The family Aeolidiidae Gray, 1827 (Gastropoda Opisthobranchia) from Brazil, with a description of a new species belonging to the genus Berghia Trinchese, 1877 Author Domínguez, Marta Author Troncoso, Jesús S. Author García, Francisco J. text Zoological Journal of the Linnean Society 2008 2008-06-30 153 2 349 368 https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2008.00390.x journal article 3188 10.1111/j.1096-3642.2008.00390.x 21e17d7f-23fe-4244-bb55-736a89096f28 0024-4082 4687689 BERGHIA MARCUSI SP. NOV. Berghia coerulescens Marcus, 1957: 477–481 , figs 237, 240, 241, 245. Type material: Holotype : one specimen, 12-mm long, 0-m depth at Praia de Armação ( Buzios , Rio de Janeiro ), 22 June 2000 , deposited in the Museo de Zoologia de la Universidad de São Paulo , Brazil (catalogue number MZUSP 64150 ). Paratype : one specimen, same data as for holotype, 11-mm long, deposited in the Museo de Zoologia de la Universidad de São Paulo , Brazil (catalogue number MZUSP 64151 ) . Other material: Praia dos Ossos (Buzios, Rio de Janeiro ), 21 March 1998 , two specimens, 0-m depth, 5–7-mm long . Praia de Armação (Buzios, Rio de Janeiro ), 22 March 1998 , one specimen, 0-m depth, 16.6-mm long . Praia de Armação (Buzios, Rio de Janeiro ), 23 March 1998 , one specimen, 0-m depth, 7-mm long . Praia dos Ossos (Buzios, Rio de Janeiro ), 26 March 1998 , two specimens, 0-m depth, 6–7-mm long . Praia dos Ossos (Buzios, Rio de Janeiro ), 27 June 1999 , one specimen, 0-m depth, 11-mm long . Praia de Armação (Buzios, Rio de Janeiro ), 25 June 2000 , one specimen, 0-m depth, 13-mm long . Etymology: This species is named after Ernst Marcus, who first described these specimens. Marcus contributed considerably to the knowledge of the Brazilian opisthobranchs. External morphology: The general body colour is whitish. The oral tentacles are elongate, their bases are translucent with orange pigment, and the apical two-thirds of each is white ( Fig. 1F ). The rhinophores bear papillae on their posterior faces. Some specimens have papillae arranged in oblique rows on the anterior and lateral faces. Approximately, the apical twothirds of the rhinophores are white–cream, and the bases are reddish. An orange band is on either side of the head, between the oral tentacles and rhinophores. The cerata are organized into between five and seven pairs of groups ( Fig. 6A ). The cerata have translucent tips, below which there is an orange ring, and the digestive gland is brown. Below the orange ring there is a whitish area that is more visible in some specimens. The cerata are arranged in horseshoe-shaped arches ( Fig. 6B ). The cerata of the middle part are larger than those at the ends. The first group contains ten or 11 cerata, and the number diminishes towards the tail. The posterior groups consist of two or three cerata, not in arches. There are oblique bright orange lines on the borders of the insertions of the cerata, especially on the first group. Near the cerata the dorsum is translucent white, and extending posteriorly from the base of the rhinophores there is an undulating opaque white band. This band may be broken, or can extend uninterrupted to the tip of the tail. On the largest specimens the dorsal band is broken on the pericardium, and there are some spots of the same colour on this area. The genital opening is among the cerata of the anteriormost group on the right. The anus is located below the second right arch. The foot is translucent, with elongated foot corners curved posteriorly ( Fig. 1G ). Internal morphology: The radula is uniseriate with between ten and 12 bi-arched teeth ( Fig. 7A ). Each tooth has a triangular median cusp with a very fine denticle on each side (in some teeth it only appears on one of the sides). In a 5-mm-long specimen the median teeth have 17–21 denticles on each side ( Fig. 6C ), and the smallest tooth has 12–13 denticles on each side ( Fig. 6D ). In an 11-mm-long specimen the smaller teeth have 19–23 denticles on each side, and the larger teeth have 24–28 denticles. Some specimens have radular teeth with denticles that are piled up ( Fig. 6E ), or with denticles ending in two or more tips ( Fig. 6F ). The masticatory border of each jaw plate shows a long row of small rounded denticles. ( Fig. 7B ). Remarks: The presence of the European species B. verrucicornis and B. coerulescens in the Western Atlantic has been an object of discussion. The Western Atlantic specimens of Berghia were assigned to a very variable species, B. coerulescens ( Engel, 1925 ; Marcus, 1957 ; Edmunds, 1964 , 1966 ; Marcus, 1976 ), although some of them were re-examined and subsequently assigned to B. verrucicornis by Tardy (1962) . However, several authors have doubted the amphiatlantism of these Berghia species, and have recognized the need for future investigations to clarify this question ( Edmunds, 1968 ; García-Gómez & Thompson, 1990 ; Muniain & Ortea, 1999 ). The study of the specimens from the Brazilian coast allows us to observe several characters, mainly related to the coloration, that differ from the specimens of the Mediterranean and Eastern Atlantic ( Table 1 ). The animals from Rio de Janeiro present orange markings at the bases of the ceratal clusters, which are especially visible in the anterior groups. This characteristic is present in all examined specimens, but is not present in the descriptions of B. verrucicornis from the Eastern Atlantic ( Tardy, 1962 ; Edmunds, 1968 ; Ballesteros, 1977 ; García-Gómez & Thompson, 1990 ; Garcia-Gomez, 2002 ). To corroborate this affirmation, we have examined specimens of B. verrucicornis from the Atlantic coast of Huelva (south-western Spain ). Variations of colour are shown in Fig. 8A and B . The back of one specimen is mostly orange ( Fig. 8A ). The other specimen has a whitish back with a translucent orange mid-dorsal band, and the pericardium has further orange pigmentation ( Fig. 8B ). These animals differ from the Brazilian Berghia specimens in the lack of an orange line on the ceratal insertion, and the opaque white band on the dorsum is also absent. Furthermore, the Western Atlantic specimens possess fewer pairs of ceratal groups than the Eastern specimens. The Brazilian animals are 5–16.5-mm long, and have between five and seven groups. Ballesteros (1977) described a 13-mm-long specimen from the Mediterranean Sea with ten ceratal clusters, and Edmunds (1968) studied a 15-mm-long specimen from Ghana with eight groups. Internal characters of Brazilian material were also compared with those from Huelva. The radular teeth of B. verrucicornis ( Fig. 7C ) have more elongated and numerous denticles than those of B. marcusi sp. nov. The teeth of B. verrucicornis have a vertical crack below the median cusp, and the lateral denticles become large gradually ( Fig. 7D ). Berghia marcusi sp. nov. radulae have larger and stouter denticles than those of B. verrucicornis , but a very fine denticle on one or both sides of the median cusp may be present. Furthermore, B. marcusi sp. nov. has the masticatory border of the jaws with rounded denticles, whereas B. verrucicornis has either smooth jaws ( Thompson, 1980 ; Garcia-Gomez, 2002 ) or jaws with hair-shaped denticles ( Tardy, 1962 ; Marcus, 1972 ). Figure 6. Berghia marcusi sp. nov. A, dorsal view of the external morphology; scale bar = 5 mm. B, lateral view of a specimen, showing the arrangement of the cerata; scale bar = 1 mm. C, D, radular teeth (Praia dos Ossos, 5-mm-long specimen); scale bars = 10 Mm. E, detail of a tooth with piled up denticles (16.5-mm-long specimen, from Praia de Aramaçao). F, detail of a tooth with denticles ending in either two or more tips (16.5-mm long specimen, from Praia de Aramaçao). Figure 7. SEM images of the radulae and jaws. A, radula of Berghia marcusi sp. nov. (12-mm-long specimen, from Praia de Aramaçao); scale bar = 20 Mm. B, masticatory border of the jaw of Berghia marcusi sp. nov. (12-mm-long specimen, from Praia de Aramaçao); scale bar = 5 Mm. C, D, radula of Berghia verrucicornis from Huelva; scale bar = 10 Mm. E, radula of Berghia columbina from Huelva; scale bar = 10 Mm. F, jaw of Berghia columbina from Huelva; scale bar = 100 Mm. Table 1. Distinctive features of our Berghia marcusi sp. nov. specimens, of Berghia marcusi sp. nov. specimens described by other authors, and of Berghia verrucicornis and Berghia columbina specimens

Length	N° of	Orange line	N° of	N° of
alive	ceratal	Marks on	on ceratal	Colour of	Colour of	radular	denticles on	Masticatory
Distribution	(mm)	groups	dorsum	insertion	the cerata	rhinophores	teeth	radular tooth	border
Berghia marcusi	Buzios (Rio	5–16	5–7	Mid-dorsal	Present	Translucent	Distal two	10–12	12–21 on	Small denticles
de Janeiro)	white band	tip, below	thirds white	(specimen	either side
(present	orange ring	or cream;	5 mm )	(specimen
paper)	and whitish	reddish base	5 mm ); 28
zone	(specimen
16 mm )
Berghia marcusi	Ubatuba	12–20	6–10	Yellow	Present	Red ring	Pink base;	–	Oldest tooth	Minutes denticles
cited as	( Sao Paulo )	mid-line and	(red	around	yellow tip	10–22 on
B. coerulescens	(Marcus,	yellow triangle	coloured)	cnidosac	either side;
1957)	between eyes	a big tooth
with 21–23
Berghia marcusi	Jamaica	12	8	Mid-dorsal	Present	Pale buff or	Orange papillae	16	40–64	With denticles
cited as	(Edmunds,	white line	greyish yellow	in the basal	per tooth
Berghia B	1968)	digestive gland.	half; creamy
Pale yellow	white tip
ring and
white band
‘ Berghia	Jamaica	18	10	Mid-dorsal	Present	Brown digestive	Orange, with	22	About 30 on	Smooth
verrucicornis ’	(Thompson,	white streak.	gland. White,	lemon-yellow	either side
1980)	Orange patch	lemon yellow	tips
over the	and white
pericardium,	bands.
and triangles
behind the
rhinophores
Berghia	île de Ré	20–30	10	Mid-dorsal	Absent	Yellowish brown	Red, with	–	44–50	Pectinate
verrucicornis	(French)	white band	digestive gland;	white tip	per tooth
( Tardy, 1962 )	red-orange ring
Berghia	Tema ( Ghana )	15	8	Mid-dorsal	Absent	Pale brown	Vermilion on	13	44–48	With denticles
verrucicornis	(Edmunds,	white line	digestive gland;	sides and	per tooth
1968)	vermilion ring	posterior
below tip.	papillae;
Distal third	cream tip and
white	distal papillae
Berghia	Florida	12	9	Opaque white	Present	There is no	Colorless	19	30–38 on	About 350
verrucicornis	(Marcus,	pattern	orange ring	rhinophore;	either side	hair-shaped
1972)	orange base	denticles

Table 1. Continued

Length	N° of	Orange line	N° of	N° of
alive	ceratal	Marks on	on ceratal	Colour of	Colour of	radular	denticles on	Masticatory
Distribution	(mm)	groups	dorsum	insertion	the cerata	rhinophores	teeth	radular tooth	border
Berghia	Medirerranean	13	10	White	Absent	White tip and	Red papillae	–	–	–
verrucicornis	Sea (Spain)	longitudinal	red-orange
(Ballesteros,	band. Orange	band below
1977)	pigment on
pericardium
Berghia	E. Atlantic	8–30	9	Mid-dorsal	Absent	Ring over	Orange	–	42–70	–
verrucicornis	(Spain)	orange line	cnidosac	rhinophoral	per tooth
(García-Gómez	orange-red	papillae
&	with a
Thompson,	gold-orange
1990)	crescent
Berghia	E. Atlantic	8–33	8–9	Mid-dorsal	Absent	Brown digestive	Orange	20–21	30–35 on	Smooth
verrucicornis	(Spain)	orange line	gland, white	papillae;	either side
(Garcia-Gomez,		tip and orange	white tip
2002)	ring below
Berghia	Mar Plata	52	6–8	Brown-red	Present	White cnidosac;	Cream-white	27	47–50 on	Smooth
rissodominguezi	(Buenos	line. Orange	orange band	with yellow	either side
Aires),	triangle on	with thin	apical
Patagonia	head	suapical red	portion
(Muniain &	ring
Ortea, 1999)
Berghia	E. Atlantic	10–30	11	Mid-dorsal	Present	White tip	Orange	24	46–76 on	Fine pointed
columbina	(Spain)	orange line	and brown	papillae	either side	denticles
(García-Gómez	digestive	on the
&	gland	posterior
Thompson,	and lateral
1990)	faces;
white tip
Berghia	E. Atlantic	12–30	9–11	Mid-dorsal	Present	White tip and	Red-orange	22–24	30–38 on	Smooth
columbina	(Spain)	orange line	brown-orange	papillae	either side
(Garcia-Gomez,		digestive	and white
2002)	gland	tip
Berghia A	Jamaica	40	11	Grey suffused	Present	Dark brown	Yellow distal	23	74–98	With denticles
(Edmunds,	with orange	digestive gland.	half, orange	per tooth
1968)	Yellow ring	basal papillae
Berghia C	Jamaica	13	8	Front half	Present	Pale olive-green	Orange middle	–	–	–
(Edmunds,	orange, rear	digestive gland;	half, and
1968)	half grey	white tip and	cream tip
white, orange
or creamy
yellow ring

The absence of data is indicated by: ‘–’. Figure 8. Living animals. A, B, Berghia verruciconnis from Huelva. C, Berghia columbina from Huelva. D, E, Spurilla neapolitana from Huelva. F, Spurilla neapolitana from El Grove. Our specimens from Rio de Janeiro show great similarity with those examined from Ubatuba by Marcus (1957) . Other authors such as García-Gómez & Thompson (1990) reported special features for these animals, which differed from the other specimens from the Western Atlantic. They stated that the specimens from Brazil identified as Spurilla coerulescens by Marcus may belong to another (new) species. Small differences exist between our material and the Ubatuba specimens (orange pigment in our animals and red pigment in those of Marcus), but both animals have orange markings on the dorsum, have oral tentacles and rhinophores that are yellow-tipped and have red bases, have red marks on the head, and have cerata with red rings. Edmunds (1968) described three specimens of Berghia from Jamaica , called A, B, and C, that showed orange markings at the ceratal insertions. He compared these animals with those from Ghana , and suggested two possibilities: first, a very variable species is present in the Western Atlantic; second, two or more new species of Western Atlantic Berghia exist. According to García-Gómez & Thompson (1990) , specimen B is distinct from any described species, and it can be distinguished from B. verrucicornis by the orange marks at the base of the cerata, and the oral tentacles with orange bases. The description of specimen B from Jamaica agrees with the description of our animals, except for the pale yellow ring and the white ring below the cnidosacs. Our specimens possess cerata with orange rings. Specimens A and C from Jamaica also possess orange markings at the base of the cerata, but differ from the Brazilian animals in that specimen A lacks the whitish dorsal band, and specimen C has the front half of the body orange, and the rear half grey. Thompson (1980) assigned to B. verrucicornis two specimens from Jamaica with orange lines at the ceratal insertions, and a mid-dorsal white band. However, these specimens are different from our animals because the cerata have three bands (white, yellow, and white) inside the orange ring, and the masticatory border of the jaws is smooth (our specimens possess denticles). The animals assigned to B. verrucicornis from Florida by Marcus (1972) are very similar to the Brazilian ones. Both have orange markings at the ceratal insertions, and a mid-dorsal white band, but the specimens from Florida have greyish coloration, and they have more ceratal groups (nine groups in a 12-mm-long specimen). Furthermore, the cerata lack rings on cnidosacs. The radulae possess more teeth and more denticles per tooth (19 teeth and 30–38 denticles on each side). The species Berghia columbina ( García-Gómez & Thompson, 1990 ) found in south-western Spain has a similar colour pattern to our specimens ( Fig. 8C ). Berghia columbina has orange markings at the ceratal insertions and on the head, orange or reddish rhinophores with white tips, and translucent orange oral tentacles with white tips. However, B. columbina possesses more pairs of ceratal groups (between nine and 11 groups according to Garcia-Gomez, 2002 ), whereas B. marcusi sp. nov. presents between five and seven groups. Furthermore, a mid-dorsal orange line exists in B. columbina inside a white band, and the cerata do not have orange rings. The radula presents more teeth and more denticles per tooth ( Fig. 7E ), and the jaw masticatory border is smooth ( Fig. 7F ). Muniain & Ortea (1999) described a new species from the Western Atlantic, Berghia rissodominguezi , which is similar to the Brazilian specimens, but there are also several differences. The specimens are larger than B. marcusi sp. nov. , having an orange triangle on the head, and a mid-dorsal reddish-brown line from the third cerata group (our specimens have a cream or opaque white spot on the head and dorsal band). The rhinophores of B. rissodominguezi are cream–white in colour, with yellow on the apical portion inside a red base and yellow tip. The translucent oral tentacles have a yellow area covering two-thirds of them (the oral tentacles of B. marcusi sp. nov. have white tips). Finally, the masticatory border of the jaw in B. rissodominguezi is smooth ( Muniain & Ortea, 1999 ; MD, JT & FG, this paper, pers. observ.), differing from the denticulate border of B. marcusi sp. nov. For all of these reasons, we consider that the Eastern Atlantic and Mediterranean specimens of B. verrucicornis are different from the Western Atlantic Berghia specimens, and possibly that there are two or more West Atlantic species assigned to B. verrucicornis . Our animals from Rio de Janeiro are very similar to the specimens cited as B. coerulescens from São Paulo ( Marcus, 1957 ), and Berghia specimen B from Jamaica ( Edmunds, 1968 ). Therefore, we suggest that our Brazilian specimens, Berghia specimen B by Edmunds (1968) and B. coerulescens by Marcus (1957) , belong to the same new taxon, which is different from B. verrucicornis . The distinguishing characteristics of the species studied in this work are compiled in Table 2 . Distribution: Jamaica ( Edmunds, 1964 ). Brazil : Ubatuba, São Paulo ( Marcus, 1957 ), Praia de Armação , Praia dos Ossos (Buzios, Rio de Janeiro ) (present paper) .