Bursatella leachii Blainville, 1817 (Mollusca, Gastropoda, Heterobranchia), a successful global sea hare

Miquel Pontes¹ & Manuel Ballesteros ^{1, 2}

¹VIMAR, Vida Marina
²Emeritus Professor, Universitat de Barcelona, mballesteros@ub.edu

Introduction

Sea hares (Gastropoda, Heterobranchia, Aplysiida) are mollusks that live in temperate, subtropical or tropical seas. No species lives in polar seas. Most of the approximately 80 known species are medium to large in size and their diet is strictly herbivorous. Many species of sea hares have well-developed lateral extensions of the body (known as parapods) that allow some species to use them as fins for swimming, as, for example, occurs with Aplysia fasciata and Akera bullata.

The species Bursatella leachii, the main character of this article, is a sea hare that does not share these traits because its parapods are fused, leaving only a small hole that communicates with the pallial cavity. In OPK (Ballesteros et al., 2023) you can delve into the curious morphology of this species, as well as different aspects of its biology. This species prefers to live in shallow bottoms and calm waters where macroscopic algae and seagrasses grow. In the Mediterranean it is usually found on sandy or muddy bottoms of bays, coastal lagoons or the interior of ports, often in meadows of Caulerpa prolifera or phanerogams such as Zostera noltei, Cymodocea nodosa and Posidonia oceanica, although it feeds mainly on diatoms, cyanobacteria and detritus, but it may occasionally feed on algae (Paige, 1988).

But Bursatella leachii has some unknowns, some of them impossible to resolve, such as, for example, the exact origin of the first captured specimens that led to its recognition as a species. Its type locality is not known, as already indicated in volume 5 of the Dictionnaire des Sciences Naturelles (Blainvile, 1817) edited by the famous French naturalist Georges Cuvier, who writes in its original language in French in the Bursatella entry : “On ignore sa patrie” (Figure 1). Later Eales & Engel (1935) indicate that the holotype, deposited in the British Museum in London, was collected in the “Indian Seas”, supposedly in some area where the Indian Ocean bathes the coasts of India, without specifying any type locality.

Figure 1. Original description of Bursatella leachii by Cuvier (1817).

On the other hand, due to its great chromatic variability and wide geographical distribution, numerous species included in the genera Aclesia, Aplysia, Barnadaclesia, Notarchus and Ramosaclesia or subspecies of Bursatella have been described, which are currently considered synonyms of B. leachii (Bazzicalupo et al ., 2020; WoRMS, 2023) (Table I).

Until very recently B. leachii was considered the only species of the genus Bursatella, with a circumtropical distribution and also present in temperate seas such as the Mediterranean Sea. However, Bazzicalupo et al. (2020) and Nimbs & Wilson (2020), by using molecular analysis methods, managed to establish two new species: B. ocelligera and B. hirsuta, whose distribution is the Central and Western Pacific Ocean, respectively. It is still unknown why B. leachii has not yet been found along the American coasts of the Pacific Ocean or along the European coasts of the Atlantic Ocean.

Another mystery about B. leachii was related to both its presence and the origin of the specimens found in the Mediterranean Sea. In this sea, the presence of B. leachii is relatively recent, with the first record of this species (O’Donoghue & White, 1940) off the coast of Israel in 1940 and, since then, there have been numerous records that seemed to indicate its expansion from the eastern basin of the Mediterranean to the western basin, so its arrival in the Mediterranean was postulated through a Lessepsian migration from the Red Sea. However, the results of a recent work (Bazzicalupo et al., 2018), when comparing different specimens from the Atlantic, the Indo-Pacific and the Mediterranean, have proven that Mediterranean specimens have more affinities in their haplotypes with those of Atlantic origin than with those of Indo-Pacific origin, thus suggesting the hypothesis of a migration of the species from the Atlantic to the Mediterranean through the Strait of Gibraltar. This has also been corroborated by another more recent work (Córdoba et al., 2023) when studying specimens collected on the Catalan coasts (NE Spain).

This article provides an extensive table with the localized records of B. leachii at a global level obtained through scientific publications, data from specimens preserved in museums and research centers, from Internet platforms confirmed by photographs or unpublished data. The presence and expansion of this species on the coasts of Catalonia (NE Spain) is discussed.

Bursatella leachii @ Laguna Fusaro, Bacoli, Napoli by Guido Villani

Bursatella leachii by Alexander Semenov

Bursatella leachii by Luis Sánchez Tocino

Bursatella leachii @ Ebro Delta (2007)

Bursatella leachii @ Delta de l’Ebre by Neus Sanmartí

Bursatella leachii @ Laguna Fusaro, Bacoli, Napoli by Guido Villani

Bursatella leachii @ Ile de NGor (Senegal), 2012, night dive by Peter Wirtz

Bursatella leachii con puesta by Ernesto Sestito

Bursatella leachii @ Sierra Leone, 1986 by Peter Wirtz

Results and discussion

Table II shows all the records of B. leachii located in the different consulted works. You can see how the oldest quotes from the 19th and early 20th centuries generally correspond to tropical or subtropical marine zones, geographically located around the original occurrence of the species (coasts of India or the Indian Ocean). From the 1930s onwards the species is recorded in numerous localities on the Pacific coast of Australia, Florida, South Africa, also in Japan, in the Caribbean, in the Gulf of Mexico, New Zealand, Brazil, the Philippines, French Polynesia and India. In the Mediterranean its presence has been known since the 1940s.

All these data indicate that its distribution area is between latitudes 40ºN and 40ºS in almost all seas. In the Atlantic Ocean, its northern limit on the coast of North America is 35ºN at North Carolina (USA) (Kruczynski & Porter, 1969) while its southern limit is 25º34’S at Pontal do Sul, Brazil. On the eastern Atlantic coasts, so far it has only been recorded in the Canary Islands, Cape Verde Islands, Senegal, Ghana, Sierra Leone and South Africa. There is no record of the species on the Atlantic coasts of the European continent. In the Indian Ocean the species has been recorded in the Red Sea, on the island of Mauritius and on the coasts of India. In the Pacific it has been recorded from 34ºN at Fukeko (Japan), to 38ºS at Flinders Reef (Australia). In this ocean there are numerous records of B. leachii on the eastern coasts of Australia, which is considered a “hotspot” for sea hares worldwide (Nimbs et al . 2016). It has also been reported in the Sea of ​​Japan, in the Philippines, New Zealand and in French Polynesia. As already indicated, there is no record of the species on the American coasts of the Pacific Ocean, possibly due to cold water currents such as the Humboldt or the California Current, as well as deep and cold water upwelling phenomena that make these coasts unfavorable for numerous species of warm or temperate waters such as B. leachii.

In recent decades, the Mediterranean Sea has become a true “hotspot” for the geographical distribution of B. leachii . Since it was first recorded in 1940, records of this species in the Mediterranean have increased almost exponentially. It is true that during these last decades studies of Mediterranean marine biodiversity have multiplied, not only on the coasts of Southern Europe, but also scientists and naturalists from countries such as Morocco, Tunisia, Algeria, Libya and Egypt, which have contributed to a much better understanding of the distribution of numerous species. It is also postulated about the phenomenon known as “tropicalization” of the waters of the Mediterranean, which are undergoing noticeable warming in general and which is allowing the arrival and colonization by numerous “thermophilous” species, whether they come from the Indo-Pacific (lessepsian migration ) and from the subtropical Atlantic, which appears to be the case for B. leachii .

Selfati et al. (2017) provide a table with all records of B. leachii by country on the Mediterranean coasts. Italy is the Mediterranean country with the most records of B. leachii , both on its western coasts and those of the Adriatic, where the species has been reported on numerous occasions since the 1970s. In the other countries bordering the Mediterranean, the presence of B. leachii has been reported and chronologically in the following order: on the coasts of Israel/Palestine it has been known since 1940 (O’Donoghue & White, 1940), in Turkey since 1959 (Swennen 1969), on the island of Malta since 1969 (Bebbington, 1970), in Egypt since 1975 (Barash & Danin, 1977), in Greece since 1975 (Barash & Danin, 1986), in Tunisia since 1982 (Antit et al., 2011), in Croatia since 1986 (Jaklin & Vio, 1989), in Lebanon since 1999 (Zenetos et al., 2004), in Libya since 2000 (Zgozi et al. (2002) [fide Bazairi et al (2013)], in Slovenia since 2001 (Lipej et al., 2008), in Algeria since 2008 (Lamouti & Bachari in Eleftheriou et al., 2011), in Montenegro since 2009 (Mačić & Kljajić, 2012) , in Albania since 2013 (Xharahi et al. in Gerovasileiou et al., 2017), in France since 2014 (Riondy et al ., 2017), on the Moroccan coasts its presence has been known since 2016 (Selfati et al., 2016) and on the island of Cyprus since 2016 (Lipej et al., 2017).

In the Iberian Peninsula there are no records of the species on the Atlantic coasts, but there are many on the Mediterranean coasts. It was reported for the first time for Spanish waters in the Balearic archipelago in 1996 by Oliver & Terrasa (2004), who found specimens up to 12 cm of length in different locations on the island of Mallorca and on different dates. They even detected the aggregation of numerous individuals in S’Arenal beach, located very close to the city of Palma de Mallorca. These authors provide a map of the Mediterranean Sea with the localities where specimens of this species had been recorded until 2004. On the peninsular coasts B. leachii appeared later, it was recorded for the first time in 2007 in the Ebro Delta (Weitzmann et al., 2007). Since then it has been reported repeatedly in the Ebro Delta (Ballesteros et al., 2016), in the Mar Menor (Izquierdo-Muñoz et al. [in] Nicolaidou et al., 2012), in Roquetas de Mar , Almería (Ibáñez-Yuste et al., 2012) and in various locations in Catalonia where, in addition to the Ebro Delta, it has been cited in L’Ametlla de Mar, Vilanova i la Geltrú, Cubelles and, starting in 2016, in the bathing area of ​​the Barcelona Forum, where there has been a stable population since then. Specimens have also been observed in Creixell, Tarragona (Carles San Vicente, personal communication). It is expected that, if the “tropicalization” process of the Mediterranean Sea continues, this species will also be able to colonize the northern coast of Catalonia, especially the Costa Brava, where the water temperature in winter drops below 15ºC, that is considered the minimum for the presence of stable populations.

With all the data from the records in Table II, it is verified that B. leachii has a great capacity to colonize very extensive marine areas of temperate or warm waters. In the near future, the effects of climate change on the oceans may further favor the expansion of the species in coastal areas where it has not yet been recorded.

Publications of the original descriptions of the synonyms

• Bursatella leachii guineensis Bebbington A. (1969) – Bursatella leachi guineensis subsp. nov. (Gastropoda, Opisthobranchia) from Ghana. Proceedings of the Malacological Society of London 38: 323-341.  Cited in Ghana
• Aplysia cirrhifera Quoy & Gaimard, 1832 – Quoy, J. R. C. & Gaimard, J. P. (1832-1835). Voyage de la corvette l’Astrolabe : exécuté par ordre du roi, pendant les années 1826-1827-1828-1829, sous le commandement de M. J. Dumont d’Urville. Zoologie. 1: i-l, 1-264; 2(1): 1-321 [1832]; 2(2): 321-686 [1833]; 3(1): 1-366 [1834]; 3(2): 367-954 [1835]; 4 [1833]; Atlas (Mollusques): pls 1-93 [1833] …etc. Paris: Tastu. In: Dumont d’Urville, J.; 1834, Voyage de Découvertes de l’Astrolabe. Paris, J. Tastu, Éditeur-Imprimeur., available online at https://doi.org/10.5962/bhl.title.2132 page(s): 311–312, pl. 24 fig. 8  Cited in Mauritius
• Bursatella savigniana Audouin, 1826 – Audouin V. (1826) Explication sommaire des planches de Mollusques de l’Egypte et de la Syrie publiées par J.C. Savigny. in: Description de l’Egypte ou recueil des observations et des recherches qui ont été faites en Egypte pendant l’expédition de l’armée française, publié par les ordres de sa majesté l’empereur Napoléon le grand. Histoire Naturelle, Animaux invertébrés 1(4): 7-56. Paris: Imprimerie impériale., available online at https://digi.ub.uni-heidelberg.de/diglit/jomard1809bd5_1_1/0548 page(s): 17-18   Cited in the Red Sea
• Aclesia glauca Cheeseman, 1878 – Cheeseman, T.F. (1878). Descriptions of three new species of opisthobranchiate Mollusca from New Zealand. Proceedings of the Zoological Society of London. 1878: 275–277. page(s): 277, pl. 15 fig. 4   Cited in New Zealand
• Notarchus laciniatus Rüppell & Leuckart, 1830 – Rüppell, E. & Leuckart, F. S. (1828-1830). Mollusca. In: Atlas zu der Reise im nordlichen Afrika von Eduard Rüppell. Neue wirbellose Thiere des Rothen Meeres. Frankfurt, H. L. Brönner. pp. 15-22, pls 4-6 [1828]; pp. 23-47, pls 7-12 [1830]. , available online at https://www.biodiversitylibrary.org/page/37140891
  page(s): 24-25, Plate 7, Figures 2 a-c  Cited in the Red Sea
• Bursatella lacinulata A. Gould, 1852 – Gould, A. A. (1852). Mollusca and shells. In: United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842 under the command of Charles Wilkes. Boston. 12: 1-510; atlas 1856: 1-16., available online at https://www.biodiversitylibrary.org/page/10991152  page(s): 223–224, pl. 16, figs 269-269a Cited in Brazil
• Aclesia africana Engel, 1926 – Engel H. (1926). Drei neue Arten der Gattung Aclesia (Rang) Bergh, 1902. Zoologischer Anzeiger. 69: 180-187. page(s): 180–184, figs 11–14   Cited in South Africa
• Aclesia rosea Engel, 1926  – Engel H. (1926). Drei neue Arten der Gattung Aclesia (Rang) Bergh, 1902. Zoologischer Anzeiger. 69: 180-187. page(s): 184–185, figs 15–16   Cited in Western Africa
• Aplysia pleii Rang, 1828 – Rang, S. (1828). Histoire naturelle des Aplysiens. Paris: Firmin Didot. 83 pp. 24 pls. , available online at https://www.biodiversitylibrary.org/page/14983674  page(s): 70, pl. 21 figs 2-4 Cited in the Antilles
• Notarchus intrapictus Cockerell, 1893 – Cockerell, T. D. A. (1893). On a new species of Aplysiidae from Jamaica. Annals and Magazine of Natural History. ser. 6, 11(63): 218–220., available online at https://www.biodiversitylibrary.org/page/24344548 Cited in Jamaica
• Notarchus cirrosus W. Stimpson, 1855 – Stimpson, W. (1855). Descriptions of some of the new Marine Invertebrata from the Chinese and Japanese Seas. Proceedings of the Academy of Natural Sciences, Philadelphia. 7(10): 375-384., available online at https://biodiversitylibrary.org/page/26299600  page(s): 378 Cited in the China Sea and Japan
• Aclesia freeri L. E. Griffin, 1912 – Griffin L.E. (1912). The anatomy of Aclesia freeri new species. The Philippine Journal of Science. section D, 7(2): 65–90, pls 1-6., available online at https://www.biodiversitylibrary.org/page/34354225  Cited in Philippines
• Ramosaclesia rex J. K. Allan, 1932 – Allan, J.K. (1932). A new genus and species of sea-slug, and two new species of sea-hares from Australia. Records of the Australian Museum. 18(6): 314–320, pls 34-35. page(s): 317–319, pl. 35 figs 1–6 Cited in Australia
• Notarchus villosus O’Donoghue, 1929 – O’Donoghue, C. H. (1929). Opisthobranchiate Mollusca collected by the South African Marine Biological Survey. Union of South Africa. Fisheries & Marine Biological Survey Report No. 7 for the year ending June 1929. pp. 1-84, pls. 1-8. Special Reports No. 1. page(s): 25–30, pl. 3 figs 27–32. Cited in South Africa
• Notarchus brevipes Hägg, 1904 – Hägg R. (1904). Two new Opisthobranchiathe Mollusca from the Red Sea. In: L. A. Jägerskiöld (ed.), Results of the Swedish Zoological Expedition to Egypt and the White Nile 1901 under the direction of L. A. Jägerskiöld, Part 1: 1–16, pl. 1. page(s): 1–5, pl. 1 figs 1–3 Cited in the Red Sea
• Bursatella leachii hirasei Eales & Engel, 1935 – Eales, N. & Engel, H. 1935. The genus Bursatella De Blainville. Proceedings of the Malacological Society of London 21: 279-303, plate 31. page(s): 298  Cited in Japan
  

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NOTES:

• This article is complementary to the work by Córdoba González, D., Enguídanos, A., Castells, O., Baños, E., Valdés, Á. & Ballesteros, M. 2023. Where do I come from? Phylogenetic analysis of Bursatella leachii Blainville, 1817 (Mollusca: Gastropoda: Heterobranchia) from the Catalan coasts (NE Spain). Cahiers de Biologie Marine, 64:325-336.
• If you have records of this species not covered in this article, better if they are supported by at least one photograph, you can send them to us by email to the address indicated above.

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