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. 2024 Dec 13;48(4):e14064. doi: 10.1111/jfd.14064

Pejerrey (Odontesthes bonariensis) Under Siege: Main Infectious Diseases and Their Role in Aquaculture and Wild Populations Amidst Environmental Change

Aarón Torres‐Martínez 1,2,✉, Miguel Mancini 3, Fabian Grosman 4, Gustavo Manuel Somoza 5,6, Carlos Augusto Strüssmann 1
PMCID: PMC11907690  PMID: 39673090

ABSTRACT

The pejerrey ( Odontesthes bonariensis ) is a key species for recreational and commercial fisheries in Argentina and holds significant aquaculture potential. It has been introduced to various countries worldwide, including Japan, where intensive aquaculture has developed. However, infectious diseases present major challenges to its cultivation, as pejerrey is susceptible to diverse pathogens, including bacteria, fungi and parasites. The primary bacterial pathogens affecting pejerrey include the genera Aeromonas, Pseudomonas and Mycobacterium (M. piscida). Fungal‐like pathogens such as Saprolegnia spp., and fungal pathogens such as Achyla racemosa and Fusarium species (F. solani and F. semitectum) are also prevalent. Additionally, pejerrey hosts external and internal parasites, primarily Lernaea cyprinacea and members of the genera Cangatiella, Gyrodactylus, Contracaecum and Diplostomum. This review explores the primary infectious diseases affecting pejerrey, focusing on their symptoms, epidemiology and causative pathogens, based on literature from multiple countries and languages. Although no new diseases have emerged, we have identified persistent challenges that have remained unsolved for decades, highlighting the need for further research. Understanding the biology and epidemiology of these pathogens is crucial for expanding the aquaculture of pejerrey. Moreover, we examine how environmental changes, such as global warming, pollution and alien species, may influence disease dynamics in wild populations, stressing the need for management measures to preserve this valuable resource.

Keywords: fungi, global warming, parasites, pollution, silverside

1. Pejerrey as an Aquaculture and Fisheries Resource

The Argentine pejerrey ( Odontesthes bonariensis , Valenciennes; hereafter ‘pejerrey’) is a freshwater fish native to Argentina and belongs to the New World silversides in the family Atherinopsidae, which includes the subfamilies Menidiinae and Atherinopsinae (FishBase 2024). In the context of Argentina's continental ichthyofauna, Odontesthes bonariensis stands out as one of the most economically important and widely distributed species across the central and northern regions of the country. Pejerrey reaches maturity in around 1 year, 10–20 cm in total length, presenting a major spawning period in spring and a smaller one in autumn (Strüssmann 1989; Chalde, Elisio, and Miranda 2014). The first spawning of a single female has been estimated to be around 4000 eggs at a standard length of 25–26 cm, which hatch between 13 and 14 days (Strüssmann 1989). Pejerrey are largely zooplanktivorous in their early life stages, later shifting to a more carnivorous diet, including fish and invertebrates (Vila‐Pinto 2011).

While the species' original distribution is debated due to extensive domestic introductions over nearly a century, it is generally thought to encompass the La Plata River basin and the lakes of the Pampa region (Tombari and Volpedo 2008). Today, pejerrey is found in various water bodies across Argentina, such as rivers, channels, lakes and reservoirs, with its current distribution possibly extending from the Río Negro in the south (or even as far south as the 46th parallel) to the northernmost regions of the country (Figure 1A). Beyond Argentina, pejerrey has been introduced to several regions worldwide, including other South American countries, Europe, North Africa, the Middle East and East Asia (Figure 1B; Tombari and Volpedo 2008; Vila‐Pinto 2011). However, the success of these introductions has been limited. Unlike other widely cultured species, such as the Nile tilapia ( Oreochromis niloticus ), common carp ( Cyprinus carpio ) and rainbow trout ( Oncorhynchus mykiss ), pejerrey introductions have predominantly failed outside of South America, with Japan being the notable exception.

FIGURE 1.

FIGURE 1

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(A) Distribution of Odontesthes bonariensis in Argentina as of 1992. This distribution includes sites of artificial stocking and does not account for more recent introductions, for which no published literature is currently available. (B) Global introductions of pejerrey: The species' native country (Argentina) is indicated in green, while countries where it has been introduced are shown in red (not necessarily distributed in the entire coloured areas). (A) Adapted from Tombari and Volpedo (2008). (B) Map created using MapChart.

Historically, pejerrey has played a significant role in Argentinian fisheries and more recently has become an important species in Lake Titicaca (Peru/Bolivia) (Amaru‐Chambilla and Flores 2021; Loubens and Osorio 1992). Marketable fish typically range from 25 to 40 cm in length and can sometimes exceed this size in fish aged between 1 and 5 years, resulting in an annual harvest of thousands of tons for food supply. The species also has great potential for aquaculture (Mancini et al. 2016; Somoza et al. 2008) and is the most popular target for sports fishing in Argentina (Grosman et al. 2019). Despite these attributes, pejerrey aquaculture development in South America has faced several challenges spanning biological, technological, sociocultural and economic factors (reviewed in Somoza et al. 2008). In contrast, Japan has made considerable advances in its cultivation (see Section 5). Nevertheless, significant obstacles remain to be addressed (Somoza et al. 2008; Strüssmann and Yasuda 2005), particularly concerning infectious diseases, which affect all life stages from eggs to adult fish (Mancini et al. 2006, 2016).

A variety of pathogens, including bacteria, fungi and parasites, have been documented in cultured and wild pejerrey. However, comprehensive data on the pathogenesis, epidemiology and management or treatment of these infectious diseases remain scarce, and notably, no viral infections have yet been reported in the scientific literature for this species. Given these knowledge gaps, this review aims to synthesise available information on the primary infectious diseases affecting pejerrey in both captive and wild environments, alongside their causal agents. To achieve this, we conducted a thorough literature search on infectious diseases impacting pejerrey across South America and Japan, covering both wild and cultured settings. Our sources included literature in Japanese, Spanish and Portuguese, as well as valuable non‐digitally available references, which are often excluded from studies published solely in English. By incorporating this broader range of materials, we provide a more comprehensive understanding of the impact of these diseases, including overlooked regional data, thus offering a detailed perspective on the health challenges faced by pejerrey populations in diverse environments. Additionally, a part of the figures was curated to improve presentation and delivery of information, while remaining faithful to the original data, as some of the literature was many years old or limited in visual clarity.

The following section will outline the known diseases affecting pejerrey, providing a detailed overview of their pathologies and impact on both cultured and wild populations.

2. Bacterial Infections

2.1. Aeromoniasis

Aeromonas hydrophila and A. salmonicida are the main pathogens responsible of aeromoniasis in fish (Austin and Austin 2016). They have been isolated from both cultured and wild pejerrey populations in Japan and Argentina, respectively (Lawhavinit, Hatai, and Kubota 1986; Mancini et al. 2006; Suzuki 1982). In pejerrey, aeromoniasis is characterised by symptoms including anorexia, abnormal swimming, pale gills and external haemorrhaging, which can develop into ulcerations around the mouth, fins or anus (Figure 2). The haemorrhages can extend into the deeper layers of the dermis and skeletal muscle. Internally, petechial haemorrhages may occur in organs such as the liver, spleen, kidney and gut (García‐Romero 2001). Aeromoniasis is often exacerbated by external parasites, such as the copepod Lernaea spp., which create skin damage and entry points for bacterial infection (Mancini et al. 2006, see Section 4.1). In cultured pejerrey in Japan, A. salmonicida has been identified as a significant cause of mortality. Chemical treatments for aeromoniasis have been explored, with studies showing that A. salmonicida is highly sensitive to the antibiotics oxytetracycline and tetracycline hydrochloride (Satoh and Yamazaki 1990). While the use of these antibiotics reduced mortality rates, notable losses persisted (Satoh and Yamazaki 1990). Environmental conditions significantly influence the prevalence of aeromoniasis in pejerrey. Factors such as poor water quality, sudden temperature changes and malnutrition can facilitate the disease. In wild populations, particularly in central Argentina, reduced surface area and water volume in lagoons contribute to poor water quality and elevated organic loads, leading to increased turbidity and phytoplankton blooms that create favourable conditions for Aeromonas spp. (García‐Romero 2001; Mancini et al. 2006).

FIGURE 2.

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Aeromoniasis in Odontesthes bonariensis . Specimens infected by Aeromonas hydrophila displaying obvious signs of external haemorrhages (arrows). Reproduced from Mancini et al. (2016).

2.2. Pseudomoniasis

Pseudomonas‐induced disease (pseudomoniasis) in pejerrey is characterised by external lesions such as hemorrhagic spots, eroded and frayed fins, exophthalmia, gill inflammation, internal bleeding and behavioural changes, including lethargy and isolation (Austin and Austin 2016). In 1995, mass mortality events were reported in pejerrey farms in central Japan (Watanabe 1996). Pseudomonas aeruginosa was isolated from the kidneys of infected fish, representing, to our knowledge, the only documented case of pseudomonas infection and treatment in this species. Two strains of P. aeruginosa , Pj9536 and TB9602, were cultured in vitro and showed differing antibiotic resistance. Pj9536 was more sensitive to oxinphosphate and oxytetracycline (Watanabe 1996).

2.3. Mycobacteriosis

Mycobacteriosis, also known as fish tuberculosis, is caused by Mycobacterium and is characterised by external inflammation, ulceration and internal granuloma formation (Austin and Austin 2016). In the early years of pejerrey aquaculture in Japan, particularly during late winter and early spring, mycobacteriosis posed significant challenges, leading to considerable mortalities (Hatai et al. 1988; Lawhavinit et al. 1988a, 1988b). Screenings at the Kanagawa Experimental Station in Kanagawa, Central Japan revealed that up to 80% of pond‐cultured pejerrey were infected with Mycobacterium sp., underscoring the severity of the issue at that time (Lawhavinit et al. 1988a). An experimental study by Hatai et al. (1988) demonstrated the pathogenicity of Mycobacterium sp. in pejerrey, isolating the agent from pond‐cultured specimens and injecting it into healthy fish. This led to granulomas in the liver, spleen, and kidney (Figure 3A–D), with mortality rates reaching 100%. Histologically, these granulomas were characterized by epithelioid cell proliferation around a necrotic nucleus containing acid‐fast bacteria, with infiltration by immune cells such as mononuclear phagocytes and macrophages (Figure 3C; Hatai et al. 1988; Lawhavinit et al. 1988b). A subsequent study identified Mycobacterium piscida as the primary pathogen responsible for these infections (Toda and Suzuki 1990). Research into therapeutic interventions found that intramuscular injections of kanamycin sulfate at 50 mg/kg were effective against this pathogen in pejerrey (Lawhavinit et al. 1988a, 1988b). Oral administration of kanamycin at doses of 100 and 200 mg/kg also proved effective (Toda and Suzuki 1990). M. piscida was characterised by weak motility, absence of spores, capsules or true branching and is Gram‐positive, acid‐fast, mesophilic, with a growth range of 15°C–36°C (Toda and Suzuki 1990). Hatai et al. (1993) reported that mycobacteriosis primarily develops when the immune system is compromised or when other pathogens create an entry point. Indeed, co‐infection with Mycobacterium sp. and Saprolegnia sp. was reported in cultured pejerrey in the same study (Figure 3E, Hatai et al. 1993). Mycobacterium also has zoonotic potential, with transmission to humans possible through direct contact with infected fish or contaminated water, particularly if skin lesions are present. Tuberculosis cases related to fish handling have been reported in Argentina, although so far these have not been linked to pejerrey consumption (Hunt et al. 2013). However, caution is needed due to the zoonotic potential of the disease.

FIGURE 3.

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Mycobateriosis in Odontesthes bonariensis . (A) Macroscopic view of internal granulomas caused by Mycobacterium sp. (B) Liver of infected pejerrey showing soft and hard granulomas (sgn and hgn, respectively, 100×). (C) Magnified view of a hard granuloma (hgn), with an internal core of necrotic cells (necrotic nucleus, nn) surrounded by mycobacteria (mb) and epithelioid cells (ep, 400×). (D) Mycobacteria (mb) present in liver tissue (necrotic nucleus, nn, 1000×). (E) Correlation between mycobacteriosis and saprolegniasis in pejerrey. Figures A–C have been faithfully reproduced and modified to colour from Hatai et al. (1993) (licence 5562811365455), as the originals were low quality and in black and white.

2.4. Other Bacteria

While Aeromonas and Pseudomonas are the most prevalent bacterial genera in pejerrey, other infections have been reported in Japan, including those caused by Vibrio, Flexibacter and Flavobacterium. Vibrio sp., responsible for vibriosis in pejerrey, lead to external hemorrhagic ulcers along the dorsal side and intestinal redness (Toda et al. 1998). Treatment with nitrofuran (1 ppm) showed efficacy in reducing symptoms in infected fish (Toda et al. 1998). However, it is important to note that nitrofuran use has been banned in many countries due to its residue in animal products and potential toxicity to humans (Vass, Hruska, and Franek 2008). Additionally, Chondrococcus spp., a member of the Flexibacter group, was isolated from cultured pejerrey in Kanagawa prefecture, manifesting as yellowish lesions, rapid body discoloration, gill destruction and high mortality rates (Toda et al. 1998). These bacteria typically colonise the gills, and outbreaks are triggered by low temperatures and poor water quality. It was reported that water exchanges and water baths with nitrofuran alleviated symptoms of Chondrococcus infection, although the dosage was not specified (Toda et al. 1998). Additionally, Flavobacterium psychrophilum , a member of the genus Flavobacterium and a well‐known pathogen responsible for bacterial cold‐water disease in fish, was reported as a pathogen of pejerrey in a review by García‐Romero (2001). However, the review did not provide primary sources or specify the location of the reported cases.

The epidemiology and pathogenesis of vibriosis, flexibacteriosis and flavobacteriosis in pejerrey remain underexplored. A potential reason for the limited data on Vibrio and Flexibacter infections is that these outbreaks were primarily documented during the establishment of pejerrey aquaculture in Japan. Differences in culture intensity, practices or specific water conditions could influence the prevalence of these pathogens in Japan compared to South America.

3. Fungal and Fungus‐Like Infections

3.1. Saprolegniasis

Saprolegniasis, caused by water moulds from the family Saprolegniaceae (Oomycetes), is characterised by cottony masses on the body, often with mild inflammation (Lindholm‐Lehto and Pylkkö 2024; Yanong 2003). This is a recurrent disease that affects both wild and farmed pejerrey (Mancini, Rodríguez, Barberis, et al. 2010), with species such as S. parasitica , S. ferax (Figure 4A), Achyla recemosa (Figure 4B,C), Aphanomyces and S. diclina (Figure 4D,E) identified in populations from Argentina and Japan (García‐Romero 2001; Kitancharoen, Yusa, and Hatai 1995; Pacheco‐Marino, Steciow, and Barbeito 2009; Mancini et al. 2016). Saprolegniasis affects all life stages of pejerrey, causing inflammation, fin fraying and body ulceration in adults (Figure 5A–C). This damage, resulting from fungal invasion into tissues (Figure 5D; Hatai et al. 1993), leads to lesions on the fins, mouth and operculum that compromise skin integrity, increasing the fish's vulnerability to bacterial and other fungal infections. In wild pejerrey infected with Saprolegnia sp. in Argentina, fluffy masses were observed on the gills, extending beyond the operculum and causing breathing difficulties (Mancini et al. 2006). Whether saprolegniasis is a primary infection or a result of stressors (e.g., abrasions, trauma, ulcers, poor water quality, temperature changes, overcrowding, parasitism) or other infections is debated. However, evidence from early studies in pejerrey suggests that saprolegniasis is a secondary infection in this species, with its occurrence increasing in the presence of other diseases, such as aeromoniasis (Hatai et al. 1993; Lawhavinit, Hatai, and Kubota 1986). In aquaculture, poor practices drive the occurrence of Saprolegnia outbreaks in pejerrey, particularly in eggs and fingerlings, while in the wild, water quality and temperature are the main factors (Mancini et al. 2016). Despite the prevalence of Saprolegnia in wild pejerrey, mortality rates are relatively low compared to cohabiting fish species (Licoff and Grosman 2008; Mancini et al. 2006; Mancini, Rodríguez, Barberis, et al. 2010).

FIGURE 4.

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Species of saprolegnia in Odontesthes bonariensis . (A) Saprolegnia ferax mycelium containing zoosporangia and oogonia in a water culture. (B) Mycelium of Achlya racemosa with oogonia. (C) Pejerrey eggs infected by A. racemosa . (D, E) Zoosporangia and pitted elongated oogonia of S. diclina , respectively. (A–C) Adapted from Pacheco‐Marino, Steciow, and Barbeito (2009), with permission from the European Association of Fish Pathologists. (D, E) Adapted from Kitancharoen, Yusa, and Hatai (1995), with permission from The Mycological Society of Japan.

FIGURE 5.

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Saprolegniasis in Odontesthes bonariensis . (A, B) Haemorrhages and fraying of the dorsal and caudal fins (indicated by arrows) caused by Saprolegnia parasitica . (C) Illustration of the main fish's body areas affected by Saprolegnia diclina . (D) Histology of the skin from an infected pejerrey, showing fungal hyphae (hy) infiltrating the dermis (de), hypodermis (hpd), and lateral musculature (skm), with the epidermis (epd) visible at 40× magnification. (A, B) Reproduced from Mancini, Rodríguez, Barberis, et al. (2010) (C, D) Adapted from Hatai et al. (1993) (licence 5562811365455).

3.2. Fusariosis

Fusariosis is a fungal disease caused by the genus Fusarium, a member of Deuteromycetes, which parasitizes various aquatic animals (Yanong 2003). In fish, Fusarium species are recognised as cutaneous or visceral pathogens. In pejerrey, F. solani, F. semitectum, F. incarnatum and F. equiseti primarily infect eggs and embryos (Pacheco‐Marino, Steciow, and Barbeito 2009; Pacheco‐Marino et al. 2016), initially targeting non‐viable or dead eggs and subsequently spreading to viable ones (Figure 6A–C), leading to significant reductions in hatching rates. The fungus begins its infection at the chorionic membrane before progressing to embryonic tissues as it grows (Figure 6B–D). Pacheco‐Marino et al. (2016) demonstrated that exposing infected eggs to sodium chloride at concentrations of 120 and 160 g/L inhibited fungal growth in vitro and fully prevented infection (Figure 6D). Unfortunately, such a high concentration of salt reduces egg survival, thereby limiting the applicability of their findings. Poor water quality, overcrowding and elevated water temperatures are key factors that promote the proliferation of Fusarium in pejerrey. While Fusarium has less impact on adult specimens than other fungal infections like saprolegniasis, they can still cause up to 90% mortality in eggs and embryos (Pacheco‐Marino et al. 2016).

FIGURE 6.

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Fusariosis in Odontesthes bonariensis . (A) Early embryos infected by Fusarium sp. (B). Histological section of an embryo displaying hyphae on the surface. (C). Magnified view of the boxed area in panel B, illustrating Fusarium sp. hyphae penetrating the chorion and invading embryonic tissue. (D). Graph depicting the average percentage of survival of Fusarium‐infected eggs following NaCl treatment for 96 h. Mycelia (Myc), Hyphae (Hyp), Developing Embryo (Eb). (A–D) Adapted from Pacheco‐Marino et al. (2016), with permission from the Asociación Española de Micología.

4. Parasite Infections

Parasitosis is prevalent in both cultured and wild pejerrey, with hosts to various parasites such as Lernaea, Cangatiella, Contracaecum, Gyrodactylus and Austrodiplostomum. Other parasitic genera identified include Hysteroda, Ichthyopthirius, Chilodonella, Trichodina, Argulus, Dactylogyrus and Tylodelphys (Flores et al. 2016; Mancini et al. 2016). However, there is a lack of information on the pathogenesis of these latter parasites, and they are not known to cause severe disease or high mortality rates in pejerrey. The main parasites affecting this species are discussed below.

4.1. Lernaea

Parasitism by the anchor worm Lernaea spp. (Crustacea), presumably Lernaea cyprinacea , is a common issue in pejerrey, particularly in central Argentina (Mancini, Rodríguez, et al. 2008; Plaul, García Romero, and Barbeito 2010; Soares et al. 2018). Most parasitological studies on pejerrey have identified this parasite only at the genus level (Lernaea spp.), as molecular identification had not yet been conducted. Although a recent analysis has confirmed the presence of L. cyprinacea in pejerrey (Soares et al. 2018), caution is warranted when interpreting earlier literature. The wide distribution of O. bonariensis across South America suggests the potential presence of other Lernaea species infecting this fish. Notably, two native lernaeid species, Lernaea argentinensis and Lernaea devastatrix, have been reported parasitizing both native and introduced fishes in South America (Waicheim et al. 2019). Therefore, in this review, we will use the term Lernaea spp. for studies that have not specifically identified the parasite as L. cyprinacea .

Lernaea cyprinacea has been widely introduced across South America through the importation of aquarium fish (Soares et al. 2018). In wild pejerrey, infestations of Lernaea spp. usually occur during the warmest months when temperatures are between 25°C and 30°C, particularly in low‐salinity waters (Bethular et al. 2014). Since the initial record of Lernaea spp. in pejerrey in Central Argentina (Mancini, Rodríguez, et al. 2008), there has been a significant rise in the incidence of L. cyprinacea (Mancini, Grosman, et al. 2019). In pejerrey, Lernaea cyprinacea attach to the fin bases, particularly the dorsal, pelvic and pectoral fins, feeding on blood and other fluids, which leads to inflammation and hemorrhagic ulcers (Figure 7A–H; Mancini et al. 2021). Additionally, it creates entry points for other pathogens, leading to secondary infections (Avenant‐Oldewage 2011; Mancini et al. 2016), although secondary infections are not always present (Mancini, Grosman, et al. 2019). The parasitic load tends to increase under conditions of warmer temperatures and lower salinity (Bethular et al. 2014; Mancini, Bucco, et al. 2008). Pejerrey are highly susceptible to Lernaea spp. infestations in aquaculture settings, which has been identified as one of the main reasons for the failure of cultivation trials for this species in Israel (Hephert and Pruginin 1981). Low loads of L. cyprinacea may not significantly impact on the overall body condition of adult pejerrey; however, they can affect the fish's appearance, diminishing its commercial value and marketability. This issue is further exacerbated as parasitic loads tend to increase in larger fish, amplifying the negative effects on their visual quality (Mancini et al. 2021). In contrast, even a few parasites can be fatal to pejerrey fingerlings (Soares et al. 2018).

FIGURE 7.

FIGURE 7

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Parasitism of Odontesthes bonariensis by Lernaea cyprinacea. (A–F) Specimens parasited by L. cyprinacea . Arrows indicate the sites of infection throughout the body. (G, H) Scanning electron micrographs of L. cyprinacea isolated from an invasive fish from Patagonia, Argentina. (A–F) Modified from Mancini et al. (2021) (license CC BY‐NC‐ND 3.0). (G, H) Reproduced from Waicheim et al. (2017) with permission from Check List‐Pensoft.

4.2. Contracaecum

Contracaecum spp. (Nematoda) are cosmopolitan parasites that infect both terrestrial and aquatic animals (Shamsi 2019). Metacercariae of these digenetic trematodes are commonly found in pejerrey, particularly in central Argentina. These parasites inhabit the intestinal mesentery surrounding the gut (Drago 2004, 2012). Due to the limitations in identifying them to the species level without molecular tools, most studies have reported Contracaecum at the genus level (Figure 8A,B). However, analysis of the final host, the Neotropic cormorant Nannopterum brasilianus, has identified two distinct Contracaecum species: C. australe and C. jorgei (Biolé et al. 2012; Sardella et al. 2020). The prevalence of Contracaecum sp. infection in pejerrey appears to be age‐dependent, with older fish (over 2 years old) being more frequently infected (Mancini et al. 2016). Higher densities of microcrustaceans, which act as intermediate hosts, and crowded fish populations are associated with increased parasitism levels by Contracaecum spp. (Mancini et al. 2014). These parasites can negatively affect the body condition of pejerrey. Moreover, Contracaecum larvae pose a potential zoonotic risk as they migrate to the muscle of infected fish and can be transmitted to humans, potentially causing human anisakidosis when consumed raw or undercooked (Shamsi 2019). Therefore, their presence should be carefully monitored, as their prevalence in pejerrey may pose a risk due to the widespread practice of consuming uncontrolled fish from recreational fisheries (Mancini et al. 2005).

FIGURE 8.

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Parasites of Odontesthes bonariensis . (A, B) Photomicrographs of the anterior and posterior ends of Contracaecum sp. larvae, respectively. (C–E) Cangatiella macdonaghi visualised through light microscopy (C, D) and Scanning Electron Microscopy (E). (F) Gyrodactylus sp. (G) Metacercaria of Austrodiplostomum mordax. (H, I) Skeletal deformities (non‐pathognomonic symptoms) in specimens infected with A. mordax . (A–D, F–G) Adapted from Flores et al. (2016), with permission from Elsevier. (E) Reproduced from Drago and Núnez (2017) (Creative Commons License 4.0).

4.3. Cangatiella

Parasitism by the flatworm Cangatiella macdonaghi (Cestoda) is prevalent in pejerrey. Like Lernaea, this parasite is more common in wild pejerrey during the warmest months (Bethular et al. 2014). C. macdonaghi primarily infects the digestive tract of pejerrey, preferably in the posterior part of the digestive tract (Figure 8C–E). According to García‐Romero (2001) the parasite does not cause histopathological alterations in the gut epithelium. Although C. macdonaghi is widespread in pejerrey populations in Argentina, its pathogenicity is low at low densities (Mancini et al. 2016). However, the parasite density can peak in conditions of reduced zooplankton, blooms of dinoflagellates (Ceratium), and low food availability (Silverio et al. 2004).

4.4. Gyrodactylus

Gyrodactylus spp. is a genus of small, leech‐like parasites that infect the skin, gills and fins of marine and freshwater fish. These parasites pose a significant concern in both cultured and wild fish due to their potential to cause disease outbreaks (Sandodden et al. 2018). However, data on Gyrodactylus parasitosis in pejerrey remains limited. In Argentina, Gyrodactylus sp. (Figure 8F) has been identified in the gills and fins of cultured fish, though the species remains unidentified. While its impact on pejerrey health appears to be minor, this may largely depend on the parasitic load (Flores et al. 2016).

4.5. Austrodiplostomum Mordax

The metacercariae of this member of the family Diplostomidae have been observed in various neural structures of pejerrey, including the telencephalon, optic nerves and spinal cord (Figure 8G; Drago 2012). While most parasites remain free within the neural ventricles, some invade neural tissue, causing inflammation and necrosis (García‐Romero 2001). Ostrowski de Nuñez (1964) provided a detailed description of the metacercariae of this parasite isolated from the brain of pejerrey. The presence of A. mordax has been associated with behavioural changes and vertebral deformities in pejerrey, including scoliosis, lordosis and kyphosis, primarily affecting the caudal and precaudal vertebrae (Figure 8H,I). Early studies suggested a causal relationship between A. mordax infection and vertebral deformities in O. bonariensis (Fuster de Plaza and Boschi 1957; Szidat and Nani 1951). However, subsequent observations indicated that skeletal deformities are not consistently found in parasitized pejerrey, and in some cases, individuals showing deformities even lack Diplostomidae parasites (Cabrera 1962). Other authors have found no association between the presence of A. mordax and the body condition or skeletal structure of the closely related Patagonian silverside, Odontesthes hatcheri (Viozzi and Flores 2002). This inconsistency arises because vertebral deformities in fish are frequently linked to adverse environmental conditions, such as hypoxia, hyperthermia, pollution or even hereditary factors (Cabrera 1962; Eissa et al. 2021). Indeed, under experimental conditions, when pejerrey larvae are reared at elevated temperatures (29°C), which are masculinizing for this species, we often observe individuals with columnar deformities and reduced size (A. Torres‐Martínez, personal communication).

However, studies conducted in other species have directly linked digenean trematodes to skeletal deformities in fish. For instance, Kent et al. (2004) observed digenean metacercariae in the vertebrae, spines, and pectoral girdles of three cyprinid fish species from the Willamette River, Oregon. The parasites were found near vertebral surfaces with lesions or embedded within bone in areas of active bone formation (Kent et al. 2004). A subsequent study in the same river strengthened this association, finding a high incidence of deformities in several fish species linked to the presence of digenean metacercariae (Villeneuve et al. 2005). This relationship was further confirmed through laboratory experiments, where fathead minnows ( Pimephales promelas ) infected with cercariae exhibited deformities identical to those observed in the wild (Villeneuve et al. 2005). To date, no studies have specifically investigated the relationship between A. mordax and spinal deformities in O. bonariensis , meaning these symptoms are not currently considered pathognomonic (Mancini, Guagliardo, et al. 2019). Further experimental research would be valuable in clarifying the potential impact of A. mordax on skeletal health in this species and related Atherinopsidae species.

4.6. Other Parasites

Several other parasites have been documented in pejerrey, affecting various regions of the body. Tylodelphys destructor, a parasite of the brain, is notable for its negative association with Austrodiplostomum mordax, in pejerrey from Patagonia, suggesting parasitic competition or resources in the brain (Viozzi and Flores 2002). Ichthyophthirius, a protozoan genus responsible for the white spot disease, has been reported in Japan; however, detailed descriptions are lacking (Murayama and Koyama 1970). In Argentina, a parasitological survey identified several additional parasites, including Goezia sp., Camallanus corderai, Steganoderma macrophallus and Derogenes patagonicus (Rauque et al. 2018). Furthermore, Myxobolus cerebralis, the myxosporean parasite that causes the whirling disease, has been reported to infect pejerrey with a prevalence of 100%; however, Reartes (1995) did not provide primary references to support this claim.

Other trematodes, such as Thometrema bonariensis, Saccocoeliodes sp. and Wolffhugelia matercula , as well as the nematode Hysterothylacium sp., have also been observed in pejerrey, although their abundance and prevalence were reported as low (Drago 2012). Wild pejerrey populations have also been found to host parasites, such as Proteocephalus (cestode), Tylodelphys cardiophilus, Ascocotyle and Phagicola (trematodes), alongside various other genera, including Chilodonella and Trichodina (protozoa), Argulus (crustacean) and Dactylogyrus (monogenean) (Flores et al. 2016; Mancini et al. 2006, 2016). Given the limited information available, the specific details of these parasites will not be elaborated further in this review.

5. Prevention and Management of Infectious Diseases in Pejerrey Aquaculture

The various bacterial, fungal and parasitic infections impacting pejerrey present considerable challenges in cultured environments. Effective disease management is therefore vital for the species' successful aquaculture. However, achieving effective disease management requires a thorough understanding of the factors driving disease emergence and spread, as well as the implementation of preventive strategies and biosecurity measures (Cain 2022). To date, these aspects have been largely overlooked in pejerrey. Given the lack of data on the treatment of infectious diseases in this species, the following section will focus on current approaches to disease prevention in pejerrey aquaculture and research facilities.

Pejerrey aquaculture faces a number of challenges that extend beyond disease management. In Argentina, aquaculture efforts have mainly concentrated on producing fry and fingerlings for repopulation programs supported by provincial governments (Somoza et al. 2008). However, development has been limited by economic, social and biological factors, with diseases being just one of many obstacles. For instance, biological challenges such as the slow growth rates compared to other commercial fish species have hindered productivity (Miranda and Somoza 2001; Somoza et al. 2008), but it is possible that the biological potential for pejerrey growth has not been fully achieved due to a lack of optimal culture conditions (Solimano et al. 2015). Additionally, the lack of genetic improvements to enhance growth and disease resistance further complicates efforts to develop efficient aquaculture practices. Ongoing research aims to address these issues by developing genetically improved lines with enhanced growth and immunity (INTECH 2023). The potential for optimising pejerrey aquaculture is not limited to genetic improvements. Strategies to overcome these challenges have included refining and adapting culture conditions across diverse environments in Argentina and other South American countries (Amaru‐Chambilla and Flores 2021; Colautti and Lenicov 2001; Colautti et al. 2010; García de Souza et al. 2021). In contrast, significant advances have been made in Japan since pejerrey was introduced in 1966, with progress primarily driven by research in seed production and intensive culture techniques (Strüssmann and Yasuda 2005; Takashima and Strüssmann 1991; Yoshino et al. 1999). However, the persistent occurrence of diseases related to environmental stressors and inadequate management practices has posed major challenges to its further aquaculture development (Murayama and Koyama 1970; Yasuda 1994). Despite efforts to manage these diseases, unresolved challenges, along with a decrease in public awareness, have contributed to the decline of pejerrey aquaculture in Japan (Takashima and Strüssmann 1991). Consequently, its cultivation is now primarily focused on academic research related to environmental sex determination (Strüssmann et al. 2021; Torres‐Martínez et al. 2023) and experimental enclosed aquaculture (Yoshino 2017). A lack of research on diagnosing and treating diseases in cultured pejerrey limits sustainable aquaculture development. A critical point of focus is the well‐documented link between environmental fluctuations, poor handling, suboptimal rearing conditions and the onset of infectious diseases in pejerrey. At the Department of Marine Bioscience at Tokyo University of Marine Science and Technology, we have established rearing protocols to optimise salinity and temperature conditions to prevent disease at various life stages of pejerrey. Research from our laboratory indicates that eggs and embryos incubated at low salinities show greater resistance to fungal infections (Tsuzuki et al. 2000). Similarly, rearing pejerrey at intermediate salinities (0.1%–0.7%, typically 0.3% NaCl) significantly enhances survival across all developmental stages, likely due to better osmotic balance, reducing stress and bolstering immunity (Tsuzuki et al. 2000, 2001). Pejerrey survival at 0% salinity depends on optimal conditions and stress absence (Strüssmann and Yasuda 2005). Temperature requirements decrease from around 26°C for larvae to below 20°C for adults, aligning with developmental changes (Strüssmann and Yasuda 2005). Effective management of temperature, salinity and disinfection is crucial to prevent cross‐contamination and disease outbreaks in this species. Additionally, implementing stringent disinfection measures for nets and containers, as well as using soft, wet towels when handling individual pejerrey to avoid skin abrasions, has proven effective in preventing skin damage that could lead to infections. This approach addresses one of the primary challenges in pejerrey culture, where skin damage and subsequent infections have been recurring issues since the early years of cultivation (Yasuda 1994). The use of UV irradiation for disinfecting incoming water is another potential approach to prevent disease outbreaks, particularly during larviculture, although it adds to production costs. Implementing these measures helps to prevent the convergence of the three primary factors contributing to infectious disease outbreaks in pejerrey: pathogen presence, environmental changes and reduced immunity (Toda et al. 1998). By managing these elements effectively, the risks of disease emergence in cultured pejerrey are significantly minimised.

6. Environmental Stressors and Infectious Diseases in Wild Pejerrey

While controlled aquaculture environments allow for management strategies that can prevent and mitigate disease outbreaks, the situation is more complex in natural ecosystems. Factors such as climate change, pollution, habitat modifications and invasive species significantly alter the ecological dynamics in areas inhabited by pejerrey (Kopprio et al. 2010; Miranda and Somoza 2022; Rauque et al. 2018), potentially increasing their susceptibility to infections, a pattern observed in other species (Marcos‐López, Gale, and Peeler 2010). Understanding these factors is crucial for developing strategies to protect wild populations and prevent disease spread beyond aquaculture systems. Although pejerrey has established self‐sustaining wild populations in Japan, such as in Lake Kasumigaura (Hanzawa, Kubota, and Hori 2004), there is limited data on the ecological challenges faced by these populations. Indeed, as an introduced alien species, wild pejerrey in Japan represents an example of global environmental change (NIES 2024). Therefore, we provide below a brief overview of the environmental stressors, with a focus on wild pejerrey populations in South America.

6.1. Global Warming

Previous research has highlighted global warming's impact on pejerrey, particularly focusing on rising temperatures affecting reproductive regulation, sterility induction and temperature‐induced sex reversal (see Strüssmann et al. 2010; Miranda et al. 2013 for a review). These temperature increases are anticipated to drive infectious diseases in pejerrey and other fish (Franke, Beemelmanns, and Miest 2024). Many habitats of pejerrey, including Argentina's shallow Pampean lakes, are characterised by fluctuating droughts and floods, making them particularly vulnerable to these climatic shifts (Kopprio et al. 2010). Environmental factors such as temperature, salinity and water quality play crucial roles in the emergence and pathogenicity of pejerrey diseases (Mancini, Rodríguez, et al. 2008; Mancini, Bucco, et al. 2008; Mancini et al. 2010). As previously discussed in this review, infections by Saprolegnia spp., Fusarium spp. and parasitism by L. cyprinacea and C. macdonaghi are more prevalent at higher temperatures (Bethular et al. 2014; Mancini, Rodríguez, et al. 2008; Mancini, Bucco, et al. 2008), whereas Contracaecum spp. is more commonly associated with elevated salinity (Mancini et al. 2005). Consequently, conditions resulting from increased water temperatures and evaporation rates (such as higher salinity and increased drought frequency) are likely to promote disease outbreaks in pejerrey, as has been observed in the past for this (Del Ponti and Galea 2018) and other species (Macnab and Barber 2011).

6.2. Anthropogenic Stressors

Studies on water bodies in Argentina, which serve as key habitats for pejerrey, have revealed elevated levels of contamination, particularly in areas of intensive agriculture or urbanisation (Avigliano et al. 2015; Carriquiriborde and Ronco 2006; Puntoriero, Cirelli, and Volpedo 2018). For instance, González et al. (2020) identified high concentrations of endocrine disruptors, including estrogens, androgens and progestogens, in sites near wastewater outfalls in Chascomús, Argentina, at levels up to 100 times higher than those reported in similar environments worldwide. These pollutants can have significant effects on pejerrey, leading to histological alterations (Romano and Cueva 1988), immunosuppression and disruptions of endocrine and reproductive functions (Miranda and Somoza 2022) in this and other species (Anderson 1996; Torres‐Martínez et al. 2017). Concurrently, eutrophication of freshwater systems has led to increased parasite prevalence and opportunistic infections, as it has occurred in three‐spine stickleback populations (Budrian and Candolin 2014). In Argentina, blooms of cyanobacteria and dinoflagellates (Ceratium sp.) have led to massive mortalities of pejerrey and other aquatic species, either through direct toxicity or water quality degradation (Grosman and Sanzano 2002; Quirós 2000; Mancini et al. 2006, Mancini, Rodríguez, Bagnis, et al. 2010). Additionally, land use changes and urbanisation have altered the ecological and hydrological dynamics of pejerrey habitats, affecting water quality, particularly through wastewater discharges (Paredes del Puerto et al. 2022). An illustrative example of environmental impact occurred during the summer of 2018, when mass mortality of pejerrey was reported in Lake Santa Rosa de La Pampa, central Argentina, due to a combination of hypoxia, high temperatures and desiccation (Del Ponti and Galea 2018; Figure 9).

FIGURE 9.

FIGURE 9

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Mass mortality of Odontesthes bonariensis in a shallow lake in central Argentina. In the summer of 2018, a mass die‐off of pejerrey occurred in Lake Santa Rosa de la Pampa, with approximately 22–25 tons of fish dying due to a combination of hypoxia and elevated temperatures. Reproduced from Del Ponti and Galea (2018).

6.3. Impact of Invasive Species

The introduction of alien species such as common carp ( Cyprinus carpio ), ornamental fish, several small Brazilian fishes and salmonids has complicated disease ecology in pejerrey populations by introducing new parasites (Waicheim et al. 2019). However, the full extent of this impact remains unclear, not only in the case of pejerrey but also for native fish in general, due to limited baseline data and the complex nature of parasitic colonisation processes (Dunn, Blakeslee, and Bojko 2023). The effect of alien parasites, known as parasite spillover, on native populations depends on their virulence and pathogenicity, as well as environmental factors like temperature, salinity, organic matter and parasite diversity, which vary across different reservoirs and lakes in Argentina and other South American countries (Llames and Zagarese 2009; Quirós and Drago 1999). Generalist alien parasites or those utilising a range of intermediate hosts pose a significant threat to pejerrey and other native fish, especially if they tolerate broader environmental conditions. For instance, helminth parasites such as digenean trematodes exhibit reduced host specificity and a remarkable ability to parasitize new species, while monogeneans are highly specific parasites (Rauque et al. 2018). The invasive L. cyprinacea , a generalist parasite capable of colonising a variety of fish hosts, both native and invasive (Waicheim et al. 2019), exemplifies this, with nutrient pollution and global warming facilitating its invasion (Maceda‐Veiga et al. 2019). In some cases, invasive fish acquire more parasites than they transmit to native fish, as observed in Argentine Patagonia (Rauque et al. 2018). In addition to transmitting parasites, invasive species can disrupt ecological balance through other mechanisms. For instance, C. carpio , which has colonised diverse environments across Argentina (Maiztegui et al. 2016), impacts native ecosystems beyond parasite spread. It contributes to eutrophication through sediment disturbance and alters water temperatures through ecosystem engineering, potentially triggering a cascade of ecological effects (Badiou, Goldsborough, and Wrubleski 2011; Colautti 2001; Matsuzaki et al. 2007). Under the current climate change scenario, the distribution of invasive species such as the common carp and other alien fish is expected to expand, as climate change creates favourable conditions that promote their dispersion (Carosi, Lorenzoni, and Lorenzoni 2023).

In summary, environmental factors influencing pejerrey disease go beyond global warming, climatic phenomena to encompass anthropogenic pollution, habitat modification and the introduction of exotic species (Neubauer and Craft 2009; Oberdorff 2022), all of which can disrupt the ecological balance and health of pejerrey populations and their vulnerability to infectious diseases (Figure 10).

FIGURE 10.

FIGURE 10

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Environmental change and infectious diseases in wild populations of Odontesthes bonariensis . Global warming, along with associated factors like elevated temperature, low oxygen, increased salinity and anthropogenic influences such as water pollution, eutrophication and the introduction of alien species, triggers a cascade of ecological impacts. These include diminished fish health and altered pathogen dynamics, which ultimately heighten the vulnerability of pejerrey to infectious diseases. Adapted from Kopprio et al. (2010).

7. Securing Pejerrey for Present and Future Generations

The spread of fish diseases has significant implications not only for pejerrey populations but also for human communities, given the species' role in fisheries and food security across South America (Salazar and Muñoz 2019). A comprehensive understanding of the interplay between environmental and biological factors is imperative for preserving wild pejerrey populations. This understanding is also critical for mitigating the physiological, behavioural and survival impacts of these stressors observed in fish species more broadly (Chapman et al. 2021). Specific management plans, such as regular water quality monitoring, are essential to mitigate the economic and health risks posed to local communities that rely on pejerrey (Kopprio et al. 2010) as emphasised by Oberdorff (2022). Preventive strategies that have proven successful in other species (Garseth, Britt, and Hilde 2024; Irwin et al. 2023) could be adapted for pejerrey, including constant monitoring of fish health, restricting access to disease‐prone areas, sterilising gear before fishing, avoiding the release of live bait fish, reducing stock density and improving overall water quality in lagoons (Mancini, Grosman, et al. 2019). These measures require balancing consumption with conservation efforts (Irwin et al. 2023). Implementation of such strategies demands collaboration between government, fisheries companies, fishing clubs, and local communities that rely on pejerrey as a food source (Grosman 2001). Governmental bodies would serve as key regulators, establishing institutional guidelines and legal frameworks for both aquaculture and wild management of pejerrey resources. A sustainable exploitation model would not only safeguard the species' health but also contribute to long‐term ecological balance and food security in regions where pejerrey is an important resource.

8. Conclusion and Prospects

The development of pejerrey aquaculture in South America has faced considerable challenges, with the management of infectious diseases remaining a critical concern. Although Japan has made significant progress in pejerrey aquaculture, particularly in understanding and optimising the requirements for maintaining healthy stocks, these advancements have not been fully adopted in other countries. Additionally, there is a lack of research on the stress response and immune system of pejerrey and their connection to infectious diseases. Understanding these mechanisms could pave the way for innovative disease management strategies that go beyond pharmacological treatments, such as boosting immunity, enhancing resilience to stress and promoting overall fish health. Furthermore, environmental changes and human activities, such as pollution and the introduction of invasive species, exacerbate health and sustainability challenges for wild pejerrey populations. To secure the future of pejerrey aquaculture and preserve natural populations, it is crucial to incorporate comprehensive disease prevention and management strategies that address both aquaculture practices and conservation efforts for wild populations. By prioritising research into non‐pharmacological approaches and addressing environmental stressors, we can mitigate risks to both pejerrey and their ecosystems, fostering the sustainable exploitation and conservation of this valuable species.

Author Contributions

Aarón Torres‐Martínez: conceptualization, investigation, visualization, writing – original draft, writing – review and editing, data curation. Miguel Mancini: writing – review and editing. Fabian Grosman: writing – review and editing. Gustavo Manuel Somoza: writing – review and editing. Carlos Augusto Strüssmann: writing – review and editing, funding acquisition.

Conflicts of Interest

The authors declare no conflicts of interest.

Access to Non‐English Literature

Should readers require further information about the cited literature not available in English, they are encouraged to contact the authors directly.

Acknowledgements

A.T.‐M. would like to thank the Ministry of Education, Sport, Science, and Technology of Japan (MEXT) for the doctoral scholarship that supported him for part of the time during the writing of this review. The authors would like to thank the scientific societies and editorials that kindly granted permission to reproduce published materials on the infectious diseases of pejerrey. We also extend our thanks to Dr. Fabiana Drago from the Museo de La Plata, Buenos Aires, Argentina, for providing the scanning electron micrograph of Cangatiella macdonaghi (Figure 8E) for this review. The first author A.T.‐M. would like to dedicate this paper to Drs. Carlos Augusto Strüssmann and Gustavo Manuel Somoza (the friends of the pejerrey), in recognition of their retirement and their remarkable careers dedicated to the study of O. bonariensis . Their commitment and contributions have laid a strong foundation in pejerrey research and serve as an enduring inspiration.

Funding: The authors received no specific funding for this work.

Data Availability Statement

Data sharing is not applicable to this article as no data sets were generated or analysed during the current study.

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