all species of freshwater fish rated in Madagascar and the Indian Ocean islands, 58% are endemic (Máiz-Tomé et al., 2018). These taxa in Madagascar are represented by about 178 species and around 35 families (Froese & Pauly, 2023). Despite the great specific richness of the local Malagasy ichthyofauna (Fricke et al., 2018; Garibaldi, 1996), close to 100% of farmed species from inland waters have been introduced (e.g., carp, goldfish, black bass, tilapia). On the other hand, overfishing, habitat fragmentation and destruction, and the potential pressures exerted by introduced species are major threats to many native freshwater fish species in Madagascar (Benstead et al., 2003; Gozlan et al., 2024; Šimková et al., 2019). These factors are negatively affecting many of the larger commercial species such as endemic cichlids (IUCN, Freshwater Biodiversity Department), the most diverse Malagasy freshwater fish family (Froese & Pauly, 2023). Among these species, we can find around 57 genera of Malagasy fish, with 5 endemic genera in the family of cichlids (Šimková et al., 2016) that like tilapia may have some interest for fish farming: *Paretroplus*, *Ptychochromis*, *Ptychochromoides*, *Oxylapia*, and *Paratilapia*.

Members of *Paratilapia* are large cichlids endemic to Madagascar (maximum size 30 cm standard length) with one apparent representative from the Congo River (*Paratilapia toddi*, Moelants, 2010), even though some authors classified this last species to the genus *Serranochromis* (Poll, 1939). The genus may include three species in Madagascar (*Paratilapia polleni*, *Paratilapia bleekeri*, and *Paratilapia typus*), and at least eight undescribed species (Loiselle & Ravelomanana, 2022). However, only *P. polleni* (Bleeker, 1868) and *P. bleekeri* (Sauvage, 1891) were previously reported on the Island, and the others belong to personal communications (Ravelomanana & Sparks, 2016) or synonyms until further genetic analysis allows for proper identification. *Paratilapia* spp. are classified as “Vulnerable” or “Data deficient” throughout Madagascar waterbodies according to IUCN (Ravelomanana, 2016; Ravelomanana & Sparks, 2016). Such status is a concern considering that phylogenetic relationships within the genus are not fully resolved, and a clear east-west disjunction is not evident (i.e., *P. polleni* and *P. sp.* “Itasy” are restricted to north-western basins, whereas *P. sp.* “East” and *P. bleekeri* were collected from eastern drainages; De Rham &
Nourissat, 2002; Sparks & Smith, 2004). In this context, some authors consider that there are gaps in knowledge and, for example, consider "hasty" the classification of individuals as *P. bleekeri* (Rakotomamonjy & Oliarinony, 2018).

Because of the lack of general information on the distribution and phylogeny of the species that make up the genus, there is also a lack of knowledge regarding its physiology and, specifically, in terms of reproduction. Indeed, successful conservation measures depend on adequate knowledge of its different biological traits, including reproductive biology and early developmental ontogeny. Linking aquaculture to conservation programs could allow faster access to species of aquaculture interest likely to better integrate into a given socioeconomic and environmental context (Fontaine et al., 2009; Thomas et al., 2021) while promoting the conservation of biodiversity. According to Comizzoli and Holt (2019), more basic and applied research in reproductive biology is needed to preserve species and design appropriate production strategies to ensure population sustainability, for example, with the implementation of cultivated biodiversity in multitrophic and integrative systems (e.g., rice-fish farming).

Therefore, to modestly remedy this lack of useful information for the domestication of *Paratilapia* spp., we present an analysis of the conditions for its reproduction, sexual dimorphism, reproductive behavior, and physiology, followed by the report of the embryonic development. Secondly, to support the sustainable development of continental aquaculture and its integration into agroecological systems, and considering that *Paratilapia* has already been the subject of breeding attempts (e.g., Kiener, 1963; Stiassny & Gerstner, 1992; Stiassny & Mezey, 1993), we discussed the species requirements and zootechnical practices followed in this study. Therefore, the major objective of this study was to provide updated knowledge on the domestication of *Paratilapia* sp., with a focus on its reproductive traits. This information will be essential to incorporate this species into sustainable production systems, as well as to the establishment of conservation programs.

## 2 | MATERIALS AND METHODS

### 2.1 | Localization of the study

The DRZVP/FOFIFA fish farming research station (Figure 1) is located inside the Analamazaotra National Park, rural commune of Andasibe, Moramanga District, in the Region Alaotra-Mangoro, 120 km from Antananarivo. The altitude of this station is 964 m, with geographical coordinates 18°56'11" S 48°25'05" E. The climate of Andasibe is very humid. The average temperature recorded was 18.50 ± 4.05°C during 2020 (Figure 2). Rainfall was 1700 mm over 210 days. From the beginning of the year until July 2021, the average air temperature observed from a weather station was 20.00 ± 1.71°C (Figure 2). Fish monitoring and measured parameters in this study were performed from December 2020 to March 2021, from the warm season to the early cool season.

### 2.2 | Biological units and sampling methods

#### 2.2.1 | Broodstock characteristics and monitoring

The *Paratilapia* sp. broodstock used in the experiment (N = 43) came from Lake Alaotra, Ambatondrazaka, but had been acclimatized for years at the Beforona fish farming station. They were transferred to the Andasibe station in October 2019 and separated into two groups (males and females) before breeding in a 100 m² pond (spawner pond). Dip, bowls, and bucket nets with mesh sizes ranging from 0.2 to 0.5 cm were used to capture the broodstock and fry.

Observation and monitoring were applied to recognize the ethology of *Paratilapia* sp., the formation of nests and egg incubation areas, as well as their movement in the breeding pond. It was carried out daily for 1 h at 1-h intervals (day and night). The capture of broodstock was performed with the identification of spawners after
<figure><img src="image_2.png" /><figcaption>**FIGURE 1** Location of the Andasibe fish farming station. (a) Topographic map of the island of Madagascar. (b) Satellite image of the station, coordinates: 18°56'11" S 48°25'05" E (Google Maps, 2023). (c) Drone picture of the station. Red square indicates 10 × 10 m (100 m²).</figcaption></figure>
<figure><img src="image_3.png" /><figcaption>**FIGURE 2** Air temperature variation and precipitation in Andasibe (year 2020–2021). Source: Weather station in the Andasibe fish station 2021. [Correction added on 30 September 2024, after first online publication: Figure 2 image has been updated in this version.]</figcaption></figure>

<table><caption>**TABLE 1** *Paratilapia* sp. stocked in the broodstock pond.</caption><thead><tr><th></th><th>Males</th><th>Females</th><th>Undetermined</th></tr></thead><tbody><tr><td>Number (individuals)</td><td>14</td><td>25</td><td>4</td></tr><tr><td>Average weight (g)</td><td>24.14 ± 12.08</td><td>20.36 ± 9.79</td><td>7.00 ± 1.41</td></tr><tr><td>Average total length (cm)</td><td>10.35 ± 2.05</td><td>9.68 ± 1.76</td><td>6.00 ± 0.47</td></tr></tbody></table>

spawning trials. Individual monitoring was carried out by inserting an RFID or Transponder electronic chip (2 mm × 12 mm) between the dorsal fin and the lateral line of *Paratilapia* sp. The VB Real trace Reader is a portable Bluetooth reader with discontinuous, instantaneous, and manual reading, with successful beep of the ID number of all the transponders injected into the fish. The transponders were injected using an injector or a syringe. Males and females were weighed, total length measured, and returned to the breeding pond. Equipment consisted of a Domo® precision scale with an accuracy of 0.01 g up to 200 g for weighing fry and fingerlings, while larger individuals such as broodstock were weighted using a Kern® scale with an accuracy of 0.10 g. The fish's total length measurements were realized using graph paper. On the other hand, the follow-up of fry was carried out before the transfer or after 1 month of pre-fattening and at the end of each test. Table 1 shows broodstock data at the time of stocking in the broodstock pond.

### 2.2.2 | Rearing techniques and zootechnical parameters

A 100 m² pond was used for broodstock breeding. Bamboo dividers were placed at the end of the pond to multiply breeding territories. Safety nets for predatory birds (especially Vintsy or *Corythornis vintsioides*) were placed above the pond. The stocking density in the broodstock pond was 0.43 individuals m⁻² including 14 males, 25 females, and 4 sexually unidentified fish (Figure 3), with a sex ratio of 1:2.

Water physical and chemical factors including temperature, dissolved oxygen, and pH were measured weekly to ensure a proper environment for the fish. A temperature sensor “Onset HOBO probes” was also used to record the
<figure><img src="image_4.png" /><figcaption>**FIGURE 3** External morphological difference of male and female of *Paratilapia* sp. From the broodstock. (a) 160 g male and (b) 56.9 g female.</figcaption></figure>

water temperature every hour. Data were recovered every 15 days using an optical USB and the corresponding HOBOware v.3.7.23 software. The average temperature was 27°C during the warm season and decreased to 24°C at the beginning of the cool season in the spawning pond. A Handy Polaris OxyGuard probe was used to measure dissolved oxygen in mg L⁻¹ (7.16 ± 0.63 mg L⁻¹, during the whole experiment). A pH meter “WTW pH 330i” with SenTix™ Epoxy combination pH electrode was used to record pH in water (7.34 ± 0.24, during the whole experiment). Lastly, a Vantage Pro2 weather station was used to record air temperature and precipitation (Figure 2).

Broodstock was fed twice a day: at 10 a.m. and 3 p.m. with 120 g day⁻¹ of earthworms (*Eisenia fetida*, *Amynthas corticis*, and *Nematogenia laccum*), 200 g day⁻¹ of rice bran, and 100 g day⁻¹ of Livestock Feed Limited (LFL) industrial feed. Earthworms were used to help adaptation of the fish to industrial pellets. These earthworms were locally produced on vermiculture pilots and fed on household waste and kitchen scraps. Fertilizers and four bags of compost were installed on the sides of the pond to increase natural productivity and organisms consumed by fish such as plankton and aquatic insects. Fertilization was applied to all livestock habitats using 500 kg ha⁻¹ of zebu dung for 3 days and then renewed if the transparency was greater than 30 cm.

### 2.2.3 | Egg counting, embryogenesis analysis, and survival

Before the breeding study, naturally, dead females found in the separated ponds were weighed and dissected to extract the ovaries. The ovarian lobes were then weighed on the Domo® precision scale to determine their fresh weight, and the gonadosomatic index (GSI) was calculated as follows: GSI = (fresh weight of the ovary/total weight of the individual) × 100. The eggs in the ovarian lobes were counted manually, 50 × 50, in a Petri dish with a little water placed over a sheet of white paper. Pasteur pipettes and Petri dishes were used for sampling and placing eggs and larvae. A binocular digital microscope with B-290 TB tablet and 4 E-PLAN objectives (×4, ×10, ×40, ×100) was used to observe the evolution of eggs and larvae. Taking egg samples every 6 h at the time of laying allowed us to
obtain the embryonic evolution of *Paratilapia* sp. The embryogenesis of *Paratilapia* sp. was described during the first 42 h post-fertilization at an average temperature of 26°C using a digital microscope with 4× objective. According to the first study carried out in 2020, a female can give 900 fry on average and 30 days after spawning, fry were already autonomous.

In this study, the fry gradually disappeared before the planned 30 days of their transfers and the start of pre-
fattening. In this context, they were transferred and separated from their parents 15 days after laying.

2.3 | Weight indices and statistical analyses

The Daily Mass Gain (DMG), consisting of the growth rate of an animal, and expressed in g day⁻¹, was calculated as DMG = (Final weight – Initial weight)/Breeding time; the Daily Length Gain (DLG) was the zootechnical index used to represent linear growth and is expressed in cm day, expressed as LMG = (Final length – Initial length)/ Breeding time.

For testing the height–weight correlation, Pearson's correlation coefficient was evaluated. Pearson's correlation assesses the linear relationship between two variables. We used this analysis to determine whether weight increases are associated with increased height. Student's t-test for independent samples was applied to compare the means of two groups of samples. The question is therefore whether the means of the two groups are statistically significantly different. Statistically significant differences were considered when *p* < 0.05.

3 | RESULTS

3.1 | Sexual dimorphism

In the genus *Paratilapia*, the male was easily differentiated from the female by phenotypic criteria, in particular during the breeding season when the pigmentation is darker (Figure 3). Indeed, the male had a black coloring of the sides, dotted with many round bluish spots as well as the dorsal and anal fins sometimes extending to the end of the caudal fin. The male may also present a developed mouth and a bump on the forehead in large specimens. The female had an olive-brown coloring of the flanks with its bluish spots and smaller fins than the male. It had a less developed mouth and the caudal part was shorter than the males.

3.2 | Reproductive frequency

During the broodstock breeding, seven females (Table 2) were captured and identified at the time of spawning. They had a reproductive frequency of one to two spawns/female by cycle (synchronous oocyte maturation; Figure 4a), although all the individuals selected for the broodstock did not reproduce. The spawning was grouped mainly during January (Figure 4b) when an average temperature of 27°C was recorded (Figure 4a). In February, the frequency of spawning underwent a drastic drop according to the average temperature (Figure 4a). New clutches appeared from the end of February to the beginning of March (Figure 4a). The first spawning is not shown in the spawning distribu- tion schema (Figure 4b) because its location was not well determined.

3.3 | Reproductive capacity

The fresh weight of the ovaries varied from 2 to 3 g (Table 3). There were between 1000 and 1500 eggs in each ovarian lobe. The average number of mature oocytes in the ovaries of *Paratilapia* sp. was therefore 2756 ± 598,
<table><caption>TABLE 2 *Paratilapia* sp. identified during the spawning.</caption><thead><tr><th>ID</th><th>Spann</th><th>Total length (cm)</th><th>Total weight (g)</th><th>No. days at broodstock pond</th></tr></thead><tbody><tr><td>967000009205089</td><td>2</td><td>13.5</td><td>38.0</td><td>81</td></tr><tr><td>967000009205585</td><td>3</td><td>10.7</td><td>24.0</td><td>82</td></tr><tr><td>967000009205797</td><td>4 and 15</td><td>12.5-13.1</td><td>31.0-44.0</td><td>82 and 134</td></tr><tr><td>967000009205587</td><td>5 and 13</td><td>12.5-13.0</td><td>30.0-37.0</td><td>134 and 87</td></tr><tr><td>967000009205992</td><td>6</td><td>11.5</td><td>20.0</td><td>87</td></tr><tr><td>967000009205893</td><td>12</td><td>12.0</td><td>40.0</td><td>96</td></tr><tr><td>967000009205151</td><td>14</td><td>15.0</td><td>64.0</td><td>129</td></tr></tbody></table>

<figure><img src="image_5.png" /><figcaption>(b)</figcaption></figure>

<figure><img src="image_6.png" /><figcaption>FIGURE 4 (a) Frequency of spawning with water temperature variation (°C). (b) Distribution of egg laying observed in the spawning pond. The date and depth at which each spawning occurred are indicated.</figcaption></figure>

<table><caption>TABLE 3 Gonad biometry and gonadosomatic index (GSI, %) of *Paratilapia* sp. females.</caption><thead><tr><th rowspan="2">Individuals</th><th rowspan="2">Body weight (g)</th><th rowspan="2">Ovary fresh weight (g)</th><th rowspan="2">GSI (%)</th><th colspan="2">Number of oocytes</th></tr><tr><th>Lobe I</th><th>Lobe II</th></tr></thead><tbody><tr><td>Female 1</td><td>38</td><td>0.8<sup>a</sup></td><td>2.1</td><td>1564</td><td>1460</td></tr><tr><td>Female 2</td><td>41</td><td>3</td><td>7.3</td><td>1013</td><td>1058</td></tr><tr><td>Female 3</td><td>54</td><td>2.4</td><td>4.4</td><td>1602</td><td>1572</td></tr><tr><td>Average (n = 3)</td><td>44.3 ± 8.5</td><td>2.1 ± 1.1</td><td>4.6 ± 2.6</td><td>1393 ± 330</td><td>1363 ± 270</td></tr></tbody></table>

although these figures varied according to their weight. The average gonadosomatic index was 4.6%. The reproductive capacity of females was high, with mass production of mature oocytes during gametogenesis.

## 3.4 | Ethology of parents during reproduction

### 3.4.1 | Battle between males and pair formation

Before the formation of a pair, field observation has identified that in some clutches, two males fought for a female. The latter kept intruders away from the battle zone. After several hours of battle, the female selected only a

<footnote><sup>a</sup>Ovaries already atrophied, preserved in alcohol.</footnote>
dominant male, and the pair formed in the pond. Both moved around the edges of the pond to prepare their nest. The females that reproduced more than once during the study did not keep the same male for each of the reproductive encounters, nor the same area.

### 3.4.2 | Preparation of the nest and encirclement of the territory

The male and the female encircled a territory with a radius of 1.5-2.0 m and looked for a favorable place for the egg laying. The couple became more aggressive and its hue darkened. Based on observations of several clutches, two to five holes were cleaned in the ponds to make a nest.

### 3.4.3 | Egg laying and hatching

After 2-3 days of pair formation, the female laid the eggs at the bottom and near the edge of the pond, at a shallow depth varying from 10 to 25 cm (Figure 4b), in an area where decaying plants were present (Figure 5a). The depth at which the eggs were found varied according to the pair and the water level. The male and female became very aggressive and surrounded a multifunctional territory: a breeding territory (0.5 m radius) followed by a vital territory (1.0-1.5 m radius).

The female ventilated the egg cluster with repeated movements of the pectoral fin until hatching. The female moved the eggs two to three times until hatching. Movement was usually a short distance, within the original breeding territory, but some females occasionally moved the eggs to a new territory. Depending on the water temperature, egg hatching took place 1.5-3.0 days after laying. The larvae were mobile but could not yet swim and concentrated on the incubation area.

### 3.4.4 | Parental care and the first swimming of the fry

In this study, parental care of the fry could last on average up to 20 days after spawning. Although the most widespread behavior was for both parents to protect the larvae, the males were not permanently accompanying them, unlike the females. Moreover, the male may immediately abandon their nest even to form a new pair if one or more

<figure><img src="image_7.png" /><figcaption>**FIGURE 5** (a) Protection and ventilation of eggs by a 44 g female; (b) Eggs of *Paratilapia* sp. a few hours after laying.</figcaption></figure>
other females were ready for reproduction. Some males abandoned their nest from the second day of laying. Females could also abandon their fry even 2 days after hatching if disturbed. However, from the 15th day, most of the fry moved around the pond in groups without the protection of their parents, although a few individuals may remain with the female. The fry's first swim took place 2-3 days after the hatching of the eggs when the fins were well developed. On the day of the first swim, the fry stayed in the incubation area near the female. On the second day of swimming, they followed the female and the male.

### 3.4.5 | Embryogenesis

After fertilization, between midnight and 6 a.m., the eggs laid by the female were very sticky to each other and yellowish in color (Figure 5b). A few hours after fertilization, the blastomeres underwent segmentation and connected with the blastocoel (Figure 6). From 6 h after fertilization, the eggs become less sticky and detached from each other. From 12 to 30 h after, the outlines of the first organs became visible. The antero-posterior and dorsoventral axes became visible. We then observed the first movements and the first differentiated cells, which corresponded to the period of segmentation. From 36 h, the larvae became mobile and a well-marked bilateral symmetry was observed. The eggs then reach the pharyngeal period, which was followed by hatching after 42 h.

### 3.4.6 | Growth and survival of individuals (spawners and juveniles)

The survival of the spawners was 100% during the experiment. The following figure illustrates the difference in linear growth between males and females of *Paratilapia* sp.

According to the analyses of the morphometric data of both sexes, we observed sexual dimorphism in the linear and weight growth of *Paratilapia* sp. At the time of stocking towards the broodstock pond, the average sizes of the two sex groups were around 10 cm (Figure 7a). The males had an average size of 10.17 ± 1.87 cm and the females were 9.93 ± 1.69 cm. After 180 days of breeding, the males had an average size of 16.70 ± 2.24 cm and the

<figure><img src="image_8.png" /><figcaption>**FIGURE 6** Embryogenesis of *Paratilapia* sp.</figcaption></figure>
<figure><img src="image_9.png" /></figure>

<figure><img src="image_10.png" /></figure>

<figure><img src="image_11.png" /></figure>

<figure><img src="image_12.png" /><figcaption>FIGURE 7 (a) Linear growth of males and females of *Paratilapia* sp.; (b) Weight growth of males and females; (c) Linear growth performance and weight growth performance of both sexes; (d) Length/weight relationship.</figcaption></figure>

females were 14.22 ± 1.61 cm. After checking the normality (Shapiro-Wilk test) of the data, Student's t test showed significant differences between males and females regarding the DLG (*p* < 0.001), which was higher in males than in females (Figure 7c).

In the beginning, the average weights of the two sex groups were close to 22 g (Figure 7b). Males had an average weight of 22.07 ± 10.99 g and females were 22.00 ± 10.08 g. After 180 days of breeding, the males had an average weight of 101.28 ± 36.24 g, while females' weight was 59.64 ± 16.69 g. Males had a significantly higher DMG than females (*p* < 0.05; 0.44 and 0.20 g, respectively; Figure 7c).

The Pearson test demonstrates that there was a significant and positive strong correlation between the total length and the weight of *Paratilapia* sp. The values of the correlation coefficient were 0.98 with R² = 0.95 for males; 0.96 with R² = 0.93 for females; and 0.96 with R² = 0.92 for undetermined individuals (Figure 7d).

4 | DISCUSSION

The successful breeding of a species for aquaculture requires the understanding of multiple factors (e.g., environmental, nutritional, zootechnical, and socioeconomic), that affect reproductive performance (e.g., sexual maturation, fecundity, and quality of larvae and juveniles) and the economic viability of production models (Berlinsky et al., 2020; Mitra et al., 2018). Considering the previous components, the process of fish domestication includes three cascading stages: (1) the adaptation of wild fish to captivity, (2) the reproduction of fish, and (3) the production and maintenance of young individuals (Shiguemoto et al., 2020). In this context, the current study illustrates for the first time various aspects of the reproductive biology of *Paratilapia* sp., but also goes a step further with its adaptation to captivity.

According to the bibliography, the main physicochemical properties of water required by *Paratilapia* sp. are optimal temperatures between 22 and 28°C (22°C for reproduction), and pH between 6.5 and 9 (6.7 for reproduction)
(Catala, 1977; Kiener, 1963). Complying with these requirements, in our study, broodstock ponds oscillated around 26.32 ± 1.16°C, and the pH was 7.34 ± 0.24. However, Stiassny and Gerstner (1992) noted that this fish can tolerate adverse conditions. For example, it can survive at a temperature of 12°C (at an altitude of 1500–1550 m), but it can also tolerate water of 40°C for a short duration according to our previous observations in the field (Makira National Park, Ambinanitelo). It should be considered that fry is more sensitive, and it cannot usually withstand temperatures below 17°C (observation at Analamazoa tra National Park, rural commune of Andasibe). It can even be found in turbid waters with a very acidic pH (Catala, 1977; Kiener & Mauge, 1966). Temperature is a crucial factor for the growth and survival of fish, as well as for the spawning frequency, as it varies depending on environmental conditions (FAO, 2016). As fish are poikilothermic animals (Lall & Tibbetts, 2009), water temperature affects the growth of fish larvae depending in particular on whether or not it is far from the thermal preference zone. This factor has effects on the heterogeneity of growth observed in farms (Fontaine & Le Bail, 2004). During February, the frequency of spawning experienced a drastic drop, which was probably not related to the temperature, according to the documented preference for this species (22–28°C, Stiassny & Gerstner, 1992). This fall in reproduction seems, on the contrary, to be linked to the density of pairs incubating, inhibiting the creation of new pairs for the delimitation of a spawning territory. The number of eggs per spawn presented a variation among individuals in our study, although on average it was 2756 ± 598. According to Kiener (1963), the number of eggs by spawn for *Paratilapia* sp. varied from 800 to 900, and it increased up to 3500 eggs at the second spawning. In contrast, Catala (1977) estimated around 1000 eggs for a mature female of average size.

During this investigation, phenomena of hierarchization within the batches of parents were noticed. The dominance of males over females, and their aggressiveness toward other fish of the same sex, increased during the breeding season. This mode of communication is mainly used during encounters between rivals or courtship displays. Baerends and Baerends-Van Roon (1950) were the first to propose a nomenclature of attitudes for the Cichlidae. Some are common to many species. For example, in the “side” position, a threatened Cichlidae may appear larger if its mid- and pelvic fins are extended. Similarly, in the “face-to-face” position, the threatened fish turns its head toward the attacker by inflating its opercula and branchiostegal membranes. Conversely, frightened fish make themselves as small as possible by folding their fins over their bodies. However, by establishing for the same family the catalog of postures, their order of appearance, their arrangement, and their respective duration, we note specific differences that can play an important role, in particular as a reproductive barrier during courtship displays (Lévêque et al., 1990). Reproductive, social, and feeding behaviors reflect the active attitudes of fish and their adaptations to external constraints to ensure their survival and propagation. To guarantee the best possible reproductive success, fish have various strategies that are genetically coded. These result in often complex sexual and reproductive behaviors (Lévêque et al., 1990). In terms of reproductive strategy, we were able to show that *Paratilapia* sp. is a successive monogamous species, observed by the mate changing by females that reproduced more than once. However, one male was observed switching between two females at the same time, suggesting that bigamy could be considered as an adaptive behavior to captivity, although success in hatching was not reached. The female selects the male before nest formation and can be based on the search for an immediate benefit or linked to the genetic potential of the partner. The search for an immediate or material benefit aims to increase the chances of dispersal of its genes and their sustainability. This generally translates into active selection involving, for example, the best nest builders or protectors (Bessa et al., 2022). Some behavioral traits resemble those of other substrate-spawning cichlids. Fryer and lles (1972) showed courtship and egg-laying in a substrate-laying Cichlidae: the female lays a first series of eggs on the previously cleaned substrate and the male remains nearby. Afterward, the female leaves the egg-laying site, and the male passes over the eggs and fertilizes them; the female then becomes darker and guards and aerates the eggs that have been collected. The two parents can remain united or not until their offspring reach the independent stage, with a well-defined distribution of tasks. Some males abandon their territory a few days after spawning, leaving the female to provide the parental care. Nevertheless, we conjecture that females abandon their fry some days after hatching if they can no longer control their territory and their nest. Parent custody of the fry does not last more than 2 weeks, which promotes their autonomy, living in small schools in several groups. Biparental guarding behavior is
widely considered to be the ancestral care state in cichlids (Goodwin et al., 1998). It is specific to benthic, substrate-laying fish species (Stiassny & Gerstner, 1992) such as *Coptodon rendalli* and *Coptodon zillii* (Bruton & Gophen, 1992), unlike mouthbrooders, which adopt the practice of maternal single parental care, which is a recent reproductive strategy of cichlids to avoid predation pressure (e.g., *Oreochromis niloticus*; Lévêque & Paugy, 2006; Ramirez, 2008). Several authors (Kop & Heuts, 1973; Russock & Schein, 1977, 1978) have shown that the fact that the young and the parents can able to recognize each other, at least as a species but also perhaps as an individual, results from their association during parental care.

Stiassny and Mezey (1993) analyzed the egg characteristics of *P. polleni* during their investigations of the reproductive biology of Asian and Madagascar cichlids. They described eggs that are adhesive to each other but not to substrates. Compared with most of the Cichlids studied (e.g., subfamily Etroplinae), the specific *Paratilapia* sp. system of attachment of the eggs to each other by tufts of filaments but free from the substrate is unusual. The ultrastructure of the eggs studied under the microscope allowed us to observe these filaments more precisely. These start from the opposite pole to the micropyle and attach to a single filament, allowing the eggs to stay between them in a single mass. Lévêque et al. (1990) also reported this in the genus *Paratilapia*, in which, according to the authors, adhesive eggs are deposited on a hard surface. However, our observation allow us to show that eggs are deposited on soil substrates together with plant debris. The eggs are fertilized and hatch after a few days during which both parents or just the female generally provide vigilant guarding. When the larvae can swim freely, they remain in groups near the substrate, under parental supervision.

The data acquired provide a fairly comprehensive view of the size/weight relationship of this species. This is marked in particular by a sexual dimorphism in favor of males beyond 15 cm 100 g⁻¹. At this size, the average daily gain can reach 1.4 g day⁻¹ (there is 38 g in 28 days). This is identical to tilapia *O. niloticus*, in which as soon as the individuals reach the age of maturity (1-3 years depending on the sex and the environment), male individuals show faster growth than the females and reach a much larger size (FAO, 2004) (growth almost twice as fast as the females; Fuentes-Silva et al., 2013). In *Paratilapia* sp., even at the time of pair formation, the male is always larger than the female. In addition, the largest individual measured to date by the research group had reached a size of 28 cm for 386 g. However, if notable sizes can be reached for this species, it is necessary to assess the corresponding age of the fish under culture conditions. Moreover, and according to the pattern in cichlids, sexual dimorphism in *Paratilapia* sp. is accentuated during the breeding season. The color pattern, pigmentation, genital papilla, and shape of the mouth allow for distinguishing between the sexes.

Adding some aspects of feeding behavior, we observed that *Paratilapia* sp. prefers live prey. The fry probably consumes planktonic organisms, like most fish larvae, whose size is adapted to the diameter of their mouth (Moreau, 1979). However, in adulthood, the diet of this fish is defined by a carnivorous toward omnivorous tendency (Stiassny & Gerstner, 1992). Nevertheless, the main components of their food bolus are represented by invertebrates, especially benthic macroinvertebrates (e.g., mollusks, insects, annelids) (Lauzanne, 1988). According to Lévêque and Paugy (2006), it does indeed appear that fish, such as African cichlids, are able to expand their feeding habits and behaviors depending on the availability of prey. We believe this is the case for *Paratilapia* sp., and we have practiced vermiculture as a source of protein for the spawners' food, as well as an adaptative strategy to acclimate them to industrial feed by mixing it with worms. Generally, many species adapt their diet to environmental constraints. Thus, for samples from different ponds of a temporary watercourse in Mali, it has been shown that, for the majority of species, regime affinities are greater within the same sample than between identical species from different samples (Paugy, 1994). This means that, in order to survive sometimes under extreme conditions, species adapt, within certain limits, their diet to the dominant resources available. We can thus show that faced with a low diversity of food, also linked to a lower quantity, the species use the most abundant resource, and conversely, when the prey becomes more numerous, the fish feed from a larger number of prey taxa. In other words, the diversity of the diet dependent on the richness of the prey ingested is probably dependent on the prey available in the environment (Lévêque et al., 1992). Further studies are thus needed on the feeding behavior and nutritional requirement of *Paratilapia* sp. spawners and fry as a cultural food item included in aquaculture as a target or complementary species
(Personal observation on the East Coast of Madagascar, from Andapa to Mahanoro). Nevertheless, we trust that this study can be a useful tool for the development of zootechnics of this Malagasy native species, especially from the reproductive perspective. This knowledge may be key to the more widespread inclusion of this species in Madagascar's sustainable aquaculture, as well as being key to its conservation.

## ACKNOWLEDGMENTS

We are grateful to the team of the experimental fish farming research station of FOFIFA Andasibe and Beforona (Centre National de Recherche Appliquée au Développement Rural, Madagascar) for experimental facility and biological material. We are also thankful to Guy Tam Hyock (Association des Producteurs Privés d'Alevins d'Andapa) for inspiring this work and the precious advices on biological requirement and behavior of the species. We are grateful to WCS Madagascar team of the SWM Programme for supporting this study and promoting knowledge transfer to the farmers in Makira National Park. This is publication ISEM 2024-210 SUD.

## FUNDING INFORMATION

The research presented in this paper was carried out as part of the Sustainable Wildlife Management (SWM) Programme, a major international initiative funded by the European Union, with co-funding from the French Facility for Global Environment (FFEM) and the French Development Agency (AFD). A CC-BY 4.0 public copyright license has been applied by the authors to the present document and will be applied to all subsequent versions up to the Author Accepted Manuscript arising from this submission, in accordance with the grant's open access conditions. The SWM Programme is being implemented by a consortium partnership, which includes the Food and Agriculture Organization of the United Nations (FAO), the French Agricultural Research Centre for International Development (CIRAD), the Centre for International Forestry Research and World Agroforestry (CIFOR-ICRAF), and the Wildlife Conservation Society (WCS). For more information, please visit https://www.swm-programme.info/homepage. We are grateful to WCS Madagascar team in MaMaBay Landscape & Seascape for supporting this study and promoting knowledge transfer to the farmers in Makira National Park greenbelt. [Correction added on 30 September 2024, after first online publication: Funding information statement has been updated in this version.]

## CONFLICT OF INTEREST STATEMENT

Mortillaro Jean-Michel has patent no. CC-BY 4.0 licensed to Creative Commons.org. All other authors declare no conflicts of interest.

## DATA AVAILABILITY STATEMENT

The data that support the findings of this study are available from the corresponding author upon reasonable request.

## ORCID

Jean-Michel Mortillaro https://orcid.org/0000-0003-1911-9338

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