Research Article |
Corresponding author: Daniel Cuadrado ( cuadradopm@hotmail.com ) Corresponding author: Nuno Simões ( ns@ciencias.unam.mx ) Academic editor: Christopher Glasby
© 2024 Daniel Cuadrado, Alejandro Hernández-Gonzalez, Carolina Noreña, Nuno Simões.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Cuadrado D, Hernández-Gonzalez A, Noreña C, Simões N (2024) Polyclads (Platyhelminthes) in the southern Gulf of Mexico: unveiling biodiversity and descriptions of two new species. ZooKeys 1221: 103-144. https://doi.org/10.3897/zookeys.1221.128260
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The order Polycladida (Platyhelminthes) in Mexico has historically received limited attention from researchers, primarily due to challenges associated with its low detectability and the scarcity of specialists. This study addresses part of the gap by conducting a comprehensive assessment of polyclad diversity in the southern Gulf of Mexico. Our investigation revealed a total of 27 distinct species, belonging to 17 genera and 12 families, within the suborders Cotylea and Acotylea. Our findings include the identification of 17 species previously undocumented in the Gulf of Mexico. This represents a significant expansion of the region’s known polyclad biodiversity. By revising the polyclad records in the Gulf of Mexico, the known species count has increased from 31 to 50. Furthermore, our research unveiled the presence of two new species, Stylochoplana sisalensis sp. nov. and Emprosthopharynx hartei sp. nov., also marking the first time a species of the genus Emprosthopharynx has been reported for the Atlantic coast of the Americas.
Campeche, flatworms, histological analysis, marine invertebrates, new record, Quintana Roo, species discovery, taxonomy, Yucatan
The Gulf of Mexico (GoM) is renowned for being one of the largest marine ecosystems worldwide, due to its unique combination of hydrographic factors, biological productivity, and population diversity. Covering more than 1.5 million km2, the GoM boasts a broad range of marine habitats, from coral reefs to fishing banks and coastal areas, making it a globally significant ecosystem (
Polyclads, marine flatworms, have not received much attention from researchers in Mexico. There are several reasons for the difficulty of observing these small organisms, including their ability to mimic their surroundings. More than 1000 species of polyclads are known worldwide according to
The present work is a significant contribution to the region as it addresses the shortage of species records and the urgent need to update the biodiversity inventory of the order Polycladida in the Gulf of Mexico (GoM). By examining 142 specimens, we identified 27 taxa that belong to 12 families and 17 genera, increasing the known species count from 31 to 50 (Suppl. material
Records of the order Polycladida in the Gulf of Mexico as reported in Suppl. material
The study material from the southern Gulf of Mexico was obtained through direct field collection using scuba diving and snorkelling in the subtidal (Fig.
To ensure proper fixation, individuals were first anaesthetised with a seawater/magnesium chloride solution (7%). A small tissue sample was extracted and preserved in absolute ethanol for future molecular studies, and the entire specimen was then fixed in Bouin’s solution (saturated picric acid solution, formaldehyde, and acetic acid in a 15:5:1 proportion) for histological studies (
Whole body specimens and the histological preparations were deposited at the Colección regional de Policladidos de la Península del Yucatán, Mexico (CRPPY), located at the Unidad Multidisciplinaria de Docencia e Investigación de Sisal, Facultad de Ciencias, Universidad Nacional Autónoma de México (UMDI-Sisal, FC-UNAM).
cg: cement glands, co: copulatory organ, ct: connective tissue, de: ejaculatory duct, dlv: Lang’s vesicle duct, e: stylet, ed: ejaculatory duct, e-gpt: epithelial-glandular prostate tissue, ev: external vagina, fa: female atrium, fg: female gonopore, i: intestine, iv: internal vagina, lv: Lang’s vesicle, m: muscle layer, ma: male atrium, mb: marginal band, mg: male gonopore, ml: marginal line, oc: cerebral ocelli/eyes, om: marginal ocelli/eyes, ot: tentacular ocelli/eyes, ov: oviduct, p: pharynx, pa: papillae, pp: penis papilla, pt: pseudotentacles, pte: pseudotentacle eyes, pv: prostatic vesicle, s: sucker, sg: shell glands, sv: seminal vesicle, t: tentacle, va: vagina, vd: vas deferens.
Total genomic DNA was extracted from each sample following the phenol-chloroform protocol (
The PCR products were observed using TBE gel electrophoresis in 1.5% agarose gel stained with SYBER Safe and visualised under UV light. PCR products were sent to Macrogen for clean-up and sequencing. Lastly, obtained forward and reverse sequences were combined using the program Geneious Prime v. 2020.2.4 (http://www.geneious.com,
All sequences obtained in the present study have been deposited in the GenBank database under the accession numbers included in Suppl. material
Suborder Cotylea
Periceloidea Bahia, Padula & Schrödl, 2017
Pericelidae Laidlaw, 1902
Pericelis Laidlaw, 1902
Campeche coast, Mexico • 1; Cayos sumergidos del Oeste; 20.9°N, 92.2°W; 0 m; 10 Sep. 2017; A. Gutiérrez leg.; CRPPY-0011 • 1; Cayos sumergidos del Oeste; 20.9°N, 92.2°W; 13 m; 14 Sep. 2017; A. Gutiérrez leg.; CRPPY-0020 • 1; Cayos sumergidos del Oeste; 21.0°N, 92.3°W; 10 m; 9 Sep. 2017; F. Márquez leg.; CRPPY-0022 • 2; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 10.8 m; 14 Sep. 2017; A. Gutiérrez leg.; CRPPY-0024 • 1; Cayos sumergidos del Oeste; 20.5°N, 92.2°W; 26 m; 13 Sep. 2017; X. Vital leg.; CRPPY-0025; Quintana Roo coast, Mexico • 1; Mahahual; 18.6°N, 87.7°W; 13.4 m; 18 Mar. 2018; A. Hernández leg.; CRPPY-0040; Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 5 m; 19 Apr. 2018; A. Hernández leg.; CRPPY-0046 • 1; Cayo Arcas; 20.2°N, 92.0°W; 4.7 m; 19 Apr. 2018; A. Hernández leg.; CRPPY-0051 • 1; Cayo Arcas; 20.2°N, 92.0°W; 6.3 m; 22 Apr. 2018; A. Hernández leg.; CRPPY-0065 • 1; Cayo Arcas; 20.2°N, 92.0°W; 3.2 m; 24 Apr. 2018; A. Hernández leg.; CRPPY-0078 • 1; Cayo Arcas; 20.2°N, 92.0°W; 7.7 m; 25 Apr. 2018; A. Hernández leg.; CRPPY-0083.
This species was previously recorded in Curaçao (
Body oval with multiple marginal folds, 4 cm in length and 2.5 cm in width. Dorsally, pattern of dark pigmentation is interrupted by spots where the white parenchyma is observed. Towards the margin, the white patches become smaller, and the space between them decreases, with scattered black dots. The tentacles are subtle marginal folds, with a clear separation between them, characteristic of the genus Pericelis (Fig.
Pericelis cata
morphology found in the Gulf of Mexico corresponds to the original description by
Yucatan coast, Mexico • 1; Punta Piedra, Sisal; 21.2°N, 90.1°W; 1 m; 30 Apr. 2018; A. Hernández leg.; CRPPY-0087 • 1; Dzilam; 21.5°N, 88.9°W; 9.3 m; 8 May 2018; A. Hernández leg.; CRPPY-0091 • 1; 12 slides; Dzilam; 21.5°N, 88.9°W; 13 m; 10 May 2018; A. Hernández leg.; CRPPY-0097.
Pericelis orbicularia
is known from the south coast of Jamaica (
Body oval-shaped, 2 cm in length and 1 cm in width, with small pseudotentacles, less than 1 mm. Dorsal surface exhibits an orange to light brown reticulated pattern on a regular creamy beige background (Fig.
Prosthiostomum Quatrefage, 1845
Quintana Roo coast, Mexico • 1; Mahahual; 18.6°N, 87.7°W; 13.4 m; 18 Mar. 2018; A. Hernández leg.; CRPPY-0041.
This species was described from Sao Sebastiao Island, Brazil (type locality;
Body shape elongated, 3 cm in length and 0.7 cm in width with a rounded anterior end and a pointed posterior end (Fig.
Prosthiostomum utarum
, originally described as Lurymare utarum (Marcus, 1952), was recently reassigned to the genus Prosthiostomum based on the work of
Yucatan coast, Mexico • 1; Bajos de Sisal; 21.2°N, 90.0°W; 1 m; 10 Sep. 2017; A. Hernández leg.; CRPPY-0033.
Recorded in São Paulo (
Body shape elongated, 3 cm in length and 1 cm in width. Body cream-coloured with brown, orange, and yellow spots arranged densely along the midline and paler towards the margins (Fig.
The spots disappear after fixation and, according to
Yucatan coast, Mexico • 1; Arrecife Alacranes; 22.4°N, 89.7°W; 1 m; 3 Nov. 2017; A. Hernández leg.; CRPPY-0001 • 1; Arrecife Alacranes; 22.4°N, 89.7°W; 1 m; 3 Nov. 2017; A. Hernández leg.; CRPPY-0003 • 2; Arrecife Alacranes; 22.4°N, 89.7°W; 1 m; 4 Nov. 2017; A. Hernández leg.; CRPPY-0005 • 1; Arrecife Alacranes; 22.4°N, 89.7°W; 2 m; 4 Nov. 2017; A. Hernández leg.; CRPPY-0007; Quintana Roo coast, Mexico • 1; Mahahual; 18.6°N, 87.7°W; 15 m; 18 Mar. 2018; A. Hernández leg.; CRPPY-0042; Yucatan coast, Mexico • 1; Dzilam; 21.5°N, 88.9°W; 13 m; 10 May 2018; A. Hernández leg.; CRPPY-0098.
The species was originally described in Thatch Island, US Virgin Islands (
Body elongated, 1.5 cm in length and 0.5 cm in width, with a rounded anterior end and a tapered posterior end. Marginal eyes densely distributed along the anterior margin; cerebral eyes in a heart-shaped cluster. Translucent white background speckled with dense brown to orange spots that gradually decrease in number towards the edges (Fig.
Euryleptidae Stimpson, 1857
Eurylepta Ehrenberg, 1831
Yucatan coast, Mexico • 1; Dzilam; 21.5°N, 88.9°W; 9.3 m; 8 May 2018; A. Hernández leg.; CRPPY-0088.
The species was recorded in Monterey Bay, California (
Body shape elliptical, 1.7 cm in length and 0.5 cm in width, with a translucent to orange-pink colouration, white spots on the dorsal side, and a reddish median line with reddish dots (Fig.
The specimen collected from the Mexican coast exhibits a pigmentation characterised by a pinkish orange hue, as illustrated in Fig.
Campeche coast, Mexico • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 18 m; 14 Sep. 2017; A. Gutiérrez leg.; CRPPY-0018 • 1; Cayo Arcas; 20.2°N, 92.0°W; 16.3 m; 20 Aug. 2018; A. Hernández leg.; CRPPY-0108.
Recorded in the east coast of Florida and the Florida Keys, Bermuda (
Oval or circular-shaped body, 2.3 cm in length and 0.8 cm in width, with a semi-transparent white-beige background with transverse wavy black lines. The lines alternately end in a black spot or an orange blotch. Dorsal surface with white spots and a submarginal semi-transparent and marginal narrow white band (Fig.
The specimen from the Gulf of Mexico aligns with the description of Prostheceraeus crozieri provided by
Prostheceraeus crozieri is documented as a primary consumer of the sea squirt Ecteinascidia turbinata. Although this ascidian species was prolific within the research areas, the occurrence of Prostheceraeus crozieri was limited.
Pseudoceros Lang, 1884
Yucatan coast, Mexico • 1; Arrecife Alacranes; 22.4°N, 89.7°W; 3 m; 4 Nov. 2017; A. Hernández leg.; CRPPY-0008; Campeche coast, Mexico • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 10 m; 14 Sep. 2017; A. Hernández leg.; CRPPY-0015 • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 11.1 m; 13 Sep. 2017; D. Ortigosa leg.; CRPPY-0027 • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 11.7 m; 14 Sep. 2017; D. Ortigosa leg.; CRPPY-0028; Yucatan coast, Mexico • 1; Bajos de Sisal; 21.2°N, 90.0°W; 1 m; 22 Feb. 2018; A. Hernández leg.; CRPPY-0032; Quintana Roo coast, Mexico • 1; Mahahual; 18.6°N, 87.7°W; 7.7 m; 17 Mar. 2018; A. Hernández leg.; CRPPY-0039; Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 6.4 m; 19 Apr. 2018; A. Hernández leg.; CRPPY-0048 • 1; Cayo Arcas; 20.2°N, 92.0°W; 2.2 m; 20 Apr. 2018; A. Hernández leg.; CRPPY-0052 • 2; Cayo Arcas; 20.2°N, 92.0°W; 2.2 m; 20 Apr. 2018; A. Hernández leg.; CRPPY-0054 • 1; Cayo Arcas; 20.2°N, 92.0°W; 2.2 m; 20 Apr. 2018; A. Hernández leg.; CRPPY-0055 • 1; Cayo Arcas; 20.2°N, 92.0°W; 2.2 m; 21 Apr. 2018; A. Hernández leg.; CRPPY-0061 • 1; Cayo Arcas; 20.2°N, 92.0°W; 2.2 m; 22 Apr. 2018; A. Hernández leg.; CRPPY-0063 • 2; Cayo Arcas; 20.2°N, 92.0°W; 6.3 m; 22 Apr. 2018; A. Hernández leg.; CRPPY-0067 • 1; Cayo Arcas; 20.2°N, 92.0°W; 7.5 m; 23 Apr. 2018; A. Hernández leg.; CRPPY-0069 • 1; Cayo Arcas; 20.2°N, 92.0°W; 5.9 m; 23 Apr. 2018; A. Hernández leg.; CRPPY-0070 • 1; Cayo Arcas; 20.2°N, 92.0°W; 5.3 m; 23 Apr. 2018; A. Hernández leg.; CRPPY-0072 • 1; Cayo Arcas; 20.2°N, 92.0°W; 5.3 m; 24 Apr. 2018; A. Hernández leg.; CRPPY-0074 • 1; Cayo Arcas; 20.2°N, 92.0°W; 9.9 m; 25 Apr. 2018; A. Hernández leg.; CRPPY-0081 • 1; Cayo Arcas; 20.2°N, 92.0°W; 7.7 m; 25 Apr. 2018; A. Hernández leg.; CRPPY-0084 • 1; Cayo Arcas; 20.2°N, 92.0°W; 3.4 m; 18 Aug. 2018; A. Hernández leg.; CRPPY-0106 • 1; Cayo Arcas; 20.2°N, 92.0°W; 3.4 m; 18 Aug. 2018; A. Hernández leg.; CRPPY-0107.
Recorded in the Birds Islands, Bermuda (
Body shape elongated with rounded anterior and posterior end, 2.5 cm in length and 1 cm in width. Dorsal pigmentation ranges from yellow to dark brown, with scattered white dots on its dorsal surface and with a yellowish or pale green marginal rim. Conspicuous dark marginal band interrupted by transverse white stripes (Fig.
The pigmentation observed in P. bicolor in Yucatan aligns with the morphotype seen in Brazil (
Pseudoceros bicolor A dorsal view B ventral view, sucker, oral pore, pharynx, male gonopore, female gonopore and intestine C detail of tentacular eyes, pseudotentacles D P. bicolor in situ E sagittal section, prostatic vesicle, seminal vesicle, male gonopore, stylet, vagina, female gonopore, female atrium Azan stained.
Yucatan coast, Mexico • 1; Bajos de Sisal; 21.2°N, 90.0°W; 1 m; 22 Feb. 2018; A. Hernández leg.; CRPPY-0029; Quintana Roo coast, Mexico • 1; Mahahual; 18.6°N, 87.7°W; 5.3 m; 17 Mar. 2018; A. Hernández leg.; CRPPY-0036; Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 9.3 m; 21 Apr. 2018; A. Hernández leg.; CRPPY-0056 • 1; Cayo Arcas; 20.2°N, 92.0°W; 5 m; 25 Apr. 2018; A. Hernández leg.; CRPPY-0085.
Pseudoceros rawlinsonae
has been recorded in the Caribbean Sea: Virgin Islands, Honduras, Jamaica, Bahamas, Curaçao; the Gulf of Mexico, Florida (
Body shape elongated with rounded anterior and posterior end, 1.2 cm in length and 0.5 cm in width. Pigmentation brownish yellow to black with scattered white dots (Fig.
Pseudoceros bicolor
and P. rawlinsonae are closely related. Externally, the primary distinguishing features between the two are the prominent white submarginal band and the orange rim that characterise P. rawlinsonae. A study by
Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 3.2 m; 24 Apr. 2018; A. Hernández leg.; CRPPY-0079.
Recorded in Heron Island and One Tree Island, Australia; Madang, Papua New Guinea (
Body shape elongated with rounded anterior margin, tapering posteriorly, 3.5 cm in length and 1 cm in width. Margins slightly wavy. Ground colour velvety black, without any specific additional pattern, but with a small stain, devoid of pigment, present in area of cerebral eyes (Fig.
Within Pseudoceros, P. bolool and P. velutinus (Blanchard, 1847) share several external and internal morphological characters, characterised by a uniform velvety black coloration, without spots, bands, or marginal lines. While both species share several external and internal morphological traits, they can be distinguished by the ventral coloration: P. velutinus has a bluish violet background whereas P. bolool is grey, and by their marginal folds, which are broader in P. velutinus and more subtly defined in P. bolool. The specimen found in the Gulf of Mexico matches the original description of P. bolool. This species has been previously reported from the Australasian region (
Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 5 m; 26 Apr. 2018; A. Hernández leg.; CRPPY-0086.
Cabo Frío, Brazil (
Elongated and elliptical body, 1.5 cm in length and 0.5 cm in width (Fig.
Specimens of Pseudoceros juani from Brazil show a darker colouration, characterised by more abundant and smaller dots distributed along the middle dorsal line. Additionally, the white marginal band with a yellow line is more conspicuous in the Brazilian exemplar. Disparities in the distribution of the dorsal dots and the lighter tones of the final brown band may be attributed to the maturity state of the individuals. Brazilian individuals are longer than those from the Gulf of Mexico.
Campeche coast, Mexico • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 13 m; 14 Sep. 2017; A. Hernández leg.; CRPPY-0014 • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 10.8 m; 14 Sep. 2017; A. Gutiérrez leg.; CRPPY-0019 • 3; Cayo Arcas; 20.2°N, 92.0°W; 4.1 m; 21 Apr. 2018; A. Hernández leg.; CRPPY-0057 • 1; Cayo Arcas; 20.2°N, 92.0°W; 2.2 m; 21 Apr. 2018; A. Hernández leg.; CRPPY-0058 • 1; Cayo Arcas; 20.2°N, 92.0°W; 9.3 m; 24 Apr. 2018; A. Hernández leg.; CRPPY-0075 • 1; Cayo Arcas; 20.2°N, 92.0°W; 4.4 m; 17 Aug. 2018; A. Hernández leg.; CRPPY-0105.
Recorded in Curaçao, Jamaica, Florida, and Honduras (
Elongated and elliptical body, 2 cm in length and 1 cm in width, Dorsal background pigmentation caramel-brown with dispersed, darker tonalities. Small white and black spots scattered across entire surface. Median longitudinal thickening traversed with two large white patches, white median line more visible in anterior ½ of body, especially between patches. One patch situated near pharynx in first 1/3 of body, and second in posterior region of body (Fig.
Yucatan coast, Mexico • 1; Bajos de Sisal; 21.2°N, 90.0°W; 1 m; 22 Feb. 2018; A. Hernández leg.; CRPPY-0030.
Originally described from Naples, Italy (
Body shape elongated with rounded anterior end and tapered posterior end, 1 cm in length and 0.5 cm in width. Velvety, wine-coloured background with a submarginal orange and marginal black band, interrupted at level of the pseudotentacles (Fig.
Specimens of Pseudobiceros splendidus studied show a colouration pattern similar to the specimens from Florida, illustrated in
Quintana Roo coast, Mexico • 1; Mahahual; 18.6°N, 87.7°W; 5.3 m; 17 Mar. 2018; A. Hernández leg.; CRPPY-0113.
Original description from Bermuda (
Body shape elongated with rounded anterior and tapering posteriorly, 1.3 cm in length and 0.7 cm in width. Body margins slightly wavy. Purple-brown background, darker at the margins, with yellow and orange spots outlined by a black circle, and tiny white spots along the entire body margin (Fig.
Campeche coast, Mexico • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 13 m; 14 Sep. 2017; A. Gutiérrez leg.; CRPPY-0021; Quintana Roo coast, Mexico • 1; Mahahual; 18.6°N, 87.7°W; 13.6 m; 18 Mar. 2018; A. Hernández leg.; CRPPY-0043.
Phrikoceros mopsus
was originally described in São Paulo, south-eastern Brazil (
Body shape oval and elongated, with an extremely delicate consistency and a wavy margin, 3 cm in length and 1.5 cm in width. Marginal pseudotentacles deeply folded. Dorsally, with the characteristic small white spots on a caramel brown background, body midline darker. Marginal black rim, interrupted in the distal region of the pseudotentacles. (Fig.
Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 6.3 m; 22 Apr. 2018; A. Hernández leg.; CRPPY-0068 • 1; Cayo Arcas; 20.2°N, 92.0°W; 13.2 m; 24 Apr. 2018; A. Hernández leg.; CRPPY-0076.
The species was described from Naples, Italy (
Body oval-shaped and firm consistency, 1.5 cm in length and 0.5 cm in width. Background colour ranges from brown to yellowish-ochre. Two stripes of paler cream spots, one longitudinally and the other perpendicular, form an inverted cross (Fig.
Thysanozoon brocchii
is noted for the abundance of papillae covering its dorsal surface and its cosmopolitan distribution. Various morphological descriptions with distinct colour patterns occur for different localities (
Discoceloidea Dittmann, Cuadrado, Aguado, Noreña, & Egger, 2019
Cryptocelididae Laidlaw, 1903
Phaenocelis Stummer-Traunfels, 1933
Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 6.2 m; 20 Aug. 2018; A. Hernández leg.; CRPPY-0109 • 1; Cayo Arcas; 20.2°N, 92.0°W; 3.3 m; 20 Aug. 2018; A. Hernández leg.; CRPPY-0112.
Phaenocelis medvedica
was recorded in Brazil (
Body shape elongated with rounded anterior end and pointed posterior end, 0.45 cm in length and 0.2 cm in width. Translucent pinkish colouration, including two longitudinal dark brown rows parallel to the main body’s axis (Fig.
Campeche coast, Mexico • 2; Cayo Arcas; 20.2°N, 92.0°W; 2.2 m; 20 Apr. 2018; A. Hernández leg.; CRPPY-0053 • 1; Cayo Arcas; 20.2°N, 92.0°W; 9.3 m; 24 Apr. 2018; A. Hernández leg.; CRPPY-0077.
Piscadera Bay, Curaçao, Caribbean Sea (
Body shape oval and tapers in the posterior region, 4 cm in length, 2 cm in width. Compact consistency. Milky-translucent colouration, with translucent margin (Fig.
The specimen of the Gulf of Mexico aligns with the description of Phaenocelis peleca provided by
Stylochoplanidae Faubel, 1983
Stylochoplana Stimpson, 1857
Holotype: Campeche coast, Mexico • 1; 9 slides; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 0 m; 11 Sep. 2017; A. Hernández leg.; CRPPY-0013. Paratypes: Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 6.2 m; 20 Aug. 2018; A. Hernández leg.; CRPPY-0110 • 1; Cayo Arcas; 20.2°N, 92.0°W; 3.3 m; 20 Aug. 2018; A. Hernández leg.; CRPPY-0111.
Found in submerged West Keys of Reef Triángulos and Cayo Arcas, Campeche coast, Mexico.
Body shape oval with rounded anterior and posterior end, 10 mm long and 5 mm wide. Whitish translucent colour with a pale brown tonality due to the gut contents. A network of independent intestinal branches, not anastomosing, extends to the body’s margin (Fig.
The name sisalensis is dedicated to the town where the research centre is located, the UNAM campus in Sisal, Yucatán province, Mexico.
Currently, the genus Stylochoplana comprises 25 valid species worldwide. This genus is one of the most species-rich within the order Polycladida and has been divided into different informal groups by several authors (
Stylochoplana sisalensis sp. nov. presents the greatest similarity with S. walsergia from Brazil, S. chilensis from Chile, S. utunomii from Japan, and S. minuta from the Palau Islands (Micronesia). These species are all characterised by the presence of a well-developed and elongated seminal vesicle, as well as an ejaculatory duct that widens and is covered by a prostatic glandular epithelium. All other species in this group present an isolated, more or less elongated and interpolated prostatic vesicle.
On the other hand, the species of Marcus’ Group BI present a very similar female copulatory apparatus directed towards the anterior region and, at the level of the internal vagina, then curving towards the posterior region. In the middle of the female duct, the oviduct opens and the internal epithelium thickens to form Lang’s duct that ends in the rounded Lang’s vesicle.
Stylochoplana sisalensis differs from S. walsergia by the location of the prostate tissue and the shape of the penis papilla. In S. walsergia, the prostatic dilation is included entirely in the penis papilla and surrounded by the male atrium, while in S. sisalensis the penis papilla encloses only 1/2 of the prostatic tissue and the common male duct. This characteristic is shared by S. chilensis, but not with S. utunomii in which the prostate tissue is practically outside the penis papilla, a short protrusion within the male atrium. As in S. suosensis, we lack data on the female apparatus, but we assume that its arrangement is like that of the entire Marcus group B of Stylochoplana.
Holotype: Yucatan coast, Mexico • 1; 18 Slides; Dzilam; 21.5°N, 88.9°W; 13 m; 10 May 2018; A. Hernández leg.; CRPPY-0095. Paratypes: Yucatan coast, Mexico • 4; Dzilam; 21.5°N, 88.9°W; 14 m; 9 May 2018; A. Hernández leg.; CRPPY-0092 • 3; Dzilam; 21.5°N, 88.9°W; 14 m; 9 May 2018; A. Hernández leg.; CRPPY-0093.
Dzilam de Bravo, Yucatan, Mexico.
Body shape elliptical, 10 mm length and 7 mm width. Body constitution solid with a translucent background (Fig.
The species name hartei is dedicated to the conservationist Edward H. Harte, in recognition of his lifelong commitment to environmental conservation and his significant contributions to marine science and the protection of marine ecosystems.
Currently, the genus Emprosthopharynx is composed of nine species: E. gracilis (Heath & McGregor, 1912); E. hancocki (Hyman, 1953); E. heroniensis Beveridge, 2018; E. lysiosquillae Oya, Nakajima & Kajihara, 2022; E. opisthoporus Bock, 1913; E. pallida (Quatrefage, 1845); E. vanhoffeni Bock, 1931; E. onubensis Pérez-García, Gouveia, Calado, Noreña & Cervera, 2024; and E. rasae Prudhoe, 1968. The genus is distributed mainly within the Pacific Ocean, except for E. pallida and E. onubensis, which are native to the Mediterranean, and E. vanhoffeni found in the Cape Verde Islands (
Regarding the internal characteristics between Emprosthopharynx hartei and E. hancocki, in both species, the distal region of the papilla peneal is covered by a series of bristles, thickened, or with pseudosclerotised cilia (the styliform development of the basal membrane mentioned by
The main difference between Emprosthopharynx hancocki and E. hartei is found at the level of the arrangement and shape of the reproductive system. The seminal vesicle and the prostatic vesicle in E. hancocki barely present a small constriction between one organ and another, while in E. hartei the transition between the seminal vesicle and the prostatic vesicle is marked by a tube-like narrowing. In addition, the prostatic vesicle in E. hartei empties through an elongated sinuous extension until the papilla peneal.
The distinction between Emprosthopharynx gracilis and E. hartei lies in the structural and morphological characteristics of the distal region of the male copulatory apparatus. Emprosthopharynx hartei has an elongated prostatic vesicle that leads to a long ejaculatory duct. The duct ends in a papilla peneal surrounded by a flattened atrium. In contrast, E. gracilis is characterised by a rounded prostatic vesicle and a short and robust papilla peneal that opens into a long and deep atrium. Within the female reproductive system, we can observe differences between E. hartei and E. gracilis. For instance, the female atrium in E. hartei is elongated and narrow, whereas in E. gracilis it is short and widened. Additionally, the thickening of the vagina is distinct in the two species: in E. hartei, the thickening is located in the proximal region, while in E. gracilis it is found in the distal region.
Emprosthopharynx hartei sp. nov. A photographed on a black background, where the intestinal branches and pharynx are patent B anterior end with tentacular eyes and cerebral eyes C histological sagittal section (Azan stained) at the level of the reproductive system D sagittal reconstruction of the reproductive system E specimen in situ, showing natural colouration. Abbreviations: ot tentacular eyes, oc cerebral eyes
Notoplana Laidlaw, 1903
Campeche coast, Mexico • 5; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 5 m; 10 Sep. 2017; A. Hernández leg.; CRPPY-0009 • 8; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 0 m; 10 Sep. 2017; A. Gutiérrez leg.; CRPPY-0010 • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 0 m; 11 Sep. 2017; A. Hernández leg.; CRPPY-0012 • 5; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 11 m; 11 Sep. 2017; A. Gutiérrez leg.; CRPPY-0016 • 2; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 13 m; 14 Sep. 2017; A. Gutiérrez leg.; CRPPY-0017 • 1; Cayos sumergidos del Oeste; 20.4°N, 92.2°W; 13 m; 14 Sep. 2017; A. Gutiérrez leg.; CRPPY-0023.
Recorded in Piscadera Bay and Fuik Bay, Curaçao and Virginia Key, Florida (
Body shape elongated and smooth, 0.7 cm in length and 0.3 cm in width. Pigmentation varies from whitish beige to greenish hues. Intestinal extensions well-branched, not anastomosing, extending to body margin, with contents visible due to transparency (Fig.
Heading as above. Male copulatory system comprises an elongated seminal vesicle with two broad sperm ducts proximally (Fig.
Notocomplana Faubel, 1983
Yucatan coast, Mexico • 1; Arrecife Alacranes; 22.4°N, 89.7°W; 1 m; 4 Nov. 2017; A. Hernández leg.; CRPPY-0004; Veracruz coast, Mexico • 6; Veracruz; 19.2°N, 96.1°W; 1 m; 31 Sep. 2018; A. Hernández leg.; CRPPY-0101 • 7; Veracruz; 19.2°N, 96.1°W; 2 m; 1 Sep. 2018; A. Hernández leg.; CRPPY-0103.
The species was described from Jamaica (
Body shape oval with rounded anterior and tapered posterior end, margins pale, wavy, 2.5 cm in length and 1 cm in width. Translucent beige colouration, darker in the middle region of the pharynx (Fig.
Research conducted by
Considering the limited morphological variation between the genera and recent molecular analyses, there is a tendency to propose eliminating the genus Melloplana, as well as the family Pleioplanidae. However, Pleioplanidae also includes other genera like Izmira, Pleioplana, and Laqueusplana, which are poorly known. Therefore, additional molecular analyses and morphological evidence are necessary to confirm the elimination of the family Pleioplanidae (
Gnesioceros Diesing, 1862
Quintana Roo coast, Mexico • 2; Mahahual; 18.6°N, 87.7°W; 7.7 m; 17 Mar. 2018; A. Hernández leg.; CRPPY-0037.
Currently, Gnesioceros sargassicola is known from Bermuda (
Dorsoventrally flattened body, anteriorly widened with a shallow constriction after the tentacles, undulated body margins, and a blunt-tailed posterior end, 0.8 cm in length and 0.3 cm in width. Yellowish grey background colouration with numerous, rounded, orange or brown spots. Narrow elongated pharyngeal pouch with a central oral opening (Fig.
Associated with Sargassum algae.
Idioplanidae Dittmann, Cuadrado, Aguado, Noreña & Egger, 2019
Idioplana Woodworth, 1898
Yucatan coast, Mexico • 3; Dzilam; 21.5°N, 88.9°W; 13 m; 10 May 2018; A. Hernández leg.; CRPPY-0096.
Idioplana atlantica
was originally recorded off St Thomas Island (USA Virgin Island, Caribbean Sea;
Body shape oval, with a rounded posterior end and a more pointed anterior end, 2 cm in length and 1 cm in width. Firm and dense consistency. Background colour ranges from yellowish-white to amber. Dorsally, is covered by dark rings with cream-pigmented inner. The shape of these rings is variable, appearing more rounded anteriorly and elongated posteriorly. Also, the central rings are larger compared to those along the body’s margin (Fig.
Yucatan specimens show a resemblance to those documented by
Although some doubts have arisen about the identity of this species, the presence or absence of pigmentation spots in the preserved specimens of Idioplana described from the Gulf of Mexico does not provide sufficient evidence to confirm the existence of a new species. Therefore, we have decided to classify the Idioplana specimens found in Yucatan as I. atlantica. Further morphological and molecular studies will be necessary to determine whether it represents a distinct species.
Stylochus Ehrenberg, 1831
Yucatan coast, Mexico • 1; Dzilam; 21.5°N, 88.9°W; 9.3 m; 8 May 2018; A. Hernández leg.; CRPPY-0090 • 1; 12 slides; Dzilam; 21.5°N, 88.9°W; 13 m; 10 May 2018; A. Hernández leg.; CRPPY-0099.
The species was originally described in Cape Verde (
Elongated oval body shape, firm and fleshy consistency. In live specimens, body measures 20 mm in length, and 7 mm in width, whereas in fixed specimens, measures are reduced to 10 mm length, and 8 mm in width. Rounded nuchal tentacles visible. Pale beige colouration, translucent near the margins. Small white spots, denser along the midline and a white dotted line along the entire body margin (Fig.
To avoid confusion between Stylochus crassus Verrill, 1893 (from the coast of Maryland, USA) and S. crassus Bock, 1931 (from the coast of Cape Verde Island),
Stylochus sixteni A live animal, photographed on a black background, tentacles B anterior end, marginal, cerebral and tentacular eyes, after fixation C sagittal section through the reproductive system (stained with AZAN) D specimen in situ E sagittal reconstruction of male and female copulatory organs.
Hoploplana Laidlaw 1902
Yucatan coast, Mexico • 1; Dzilam; 21.5°N, 88.9°W; 9.3 m; 8 May 2018; A. Hernández leg.; CRPPY-0089.
Hoploplana inquilina
has been observed off St. Thomas Island, Caribbean (
Body shape oval, 5 mm in length and 3 mm in width. Translucent bluish-to-grey background colour with a distinctive white reticulum on the dorsal surface that does not correspond to the intestine (Fig.
Campeche coast, Mexico • 1; Cayo Arcas; 20.2°N, 92.0°W; 5.3 m; 23 Apr. 2018; A. Hernández leg.; CRPPY-0073 • 1; Cayo Arcas; 20.2°N, 92.0°W; 13.2 m; 24 Apr. 2018; A. Hernández leg.; CRPPY-0080.
Originally described in São Paulo, Southeast of Brazil (
Body shape oval with rounded anterior and posterior end, 1.2 cm in length and 0.6 cm in width. Two small cylindrical nuchal tentacles (Fig.
This study provides a valuable contribution to our knowledge of Polycladida diversity in the southern regions of the Gulf of Mexico. Our research reveals the presence of 27 polyclad species, belonging to 17 genera and 12 families. By revising the polyclad records in the Gulf of Mexico, we have increased the known species count from 31 to 50. It is noteworthy that this is the first known report of marine flatworms along the coasts of Campeche, Yucatan, and Quintana Roo.
This study has identified 17 species that were previously unknown in the Gulf of Mexico, thus expanding their known distribution ranges. Some of the notable findings include the extension of distribution ranges for Enchiridium evelinae, Pseudoceros juani, Phaenocelis peleca, Stylochus sixteni, and Hoploplana divae. Additionally, this study has introduced two new species, Stylochoplana sisalensis sp. nov. and Emprosthopharynx hartei sp. nov. The latter marks the first report of its genus on the Atlantic coast of the Americas.
This work highlights the rich diversity of Polycladida along the Atlantic coastline of Mexico. It also emphasises the importance of exploring and documenting under-researched species, particularly in regions home to abundant fauna. Ultimately, our study contributes to the development of a comprehensive atlas of unrecorded species, which will help to enhance conservation efforts and advance our knowledge of marine biodiversity in the Gulf of Mexico.
We thank Patricia Guadarrama from the Ecology and microscopy laboratory, UMDI-Sisal, UNAM, México; Diana Ugalde and Tonali Mendoza for support on field trip logistics and project administration; to Efrain Chavez and Quetzalli Hernandez for help during SCUBA diving collections; to Pedro Homa, Raul Castillo, and Diana Ugalde for their work on the UNAM-Sisal scientific collection. We thank Gregory Arjona and Danilú Couoh from Aquatic Pathology Laboratory of CINVESTA-IPN Mérida for their help with histology preparations. We also thank the anonymous reviewers for their valuable comments and suggestions, which greatly improved the quality of this manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
Field and lab work was financed by grants to NS by the Harte Research Institute with Harte Charitable Foundation funds and CONABIO-NE018. AHG received CONACyT scholarship number #629884.
DC and AHG took the lead in writing the manuscript, developed the conception of the research, contributed to data interpretation. NS wrote respective parts of the manuscript and acquired funds. CN analysed the samples, wrote parts of the manuscript, and mostly contributed to histological interpretation. All authors edited and reviewed the manuscript.
Daniel Cuadrado https://orcid.org/0000-0001-5565-3023
Alejandro Hernández-González https://orcid.org/0000-0002-1557-5582
Carolina Noreña https://orcid.org/0000-0002-4580-7460
Nuno Simões https://orcid.org/0000-0001-7490-3147
The samples were collected under the collection permit N. PPF / DGOPA-295/17 and PFF / DGOPA-079/19, issued by Mexico’s State Secretariat of Agriculture, Livestock, Rural Development, Fisheries, and Food (SAGARPA). Access to Alacranes Reef National Park was authorised by the Park authorities and we appreciate the possibility of using their facilities in Perez Island.
Whole body specimens and the histological preparations of those specimens under study were deposited at the 'Colección regional de Policladidos de la Península del Yucatán, Mexico' (CRPPY), located at the Multidisciplinary Teaching and Research Unit of Sisal, Faculty of Sciences, National Autonomous University of Mexico (UMDI-Sisal, FC-UNAM). All sequences obtained in the present study have been deposited in the GenBank (Suppl. material
Supplementary information
Data type: docx
Explanation note: table S1. Polycladida of the Gulf of Mexico, updated and with species studied, new records for the Gulf of Mexico and new species added. table S2. GenBank accession numbers of the molecular sequences generated in this study of 28S from Polycladida of the Gulf of Mexico.