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Anadara Inaequivalvis in the Footsteps of Anadara Kagoshimensis in the Mediterranean Sea - (Bivalvia, Arcidae)

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Anadara Inaequivalvis in the Footsteps of Anadara Kagoshimensis in the Mediterranean Sea - (Bivalvia, Arcidae) Anadara inaequivalvis in the footsteps of Anadara kagoshimensis in the Mediterranean Sea - (Bivalvia, Arcidae) By M.C. van Veen, August 2020 Introduction It is well known that Anadara kagoshimensis (Tokunaga, 1906) has become an alien invasive species in the Mediterranean Sea, extending its range all the way up the southern part of Bretagne, France. It is assumed it was taken to the Mediterranean by East Asian ships discharging their ballast water. Numerous articles have been written about this invasive and highly successful species, often erroneously recognised as Anadara inaequivalvis (Bruguière, 1789) and sometimes even Anadara cornea (Reeve, 1844). Both of these species did not reside in the Mediterranean; two other invasive migrants did however: the Anadara transversa (Say, 1822) (often put forward as Anadara demiri) and the Lessepsian migrant Anadara natalensis (Krauss, 1848), which travelled through the Suez Canal to the northern Egyptian shores and further north to the Israeli coast. A number of years ago specimens of true Anadara inaequivalvis began to appear in and around the Iskenderun Gulf, Turkey. Even though previous publications were incorrect in their assessment concerning Anadara kagoshimensis, it seems like Anadara inaequivalvis does reside in the Mediterranean Sea after all. Morphological analysis of the Mediterranean Anadara species Comparison between species is required in order to obtain correct determination. It is possible to recognise each Anadara species by its morphological characteristics, even though it takes a trained eye to see the differences between these. Even though morphological characteristics can be analysed separately and different elements between species can be compared, the whole will get fragmented into unrelated parts that do not tell much about the total appearance of a species. This method aims at making an objective comparison, but it requires correct species determination beforehand in order to be valid. Only with correct species determination any morphological or genetic analysis can be validated. One problem with comparison of morphological characteristics between species is that these characteristics are required to be fixed in order for the comparison to work. Sometimes these characteristics are based on the analysis of a limited number of specimens and do not properly represent the intraspecific variability. Assumed fixed morphological elements might be incorrect. In case of comparing specimens, it must be noted that there is a difference between a match with the fixed characteristics as given by the author's description, and a match with assumed fixed characteristics by later researchers. The fixed characteristics by the author are core elements by which a species has been validated, even though these can be expanded upon at a later date, while the assumed fixed characteristics by later researchers serve the purpose of comparison between species, which might be incorrect. Anadara kagoshimensis (Tokunaga, 1906) Tokunaga gave the following descrption and picture in 1906: "Shell small, solid, oblong, tumid; anterior side rounded; posterior subtruncated; surface sculptured with 31 to 34 finely granulated radial ribs, with interstices broader than the ribs; umbones slightly directed anteriorly; teeth numerous." Fig 1. Anadara kagoshimensis (Tokunaga, 1906). 53 mm. Italy, Venice, Caorle, March 1997. Fig 2. Anadara kagoshimensis (Tokunaga, 1906). 35 mm. Italy, Venice, Caorle, March 1997. Fig 3. Anadara kagoshimensis (Tokunaga, 1906). 56 mm. France, Bretagne, Saint-Philibert, 2003. Fig 4. Anadara kagoshimensis (Tokunaga, 1906). 45 mm. Italy, Grado, 10-20 m deep on sand, 1983. Fig 5. Anadara kagoshimensis (Tokunaga, 1906). 59 mm. Italy, Ravenna, Adriatic Sea, Sept 1978. Anistratenko et al. (2014) incorrectly named this species Anadara inaequivalvis. An extra note regarding Arca loricata Reeve, 1844: there are three type specimens of unknown origin residing in the Natural History Museum London (reg. nr. 1969253). One of these specimens is Anadara cornea (Reeve, 1844), the other two specimens are most likely Anadara kagoshimensis and are very similar to Fig 2 and Fig 4. The locality of these shells could well be Hong Kong. Anadara inaequivalvis (Bruguière, 1789) Bruguière's description of this species seems unclear at first glance, but it clarifies the distinction with other species well. In his description he referred to a picture in Chemnitz & Martini (1784), which shows Anadara rhomboidalis however, a species now assigned to Shumacher (1817). Chemnitz & Martini's Neues Systematisches Conchylien-Cabinet is a work which is not recognised by the ICZN (International Commission on Zoological Nomenclature). Anadara inaequivalvis, in contrast to Anadara kagosimensis, hasn't got granulated ribs on either valve, but it does have a strongly overlapping left valve. The length is up to about 65 mm. It looks and feels more delicate than Anadara kagoshimensis, less robust. Reeve (1844) depicts this species well, as shown below. Fig 6. Anadara inaequivalvis (Bruguière, 1789). 61 mm. India, Rameshwaram, Aug. 2014. Fig 7. Anadara inaequivalvis (Bruguière, 1789). 54 mm. Indonesia, Sumatra, Tanjung Tiram. Strafella et al. (2017) made a morphological comparison between Anadara kagoshimensis and another species that was named Anadara inaequivalvis (from the Philippines). However, the other species was likely Anadara uropigimelana (Bory de St-Vincent, 1827) instead. Below the picture of Anadara uropigimelana that was provided by Bory de St.-Vincent. Fig 8. Anadara uropigimelana (Bory de St-Vincent, 1827). 76 mm. Madagascar, Mahavelona, 2015. Anadara cornea (Reeve, 1844) Fig 9. Anadara cornea (Reeve, 1844). 50 mm. Australia, Queensland, Dingo Beach, July 2002. Reeve (1844) writes that the number of ribs in Anadara cornea is about 29; the ribs of the left valve are nodulously crenated, but the ribs of the right valve are flat and smooth. Anadara kagoshimensis shows a similar rib sculpture in the left and right valves. Reeve also mentions that Anadara cornea is inequivalve, although this becomes less obvious as the age progresses and the size increases. Also the degree of inequivalvity shows similarity to Anadara kagoshimensis. These two corresponding aspects might be reasons why both species were confused in the past. Anadara natalensis (Krauss, 1848) Krauss gave the following description and picture in 1848: "The ribs, also in a smaller specimen, are always 30 in number, and are striking, flat, not granular or notched, but indistinctly grooved transversally. At the anterior and in the middle the ribs are angular, posteriorly the ribs are more rounded." Although this is only part of the description, it gives some key elements by which this species can be recognised. In reality the number of ribs have some variabilty, just like e.g. its overall shape and its thickness, and the number of ribs should lie between approximately 28 and 31. One remarkable feature of this species is that its shell is almost orthogyrate, whereas most other Anadara species are obviously prosogyrate. The left valve mainly overlaps ventrally, in contrast to Anadara inaequivalvis, which has got a strong overlap that is orientated posteriorly as well. Fig 10. Anadara natalensis (Krauss, 1848). 55 mm. Israel, Haifa. 30 ribs. Fig 11. Anadara natalensis (Krauss, 1848). 63 mm. Egypt, North Sinaï, just east of Port Said, Mediterranean. Beached shell, Oct 2002. 30 ribs. The figure below shows a thick and inflated specimen of Anadara natalensis from the southeastern part of the Arabian Peninsula, with only 28 ribs. Fig 12. Anadara natalensis (Krauss, 1848). 60 mm. SE. Arabian Peninsula, Gulf of Oman, 1987. Anadara inaequivalvis in the eastern Mediterranean Sea During the past decades numerous developing countries have strengthened their economy and have expanded their trade routes significantly. One of these major economies is India. The reason India is highlighted instead of other countries is because true Anadara inaequivalvis is well known from southern India, or the "Coromandel Coast," as Bruguière wrote in his description of this species in 1789. Shipping routes from India to Europe go through the Arabian Sea, the Red Sea and the Mediterranean Sea, thereby potentially facilitating an artificial Lessepsian spread, just like the East Asian ships in the 1970s. It seems plausible Anadara inaequivalvis has also become a new invasive species due to the discharge of ballast water. Fig 13. Anadara inaequivalvis (Bruguière, 1789). 30.2 mm. Turkey, Taşucu, dived at 6 m depth, dead on sand, August 2012. The specimen has got 32 ribs and a typical narrow arrow- shaped cardinal area. See also Lutaenko (2006) plate 3, specimens G-I. Anadara transversa (Say, 1822) Anadara transversa is known from the Mediterranean Sea, although specimens in this region usually remain much smaller than 30 mm. Naturalis Biodiversity Center provided several photographs of Anadara transversa on Wikimedia Commons, one of which is catalog number ZMA.MOLL.411489. One of the specimens resembles Anadara inaequivalvis in Fig 13, as shown below. Fig 14. Anadara transversa (Say, 1822). approximately 30 mm. USA, Massachusetts, Osterville, exchanged in 1948. Provenance: Naturalis Biodiversity Center, ZMA.MOLL.411489. There are some notable differences with the specimen in Fig 13: 1. The interstices are much wider, those have the same width or are even wider than the ribs. 2. On the inside at the ventral margin the negative imprints of the interstices extend much further inward,
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