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Article

Diversity and Distribution of the Order Tetraodontiformes in Spain: New Records, Biological Insights and Ecological Implications

by
Rafael Bañón
1,*,
Bruno Almón
1,2,
Begoña Ben-Gigirey
1,
Andrés Villaverde
1,
Mónica González-Castrillón
1,
Rosario Domínguez-Petit
1,
Carlos García Soler
3 and
Alejandro de Carlos
4,5
1
Instituto Español de Oceanografía (IEO-CSIC), Centro Oceanográfico de Vigo, Subida a Radio Faro, 50-52, Cabo Estai, 36390 Vigo, Spain
2
Grupo de Estudo do Medio Mariño (GEMM), Puerto Deportivo s/n, 15960 Ribeira, Spain
3
Aquarium Finisterrae, Pº Marítimo Alcalde Francisco Vázquez, 34, 15002 A Coruña, Spain
4
Departamento de Bioquímica, Xenética e Inmunoloxía, Facultade de Bioloxía, Universidade de Vigo, Rúa Fonte das Abelleiras s/n, 36310 Vigo, Spain
5
Centro de Investigación Mariña, Universidade de Vigo, 36310 Vigo, Spain
*
Author to whom correspondence should be addressed.
Fishes 2026, 11(3), 157; https://doi.org/10.3390/fishes11030157
Submission received: 22 January 2026 / Revised: 26 February 2026 / Accepted: 2 March 2026 / Published: 9 March 2026
(This article belongs to the Section Taxonomy, Evolution, and Biogeography)
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Abstract

This study documents the presence of two uncommon tetraodontiform fishes and reviews the occurrence of species from this order in Spanish marine waters. Two tetraodontid specimens (Family Tetraodontidae) were caught in the Atlantic waters off the coast of Galicia, northwestern Spain. A specimen of Sphoeroides pachygaster was collected in 2021 off the Costa da Vela, while a specimen of Ephippion guttifer was captured in 2025 in the Ría de Pontevedra, both locations situated in southern Galicia. Morphological analyses, supported by photographic evidence and DNA barcoding, confirmed the preliminary taxonomic identification of the two species. Histological reproductive analysis of the Ephippion guttifer specimen revealed a female in the spawning-capable phase. These findings constitute the first verified record of S. pachygaster and the second of E. guttifer in Galician waters. An updated comprehensive list of tetraodontiform species found in Spanish waters across five geographical demarcations was compiled. Historically, a total of 26 species across five families have been reported in Spanish waters, with 22 in the Canary Islands and 15 in the Spanish Iberian Peninsula and Balearic Islands. Additionally, a review of the presence of neurotoxic tetrodotoxins (TTXs) or paralytic shellfish toxins (PSTs) in each species is included, providing an up-to-date overview of a largely unexplored field in European waters. The increasing occurrence of tetraodontiform fishes in Spanish waters provides further evidence of the progressive tropicalization of the Spanish marine environment.
Keywords:
Galicia; pufferfish; barcoding; morphology; distribution; tropicalization; tetrodotoxins; paralytic shellfish toxins
Key Contribution: The first histological reproductive analysis of a female of Ephippion guttifer and a new northern limit of distribution for this species in the eastern Atlantic are provided. Climate change and tropicalization processes increase the number of tetraodontiform species and the risk of intoxication. Detailed knowledge of regional species distribution and potential toxicity is useful for risk assessment, as well as for developing alert campaigns and preventing accidental consumption.

1. Introduction

Although the order Tetraodontiformes has long been considered a valid category, recent phylogenetic studies on ray-finned fishes have proposed a reorganization of the group. Under this new classification, Tetraodontiformes would be considered a suborder, Tetraodontoidei, within the order Acanthuriformes [1]. However, this proposal has not yet been fully accepted by the wider scientific community. Therefore, until this new classification is confirmed or refuted, the traditional classification is considered herein for the sake of clarity.
There are 412 extant species in the 10 families of living tetraodontiform fishes distributed in tropical to temperate marine and freshwater environments worldwide, showing remarkable diversity in shape, size, and ecological traits [2]. Tetraodontiform fishes are frequently distinguished by their diverse shapes, some of which are highly inflatable. The mouth is small and equipped with relatively few modified, enlarged teeth or substantial, beak-like tooth plates. Moreover, the reduced gill openings are located near the pectoral fin base, and the scales are typically modified into spines, ossicles, or fused bony plates, with pelvic fins either reduced or absent.
The family Tetraodontidae is the most speciose within the order Tetraodontiformes, comprising 193 valid species across 27 genera of pufferfish [3]. These species predominantly inhabit shallow, warm, tropical seas and freshwater environments globally, although certain species, such as the blunthead puffer Sphoeroides pachygaster (Müller & Troschel, 1848), have been recorded at depths exceeding 350 m [2]. Tetraodontid species have been extensively studied in genomic research to elucidate their genome architecture and evolutionary mechanisms [4], toxin ecology and physiology [5], and invasive processes impacting ecosystems and fisheries [6].
The classification and identification of tetraodontiform fishes have primarily relied on morphological characteristics, such as body shape, fin ray counts, stripes and spots, body color, and mouth shape. However, differentiating closely related species within this order is often challenging when based solely on morphology, as many species have very similar characteristics, and limited information is available on the intraspecific variability of relevant features [7]. Consequently, the use of molecular tools coupled with morphological taxonomy in an integrative context has gained importance in recent decades. Several molecular markers have been employed in Tetraodontiformes for phylogenetic [8] and taxonomic analysis [9]. Among them, the cytochrome c oxidase subunit I (COI) stands out as the most widely used due to its proven reliability for fish barcoding and specifically for the differentiation of tetraodontiform species [7,10]. The widespread use of this marker is currently facilitating further progress in clarifying the taxonomy of this order, detecting misidentifications, synonyms, and screening for cryptic species.
Preliminary identification of the specimens covered in this study indicated that they belonged to the genera Ephippion Bibron in Duméril 1855 and Sphoeroides Anonymous 1798. The genus Ephippion is monospecific, with the spiny pufferfish Ephippion guttifer (Bennett 1831) as its sole representative. This species is distributed in the eastern Atlantic Ocean, from Galicia (northwestern Spain) to Angola, and in the Mediterranean Sea, in Algeria, Tunisia, and Italy [11,12].
Considering the growing number of reports of species beyond their known distribution range, the term ‘neo-native’ has recently been defined to include species that change their range and establish themselves beyond their historical boundaries as a result of human-induced environmental changes [13]. The arrival of neo-native species can pose significant ecological risks, including displacing native species, introducing new parasites, altering habitats, and causing local extinctions. In addition, pufferfish are notorious for their toxicity and may display aggressive behavior, posing a threat to human health [14]. However, biological information on these species is frequently scarce and relies on information obtained from their natural areas of distribution, often with very different ecological conditions.
Knowledge of the reproductive ecology and potential of fish provides key parameters for understanding population dynamics. Only information on the reproductive phase of a male specimen of E. guttifer has been reported [11]. Among pufferfish, reproductive strategies show great diversity. Some marine species, such as Torquigener sp., construct elaborate geometric nests on sandy seabeds, and males invest significant energy in building these structures, even providing paternal care in some species [15,16]. Other species, such as Sphoeroides nephelus (Goode & Bean, 1882) and Sphoeroides annulatus (Jenyns, 1842), exhibit batch spawning with asynchronous oocyte development and protracted reproductive seasons, often influenced by environmental factors such as temperature, photoperiod, and lunar cycles [17,18,19]. Freshwater and estuarine species, including Carinotetraodon travancoricus (Hora & Nair, 1941) and Tetraodon schoutedeni Pellegrin, 1926, exhibit batch spawning, low fecundity, and specific courtship behaviors [20,21,22]. However, studies on the reproductive ecology of pufferfish remain scarce.
Tetraodontiformes, mainly pufferfish of the family Tetraodontidae, can be toxic, containing tetrodotoxins (TTXs) and/or paralytic shellfish toxins (PSTs). However, the toxin profile differs depending on the genus or species [23,24]. In marine ecosystems, PSTs are mainly produced by certain dinoflagellates and thereafter transferred and bioaccumulated through the aquatic food web [25], reaching several invertebrate and vertebrate groups, including fish [26,27]. TTXs are believed to originate from bacteria belonging to several bacterial phyla; however, several studies suggest their association with specific dinoflagellate blooms [28]. PSTs and TTXs are potent neurotoxins that block sodium channels in nerve cells and skeletal muscles and produce a blockade of ion conductance, resulting in the occurrence of typical symptoms and signs such as paresthesia of the lips, tongue, and pharynx; weakness; dizziness; and gastrointestinal and neurological symptoms [29,30]. Ingestion of toxic fish tissues containing PSTs and/or TTXs can lead to severe illness and death.
Considering the scenario described above, the study of two new records of tetraodontids identified through integrative taxonomy provides the basis for undertaking a comprehensive review of the records within the entire order in Spain. Therefore, the aim of this study was to update relevant information on species of this order with a proven presence in the study area, with special attention to the two new records and their associated biological traits, as well as to gather information on the potential risk posed by the presence of toxins.

2. Materials and Methods

2.1. Study Area

Spain, with a total coastline extending over 7879 km, represents the southernmost point of continental Europe, with most of its territory situated on the Iberian Peninsula, including the Canary and Balearic Islands and the North African cities of Ceuta and Melilla [31]. To improve the management and conservation of the species, the Spanish government established five marine regions or demarcations based on biogeographic, oceanographic, and hydrological characteristics. The Atlantic region includes the North Atlantic (NOR, Spanish north coast), South Atlantic (SUD, Spanish coast of the Gulf of Cádiz), and the Canary Islands (CAN, the Canary Islands) demarcations, whereas the Mediterranean region comprises the Levantine–Balearic (LEBA, East coast of Spain and Balearic Islands) and the Strait of Gibraltar and Alboran Sea (ESAL) demarcations (Figure 1).

2.2. Sampling Data and Morphological Analysis

The two specimens described herein were caught as bycatch by professional artisanal fishers working in Galician waters, northwest of Spain (Figure 1). The specimens were first frozen and later thawed and measured to the nearest millimeter. Morphometric and meristic characters were determined according to Shipp [32]. The specimens were deposited in the fish collection of the Museo de Historia Natural Luis Iglesias in Santiago de Compostela, with reference codes MHN USC 25103-1 for E. guttifer and MHN USC 25234 for S. pachygaster. Total length (TL) and standard length (SL) were used throughout.

2.3. Molecular Analysis

To generate DNA barcodes for both tetraodontiforms, a partial sequence of the 5′ end of the mitochondrial genome locus COI was first obtained. DNA was extracted from a muscle sample from each specimen using the E.Z.N.A. Tissue DNA Kit (Omega Bio-TeK, Norcross, GA, USA), following the manufacturer’s instructions. Amplification was performed by PCR using the universal fish primer sets C_FishF1t1-C_FishR1t1 and the recommended reaction conditions [33], employing the Horse-Power Taq DNA polymerase MasterMix from Canvax Reagents (Valladolid, Spain). In both cases, amplicons of the expected size were obtained and sequenced using the Sanger method [34] at the Centro de Apoyo Científico y Tecnológico a la Investigación of the University of Vigo (https://cactiweb.uvigo.es/). The nucleotide sequences and their corresponding trace files, together with images of the specimens and the geographical locations of the catches, were deposited in the Marine Fishes from Galicia project created in the BOLD database (https://boldsystems.org/), from where they were also submitted to the GenBank database (https://www.ncbi.nlm.nih.gov/genbank/). Sequence analysis was performed using MEGA11 [35]. To this end, an alignment was performed with all publicly available E. guttifer sequences and a selection of S. pachygaster sequences, provided that their geographical origins were known. Next, a clustering analysis was performed using the neighbor-joining algorithm in pairwise comparison mode with 1000 iterations. The resulting dendrogram was edited using the open-source program Inkscape (https://inkscape.org/). The proportion of nucleotide differences between sequences was measured as uncorrected p-distance.

2.4. Reproductive Analysis

The total weight of the thawed ovary was recorded. One lobe was fixed in 4% buffered formaldehyde solution. One subsample from the central part of the ovary was histologically processed (embedded in paraffin, sectioned, and stained with hematoxylin–eosin) for maturity stage determination. Additionally, four subsamples (20–46 mg in weight) from the anterior, central, and posterior parts of the lobe were taken to estimate fecundity by applying the gravimetric method [36] combined with the whole-mount method [37]. Oocytes were separated from connective tissue using a washing process (modified from Lowerre-Barbieri and Barbieri [38]) and by size using a 500 µm sieve. All subsamples were photographed using a stereo microscope Nikon SMZ18 (Nikon Corporation Japan) connected to a video camera (Nikon Fi13). The oocytes were counted and measured using the Fiji ImageJ 1.54g software [39]. Oocyte density per subsample (oocytes per gram of ovary weight, OD), as well as potential (PF) and relative (PFrel) fecundity (oocytes and oocytes per gram of female weight, respectively), were estimated using the package stats of the software R 4.3.3 [40] and RStudio console (2024-02-29 release) [41], and the results were plotted with the package ggplot2 v.4.0.0 [42].

2.5. Spanish tetraodontiform Species and Vulnerability

A list of tetraodontiform species was compiled for each geographical area based on exhaustive bibliographic research and complemented with GBIF records containing relevant information permitting validation of the original identification [43]. The most recent previous list [44] was used to determine the temporal and spatial changes. Records found outside the five demarcations in the Spanish African territory were included by proximity to the nearest demarcation. Thus, records of Aluterus monoceros (Linnaeus, 1758) from the Chafarinas Islands [45] and Lagocephalus sceleratus (Gmelin, 1789) from Ceuta [46] were included in the ESAL demarcation. The vulnerability status of species recorded in Spanish waters was analyzed following the International Union for Conservation of Nature (IUCN) Red List [47]. The IUCN assesses species for their vulnerability status, which indicates the risk of extinction. According to the IUCN criteria, species are considered threatened if they are categorized as critically endangered (CR), endangered (EN), or vulnerable (VU), and non-threatened if categorized as near threatened (NT), least concern (LC), or data deficient (DD). Species without evaluation were categorized as not evaluated (NE), and the not applicable (NA) category was employed for species of marginal occurrence in Europe (i.e., species whose population in Europe represents less than 1% of the total population) [48].

2.6. Marine Biotoxins

The presence of two groups of marine biotoxins normally associated with tetraodontiform fishes, TTXs and PSTs, in Spanish species was explored in the literature. Therefore, a comprehensive literature search was conducted. The keywords combined the name of each of the species reported in Spain, entered individually, with these others: intoxications, tetrodotoxins, TTXs, paralytic shellfish poisoning, PSP, paralytic shellfish toxins, PSTs, saxitoxins, and STX.
The presence of any of these toxin groups in a fish species, independent of the fish capture location and toxicity levels, was considered. Presence/absence was categorized as detected (D, when they were present and described as detected or quantified in a manuscript), not detected (ND, the species was examined and no toxin was detected), and not evaluated (NE, when there was no information).

3. Results

3.1. New Tetraodontidae Records

Two tetraodontid specimens were examined (Figure 2): Sphoeroides pachygaster MHN USC-25234, 220 mm LT, 26 October 2021, 42.306, −8.890, 15 m depth; Ephippion guttifer MHN USC-25103-1, 567 mm LT, 14 October 2025, 42.409, −8.688, 6 m depth. The main meristic and morphometric characters of both specimens are summarized in Table 1.

3.1.1. Molecular Analysis

The molecular dendrogram (Figure 3) shows how the new sequences grouped unequivocally with those corresponding to their respective species in separate clades, with a minimum distance of 17.64%. The mean intraspecific distance was 0.44% (0–0.83%) for E. guttifer and 0.64% (0–2.23%) for S. pachygaster. The wide range of distances between sequences of the latter, compared to its relatively low average value, is due to the values between the sequences from the United States and Uruguay in relation to the others, whose subclade can be seen in the dendrogram, and which corresponds to values ranging from 1.99% to 2.23%.

3.1.2. Reproductive Analysis

Oocytes in advanced vitellogenesis predominated in the histological sections of E. guttifer, whereas oocytes in early vitellogenesis, cortical alveoli, and atretic oocytes were present, although in low numbers. Some melanomacrophage centers were also observed, likely linked to the atretic process. No postovulatory follicles were detected in the sample; however, because the gonads were frozen and thawed prior to fixation, the structures were not preserved with sufficient quality to confirm this with certainty (Figure 4).
Oocyte diameter ranged between 214 and 1134 µm (692.4 ± 192.3 µm, mean ± s.d.). Subsample 1 (sub1) showed a bimodal distribution (371 and 851 µm), while the other subsamples presented a single mode of 407, 793, and 772 µm for subsamples 2, 3, and 4, respectively. When all diameter data were pooled, a bimodal distribution was detected with a main mode of 782 µm and a secondary mode of 383 µm (Figure 5).
The number of oocytes in each subsample ranged between 85 and 197 oocytes (mean 141 oocytes), which corresponds to an OD between 1891 and 9850 oocytes per ovary gram (Table 2).
Given the large difference observed in OD among subsamples—up to five times higher in sub3 compared to sub2—the potential fecundity was calculated by pooling data from all subsamples. This resulted in a PF of 1,335,584 oocytes (95% CI: 635,227.3–2,593,367.3 oocytes) and a relative PF of 533.4 oocytes (95% CI: 253.685–1035.690 oocytes) per gram of female gutted weight. If fecundity was estimated as the average fecundity from the four subsamples, it resulted in a PF of 1,661,950 oocytes (95% CI: 636,206–2,687,694 oocytes) and PFrel of 663.7 oocytes (95% CI: 254.08–1073.36) per gram of female body weight.

3.2. Spanish tetraodontiform Species

A list of Tetraodontiformes recorded by Spanish marine demarcations is shown in Table 3, including 26 species belonging to five families.
The most speciose family was Tetraodontidae, with seven species, while the least represented was Molidae, with four species. Following the latest review [44], five additional species were added: Cantherhines pullus (Ranzani, 1842) and Cantherhines macrocerus (Hollard, 1853) (Monacanthidae), Chilomycterus mauretanicus (Le Danois, 1954) (Diodontidae), Mola alexandrini (Ranzani, 1834) (Molidae), and Melichthys niger (Bloch, 1786) (Balistidae).
In terms of spatial distribution, CAN, with 22 species (85% of the total), was the most speciose region, followed by SUD (13 species, 50%), ESAL (12 species, 46%), NOR (9 species, 35%), and LEBA (8 species, 31%).

3.2.1. Vulnerability Status

Table 4 shows the IUCN vulnerability status of Spanish species of Tetraodontiformes at the global and European levels.
The global assessment covered 25 of the 26 species (96.2%), and most of them (84.6%) were assessed as LC, with only three species (11.5%) considered vulnerable. At the European level, the assessment was reduced to 14 species (53.8%), of which seven (26.9%) were assessed as LC, and none were in the vulnerable category.

3.2.2. Report of TTXs and/or PSTs in Spanish tetraodontiform Species

The compiled information on the reporting of either TTXs or PSTs presence in the tetraodontiform species described in Spain is summarized in Table 5. Extended information, including details on toxins found, analysis methods employed, country of study, and interesting remarks, is shown in Table S1.

4. Discussion

Morphometric, meristic, and genetic analyses confirmed the identification of E. guttifer and S. pachygaster [2,12,32]. Molecular analysis supported the reliability of morphological identification, showing newly generated sequences clustered in the same clades as those retrieved from public databases and assigned to the same species, although obtained from different geographical areas.
Molecular techniques based on voucher specimens identified by expert taxonomists are highly recommended for accurate identification of fish. This is particularly useful in the case of potentially toxic species that may be consumed, as it provides the necessary certainty in identification when assessing and managing the risks associated with their accidental capture and subsequent entry into the food chain, thus ensuring public health [70]. DNA barcoding is a powerful technique used for food safety and to prevent commercial fraud. It has been used successfully to detect fillets of E. guttifer imported from African countries into the European Union (Germany and Italy) labeled as African monkfish (Lophius spp.) [71] and also a toxic pufferfish illegally imported, which was mislabeled as headless monkfish in the Chicago market [72].
Based on the inspection of the histological section and the frequency distribution of oocyte diameters, the species could exhibit either asynchronous or group-synchronous oocyte development. The fact that most of the observed oocytes were in advanced vitellogenesis and that the cohort of oocytes smaller than 500 µm was very small leads us to suspect group-synchronous development; however, because hydrated oocytes were not observed and the sample quality was insufficient to detect post-ovulatory follicles, we cannot rule out that eggs have already been released and that, therefore, the species has asynchronous development. For the same reason, it is not possible to determine whether spawning is partial or complete. Previous studies on other genera of the family Tetraodontidae have reported that pufferfishes have asynchronous oocyte development with partial spawning [19,20,22], so it is expected that E. guttifer follows the same reproductive strategy.
The females analyzed were in the spawning-capable phase, according to Lowerre-Barbieri et al. [73]. Although there is no information on the spawning season of this species, reproductive studies of other genera within the family Tetraodontidae show a wide variability in patterns depending on species and location, ranging from a protracted spawning season in S. nephelus [19] and Lagocephalus lunaris (Bloch & Schneider, 1801) [74], to distinct seasonal peaks, for example, spring for Marilyna pleurosticta (Günther, 1872) [75], spring/summer for C. travancoricus [20], summer for Torquigener flavimaculosus Hardy & Randall, 1983 [76], or autumn/winter for Tetractenos hamiltoni (Richardson, 1846) [75] and Sphoeroides testudineus (Linnaeus, 1758) [77].
The distribution of oocytes was not homogeneous within the ovary, which may be due either to the progressive development of oocytes along the ovary or the fact that oocytes do not hydrate or are not released homogeneously throughout the ovary but rather by zones. This second hypothesis would imply partial spawning. In any case, it is recommended that several subsamples be taken from different regions of the ovary in future fecundity studies. Likewise, to properly assign maturity status based on histology, it is recommended to analyze sections from the anterior, central, and posterior regions of the ovaries.
The potential fecundity estimated in the present study was 1,335,584 oocytes, and the relative fecundity reached 533.4 oocytes per gram of female gutted weight, although it could be higher, since we were unable to confirm whether the female had already released at least one batch. No previous references on fecundity were found for this species; however, there are data on the partial fecundity of S. nephelus, ranging between 59,087 and 367,022 hydrated oocytes (mean 176,456) [19], and fecundity data (without specification of type) for L. lunaris, ranging between 103,355 and 298,795 oocytes [74]. In both cases, the values are one order of magnitude lower than those estimated in the present study, likely because both refer to partial fecundity.
The previous record of E. guttifer in Galician waters, a male specimen caught in January, was also in the spawning-capable phase [11]. The occurrence of specimens both in maturation and/or mature would testify to the presence of a stable population with active reproductive individuals [78].
Owing to their primarily tropical distribution and non-commercial nature, the species composition of tetraodontiform fishes has historically been poorly understood in Spain. Only three species were initially recorded by Cornide (1734–1803) and Cabrera (1763–1827). Mola mola (Linnaeus, 1758) was recorded by Cornide [79] and Cabrera [80] in the NOR and SUD demarcations, respectively, while Diodon hystrix (Linnaeus, 1758) and Balistes capriscus (Gmelin, 1789) were only recorded in the SUD [80]. Subsequently, E. guttifer [81], Lagocephalus lagocephalus (Linnaeus, 1758) [82], and Ranzania laevis (Pennant, 1776) [56] were added to the list, bringing the total number of species to six by the mid-twentieth century. Since then, this number has increased considerably, reaching 21 species in the latest update [44].
Our review increases the total number of species to 26 (an 18% increase) and the total number of new demarcation records to 17 (a 36% increase) compared with Báez et al. [44]. Five new species were recorded: C. pullus [53], M. alexandrini [57], C. mauretanicus, C. macrocerus, and M. niger [54].
CAN had the highest number of species of any demarcation (22 species), but it also experienced the greatest increase, with five new species recorded. The geographical location of the Canary Islands—at a lower latitude than the Iberian Peninsula and close to the African coast, where a similar number of tetraodontiform species are found—probably explains their greater species richness. Along the Iberian coast, there are more species per demarcation in the south (SUD and ESAL, with 13 and 12 species, respectively) than in the north (NOR and LEBA, with 9 and 8 species, respectively). This suggests a general gradient in species abundance from south to north, with fewer species present at higher latitudes with lower temperatures. This is in accordance with the tropical nature of this order.
Although an increase in the number of known species is a normal process in ichthyological knowledge, climate change and warming waters appear to have played an important role in increasing Spanish fish biodiversity in recent decades [44,83,84]. Global warming can cause fish to adapt or migrate in search of optimal thermal conditions, resulting in changes in their distribution over time and space, typically northward [85]. Consequently, the tropicalization of temperate marine ecosystems is evident, a phenomenon that has been well documented in different Atlantic and Mediterranean Spanish regions [51,54,83,86].
Aside from a few exceptions (e.g., molids), most tetraodontiform species occur in tropical and subtropical waters. Therefore, new records from this order at higher latitudes provide further evidence of the evolution of this process. In pioneering works on the tropicalization of North Atlantic waters, S. pachygaster is mentioned as one of the species that has spread more rapidly northward [87,88]. The evolution of the Atlantic French fauna of Tetraodontiformes was also related to global warming [89]. In Spanish waters, records of Lagocephalus laevigatus (Linnaeus, 1766) [90] and E. guttifer ([11]; this study) in NOR, and C. pullus [53] and C. macrocerus [54] in CAN, represent a new northern limit of distribution for these species in the East Atlantic, which also supports the tropicalization hypothesis.
Another record of biogeographical interest is the westernmost occurrence of the Lessepsian L. sceleratus in the Mediterranean Sea, at the Strait of Gibraltar [46]. This Indo-Pacific species was introduced to the Mediterranean via the Suez Canal in 2003 and has since spread across the entire basin. As climate models predict, global warming is weakening the climatic barriers that have historically prevented Lessepsian species from expanding further and moving into the Atlantic Ocean in the coming years [91].
Only two species on the list, Balistes punctatus (Gmelin, 1789) and C. pullus from CAN, have been included in the national reference inventories of non-native marine species [92]. Both species have been classified as “crypto-expansive species” by these authors, defined as species with some evidence of their non-native status, but uncertain due to an unclear mode of introduction from their native range (i.e., natural range expansion versus human-mediated expansion). However, following these criteria, the authors omitted many other species, such as the aforementioned E. guttifer and L. laevigatus, from the NOR demarcation. According to this criterion, many tetraodontiforms recorded in Spain should be added to this list.
In terms of vulnerability status, only C. reticulatus is listed as vulnerable in the Spanish Catalogue of Threatened Species [93]. However, the European IUCN Red List assessment does not include any species in the vulnerable category, which is consistent with a previous report [48]. These authors only assessed tetraodontiform species in the LC or DD categories, the latter often including species with low occurrence and/or taxonomic uncertainty [48]. The last IUCN assessment of this species was conducted in the 2010s. Since then, new taxonomic data and regional changes in abundance have prompted a re-evaluation of the IUCN assessment. Although LC remains the dominant category, an increase in the percentage of threatened species has been observed in recent reviews [94,95]. About 8% of tetraodontids are considered threatened or near-threatened [94], and three of the five species in the Molidae family have changed their status from LC to VU [96].
There is clear evidence that many Tetraodontiformes may benefit from global warming and other human-induced environmental changes, expanding their distribution range, which could have severe socio-economic and ecological impacts. Of the species listed, L. sceleratus has been considered the worst invasive species in the Mediterranean Sea due to its neurotoxins (TTXs), its impact on marine biodiversity, and the increased costs and labor it entails for fishermen [97].
Several studies worldwide have reported the presence of either TTXs or PSTs in certain tetraodontiform species inhabiting Spanish waters. However, the information available on the potential presence of toxins in the species included here is still scarce, with some fish species evaluated for either TTXs or PSTs. Moreover, for 16 of the species evaluated, we could not find any study on the analysis of TTXs or PSTs. While Rambla-Alegre et al. [64] reported the first evidence of high amounts of TTXs in L. sceleratus specimens caught off the Spanish Mediterranean coast, they did not find TTXs in S. pachygaster or L. lagocephalus from the same area. However, Alkassar et al. [66] detected PSTs in L. lagocephalus from the same coast. In Portugal, which shares waters with Spain, TTXs have already been reported in S. marmoratus [65,68], and PSTs have been found in L. lagocephalus [65]. The presence of strong neurotoxins such as TTXs or PSTs in pufferfish from the Spanish coast, combined with the scarcity of data, points to the need to extend studies to more Spanish tetraodontiform species.
The arrival of E. guttifer to the Atlantic European waters also showed a high new parasitic nematode load [98]. The introduction of parasite species into a new marine environment could negatively affect native fauna because they have not had the evolutionary time to reach an equilibrium relationship [99].
Species belonging to the order Tetraodontiformes can be toxic when consumed. However, the number and abundance of toxic species in Spanish waters are relatively low; there is neither a target fishery nor a traditional culinary history, and the current European food legislation prohibits their commercialization [100]. Therefore, the probability of potential intoxication is currently low. However, under the present scenario of climate change and tropicalization, this risk is growing. Knowing the composition, geographical distribution, and potential toxicity of species is particularly important for the early detection of toxic fish species and alerting the population to potential risks. In this way, food safety issues with potentially severe consequences resulting from accidental consumption can be prevented.

5. Conclusions

The order Tetraodontiformes in Spanish waters currently comprises 26 species in five families, reflecting moderately rich but shifting marine biodiversity. Research indicates a clear south-to-north diversity gradient, with the Canary Islands hosting the highest number of species in the region. However, climate change and rising sea temperatures are driving the progressive tropicalization of the marine environment, causing tropical species to expand their ranges toward higher latitudes. The recent findings of S. pachygaster and E. guttifer in Galician waters represent an example of this process.
Histological analysis of the E. guttifer specimen revealed a spawning-capable female, which increases the chances of new colonization. Given that tetraodontiform species may contain potent neurotoxins, such as TTXs or PSTs, it is essential to identify them accurately to prevent their appearance in the commercial market. Integrative taxonomy, which combines morphology with DNA barcoding, enables the correct identification of species. Although current risks to human health remain low owing to existing trade bans in the EU, the increasing frequency of these fish requires continuous monitoring to mitigate potential socio-economic, ecological, and human health impacts.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/fishes11030157/s1, Table S1. Tetraodontiform species described in Spain, together with information on studies reporting the presence of TTXs or PSTs, including toxins reported, analytical methods and country [58,59,60,61,62,63,64,65,66,67,68,69,101,102]. Analytical techniques abbreviations: liquid chromatography-tandem mass spectrometry (LC-MS/MS); Liquid Chromatography-High Resolution Mass Spectrometry (LC-HRMS); Liquid Chromatography/Matrix-Assisted Laser Desorption/Ionization-Time of Flight Mass Spectrometry (LC MALDI-TOF MS); High Performance Liquid Chromatography with Fluorescence Detection (HPLC-FLD); Ultraviolet Spectrophotometry (UV) and Gas Chromatography-Mass Spectrometry (GC-MS).

Author Contributions

Conceptualization, R.B.; methodology, R.B., B.A., R.D.-P., B.B.-G. and A.d.C.; formal analysis, R.B., B.A., R.D.-P., B.B.-G. and A.d.C.; investigation, R.B., A.V., M.G.-C. and A.d.C.; resources, R.B., C.G.S., R.D.-P. and A.d.C.; writing—original draft preparation, R.B., B.A., R.D.-P., B.B.-G. and A.d.C.; writing—review and editing, R.B., B.A., R.D.-P., B.B.-G. and A.d.C.; visualization, R.B.; supervision, R.B. and B.A.; project administration, R.B.; funding acquisition, R.D.-P. and A.d.C. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The sequences generated and used in the current study are available in the BOLD systems (https://www.boldsystems.org/, accessed on 25 November 2025) and GenBank (https://www.ncbi.nlm.nih.gov/genbank/, accessed on 25 November 2025) repositories. The specimens used in this study for a taxonomic purpose have been deposited in the fish collection of the Museo de Historia Natural, Universidade de Santiago de Compostela (MHNUSC) in Santiago de Compostela, Spain.

Acknowledgments

We would like to thank the crew of the vessel “A Entallada” and the Combarro fishermen’s association for providing the specimen and data of E. guttifer. We are also grateful to the crew of the vessel “Dous mil” and the Bueu fishermen’s association for providing the specimen and data of S. pachygaster. We also want to thank the Aquarium Finisterrae staff, who helped us manage some of the specimens examined.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Map of the study area showing the different geographical demarcations and the number of species recorded in each of them. The close-up views in red show the detailed locations of the new records. This map is for technical use only and does not reflect the official boundaries with neighboring states.
Figure 1. Map of the study area showing the different geographical demarcations and the number of species recorded in each of them. The close-up views in red show the detailed locations of the new records. This map is for technical use only and does not reflect the official boundaries with neighboring states.
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Figure 2. Lateral (left) and dorsal (middle) views of Sphoeroides pachygaster (A) and Ephippion guttifer (B).
Figure 2. Lateral (left) and dorsal (middle) views of Sphoeroides pachygaster (A) and Ephippion guttifer (B).
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Figure 3. Neighbor-Joining dendrogram representation of uncorrected p-distances based on the analysis of 652 nucleotides of the cytochrome c oxidase subunit I (COI) gene in two taxa of Tetraodontidae. The sequences of E. guttifer and S. pachygaster are accompanied by their respective GenBank accession numbers, except for the specimens covered in this study (in bold), which are listed with their BOLD process IDs. Geographical locations are also included when available. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test is shown below each branch. The bar indicates the number of nucleotide differences per site.
Figure 3. Neighbor-Joining dendrogram representation of uncorrected p-distances based on the analysis of 652 nucleotides of the cytochrome c oxidase subunit I (COI) gene in two taxa of Tetraodontidae. The sequences of E. guttifer and S. pachygaster are accompanied by their respective GenBank accession numbers, except for the specimens covered in this study (in bold), which are listed with their BOLD process IDs. Geographical locations are also included when available. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test is shown below each branch. The bar indicates the number of nucleotide differences per site.
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Figure 4. Histological section of the ovary of Ephippion guttifer: (A) general view (2×) of the ovary; (B) detailed view (4×) of oocyte developmental stages; (C) detailed view (10×) of atretic oocytes; (D) melanomacrophage center (40×). PG: primary growth, CA: cortical alveoli, AV: advanced vitellogenic, AT: atretic, MC: melanomacrophage center.
Figure 4. Histological section of the ovary of Ephippion guttifer: (A) general view (2×) of the ovary; (B) detailed view (4×) of oocyte developmental stages; (C) detailed view (10×) of atretic oocytes; (D) melanomacrophage center (40×). PG: primary growth, CA: cortical alveoli, AV: advanced vitellogenic, AT: atretic, MC: melanomacrophage center.
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Figure 5. Density plot of oocyte diameter frequency in each subsample: sub1 in red, sub2 in green, sub3 in blue, and sub4 in purple (left) and pooling all subsamples together (right).
Figure 5. Density plot of oocyte diameter frequency in each subsample: sub1 in red, sub2 in green, sub3 in blue, and sub4 in purple (left) and pooling all subsamples together (right).
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Table 1. Morphometric and meristic data of Ephippion guttifer and Sphoeroides pachygaster caught in Galician waters.
Table 1. Morphometric and meristic data of Ephippion guttifer and Sphoeroides pachygaster caught in Galician waters.
Ephippion guttifer
MHN USC 25103-1		Sphoeroides pachygaster
MHN USC 25234
Total length (mm)567220
Standard length (mm)462187
As % Standard length
Head length29.628.9
Pre-orbital length1419.7
Post-orbital length14.913.5
Horizontal eye diameter5.88.9
Interorbital width13.517.8
Length of gill opening10.29.8
Pre-dorsal-fin length68.475.5
Dorsal-fin base length7.211.3
Pre-anal-fin length71.281.3
Pre-pectoral-fin length31.843.9
Anal-fin base5.33.9
Pectoral-fin length14.815.3
Caudal peduncle depth10.29.8
Body depth28.429.1
Body width28.824.9
Meristic
Dorsal-fin rays108
Anal-fin rays98
Pectoral-fin rays1914
Caudal-fin rays1010
Table 2. Weight in grams (SW), number of oocytes (NO), and oocyte density (OD, oocytes/ovary g) of the four analyzed ovary subsamples of Ephippion guttifer.
Table 2. Weight in grams (SW), number of oocytes (NO), and oocyte density (OD, oocytes/ovary g) of the four analyzed ovary subsamples of Ephippion guttifer.
SWNOOD
Subsample10.046871891
Subsample20.042852024
Subsample30.021979850
Subsample40.0291946690
Table 3. Presence of tetraodontiform fishes listed by family and geographical demarcations. Previous records reported by Báez et al. [44] are consigned as Present (P). The asterisk indicates species that are new to the list. The new occurrence by demarcation is indicated with its corresponding reference number. NOR: Spanish north coast, SUD: Gulf of Cádiz, ESAL: Alboran Sea, LEBA: Levantine–Balearic, CAN: Canary Islands.
Table 3. Presence of tetraodontiform fishes listed by family and geographical demarcations. Previous records reported by Báez et al. [44] are consigned as Present (P). The asterisk indicates species that are new to the list. The new occurrence by demarcation is indicated with its corresponding reference number. NOR: Spanish north coast, SUD: Gulf of Cádiz, ESAL: Alboran Sea, LEBA: Levantine–Balearic, CAN: Canary Islands.
FamilySpeciesNORSUDESALLEBACAN
TetraodontidaeSphoeroides marmoratus (Lowe, 1838) PP[49]P
TetraodontidaeSphoeroides pachygaster (Müller & Troschel, 1848)PPP[50]P
TetraodontidaeCanthigaster capistrata (Lowe, 1839) P[43] P
TetraodontidaeLagocephalus laevigatus (Linnaeus, 1766)P[51]
TetraodontidaeLagocephalus lagocephalus(Linnaeus, 1758)PPPPP
TetraodontidaeLagocephalus sceleratus (Gmelin, 1789) [52]P
TetraodontidaeEphippion guttifer (Bennett, 1831)[11][43]P
MonacanthidaeAluterus monoceros (Linnaeus, 1758)PP[45] P
MonacanthidaeAluterus scriptus (Osbeck, 1765) P
MonacanthidaeCantherhines pullus (Ranzani, 1842) * [53]
MonacanthidaeCantherhines macrocerus (Hollard, 1853) * [54]
MonacanthidaeStephanolepis hispida (Linnaeus, 1766) P
DiodontidaeChilomycterus reticulatus (Linnaeus, 1758) P
DiodontidaeChilomycterus mauretanicus (Le Danois, 1954) * [55] [54]
DiodontidaeDiodon eydouxii (Brisout de Barneville, 1846) PP P
DiodontidaeDiodon holocanthus (Linnaeus, 1758) [51] P
DiodontidaeDiodon hystrix (Linnaeus, 1758)P[56] P
MolidaeMasturus lanceolatus (Liénard, 1840) P
MolidaeMola mola (Linnaeus, 1758)PPPPP
MolidaeMola alexandrini (Ranzani 1834) * [57]
MolidaeRanzania laevis (Pennant, 1776)[43]PPPP
BalistidaeCanthidermis sufflamen (Mitchill, 1815) P
BalistidaeCanthidermis maculata (Bloch, 1786) P
BalistidaeMelichthys niger (Bloch, 1786) * [54]
BalistidaeBalistes capriscus (Gmelin, 1789)PPPPP
BalistidaeBalistes punctatus (Gmelin, 1789) P
Table 4. IUCN Red List status at the global and European levels of Spanish tetraodontiform species. LC: least concern; DD: data deficient; VU: vulnerable; NA: not applicable; NE: not evaluated.
Table 4. IUCN Red List status at the global and European levels of Spanish tetraodontiform species. LC: least concern; DD: data deficient; VU: vulnerable; NA: not applicable; NE: not evaluated.
FamilySpeciesIUCN GlobalIUCN Europe
BalistidaeBalistes capriscusVUDD
BalistidaeBalistes punctatusVUNE
BalistidaeCanthidermis maculataLCLC
BalistidaeCanthidermis sufflamenLCNA
BalistidaeMelichthys nigerLCNE
DiodontidaeChilomycterus mauretanicusLCNA
DiodontidaeChilomycterus reticulatusLCLC
DiodontidaeDiodon eydouxiiLCNA
DiodontidaeDiodon holocanthusLCNE
DiodontidaeDiodon hystrixLCDD
MolidaeMasturus lanceolatusLCNA
MolidaeMola alexandriniNENE
MolidaeMola molaVUDD
MolidaeRanzania laevisLCDD
MonacanthidaeAluterus monocerosLCDD
MonacanthidaeAluterus scriptusLCDD
MonacanthidaeCantherhines macrocerusLCNE
MonacanthidaeCantherhines pullusLCNE
MonacanthidaeStephanolepis hispidaLCLC
TetraodontidaeCanthigaster capistrataLCLC
TetraodontidaeEphippion guttiferLCDD
TetraodontidaeLagocephalus laevigatusLCNE
TetraodontidaeLagocephalus lagocephalusLCLC
TetraodontidaeLagocephalus sceleratusLCNE
TetraodontidaeSphoeroides marmoratusLCLC
TetraodontidaeSphoeroides pachygasterLCLC
Table 5. Detected (D), not detected (ND), not evaluated (NE) tetrodotoxins (TTXs) and paralytic shellfish toxins (PSTs) in Spanish tetraodontiform fishes according to the literature.
Table 5. Detected (D), not detected (ND), not evaluated (NE) tetrodotoxins (TTXs) and paralytic shellfish toxins (PSTs) in Spanish tetraodontiform fishes according to the literature.
FamilySpeciesTTXPSTSource
BalistidaeBalistes capriscusNENE—
BalistidaeBalistes punctatusNENE—
BalistidaeCanthidermis maculataNENE—
BalistidaeCanthidermis sufflamenNENE—
BalistidaeMelichthys nigerNENE—
DiodontidaeChilomycterus mauretanicusNENE—
DiodontidaeChilomycterus reticulatusNENE—
DiodontidaeDiodon eydouxiiNENE—
DiodontidaeDiodon holocanthusDNE[58]
NDNE[59]
DiodontidaeDiodon hystrixDNE[60]
DNE[61]
MolidaeMasturus lanceolatusNENE—
MolidaeMola alexandriniNDNE[62]
MolidaeMola molaNDNE[58]
NDNE[62]
MolidaeRanzania laevisNENE—
MonacanthidaeAluterus monocerosNENE—
MonacanthidaeAluterus scriptusNENE—
MonacanthidaeCantherhines macrocerusNENE—
MonacanthidaeCantherhines pullusNENE—
MonacanthidaeStephanolepis hispidaNENE—
TetraodontidaeCanthigaster capistrataNENE—
TetraodontidaeEphippion guttiferNENE—
TetraodontidaeLagocephalus laevigatusDNE[63]
TetraodontidaeLagocephalus lagocephalusNDNE[64]
NDD[65]
NED[66]
TetraodontidaeLagocephalus sceleratusDNE[67]
DNE[64]
TetraodontidaeSphoeroides marmoratusDNE[68]
DND[65]
TetraodontidaeSphoeroides pachygasterNDNE[64]
NDND[69]
NEND[66]
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Bañón, R.; Almón, B.; Ben-Gigirey, B.; Villaverde, A.; González-Castrillón, M.; Domínguez-Petit, R.; García Soler, C.; de Carlos, A. Diversity and Distribution of the Order Tetraodontiformes in Spain: New Records, Biological Insights and Ecological Implications. Fishes 2026, 11, 157. https://doi.org/10.3390/fishes11030157

AMA Style

Bañón R, Almón B, Ben-Gigirey B, Villaverde A, González-Castrillón M, Domínguez-Petit R, García Soler C, de Carlos A. Diversity and Distribution of the Order Tetraodontiformes in Spain: New Records, Biological Insights and Ecological Implications. Fishes. 2026; 11(3):157. https://doi.org/10.3390/fishes11030157

Chicago/Turabian Style

Bañón, Rafael, Bruno Almón, Begoña Ben-Gigirey, Andrés Villaverde, Mónica González-Castrillón, Rosario Domínguez-Petit, Carlos García Soler, and Alejandro de Carlos. 2026. "Diversity and Distribution of the Order Tetraodontiformes in Spain: New Records, Biological Insights and Ecological Implications" Fishes 11, no. 3: 157. https://doi.org/10.3390/fishes11030157

APA Style

Bañón, R., Almón, B., Ben-Gigirey, B., Villaverde, A., González-Castrillón, M., Domínguez-Petit, R., García Soler, C., & de Carlos, A. (2026). Diversity and Distribution of the Order Tetraodontiformes in Spain: New Records, Biological Insights and Ecological Implications. Fishes, 11(3), 157. https://doi.org/10.3390/fishes11030157

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