Sphoeroides camila Carvalho-Filho, Rotundo, Pitassy & Sazima, 2023

Araujo, Gabriel S., Kurtz, Yan R., Sazima, Ivan, Carvalho, Pedro Hollanda, Floeter, Sergio R., Vilasboa, Anderson, Rotundo, Matheus M., Ferreira, Carlos E. L., Barreiros, João Pedro & Pitassy and Alfredo Carvalho-Filho, Diane E., 2023, Evolutionary history, biogeography, and a new species of Sphoeroides (Tetraodontiformes: Tetraodontidae): how the major biogeographic barriers of the Atlantic Ocean shaped the evolution of a pufferfish genus, Zoological Journal of the Linnean Society 199 (4), pp. 978-993 : 982-986

Sphoeroides camila Carvalho-Filho, Rotundo, Pitassy & Sazima sp. nov.

Figs 3–4 ; Tables 1–2

Zoobank registration: urn:lsid:zoobank.org:act:A7CE6DBB-CFB0-4006-B9F8-F80FEB5E4A01 .

English proposed common name: Southern bandtail puffer.

Portuguese ( Brazil) proposed common name: Baiacu-pinima.

Synonymy (in part): Tetraodon marmoratus : Ranzani 1839.

Sphoeroides spengleri : Jordan and Evermann 1898, Miranda Ribeiro 1915, Carvalho-Filho 1999, Rocha and Rosa 2001 Menezes 2003, Sampaio and Noưingham 2008, Garcia et al. 2015, Pinheiro et al. 2015, 2018, Nóbrega et al. 2015, Gasparini 2017, Guimarães-Costa et al. 2020, Rotundo et al. 2021.

Holotype: MZUSP 126947 (Field number: ACF82020001). 92.2 mm Standard Length (SL), Arraial do Cabo , Rio de Janeiro, Brazil. 22°52ʹS, 42°01ʹW, collected by A. Carvalho-Filho, February 1982 ( Fig. 3 ).

Paratypes (29 specimens): AZUSC 6924 (6 specimens, 79.0– 97.4 mm SL), Praia do Forno ou Praia Grande, Arraial do Cabo, Rio de Janeiro, Brazil, 22°57ʹS, 42°1ʹW, between the surface and 5 m, col. Pedro Hollanda Carvalho and Carlos E.L. Ferreira, 21 Mar. 2015 ; MCZ 12129, 36.5 mm SL, Vigia , Baía de Marajó, Pará, Brazil, 0°50ʹN 48°7ʹW, col. Thayer Expedition to Brazil: Louis Agassiz and party, 28 Feb. 1866 ; MNRJ 7717, 97.2 mm SL, Praia Santa Helena, Vitória, Espírito Santo, Brazil, 20°18ʹS, 40°17ʹW, col. V. Carvalho, 25 Jan. 1945 ; MNRJ 53329 (2 specimens, 103.5–115 mm SL), same data as AZUSC 6924; MZUSP 45607, 85 mm SL, Arquipélago dos Alcatrazes, São Paulo, Brazil, 24°06ʹS, 45°42ʹW, 6 m, col. R.L. Moura, 1 Nov. 1992 ; MZUSP 46633 (2 specimens, 31.1–68.8 mm SL), Porto Belo , Santa Catarina, Brazil, 27°9ʹS, 48°33ʹW, 5 m, col. A. Carvalho-Filho, Jan. 1988 ; MZUSP 49079, 99 mm SL, Ilha dos Frades , Vila de Paramana, Salvador, Bahia, Brazil, 12°58ʹS, 38°30ʹW, 28 Jul.1993 ; MZUSP 81099 (2, 82.3–91.9 mm SL), Arraial do Cabo , Rio de Janeiro, col. A. Carvalho-Filho, Feb. 1982 ; MZUSP 126949 (2, 79.6–103.9 mm SL), same data as AZUSC 6924; NPM 7080 (2, 71.0–79.0 mm SL), same data as AZUSC 6924; UF 19203 (4, 56.4–79.2 mm SL), Praia de Itapoã , Salvador, Bahia, Brazil, 12°57ʹS, 38°22ʹW, col. Virginia Almeida, 7 Jul. 1970 ; USNM 83162 (2, 71.8–104 mm SL), Rio de Janeiro, Brazil, col. C. Wilkes, 1838–1842 ( Fig. 3 ); USNM 104232, 90.4 mm SL, Recife , Pernambuco, Brazil, R. Von Ihering, 1932 ; ZUEC 4957, 84.9 mm SL, Praia da Fortaleza, Ubatuba, São Paulo, Brazil, 23°25ʹS, 45°4ʹW, 0.5 m, col. I. Sazima, 14 Jun. 2000 ; ZUEC 5665 78.6 mm SL, Ilhote , Ilha Bela, São Paulo, Brazil, 23°46ʹS, 45°20ʹW, col. C. Sazima and I. Sazima, 27 Mar. 2002 .

Diagnosis: Sphoeroides camila sp. nov. is distinguished from its congeners by the following combination of characters: pectoral-fin rays 13–14 (rarely 11, 12, 15, or 16), not including a rudimentary one on upper fin sheath; dorsal-fin rays 7–8 (rarely 6); anal-fin rays usually 6–8, rarely 5 (unbranched rays 0–2, branched rays 4–6); dorsum with two pale lappets before opercle, none black; 24–34 lappets on less sagiưal section of body; 1–8 lappets on head before opercle; adults without prickles on dorsum or sides of body (rarely a small patch asser interorbital space), prickles occasionally present in juveniles from interorbital to middle back; very distinct row of black spots on ventrolateral body from chin to end of caudal-fin peduncle, the mid-ones vertically oblong, those on extremities rounded and considerably smaller; no black rounded spots above ventrolateral row ( Fig. 4A, B ); caudal fin with a dark bar at base and a very wide yellow-greenish bar posteriorly. Size up to about 14 cm SL, usually 10 cm SL.

Description: Based on holotype and 29 paratypes. Morphometrics and meristic counts are shown in Table 1 .

Body oblong; head large, 33.3–38.4% of SL, longer in specimens up to 50 mm SL; mouth terminal, upper jaw projects slightly beyond lower; snout conical, long; nostrils paired, nares at the end of the moderate tube; eye moderate, 15–30% on Head Length (HL), (longer in specimens smaller than 50 mm SL); interorbital moderate, 8–14% of HL, concave (more evident in specimens up to 50 mm SL) to somewhat flat; gill slit arch-shaped, slightly shorter than the pectoral-fin base. Dorsal-fin rays 7–8 (rarely 6); anal-fin rays usually 6–8, rarely 5 (unbranched rays 0–2, branched rays 4–6); dorsal and anal fins nearly opposite, base lengths similar; pectoral fin rounded, moderate, 32–41% of HL, with 13–14 (rarely 11, 12, 15, or 16) rays, not including a rudimentary one on upper fin sheath; caudal fin truncate to slightly rounded with 11 rays, one upper and two lower unbranched. Lappets usually in two rows from the dorsum to the anterior dorsal-fin area, one or two pairs before pectoral-fin insertion; ossen one lappet between each of the black blotches of the ventral row; 1–8 lappets on head before opercle; 24–34 lappets on less sagiưal section of body. Prickles on dorsum and ventral areas, more evident in specimens up to 50 mm SL, those of the dorsum ossen absent in adults. Vertebrae 17 [base on X-rays of seven specimens ( Fig. 5 ) and 18 dissected and discarded specimens].

Coloration in life: dorsolateral head and body light reddish to greyish or olivaceous brown, sides and belly whitish. Upper snout pale with one dark irregular area in front of nares; a conspicuous lateral dark band from lower maxilla to eye; usually peppered with black spots and blue ocelli; interorbital dark blotch, ossen with anterior and posterior prolongations, resulting in an almost cross-shaped mark; dorsum with a pair of black blotches, ossen peppered with white spots and iridescent blue circles and/or ocelli ( Figs. 4B , 7 ). A conspicuous lateral row of 11–14, usually 13, black blotches, rounded from chin to pectoral fin, followed by tear-shaped oblong marks from pectoral fin to end of caudal peduncle, the largest blotch at middle body. A light horizontal band is ossen present between dark dorsum and ventral row of blotches. Pectoral and dorsal fins light yellowish to greenish, their bases dark; anal fin usually whitish to yellowish; caudal fin with distinct dark brown to dusky black bar on its base, followed by a relatively thin pale one and a wide, yellowish-green distal bar, usually at least 40% of fin-length. Lappets usually pale, conspicuous, occasionally with sparse dark pigment; one type specimen with dark lappets on upper part of body posteriorly to dorsal-fin base. Short row of black spots above ventral row lacking. Eyes yellow. Preserved specimens with dark dorsum, all dark blotches prominent over whitish to tan ground coloration, blue circles/ocelli absent ( Fig. 3 ). Occasionally a few with sparse to overall dark pigment asser dorsal-fin base. Caudal-fin distal bar dark, grayish, never black.

Etymology: A noun in apposition. The specific name honours Camila Carvalho, one of the daughters of the first author of the new species. We do not use the genitive camilae; this name was inadvertently used in a presentation at an ichthyological meeting.

Distribution: Sphoeroides camila is currently recorded from Northern to South-eastern Brazil; vagrants may occur in the Southern Caribbean.

Natural history: Sphoeroides camila is common in shallow, clear reef waters, up to 20 m deep along the Brazilian coast, occasionally to 70 m depth (AZUSC 6925). It is usually observed alone, in pairs or small groups, and ossen several individuals are found in a small area, hovering over the substrate. It is rarely found in seagrass beds, a behavioural difference between S. camila and S. spengleri , which is typically observed in seagrass plains. It is a generalist zoobenthivore ( Fig. 4 ), but frequently consumes zooplankton and driss material in the water column or at water surface. Diet is composed of crustaceans, molluscs, worms, echinoderms, and zooplankton larvae, which agree with findings for its congener S. spengleri in Florida ( Targeư, 1978). Several individuals of Sphoeroides camila were aưracted to mussels opened by a diver ( Fig. 4D ), which indicates opportunistic foraging, a feature observed for several other pufferfish species (e.g. Duncan and Szelistowski 1998, Kalogirou 2013, Sazima 2019). An individual was observed following a hunting gold spoưed eel Myrichthys ocellatus Lesueur, 1825 and snatching two small crabs flushed by the eel, which again demonstrates its opportunistic foraging. Sphoeroides camila is diurnal, spending the night in reef crevices or partially buried on the adjacent sand/gravel boưom, with the eyes and upper dorsum exposed ( Fig. 4C ).

Remarks

For more than a hundred years, S. camila was identified as S. spengleri . Both are morphologically very similar and, with the Eastern Atlantic S. marmoratus , compose a complex of similar species characterized by a conspicuous lateral row of black marks. These three species have a common ancestor and as a result of geographic isolation evolved into different species. Sphoeroides marmoratus , given an ocean of distance, differs unmistakably from both American species by the presence of a pair of black lappets on the dorsum and a row of distinct spots beneath the eye, characters lacking in its Americas congeners. Additionally, S. marmoratus has 10–12 blotches (usually 11) in the lateral-ventral row vs. 11–14 (usually 12) in American congeners. On the other hand, S. camila differs from S. spengleri , other than in molecular loci, by several characters: the new species presents at least one pair of pale, well-developed lappets on dorsum (usually absent in S. spengleri ); 24–34 lappets on less sagiưal section of body (10– 18 in S. spengleri ); oblong blotches of the lateral-ventral row vertically tear-shaped at midbody (all spots of S. spengleri rounded, seldom horizontally elongated, Fig. 4A, B ); distal bar of caudal fin yellowish-green and covers at least 40% of fin-length (black and covering not more than 30% of fin-length in S. spengleri ). Preferred habitats also differ between the two American species: S. camila prefers hard substrates such as rocky and coralline reefs, whereas S. spengleri selects seagrass beds. Number of vertebrae in both species, 17 (this paper and Tyler 1980). Specimen MZUSP 7695, examined by Shipp in 1969 and identified by him as S. spengleri , actually is the new species described here: presence of two pale lappets on dorsum before the pectoral-fin base; vertically oblong black spots on ventrolateral row before and asser pectoral-fin insertion; absence of rounded black spots above the ventrolateral row; a wide bar on posterior margin of caudal fin, not black and paler than the caudal-fin base bar, undoubtedly characterise a specimen of S. camila .

Figures

Figure 3. (A) Sphoeroides camila holotype (MZUSP 126947), 92.2 mm SL, collected by A.Carvalho-Filho at Arraial do Cabo, Rio de Janeiro, Brazil.(B) Paratype and oldest known specimen of S. camila (USNM 83162), 104 mm SL, collected by C. Wilkes between 1838 and 1842.

Figure 4. Lateral views of (A) S. spengleri and (B) S. camila specimens; red arrows indicate a pair of black blotches on the dorsum of S.spengleri, absent in S. camila. (C) Sphoeroides camila individual partly buried in sandy boưom. (D) Sphoeroides camila individuals eating mussels opened by a diver, an opportunistic foraging behaviour elicited by human interference. Figure (A) by Robertson & Van Tassell (2019). Figure (C) by Peter Wirtz.

Figure 5. X-rays of selected specimens of S.camila and S. spengleri. (A) Sphoeroides camila (AZUSC 6924). (B) Sphoeroides camila (AZUSC 6925). (C) Sphoeroides camila (USNM 83162). (D) Sphoeroides spengleri (USNM 104232).

Figure 7. Diversity of the genus Sphoeroides. On the less, yellow line represents Sphoeroides species from the Western Atlantic (WA); green line represents the only species from Eastern Atlantic (EA), S. marmoratus. On the right, pink line represents species of the former Colomesus genus; blue line represents species of the Eastern Pacific.Lower centre, S. pachygaster.