Mobula yarae, Bucair & Marshall, 2025

Bucair, Nayara, Hinojosa-Alvarez, Silvia, Marshall, Andrea Denise, Pate, Jessica, Francini, Carlo Leopoldo Bezerra, Garrido, Amana Guedes, Capel, Katia Cristina Cruz, Loboda, Thiago Silva, Monteiro, Jhonatas Sirino, Bruno, Carlos Eduardo Malavasi, Vaga, Claudia Francesca, Dove, Alistair D. M., Hoopes, Lisa A., Perry, Cameron & Kitahara, Marcelo V., 2025, An integrative taxonomy investigation unravels a cryptic species of Mobula Rafinesque, 1810 (Mobulidae, Myliobatiformes), from the Atlantic Ocean, Environmental Biology of Fishes 108, pp. 1-35 : 10-26

Mobula yarae sp. nov. Bucair & Marshall.

( Figs. 4 , 5 , 6 , 7 , 8 , 9 , 10 , 11 , 12c and f , 13 , 14 , 15 , 16a and b , and 17 ).

urn:lsid:zoobank.org:act:1B93E682-1D6A-45BA-87FF-81BAE5BC791D.

Manta birostris . - Bigelow and Schroeder (1953), p. 482 (footnote 62), 483, 484, 502, 503, 504, 505, 506, 508, figs. 111, 112, 116 (except F, G and H), and 117, morphological description of body parts of the specimen MCZ 37005. Ari (2014), p. 181 - 185, figs. 1 and 2 (identified as Manta 1). Ari (2015), p. 407, 408, 412, fig. 5 (identified as Manta 1).

Manta sp. cf. birostris . - Marshall et al. (2009), p. 1, 3, 12, 22-25 figs. 8, 16. White et al. (2018), p. 51.

Mobula cf. birostris . Stewart et al. (2018b), p. 1-6, fig. 2. Stevens et al. (2018a), p. 110-111, figures of Caribbean manta ray 1 and 2, and distribution map. Pate and Marshall (2020), p. 51-53, 55-58, 62, figs. 1, 4c-f, and 6. Bucair et al. (2021a), p. 1, 3-10, figs. 1c and e, 3b, 6c, and 7. Bucair et al. (2021b), p. 1, 3-7, 9, figs. 2c, 3, and 4. Medeiros et al. (2021), p. 203, 204, 208, 210, 213, 215, fig. 2. Medeiros et al. (2022), p. 775-776, 778-782, figs. 1, 2a,b,c. Bucair et al. (2023), p. 1611-1615, fig. 1. Bucair et al. (2024b), p. 2-4, 6-8, figs. 2, 4, 5d and e. Pate (2024), p. 5, fig.1.

Mobula birostris . Hinojosa-Alvarez et al. (2016), p. 1-16, figs. 4e-g, 5, 6, 7 (referred to as Yucatán Manta Ray). Hosegood et al. (2020), p. 4784-4786, 4788-4792, figs. 1, 2, 3, 4 (referred to as putative new Mobula species).

Mobula sp. cf. birostris . White et al. (2018), p. 65 .

Holotype

Female 248 cm DW found on July 19th, 2017, at Pompano Beach , Florida, USA (26°14′3″N, 80°07′19.20″W), is housed in the Vertebrate Zoology, Fishes Division of the Smithsonian Institution ( USNM collection number: 443866 ), hereafter identified as manta M10. The specimen was preserved in its integrity, except for the caudal bulb and tooth band, which were photographed, but lost before storage. The complete mitogenome sequence and the partial nuclear genome of the holotype are deposited in GenBank (accession numbers PV339738 and PRJNA1147742 respectively) .

Type locality

Pompano Beach (26°14′3″N, 80°07′19.20″W), Florida, USA.

Paratypes

Parts of a juvenile female (manta M1—chondrocranium, mandible, vertebrae, dorsal and ventral dermal denticles, and gill plates), a fairly well-decomposed carcass, found stranded on April 5th, 2019, at Ilha Comprida , São Paulo, Brazil, are housed in the Fish Collection of the Museum of Zoology of the University of São Paulo ( MZUSP collection number: 129738 ). Tissue samples, dermal denticles, and gill plates from the same specimen are also frozen at the Biological Collection “Prof. Edmundo F. Nonato” of the Institute of Oceanography of the University of São Paulo (ColBIO collection numbers: FISH00213). The complete mitogenome sequence and the partial nuclear genome of this specimen are deposited in GenBank (accession numbers PP316123 and PRJNA1147742 respectively) . Chondrocranium of a juvenile female (manta M12) captured at Fernando de Noronha Archipelago (3°51′13″S, 32°25′25″W), Brazil, is housed in the Fish Collection of the Museum of Zoology of the University of São Paulo (manta M12— MZUSP collection number: 129740 ). Fragments of skin, including the dermal denticles of a juvenile female (manta M3) captured at Natal (5°47′40″S, 35°12′40″W), Brazil, are housed in the Fish Collection of the Museum of Zoology of the University of São Paulo ( MZUSP collection number: 129739 ). The mitogenome sequence of this specimen is deposited in GenBank (accession number PP329119 ) .

Additional material

Measurements and meristic analyses of juvenile male (manta M9) collected in Bimini, Bahamas, originally identified as Mobula birostris ( Bigelow and Schroeder 1953) . This specimen’s body parts, including the head, dermal denticles, tail, dorsal and pelvic fins, are housed in the Harvard Museum of Comparative Zoology (MCZ 37005). Measurements, tooth band and skin of a female from Florida (manta M11); and mitogenomes of living specimens, including a juvenile male (manta M2) sampled at Fernando de Noronha, Brazil, and three adult females (manta M4, M5 and M6) (Isla Cotoy and Cabo Catoche, Mexico).

Etymology

Named after a Brazilian legend in which Yara (Iara or Uiara), a beautiful Amazonian indigenous and excellent warrior, is thrown into the water as a punishment. Yara, saved by the fishes, assumed a chimerical form (part fish and part woman), empowered with enchantment. Of dazzling beauty, the noun Yara, in the indigenous Tupi mythology, means “mother of water” or “mãe d’água” in Portuguese. The specific epithet, yarae , is in reverence to “Yara”, mother of the waters, from ancient Tupi and Guaraní mythology.

Suggested common names

Atlantic manta ray.

Diagnosis

Mobula yarae sp. nov. is a large-sized mobulid, with terminal mouth and tooth band only in the lower jaw, like the closest congeners— M. alfredi and M. birostris (vs. M. eregoodoo , M. hypostoma , M. kuhlii , M. mobular , M. munkiana , M. tarapacana , and M. thurstoni , with subterminal mouth), and is characterised by the following combination of characters: small cusped teeth in 9–13 transversal rows in the tooth band (vs. 12–16 rows in M. birostris and 6–8 rows in M. alfredi ); filter lobes positioned anterior to branchial filaments, laterally fused and with transverse projected bristles; disc width 2.2 times larger than disc length; ellipsoidal calcified mass posterior to the dorsal fin with a longitudinal length of 4.3% of DL (vs. 6% DL in M. birostris ), with an embedded residual serriform caudal spine in the posterior portion facing upwards (vs. facing backwards in M. birostris ); stellate-shaped dermal denticles, ramifications with non-bifurcated tips, arranged in raised ridges longitudinally aligned in dorsal and ventral body surfaces (vs. cross-shaped dermal denticles with bifid cusps arranged randomly in M. birostris and stellate base denticles with the tips of the star base bifurcated and arranged uniformly in M. alfredi ); and dorsal surface predominantly black, with symmetrical white supra-branchial patches diagonally aligned.

Morphometric description

Morphometric comparisons are based on juvenile specimens due to the limited availability of adult specimens examined. While this introduces certain limitations, it reflects the best available data and follows established precedent for elasmobranch descriptions under similar constraints.

In addition to measurements of a preserved chondrocranium (M12), measurements from four other specimens (M1, M9, M10 and M11) were evaluated. Due to the absence of pectoral fins in large mobulid landings, carcasses, and the integrity of preserved specimens, the body morphometric proportions were calculated in relation to the disc length (DL). Table 3 shows the morphometrics evaluation of M. yarae sp. nov. Additionally, the head is dorsally flat, with the margin of the rostrum concave, with broad sides towards the origins of cephalic fins. When unfolded, the cephalic fins meet in the centre of the mouth. The mouth opening is 62.67% of the head width, and the tooth band, present only in the lower jaw, comprises a length of ~ 70% of mouth width. The distance between the nostrils totals 92% of mouth width. Spiracles are located above the pectoral fin insertion; the distance between them is ~ 85% of head width. The whip-like tail is thicker at the base near the dorsal fin.

Morphologic description

The skin is covered with stellate-shaped dermal denticles with high ridges that do not overlap and are longitudinally aligned with the body disc length orientation on the dorsal and ventral surfaces. The dermal denticle structure of Mobula yarae sp. nov. ( Fig. 4 ) is different from that of M. birostris and M. alfredi (see Marshall et al. 2009 for comparison). In general, the structure and shape of the dermal denticles of Mobula yarae sp. nov. are similar on the ventral and dorsal body surfaces. However, the ventral denticles are larger and more spaced. The distribution of the dermal denticle is also denser in the dorsal, pelvic fins and trailing edge of the pectoral fins, close to the pelvic fins. Variations in size and number of ramifications of the dermal denticles have also been observed ( Fig. 4b ).

Similarly to Mobula birostris , a calcified cartilaginous mass with an embedded caudal spine is located posteriorly to the dorsal fin. The mass is covered with a skin layer with dermal denticles (similar to those from the rest of the body’s dermal surface) but lacks the collagenous connective tissue to the tail (as detailed by Marshall et al. 2009). As in M. birostris , only a small part, the tip, of the residual caudal spine projects out of the calcified mass in Mobula yarae sp. nov. ( Fig. 5 ), which is similar in structure to that found in Mobula mobular (dorsally flat and sawed sideways), the only species of the genus to have a functional spine. The calcified structure is ellipsoidal and slightly dorsally flattened, with the tip of the spine facing upwards ( Fig. 5b, d ) and not backwards, as in M. birostris ( Marshall et al. 2009) . The anterior portion is thinner, while the posterior portion presents a sharp slope towards the tail. Unlike M. birostris , the embedded caudal spine of Mobula yarae sp. nov. is inclined within the calcified mass, and although reduced, it has lateral serrations, with the protrusions more well-developed in the internal part of the calcified mass ( Figs. 5d and 6b ). The dorsal surface of the embedded spine is slightly concave, with at least three longitudinal dorsal grooves ( Fig. 6 ). The ventral face of the spine is convex with the presence of a well-developed longitudinal ventral ridge at its base that supports the spine ( Fig. 6 ). In addition to the juvenile specimens examined in this study, the caudal bulb was observed in adults in the wild.

Like in other Mobula species, the teeth in Mobula yarae sp. nov. are arranged in narrow bands alongside the edge of the jaw, composed of a variable number of rows (9–13) and files (170–230; Fig. 7 ). The total count of cusped teeth varies from 1680–2811 ( Table 4 ). The upper jaw lacks tooth bands. It is important to note that the number of tooth rows has been considered a taxonomic character (e.g. Bigelow and Schroeder 1953). Although the overall tooth feature is changeless, the total number of teeth and files increases with the individual’s growth.

The spiracles are small oval openings located on the anterior edge of the white supra-branchial patch, just behind the eyes, above the pectoral fin insertion ( Fig. 8 ). Branchial slits are arranged in a branchial arch, which contains several filter lobes on the anterior part ( Figs. 9a and b ), as well as the branchial plates in the posterior part of the arch ( Fig. 9a ). The filter lobes are laterally fused ( Fig. 9c ), uniformly light-grey, and their terminal lobe is rounded ( Fig. 9b ) and greyish-white in colour. Morphologically, the filter lobes have transverse bristles—lobelets—along the face towards the pharynx, rendering an even finer filter lobe. In an enlarged view ( Fig. 9d, e ), the base of the lobelets comprises small fibrils scattered irregularly (in part described and illustrated by Bigelow and Schroeder 1953, p. 482, figs. 111 and 112).

The large chondrocranium ( Fig. 10a ) contains a substantial brain and a large number of blood vessels and arteries ( Fig. 10b, c ) in the brain cavity (BC) just below the fontanelle (F), as described for other Mobula species (Schweitzer and Notarbartolo ‐ di ‐ Sciara 1986; Alexander 1996; Ari 2011). The uncertainty regarding the tangle of vessels’ functionality remains, although the hypotheses of brain temperature regulation would allow the species to be resilient to environmental adversities.

The siphonal stomach (J-shaped) presents longitudinal folds responsible for absorbing food as well as enlarging the stomach when full, on its cardiac portion, increasing the surface area for digestion. Between the stomach and intestine, there is the pylorus sphincter, which controls the digestion flux. The intestine is divided into three segments: the initial portion is the duodenum; followed by the iliumspiral valve intestine (internally coiled, increasing the surface area for absorption); and then the colon, which ends with the insertion of the rectal gland ( Fig. 10d–f ).

Lastly, the sensory units pattern outlined by Notarbartolo-di-Sciara (1987) for devil ray species was also observed in M. yarae sp. nov. and is composed of two different systems: large pores widely spaced, oval in shape with its major axis antero-posteriorly oriented ( Fig. 11a ), and dense tiny pores ( Fig. 11b ). The presence of a viscous liquid in the pores was noted ( Figs. 11c and d ). Pit organs (lateral line and Lorenzini ampullae) are scattered over the skin in species-specific patterns ( Peach and Marshall 2009); in M. yarae sp. nov., they occur mainly on the head (dorsal, ventral, around the eye and cephalic fins). It was not possible to observe the density or distribution of pores in the pectoral fins (which is quite significant in the design of the sensory structure of batoids, see Meyer and Seegers 2012) due to the lack of the body part on the examined specimen.

Colouration pattern

A wide range of colour variation has been observed in this species, which may overlap with characteristics of M. birostris and M. alfredi . Thus, caution must be taken when considering colouration as a standalone feature to identify Mobula yarae sp. nov. Overall, although the dorsal body surface of the Mobula yarae sp. nov. is predominantly black as in M. birostris , it does have significant differences. The white supra-branchial triangle marks align diagonally with the opposite symmetric mark (not parallel as in M. birostris , Fig. 12a ) towards the posterior edge of the symmetrical patch on the opposite side ( Fig. 12c ). Also, the white marks on the tips of the pectoral fins have a protruding colour on the posterior edge, and the white “V” shaped mark located anteriorly to the dorsal fin is larger, reaching the middle point of the body where it extends sidewards in a straight line (white or light grey) to the tips of the pectoral fins.

The ventral colouration of Mobula yarae sp. nov. ( Fig. 12f ) is very variable in the regular colour spectrum and can be predominantly light or dark. A grey shadow on the trailing edge of the pectoral fin may be prominent or almost imperceptible. The dark spots and patches may be present on the abdominal region and pectoral fins, but not between the gills as in M. alfredi . The posterior 5th-gill cover shade is quite diminished compared to both M. birostris and M. alfredi , or may even be absent. The face colouration is also lighter (including regions around the eyes and mouth, and anterior to the 1st-gill slits). The dorsal and ventral colour variation in the type series can be observed in Fig. 13 , as well as the variation observed in the regular colour range in nature in Fig. 14 .

Although there is considerable intraspecific variation in the colouration of the new species, the analysis revealed specific colouration traits that meet the> 95% or <5% thresholds necessary to be used as distinction characters between M. birostris and M. yarae sp. nov. In M. birostris , the alignment angle of the white supra-branchial patches (criteria 4 and 5, Table 5 ), which extends medially from the spiracle, turns ~ 90° posteriorly and remains roughly parallel, not touching the opposite triangular white patch. This forms a “T” shape within the black area in the middle of the white supra-branchial colouration. In contrast, in M. yarae sp. nov., the white markings extend from the spiracles at an obtuse angle, continuing diagonally towards the opposite supra-branchial patch, rarely touching it (11.8%). In the new manta ray species, these white patches form a “V” shape within the black colour in the middle of the white patches. Mobula birostris generally exhibits sharper and more contrasting transitions between the black and white colouration (100%) compared to M. yarae sp. nov. (20.5%), which typically displays softer transitions. Mouth colouration in M. birostris was dark in 98% of the specimens evaluated, while in M. yarae sp. nov., it was light-coloured (96.7%). Furthermore, 99% of M. birostris specimens exhibited uniform, dark, V-shaped shading along the trailing edge of the ventral surface of the pectoral fins. In contrast, 64.5% of the new species displayed grey pectoral fin shading, which was not uniform, as observed in M. birostris . Table 5 highlights the results of each criteria evaluated.

The species description and colour variation explored herein were based on the regular colour morph range of Mobula yarae sp. nov., in which lighter and darker colours were observed in both juvenile and adult individuals. However, as with M. alfredi and M. birostris , melanistic and leucistic individuals may occur. It is important to note that manta ray specimens with a greyish (carbon) tone ( Fig. 15 ) covering almost the entire pectoral fins, gills, and face, have been observed, presumably due to an excess of melanin content ( Fig. 15 ). Nevertheless, further studies, including genetic evaluations, are required for a more comprehensive understanding of the variability of colour morphs in Mobula yarae sp. nov., as some variants may overlap with those of closely related species.

Primary geographic distribution and threats

Our dataset highlights the species occurrence from the northeastern USA to southeastern Brazil, including the Gulf of Mexico, the Caribbean islands, the Amazon continental shelf and estuaries, and the oceanic systems of Saint Peter and Saint Paul and Fernando de Noronha archipelagos. The M. yarae sp. nov. records in estuaries, oceanic islands, and coastal regions suggest an association with high-productivity areas. The distribution map ( Fig. 16 ) highlights the species’ effective occurrence in the Western Atlantic Ocean. However, speculative occurrences were observed and debated for the Eastern Atlantic but require further analysis and genetic evaluation for accurate conclusions.

Remarks

Considering the cephalic fins as the synapomorphic character of the genus Mobula , Mobula yarae sp. nov. represents a new species of mobulid, recognised to date only from the Atlantic waters in both neritic and oceanic environments. Differently from M. tarapacana , M. mobular , M. thurstoni , M. hypostoma , M. eregoodoo , M. munkiana , and M. kuhlii , all of which have their mouth positioned sub-terminally, Mobula yarae sp. nov. has a terminal mouth (oriented forward), like M. birostris and M. alfredi . Indeed, M. yarae sp. nov. is also similar to M. alfredi regarding their light-coloured or even white face (including regions around the eyes and mouth, as well as anterior to the 1st-gill slits), small or even absent dark mark posterior to the 5th-gill slits, large cephalic fins, and filter lobes laterally fused, with rounded terminal lobes displaying transversal lobelets. Nevertheless, they differ in the pectoral fin shape and curvature angle, the absence of a calcified mass located posterior to the dorsal fin in M. alfredi , position of the ventral dark spots and patches (absent between the gill slits in M. yarae sp. nov.), and dermal denticles shape. Regarding the sympatric species M. birostris , M. yarae sp. nov. also shares several similarities such as pectoral fin shape; large cephalic fins; presence of a calcified mass posterior to the dorsal fin with an encapsulated residual spine; and overall characteristics of filter lobes. However, differently from M. birostris , M. yarae sp. nov. displays a different supra-branchial patch pattern, white markings on the pre-dorsal fin region and dorsal tips of the pectoral fin, semblance of ventral posterior edge of the pectoral fin mark, posterior 5th-gill cover shade, position of the ventral dark spots and patches (concentrated on the abdominal region in M. birostris ), and dermal denticles shape. The morphometric comparison among the three manta ray species also highlights differences between them ( Table 6 ).

Historically, once Lacépède (1803) original description of Raja giorna mentions “two large appendages on the front of the head; each pectoral fin forming an isosceles triangle, the base of which is attached to the body of the fish; a dorsal fin placed in front of a strong spine, serrated on both sides, which terminates the body; the tail very long, very loose, and devoid of fins”, we can discard the possibility that R. giorna might be M. yarae sp. nov. Furthermore, the potential synonymous status of M. yarae sp. nov. with Raja giorna Lacépède 1803 , Raja fabroniana Lacépède 1800 , and Pterocephalus massena Swainson 1838 were disregarded due to the mouth position (subterminal) and mention of one or two barbs in their species descriptions, which were later accepted as synonyms of Mobula mobular (see White et al. 2018; Notarbartolo-di-Sciara et al. 2020). However, Cephaloptera giorna Lesueur 1824 was described as having a terminal mouth (different from the subterminal mouth of the other Cephaloptera Cuvier 1817 or Cephalopterus Risso 1810 ) and was subsequently grouped in the genus Ceratoptera Müller and Henle 1837 (illustration 20 on Swainson 1838). Although the overall description is somewhat compatible with M. birostris or M. yarae sp. nov., the name is unavailable since Lesueur (1824, p. 119–121) adopted it after Lacépède’s Raja giorna , where the author states: “I am moreover inclined to think that they are no other than the Raja giorna, Lacep. v. 20. 3, to which Cuvier refers to the R. fabroniana, Lacep. vii. pl. 5, f.3; … I have adopted for this species the name of the celebrated Giorna”. The comparison with the description of Cephalopterus vampyrus Mitchill 1824 is, in part, in agreement with that from M. yarae sp. nov. (i.e., terminal mouth, dark dorsal body surface, presence of a knob posterior to the dorsal fin); however, the measurements and proportions of the body (DL= 326 cm, DW = 487 cm, mouth width = 83.82 cm, distance between eyes = 127 cm, distance between the 1 st and 5th gill openings were 50.8 cm and 114.3 cm respectively) differ greatly from Mobula yarae sp. nov.

A large similarity is noted to the description of a “fish of extraordinary dimension” (male 4.27 m of DW), Manta americana Bancroft 1829 (no type known), based on a specimen captured on May 8th in Jamaica, which highlights: the head “furnished at each extremity with a tongue-shaped lobe”, terminal mouth, dorsal colouration “brownish black, changing into a dark grey over the pectoral fins, which became lighter towards their anterior edges and on the side of the thorax and eyes”, ventral colouration “generally very white, but over the abdomen were several spots of irregular shapes and sizes” cephalic lobes “white, except at their inner and anterior edge, which was bordered with a broad black stripe proceeding from the upper jaw”, “a knob on the root of the tail”. In a footnote the author states that “the name here given must be regarded as synonymous with the Ceph. Giorna of Le Sueur (Journ. Acad. Nat. Sci. Phil. iv. 100.)” … which “corresponds in almost every particular”. The features detailed by Bancroft (1829) closely resemble the characteristics of M. birostris , but some features are also in agreement with M. yarae sp. nov. The measures and metrics coincide in many aspects but overlap in the range between the two species, making it impossible to distinguish between M. birostris and M. yarae sp. nov. based on the original description and the non-existence of a type.

Lastly, the description of Cephaloptera elliotti Holmes 1856 (no type known), a large fish accidentally captured near Sullivan’s Island in June 1854, refers to a manta ray description, with 12–15 rows of teeth. The size and illustration of the knob posterior to the dorsal fin differs from the characteristics of M. yarae sp. nov. See the complete taxonomic evaluation undertaken in this study in the Online Resource 1 (Appendix 2).

Figures

Fig. 4 Dermal denticle of Mobula yarae sp. nov.: a general view of skin denticle configuration and alignment, highlighting apical and lateral view of isolated dermal denticles, specimen from Rio Grande do Norte, Brazil (M3); b dermal denticle size and ramifications array, specimen from Florida, USA (M11); c dermal denticles of a specimen from the Bahamas (M9, MCZ 37005, © President and Fellows of Harvard University); and d dermal denticles of the holotype (USNM 443866)

Fig. 5 Calcified cartilaginous mass at the tail base and embedded residual spine details of Mobula yarae sp. nov. (Brazilian specimen—M1, 47 mm length, 22 mm broad, and 14 mm high): a calcified mass attachment with missing ligaments or connec- tions between the bulb and the tail structure; b different angles of the bulb and sting; c CT scan of the calcified mass on lateral view; d serrated barb outer tip

Fig. 6 CT scan image focusing exclusively on the serrated caudal spine of Mobula yarae sp. nov. (Brazilian specimen – M1): a lateral view, underlining the well-developed ventral ridge (VR) and the caudal spine tip (CTS, the outer part of the spine on the calcified mass); b tilted lateral view, highlighting the lateral serrations (LS); and c dorsal view, accentuating the dorsal grooves (DG) of the embedded part of the spine

Fig. 7 Dental band and teeth of the Mobula yarae sp. nov.: a entire tooth band (Brazilian specimen— M1), tooth alignment in specimens from b the USA (holotype—M10) and c the Bahamas (MCZ 37005, © President and Fellows of Harvard University—M9); d, e enlarged view of the teeth formats from the same specimen in (a)

Fig. 8 Spiracle size and position of Mobula yarae sp. nov. The yellow arrows indicate the location of the structure. a, c Records from Fernando de Noronha, Brazil, and b Ceará, Brazil. Photo credits: a Leo Veras, b Jones Santander-Neto, and c Bruno Galvão

Fig. 9 Mobula yarae sp. nov. gill plate. a Complete branchial arch (GF: gill filament, FL: filter lobe, and TL: terminal lobe of filter filament); b anterior view of filter lobe; c enlarged view of filter lobe (BT: bristles, RL: ridgeline, and LF: lateral fusion of filter lobe); d enlarged view of the filter lobe highlighting the transverse lobelets; and e the fibrils on its base. a, d, e Specimen from southeast Brazil (M1) and b, c) specimen from Fernando de Noronha, Brazil (M12)

Fig. 10 Organs and structures of Mobula yarae sp. nov.: a chondrocranium, highlighting the brain cavity (BC) and fontanelle (F) (specimen from Fernando de Noronha, Brazil, MZUSP 129740, M12); b tangle of vessels in the brain cavity; c brain (BR) and tangle of vessels; d digestive system, highlighting the J-shaped stomach, pylorus contraction, and intestine; e oesophagus (ES), the cardiac portion of the stomach, and the pylorus sphincter; and f small duodenum portion (DU), the spiral intestine, and the intestine’s last portion, the colon (CL). Integral structures are represented in the smaller rectangle in (e and f). b–f organs from the (M1)

Fig. 11 Sensory units pattern. Non-uniform pores with viscous liquid distributed over the body surface of the Brazilian type specimen (M1) of Mobula yarae sp. nov.: a larger and b smaller pores on dorsal surface; c, d larger pores with the viscous liquid on both, c dorsal and d ventral surface (close to the eye’s region)

Fig. 12 Distinct colouration of manta ray species. Dorsal and ventral colouration pattern of Mobula birostris a, d, Mobula alfredi b, e, and Mobula yarae sp. nov. c, f. Photo credits: Leo Francini a; Guy Stevens | Manta Trust b, e; Rawany Porfilho c; Mauricio Andrade d; and Nayara Bucair f

Fig. 13 Colour morphs of the type series: a, b Atlantic coast of Florida (USNM 443866, GenBank accession number PV339738); c, d southeastern Brazil (MZUSP 129738, Gen- Bank accession numbers PP316123); and e, f northeastern Brazil (MZUSP 129739, GenBank accession numbers PP329119)

Fig. 14 Colour morphs variation on the regular spectrum of Mobula yarae sp. nov. Dorsal (column a and b) and ventral (column c and d) colouration pattern disparity. Photo credits/sources: Projeto Mantas do Brasil and Marine Megafauna Foundation database

Fig. 15 Manta rays with greyish-carbon colouration pattern: a, c, f specimens from Mexico, and b, d, e from Brazil. Photo credits/ sources: a Guy Stevens | Manta Trust; b, d, e Projeto Mantas do Brasil database; and c, f Marine Megafauna Foundation database

Fig. 16 Primary distribu- tion of Mobula yarae sp. nov. based on effective occurrence in the Atlantic Ocean

Fig. 17 Fresh (top) and dried (bottom) gill plates of Mobula birostris (right) and Mobula yarae sp. nov. (left). Photo credits/sources: a, b Projeto Mantas do Brasil database, c Thomas P. Pechak, and d Paul Hilton