Aquatic Invasions (2012) Volume 7, Issue 3: 425–431 doi: http://dx.doi.org/10.3391/ai.2012.7.3.013 Open Access

© 2012 The Author(s). Journal compilation © 2012 REABIC

Short Communication

New record of the planktonic davisae Ferrari and Orsi in the Black Sea with notes on the identity of “Oithona brevicornis”

Alexandra Temnykh1* and Shuhei Nishida2 1 Institute of Biology of the Southern Seas, 2, Nakhimov Ave., Sevastopol 99011, Ukraine 2 Atmosphere and Ocean Research Institute, University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa 277-8564, Japan E-mail: [email protected] (AT), [email protected] (SN) *Corresponding author

Received: 1 December 2011 / Accepted: 10 February 2012 / Published online: 27 February 2012

Abstract

Specimens of small (< 1 mm) cyclopoid copepods of the genus Oithona were collected from Sevastopol Bay and an offshore area of the Black Sea. On the basis of a combination of characteristics such as those in the maxillule and the genital double-somite, the specimens were identified as O. davisae Ferrari and Orsi, 1984. The identity of “O. brevicornis” that was reported in the Black Sea and the possible origin of O. davisae in the Black Sea are discussed. Key words: Oithona davisae, species identification, ballast water, synanthropic introduction

identify the species with notes on their Introduction distributions. Possible causes for the introduction Reports since 2001 outline the occurrence of the of the species into the Black Sea are also cyclopoid copepod Oithona brevicornis Giesb- discussed. recht, 1891 (see also Giesbrecht 1893 ["1892"]), in the Black Sea (Zagorodnyaya 2002; Altukhov Materials and methods and Gubanova 2006; Gubanova and Altukhov 2007; Selifonova et al. 2008; Selifonova 2009). samples were collected by vertically- The identification was based on the examination stratified tows of a Juday net (mouth area, 0.1 of a few morphological characters, such as the m2; mesh size, 115 µm) from depth layers of shape of the head (rounded anteriorly) and 125–65, 65–40, 40–25, 25–10, and 10–0 m in an rostrum (sharply pointed) and the numbers of offshore area of the Black Sea (around 44°21′N, spines on segments 1–3 of legs 1–4 (with the 33°05′E; depth, 1250–1800 m) on November 12 formula of 1, 1, 3; 1, 1, 3; 1, 1, 3; 1, 1, 2) in the 2010. Samples were also collected by vertical female (Zagorodnyaya 2002; Gubanova and tows of the same net (but with mesh size, 85 µm) Altukhov 2007). However, according to our from 5 m depth to the surface in Sevastopol Bay recent examination of copepod specimens from (44°37′13″N, 33°32′53″E; depth, 14 m) on 18 the Black Sea, there is another species of November 2008. Immediately after collection the Oithona that is similar to O. brevicornis with samples were fixed and preserved in 4% respect to the above features, but shows marked formaldehyde/seawater solution and buffered differences in some other characters. This further with sodium tetraborate. Temperature and salini- led us to suspect that the cyclopoid species ty of the sampling sites were measured with a recently recorded as new to the Black Sea may hydrobiophysical complex “SALPA” (Vasilenko not be O. brevicornis, or include one or more et al. 1997). In the laboratory, small adult species that resemble O. brevicornis. Here we females (< 1 mm) Oithona copepods that had a describe the characteristics of the present rounded anterior margin of the prosome and a specimens and compare them with those of sharply pointed rostrum were sorted from each O. brevicornis and other related species, and sample. These specimens were put in a drop

425 A. Temnykh and S. Nishida

Figure 1. Photographs of Oithona davisae collected from the Black Sea in the present study. A, rostrum, lateral view; B, mandible, focusing on setae on the endopod (arrows); C, maxillule, focusing on long distal-most spine (arrow) on the first inner lobe and the single seta (arrowhead) on the endopod; D, genital-and next somite, lateral view, showing absence of hair-rows on the dorsal surfaces (arrows).

of 10% glycerine/distilled water on a depression m was well mixed with temperature and salinity slide, and total length measured from the anterior of ca.16.5°С and 17.5 psu. A marked end of the prosome to the posterior end of the thermocline existed in the 40–65 m layer, where caudal ramus, excluding caudal setae, with a the temperature decreased with depth to 9°С and calibrated ocular micrometer, under a compound salinity increased to 18.3 psu. The targeted microscope. For detailed examination, Oithona specimens were found in the samples appendages including the mandible, maxillule, from Sevastopol Bay and from the upper 65 m of and swimming legs, were dissected from the the offshore area, but they were not recorded in body trunk and examined at a magnification of samples from 125–65 m. ×1000 with a ×10 eyepiece and an oil-immersion The following characteristics were common to objective lens (×100); the Humes and Gooding's all the present specimens: total length ranged (1964) method was applied. from 0.50 to 0.59 mm (n=20); anterior margin of prosome was rounded in lateral view and ventrally terminated in a sharply pointed rostrum Results (Figure 1A; see also Figure 2H); the endopod of the mandible bore 4 setae (Figure 1B, arrows; The water column at the sampling site in see also Figure 2I); the distal spine on the first Sevastopol Bay was vertically mixed with inner lobe of the maxillule was very long, ca. 2.5 temperature and salinity of ca.16.5°С and 18 psu, times as long as the next spine (Figure 1C, respectively. In the offshore water the upper 40 arrow; see also Figure 2J, arrow); the endopod of

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Figure 2. Character comparison between Oithona brevicornis (A-D), O. aruensis (E-G), and O. davisae (H-K). A, E, H: rostrum. B, F, I: mandible. C, G, J: maxillule; indicating distal-most spine (thin arrow) of first inner lobe, endopod (arrowhead), seta on second inner lobe (dotted arrow; absent in J) and seta on second basal segment (thick arrow; absent in J). D: genital double-somite and next somite, lateral view; indicating rows of hairs (arrows) in O. brevicornis. K: genital double-somite, lateral view. After Nishida (1985: A-D), Nishida and Ferrari (1983: E-G) and Ferrari and Orsi (1984: H-K). Not to scale.

427 A. Temnykh and S. Nishida

Table 1. Comparison of characteristics in adult females among four species and the present specimens of Oithona.

Species Oithona brevicornis Oithona Oithona Oithona davisae Present wellershausi aruensis specimens Total length (mm) 0.48-0.71 0.45-0.50 0.44-0.46 0.49-0.62 0.50-0.59

Rostrum curved and sharply curved and curved and curved and sharply pointed curved and pointed sharply pointed sharply sharply pointed pointed Number of setae on 3 3 3 1 1 endopod of maxillule

Distal-most spine on shorter than next spine > 2 times as ca. 3 times as ca. 2.5 times as long as next ca. 2.5 times as first inner lobe of long as next long as next spine long as next maxillule spine spine spine

Number of seta on 1 0 1 0 0 second inner lobe of maxillule

Number of seta on 1 1 1 0 0 second basal segment of maxillule

Hair-rows on present absent absent absent absent dorsolateral surface of genital double-somite and next somite Geographic Indo-West Pacific Pearl River Indo-West East Asia, California, Chile, Black Sea distribution estuary (China) Pacific Mediterranean

Fine-scale distribution outer part of enclosed not available estuaries and estuaries and inner part of Sevastopol bays inner part of enclosed bays Bay and enclosed bays offshore water

References Nishida et al. (1977), Ferrari (1981) Nishida and Nishida et al. (1977), Ferrari This study Ferrari (1981: as O. Ferrari (1983), and Orsi (1984), Nishida spinulosa), Nishida Nishida (1985) (1985), Hirakawa (1988), and Ferrari (1983), Saiz et al. (2003) Nishida (1985)

the maxillule bore only one seta (Figure 1C, Table 1 compares the characteristics of the arrowhead; see also Figure 2J, arrowhead); there present specimens with those of the four Oithona were no setae on the second inner lobe and the species, namely O. brevicornis (Figure 2A–D; second basal segment of the maxillule (see Nishida 1985), O. wellershausi Ferrari, 1981, Figure 2J, dotted- and thick arrows); the exopod O. aruensis Früchtl, 1923 (Figure 2E–G; Nishida segments 1–3 of legs 1–4 bore 1, 1, 3; 1, 1, 3; 1, and Ferrari 1983) and O. davisae Ferrari and 1, 3; 1, 1, 2 lateral spines; there were no hair Orsi, 1984 (Figure 2H–K). Characteristics such rows on the dorsolateral surface of the genital as body form (oval prosome with sharply pointed double-somite and/or the next somite (Figure 1D, rostrum) and swimming legs (number of exopod arrows). The mean total length of the specimens spines on legs 1–4: 1, 1, 3; 1, 1, 3; 1, 1, 3; 1, 1, from the Sevastopol Bay [0.568±0.018 mm 2) are similar. In all the characteristics listed in (mean ±SD), n=10] was slightly larger than that Table 1, the present specimens correspond with of the offshore specimens (0.533±0.029 mm, O. davisae, but not with the other species. n=10) (t-test, p<0.01), while there were no Oithona brevicornis differs from the other marked differences in other characters between species in that the distal-most spine on the first the specimens from the two localities. inner lobe of the maxillule is much shorter than

428 Oithona in Black Sea the next spine (much longer than the next spine O. wellershausi is still unknown except for the in the other species: Figure 2C, G, J, thin arrows) type locality (Pearl River estuary, China: Ferrari and rows of fine hairs are present on the 1981). Records of the occurrence of posterior dorsolateral surface of the genital O. brevicornis in the Mediterranean (e.g. Pesta double-somite and the anterior dorsolateral 1920; Vaisierre and Seguin 1980; Shuvalov surface of the next somite (such rows are absent 1980) should be considered with caution, since in the other species: Figure 2D, K, arrows). these records are not accompanied with any Oithona davisae differs from O. aruensis and descriptions of features that distinguish O. wellershausi in having only one seta on the O. brevicornis from other related species endopod [three setae in O. aruensis (Figure 2G, (Nishida 1985). A revision of actual species J, arrowheads) and O. wellershausi (figure not should be carried out in the Mediterranean. shown)] and lack of a seta on the second basal There is also evidence that shows O. aruensis segment of maxillule [one seta present in is an estuarine species while O. brevicornis is O. aruensis (Figure 2G, J, thick arrows) and more stenohaline and distributed in more saline O. wellershausi (figure not shown)]. Oithona waters than the former, although their davisae also lacks a seta on the second inner lobe distributions may partially overlap (Table 1; of maxillule while it is present in O. brevicornis Nishida 1985; Chew and Chong 2011). The and O. aruensis (Figure 2C, G, J, dotted arrows). water temperature and salinity at the collection From these observations we conclude that the sites (16–17°С, 17–18 psu) and their yearly present Oithona specimens are O. davisae. fluctuation in the surface waters of the Black Sea (6–25°С, 17–18 psu: e.g. Ovsyaniy et al. 2000; Repetin et al. 2003; Ivanov et al. 2006; Discussion Kondrat’yev 2010) are consistent with those in the temperate embaymental waters where Oithona davisae has been reported from the O. davisae has been recorded (rather than in coastal waters of Japan (e.g. Nishida et al. 1977, tropical-subtropical waters). as “O. brevicornis f. minor”; Nishida and Ferrari In the Black Sea the occurrence of 1983, as “O. aruensis”; Nishida 1985; Ohtsuka et O. brevicornis has been reported mostly from al. 2008) and Korea (Lee et al. 2001; Orui- near-shore waters, such as Sevastopol Bay Sakaguchi et al. 2011), with its occurrences also (Zagorodnyaya 2002; Gubanova and Altukhov being reported from California, the type locality 2007) and Novorossiysk and Tuapse Ports (Ferrari and Orsi 1984), Chile (Hirakawa 1988), (Selifonova 2009). Selifonova et al. (2008) and the northwestern Mediterranean (Nishida’s reported the occurrence of O. brevicornis from unpublished observation cited by Saiz et al. open waters of the western Black Sea, but did 2003). Kasyan (2010) reported the occurrence of not indicate exact localities nor abundance. O. davisae in the ballast water of ships that Temnykh et al. (in press) observed, for the first arrived at Vladivostok from Chinese coasts. With time, high abundances (> 9000 ind. m-3) of these records, it is widely accepted that O. brevicornis in an offshore area in the Black O. davisae is originally endemic to the temperate Sea off the Crimean Peninsula (depth: ca. 1700 coastal waters of East Asia and its occurrence in m). However, the specimens identified as other remote regions are due to synanthropic O. brevicornis were re-examined and proved to introduction, mainly in ship’s ballast waters, as be O. davisae. It is unknown whether the proposed by Ferrari and Orsi (1984), Nishida significant size difference in the specimens (1985), and Hirakawa (1988) (e.g. Carlton 1987; (collected during this research) between the two Hooff and Bollens 2004; Cordell et al. 2008). In collection sites represents infraspecific variation Puget Sound, US, O. davisae was found to be due to different environmental conditions during one of the most common and abundant non- development, or distinct populations with genetic indigenous organisms found in ships from both differences; this invites a further study applying foreign and domestic ballast discharges (Cordell molecular genetics. et al. 2009; Lawrence and Cordell 2010). In In summary, this research and previous contrast, O. brevicornis and O. aruensis are records strongly suggest that O. brevicornis that distributed widely in the subtropical-tropical has been reported to be recently introduced into coastal waters of the Indo-Pacific (Giesbrecht the Black Sea is indeed O. davisae. The true 1891; Früchtl 1923; Wellershaus 1969; Nishida absence of the species before 2001 should be 1985), while the geographic distribution of confirmed on the basis of a retrospective analysis

429 A. Temnykh and S. Nishida of samples covering areas of the Black Sea from References bays to offshore waters. If we assume the absence of O. davisae from Atukhov DA, Gubanova AD (2006) Oithona brevicornis Giesbrecht in the Sevastopol Bay in October, 2005-March, the original Black Sea fauna, the following 2006. Morskoy Ekologicheskiy Zhurnal (Marine Ecology hypotheses may be proposed for the introduction Journal) 5: 32 of O. davisae from the coastal waters of East Carlton JT (1987) Patterns of transoceanic marine biological Asia: (1) direct introduction from East Asia in invasions in the Pacific Ocean. Bulletin of Marine Science 41: 452-465 ships’ ballast waters (as in the cases of, e.g. the Chew LL, Chong VC (2011) Copepod community structure and veined Rapa whelk Rapana venosa, the shrimp abundance in a tropical mangrove estuary, with comparisons Palaemon macrodactylus and the blood cockle to coastal waters. Hydrobiologia 666: 127-143, Anadara inaequivalvis: Skolka and Preda 2010) ; http://dx.doi.org/10.1007/s10750-010-0092-3 Cordell JR, Bollens SM, Draheim R, Systema M (2008) Asian (2) secondary introduction of the Mediterranean copepods on the move: recent invasions in the Columbia– populations, which were introduced and settled Snake River system, USA. ICES Journal of Marine Science in some regions of the Mediterranean (such as 65: 753-758, http://dx.doi.org/10.1093/icesjms/fsm195 those in Barcelona, Saiz et al. 2003), either by Cordell JR, Lawrence DJ, Ferm NC, Tear LM, Smith SS, Herwig RP (2009) Factors influencing densities of non-indigenous the copepods’ dispersal along coastal waters species in the ballast water of ships arriving at ports in Puget through the Turkish Straits System, which act as Sound, Washington, United States. Aquatic Conservation: a strong barrier for many species (Oguz and Marine and Freshwater Ecosystems 19: 322-343, http://dx.doi.org/10.1002/aqc.986 Öztürk 2011) (e.g. the pelagic copepod Acartia Ferrari FD (1981) Oithona wellershausi, new species, and O. tonsa: Kovalev et al. 1998; see also Oguz and spinulosa Lindberg, 1950 (Copepoda: : Öztürk 2011 for examples from other ), ) from the mouth of the Pearl River, China. or in ships’ ballast waters (see Selifonova 2011 Proceedings of the Biological Society of Washington 94: 1244-1257 for copepod examples). However, the possibility Ferrari FD, Orsi J (1984) Oithona davisae, new species, and cannot be ruled out that O. brevicornis and Limnoithona sinensis (Burckhardt, 1912) (Copepoda, O. aruensis (or one of them) may have also been Oithonidae) from the Sacramento San-Joaquin Estuary, introduced into the Black Sea, if we consider the California. Journal of Biology 4: 106-126, http://dx.doi.org/10.2307/1547900 wide distribution of these species in the coastal Früchtl F (1923) Cladocera and Copepoda der Aru-Inseln. waters of the Indo-Pacific region and the Abhandlungen der Senckenbergischen Naturforschenden potential for their introduction in ships’ ballast Gesellschaft 35: 449-457 waters from the countries where these species Giesbrecht W (1891) Elenco dei Coepodi pelagici reccolti dal tenente di bascello Gaetano Chierchia durante il viaggio della are distributed and where the ports of origin may R. Corvetta “Vettor Pisani” negli anni 1882-1885, e dal be located. tenente dal vascello Francesco Orsini nel Mar Rosso, nel For a better understanding of the current 1884. 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Oithona davisae (Oithonidae) from southern Chile. Bulletin of Marine Science 42: 337-339 Acknowledgements Hooff RC, Bollens SM (2004) Functional response and potential predatory impact of Tortanus dextrilobatus, a carnivorous copepod recently introduced to the San Francisco Estuary. The authors would like to express their sincere gratitude to Dr. Marine Ecology Progress Series 277: 167-179, Yuri N. Tokarev, Dr. Victor V. Melnikov and Dr. Julia A. http://dx.doi.org/10.3354/meps277167 Zagorodnyaya for providing the information on temperature and Humes AG, Gooding RU (1964) A method for studying the salinity, and for their help in sampling and other field activities. external anatomy of copepods. Crustaceana 6: 238-240, Special thanks are due to Dr. Frank D. 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