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Rediscovery of Sanderia malayensis and remarks on Rhopilema nomadica record in Pakistan (Cnidaria: Scyphozoa)

ABSTRACT

In this report, we record the rediscovery of Sanderia malayensis in Pakistani waters since the John Murray Expedition (1933-34) who collected two small specimens off-Karachi, Pakistan. The specimens were described and we discuss the shape of gastric pouches contour. We also comment upon the newly recorded Rhopilema nomadica jellyfish from the region, and assume it to be previously recorded as Rhopilema hispidum.

Key-Words:
Scyphomedusae; Sanderia; Rhopilema; Karachi; Northern Arabian Sea

RESUMO

Esta nota registra a redescoberta de Sanderia malayensis em águas paquistanesas desde a Expedição John Murray (1933-34) que coletou dois pequenos exemplares na costa de Carachi. Os espécimes foram descritos e nós discutimos a forma do contorno das bolsas estomacais. Nós também comentamos sobre a medusa Rhopilema nomadica recentemente registrada para a região, e acreditamos tratar-se de uma espécie do gênero previamente registrada, Rhopilema hispidum.

Palavras-Chave:
Scyphomedusae; Sanderia; Rhopilema; Carachi; Mar Arábico norte

INTRODUCTION

SanderiaGoette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837. is a genus of scyphozoan jellyfishes that includes two species: Sanderia malayensisGoette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837. and S. pampinosusGershwin & Zeidler, 2008GERSHWIN, L. & ZEIDLER, W. 2008. Two new jellyfishes (Cnidaria: Scyphozoa) from tropical Australian waters. Zootaxa, 1764:41-52.. Members of the genus are characterized by having 32 marginal lappets, 16 tentacles alternating with 16 rhopalia, 32 radial stomach pouches (all alike) and exumbrella presenting numerous nematocyst warts (Kramp, 1961KRAMP, P.L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom, 40:7-469.:330). Sanderia malayensis occurs in Indo-Pacific waters, while S. pampinosus is restricted to northwestern Australia (Kramp, 1961KRAMP, P.L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom, 40:7-469.; Gershwin & Zeidler, 2008GERSHWIN, L. & ZEIDLER, W. 2008. Two new jellyfishes (Cnidaria: Scyphozoa) from tropical Australian waters. Zootaxa, 1764:41-52.).

The knowledge of scyphomedusae from Pakistan is growing in the last years with several new records being reported (Gul & Morandini, 2013GUL, S. & MORANDINI, A.C. 2013. New records of scyphomedusae from Pakistan coast: Catostylus perezi and Pelagia cf. noctiluca (Cnidaria: Scyphozoa). Marine Biodiversity Records, 6(e86):1-6., 2015; Gul et al., 2014GUL, S.; MOAZZAM, M. & GALIL, B.S. 2014. Occurrence of Marivagia stellata (Scyphozoa: Rhizostomeae: Cepheidae) along the coast of Pakistan, northern Arabian Sea. Marine Biodiversity Records , 7(e112):1-2., 2015aGUL, S.; MOAZZAM, M. & MORANDINI, A.C. 2015a. Crowned jellyfish (Cnidaria: Scyphozoa: Rhizostomeae: Cepheidae) from waters off the coast of Pakistan, northern Arabian Sea. Check List, 11(1551):1-4., bGUL, S.; MORANDINI, A.C. & MOAZZAM, M. 2015b. First record of the crowned jellyfish Netrostoma coerulescens (Cnidaria: Scyphozoa) from Pakistani waters. Marine Biodiversity Records , 8(e156):1-4.), and added to the cnidarian fauna (Gul et al., 2015cGUL, S.; MORANDINI, A.C.; HÄUSSERMANN, V. & PÖRSCHMANN, U. 2015c. Checklist of cnidarians from Pakistani waters. Check List, 11(1609):1-8.). To date, there are nine scyphozoan jellyfishes recorded from Pakistani waters (Gul et al., 2015bGUL, S.; MORANDINI, A.C. & MOAZZAM, M. 2015b. First record of the crowned jellyfish Netrostoma coerulescens (Cnidaria: Scyphozoa) from Pakistani waters. Marine Biodiversity Records , 8(e156):1-4.), a number that corresponds to less than 5% of the total biodiversity of the class Scyphozoa (estimated in ~200 species by Daly et al., 2007DALY, M.; BRUGLER, M.R.; CARTWRIGHT, P.; COLLINS, A.G.; DAWSON, M.N.; FAUTIN, D.G.; FRANCE, S.C.; MCFADDEN, C.S.; OPRESKO, D.M.; RODRIGUEZ, E.; ROMANO, S.L. & STAKE, J.L. 2007. The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus. Zootaxa, 1668:127-182.). Out of these records, the only ones of the family Pelagiidae Gegenbaur, 1856 are: Pelagia noctiluca (Forskål, 1775) and Sanderia malayensisGoette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837. (Stiasny, 1937STIASNY, G. 1937. Scyphomedusae. In: John Murray Expedition (1933-34), Scientific Reports. Zoology. London, British Museum of Natural History. v. 4, p. 203-242.; Gul & Morandini, 2013GUL, S. & MORANDINI, A.C. 2013. New records of scyphomedusae from Pakistan coast: Catostylus perezi and Pelagia cf. noctiluca (Cnidaria: Scyphozoa). Marine Biodiversity Records, 6(e86):1-6.); interestingly the last report of S. malayensis was documented 79 years ago by Stiasny (1937STIASNY, G. 1937. Scyphomedusae. In: John Murray Expedition (1933-34), Scientific Reports. Zoology. London, British Museum of Natural History. v. 4, p. 203-242.). Our goal is to document a new record of the species in Pakistani waters. In addition, we also provided comments on the recently recorded Rhopilema nomadicaGalil, 1990GALIL, B.S.; SPANIER, E. & FERGUSON, W.W. 1990. The scyphomedusae of the Mediterranean coast of Israel, including two lessepsian migrants new to the Mediterranean. Zoologische Mededelingen, 64:95-105. (Tahera & Kazmi, 2015TAHERA, Q. & KAZMI, Q.B. 2015. Occurrence of Rhopilema nomadica Galil, 1990 (Cnidaria: Scyphozoa: Rhizostomeae: Rhizostomatidae) in Pakistani waters. International Journal of Biological Research, 3(1):49-51.) from Pakistan and consider it to be erroneously identified Rhopilema hispidum jellyfish previously recorded from this area (Gul & Morandini, 2015GUL, S. & MORANDINI, A.C. 2015. First record of the jellyfish Rhopilema hispidum (Cnidaria: Scyphozoa) from the coast of Pakistan. Marine Biodiversity Records , 8(e30):1-4.).

MATERIALS & METHODS

On 26 February 2015, 37 specimens of pelagiid scyphomedusa were collected as by-catch during several bottom trawlings (30 min, 12-80 m depth) off-Karachi (24°51’41.264”N, 67°0’35.779”E), Pakistan, northern Arabian Sea. All specimens were photographed, examined and then preserved in 4% formaldehyde solution. The specimens were studied in detail and deposited in the Museum of Department of Zoology, Jamia Millia Government Degree College (JMGDC MDZ), Malir, Karachi, Pakistan. Identification of the specimens followed Kramp (1961KRAMP, P.L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom, 40:7-469.).

The specimens were compared to the holotype held at the Museum für Naturkunde (ZMB), Berlin (Germany), and also to young medusae reared in the laboratory from Japanese polyp cultures.

RESULTS AND DISCUSSION

Systematics Class Scyphozoa Goette, 1887

Subclass Discomedusae Haeckel, 1880HAECKEL, E. 1880. System der Acraspeden. Zweite Halfte des System der Medusen. In: Haeckel, E. Das system der medusen. Jena, G. Fischer. Tomo 1 , pt. 2, p. 361-672.

Order Semaeostomeae L. Agassiz, 1862

Family Pelagiidae Gegenbaur, 1856

Genus Sanderia Goette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837. Sanderia malayensis Goette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837.

( Figs. 1 , 2 )

Figure 1:
Freshly collected specimen of Sanderia malayensisGoette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837. from Pakistani coast, off-Karachi. Bell diameter 11 cm: (A) Exumbrellar view showing transparent umbrella with numerous nematocyst warts, (B) Subumbrellar view showing gastric pouches in close-up, note the heart-shape contour of gastric pouches.

Figure 2:
Sanderia malayensisGoette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837. specimens in subumbrellar view: (A) Type specimen ZMB 2622, (B) Type specimen ZMB 2621, (C, D) Specimens cultivated from ephyra. Note the shape of gastric pouches in all individuals.

Comparison material

Holotype ZMB 2622; Singapore, 23 April 1884; 1 specimen sex undetermined, but gonads present; 2 cm bell diameter (original description mentions 2.5 cm). ZMB 2621; Singapore, no date specified; 1 specimen sex undetermined, but gonads present; 2.5 cm bell diameter. Unnumbered specimens (private collection of first author): 3 immature specimens reared in laboratory from polyp cultures (1, 1.5, and 1.7 cm bell diameter), strobilation occurred on 27 March 2013, preserved in 4% formaldehyde solution on 26 April 2013; 2 immature specimens reared in laboratory from polyp cultures (1.4 and 1.6 cm bell diameter), strobilation occurred on 29 September 2014, preserved in 4% formaldehyde solution on 05 November 2014.

Pakistani specimens

JMGDC MDZ CN 23; off-Karachi, 26 February 2015; 37 specimens, sex undetermined, but gonads present; 5.5-13.5 cm bell diameter.

Description of Pakistani specimens

Medium sized pelagiid scyphomedusae of saucer-shaped, transparent umbrella with prominent nematocysts warts on exumbrellar surface (Fig. 1A); some specimens with violet pigments. Tentacles 16, up to 29 cm long, ribbon-like, alternating with 16 rhopalia. Oral arms up to 16 cm long, bearing violet shade pronounced on edges. Gastric pouches contour horseshoe to heart-shaped; bordered externally by up to 35-40 long, light violet, finger-like gonadal papillae (Fig. 1B).

Remarks on Sanderia species

Sanderia malayensisGoette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837. was originally described from Singapore, Malayan Archipelago (Goette, 1886GOETTE, A. 1886. Verzeichniss der Medusen, welche von Dr. Sander, Stabsarzt auf S.M.S. ‘Prinz Adalbert’ gesammelt wurden. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 39:831-837.); later it was reported from several places like Japan (as Neopelagia eximiaKishinouye, 1910KISHINOUYE, K. 1910. Some medusae of Japanese waters. Journal of the College of Science Tokyo, 27:1-35.), Philippines and Suez Canal (Kramp, 1961KRAMP, P.L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom, 40:7-469.). Stiasny (1937STIASNY, G. 1937. Scyphomedusae. In: John Murray Expedition (1933-34), Scientific Reports. Zoology. London, British Museum of Natural History. v. 4, p. 203-242.) reported the species from northern Arabian Sea (off-Karachi along Pakistani coast and Gulf of Oman), and Gulf of Aden. Two small specimens (1 and 1.5 cm bell diameter) wrongly stated as in Pelagia-stage were collected off-Karachi on 8 November 1933 during the John Murray Expedition (1933-34). According to Kramp (1961KRAMP, P.L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom, 40:7-469.:331) the species has heart-shaped gastric pouches each bordered externally by about 24-30 finger-like gonadal papillae; but our specimens although agreeing with the description of the species, have more papillae (35-40) than stated, and the shape of the gastric pouches vary slightly. Vanhöffen (1902VANHÖFFEN, E. 1902. Die Acraspeden Medusen der deutschen Tiefsee-expedition 1898-1899. In: Wissenschaftliche Ergebnisse der deutschen Tiefsee-expedition auf dem dampfer Valdivia, 1898-1899. v. 3:3-52.:38) while observing specimens from the Gulf of Aden mentioned the number of gonadal papillae to be 24, tentacles compressed laterally and nematocysts concentrated on the outer portion of tentacles. Browne (1926BROWNE, E.T. 1926. Report on the Medusae. Transactions of the Zoological Society of London, 22:105-115.:110-112) observed specimens collected at the Suez Canal area ranging from 1.2-3.5 cm bell diameter and having variation in rhopalia number. Stiasny (1937STIASNY, G. 1937. Scyphomedusae. In: John Murray Expedition (1933-34), Scientific Reports. Zoology. London, British Museum of Natural History. v. 4, p. 203-242.:226) however, was not correct while identifying Pelagia-stage specimens of Sanderia from off-Karachi, Pakistan. Such specimens must correspond to a Pelagia species, based on the information about the life cycle of Sanderia malayensis (Uchida & Sugiura, 1975UCHIDA, T. & SUGIURA, Y. 1975. On the ephyra and postephyra of a semaeostome medusa, Sanderia malayensis Goette. Journal of the Faculty of Science, Hokkaido University, Series VI, Zoology, 19:879-881., 1978UCHIDA, T. & SUGIURA, Y. 1978. On the polyp of the scyphomedusa, Sanderia malayensis and its reproduction. Journal of the Faculty of Science, Hokkaido University, Series VI, Zoology , 21:279-287.) and our own observations that Sanderia ephyrae are released with already 16 rhopalia and 32 lappets. For other specimens observed by Stiasny (1937STIASNY, G. 1937. Scyphomedusae. In: John Murray Expedition (1933-34), Scientific Reports. Zoology. London, British Museum of Natural History. v. 4, p. 203-242.:225-227), the author did not mention the number of gonadal papillae, but if we focus on his pl. 1, fig. 1, the two specimens are showing to have approximately 40 papillae on each gastric pouch, similar to our observations.

Sanderia malayensis has only one congener, S. pampinosus, which bears horseshoe-shaped gastric pouches with 34-44 gonadal papillae (Gershwin & Zeidler, 2008GERSHWIN, L. & ZEIDLER, W. 2008. Two new jellyfishes (Cnidaria: Scyphozoa) from tropical Australian waters. Zootaxa, 1764:41-52.). For the authors, these are the main features that distinguish both species. However, by inspection of the holotype specimen (ZMB 2622, Fig. 2A) it is possible to observe that the contour of the gastric pouch is variable and the gonadal papillae are thicker than in other specimens. Additionally, another specimen observed by Goette (ZMB 2621, Fig. 2B) and in young specimens of S. malayensis we cultivated in the laboratory of the first author (Fig. 2C-D), it is possible to state that the gastric pouch contour is horseshoe-shaped while, horseshoe to heart-shaped with 35-40 gonadal papillae in the Pakistani specimens (Fig. 1C-D). These observations raise the question whether, the shape of the gastric pouch can pass through developmental changes during growth and that we might be dealing with a single species of the genus with a wider distributional area. Our suggestions, to solve this issue, are to cultivate this jellyfish species to maturity and perform comparative molecular studies with specimens from different areas.

Remarks on Rhopilema nomadica record in Pakistan

Currently the genus RhopilemaHaeckel, 1880HAECKEL, E. 1880. System der Acraspeden. Zweite Halfte des System der Medusen. In: Haeckel, E. Das system der medusen. Jena, G. Fischer. Tomo 1 , pt. 2, p. 361-672. (Scyphozoa: Discomedusae: Rhizostomeae: Rhizostomatidae) is composed of five valid species which differ mainly in their exumbrella morphology (smooth/rough) and the kind of appendages on mouth arms (Gul & Morandini, 2015GUL, S. & MORANDINI, A.C. 2015. First record of the jellyfish Rhopilema hispidum (Cnidaria: Scyphozoa) from the coast of Pakistan. Marine Biodiversity Records , 8(e30):1-4.). From these five species, only two have rough exumbrellar surface: (1) Rhopilema hispidum, with two kinds of warts: a) minute, blunt, colourless, and b) tiny, conical, pointed, reddish-brown warts (pronounced on marginal area); (2) Rhopilema nomadica, with only minute, blunt warts (“blunt tuberculation”, Galil et al., 1990GALIL, B.S.; SPANIER, E. & FERGUSON, W.W. 1990. The scyphomedusae of the Mediterranean coast of Israel, including two lessepsian migrants new to the Mediterranean. Zoologische Mededelingen, 64:95-105.:103) (Omori & Kitamura, 2004OMORI, M. & KITAMURA, M. 2004. Taxonomic review of three Japanese species of edible jellyfish (Scyphozoa: Rhizostomeae). Plankton Biology and Ecology, 51:36-51.; Kitamura & Omori, 2010KITAMURA, M. & OMORI, M. 2010. Synopsis of edible jellyfishes collected from Southeast Asia, with notes on jellyfish fisheries. Plankton and Benthos Research, 5(3):106-118.; Gul & Morandini, 2015GUL, S. & MORANDINI, A.C. 2015. First record of the jellyfish Rhopilema hispidum (Cnidaria: Scyphozoa) from the coast of Pakistan. Marine Biodiversity Records , 8(e30):1-4.). Recently, Tahera & Kazmi (2015TAHERA, Q. & KAZMI, Q.B. 2015. Occurrence of Rhopilema nomadica Galil, 1990 (Cnidaria: Scyphozoa: Rhizostomeae: Rhizostomatidae) in Pakistani waters. International Journal of Biological Research, 3(1):49-51., Figs. 1-3) surprisingly reported R. nomadica from Keti Bundar, Pakistan with rough exumbrellar surface bearing minute, blunt tuberculation and warts. However, the Rhopilema jellyfish in their figures is clearly showing presence of minute reddish-brown warts on marginal area of umbrella, which are however, characteristic only of R. hispidum. Moreover, the authors mentioned the presence of vermicular filaments terminal to the mouth arms in their specimens though, not visible in the figures.

According to the literature, although Rhopilema nomadica was described from the Red Sea (Galil et al., 1990GALIL, B.S.; SPANIER, E. & FERGUSON, W.W. 1990. The scyphomedusae of the Mediterranean coast of Israel, including two lessepsian migrants new to the Mediterranean. Zoologische Mededelingen, 64:95-105.) no further records provide solid evidence of its presence in the Indian Ocean so far. In fact, the species is expanding its distribution into the Mediterranean Sea (Yahia et al., 2013YAHIA, M.N.D.; YAHIA, O.K.D.; GUEROUN, S.K.M.; AISSI, M.; DEIDUN, A.; FUENTES, V. & PIRAINO, S. 2013. The invasive tropical scyphozoan Rhopilema nomadica Galil, 1990 reaches the Tunisian coast of the Mediterranean Sea. BioInvasions Records, 2(4):319-323.). In contrast, Rhopilema hispidum is widespread in Indo-Pacific waters (Gul & Morandini, 2015GUL, S. & MORANDINI, A.C. 2015. First record of the jellyfish Rhopilema hispidum (Cnidaria: Scyphozoa) from the coast of Pakistan. Marine Biodiversity Records , 8(e30):1-4.).

We consider that the jellyfish reported by Tahera & Kazmi (2015TAHERA, Q. & KAZMI, Q.B. 2015. Occurrence of Rhopilema nomadica Galil, 1990 (Cnidaria: Scyphozoa: Rhizostomeae: Rhizostomatidae) in Pakistani waters. International Journal of Biological Research, 3(1):49-51.) as R. nomadica is possibly misidentified R. hispidum previously reported from Pakistan (Gul & Morandini, 2015GUL, S. & MORANDINI, A.C. 2015. First record of the jellyfish Rhopilema hispidum (Cnidaria: Scyphozoa) from the coast of Pakistan. Marine Biodiversity Records , 8(e30):1-4.).

Although the life cycle of three Rhopilema species were formally described (Calder, 1973CALDER, D.R. 1973. Laboratory observations on the life history of Rhopilema verrilli (Scyphozoa: Rhizostomeae). Marine Biology, 21:109-114.; Ding & Chen, 1981DING, G. & CHEN, J. 1981. The life history of Rhopilema esculenta Kishinouye. Journal of Fisheries of China, 5:93-104.; Lotan et al., 1992LOTAN, A.; BEN-HILLEL, R. & LOYA, Y. 1992. Life cycle of Rhopilema nomadica: a new immigrant scyphomedusan in the Mediterranean. Marine Biology , 112:237-242.), none of these studies focused on the recognition of developmental changes in characters used for the systematics of the genus. That is the next step for a better understanding of morphology of the species in this genus, to clear if we are dealing with species that present a huge variation in general colour pattern and also in the shape of appendages.

ACKNOWLEDGEMENTS

We thank Dr. C. Lüter (Museum für Naturkunde, Berlin, Germany) for providing access to the scyphozoan specimens at the museum, Prof. Dr. Gerhard Jarms (Universität Hamburg, Germany) for providing living polyp specimens and Mr. Hamid B. Osmany (Marine Fisheries Department, Karachi, Pakistan) for help in photography of Pakistani specimens. We also thank anonymous reviewers and editors for additional comments on the manuscript. ACM was supported by Alexander von Humboldt Foundation, and grants 2010/50174-7, 2011/50242-5 São Paulo Research Foundation (FAPESP), and by CNPq (301039/2013-5). This is a contribution of NP-BioMar, USP.

REFERENCES

  • BROWNE, E.T. 1926. Report on the Medusae. Transactions of the Zoological Society of London, 22:105-115.
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  • DALY, M.; BRUGLER, M.R.; CARTWRIGHT, P.; COLLINS, A.G.; DAWSON, M.N.; FAUTIN, D.G.; FRANCE, S.C.; MCFADDEN, C.S.; OPRESKO, D.M.; RODRIGUEZ, E.; ROMANO, S.L. & STAKE, J.L. 2007. The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus. Zootaxa, 1668:127-182.
  • DING, G. & CHEN, J. 1981. The life history of Rhopilema esculenta Kishinouye. Journal of Fisheries of China, 5:93-104.
  • GALIL, B.S.; SPANIER, E. & FERGUSON, W.W. 1990. The scyphomedusae of the Mediterranean coast of Israel, including two lessepsian migrants new to the Mediterranean. Zoologische Mededelingen, 64:95-105.
  • GERSHWIN, L. & ZEIDLER, W. 2008. Two new jellyfishes (Cnidaria: Scyphozoa) from tropical Australian waters. Zootaxa, 1764:41-52.
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  • GUL, S. & MORANDINI, A.C. 2013. New records of scyphomedusae from Pakistan coast: Catostylus perezi and Pelagia cf. noctiluca (Cnidaria: Scyphozoa). Marine Biodiversity Records, 6(e86):1-6.
  • GUL, S. & MORANDINI, A.C. 2015. First record of the jellyfish Rhopilema hispidum (Cnidaria: Scyphozoa) from the coast of Pakistan. Marine Biodiversity Records , 8(e30):1-4.
  • GUL, S.; MOAZZAM, M. & GALIL, B.S. 2014. Occurrence of Marivagia stellata (Scyphozoa: Rhizostomeae: Cepheidae) along the coast of Pakistan, northern Arabian Sea. Marine Biodiversity Records , 7(e112):1-2.
  • GUL, S.; MOAZZAM, M. & MORANDINI, A.C. 2015a. Crowned jellyfish (Cnidaria: Scyphozoa: Rhizostomeae: Cepheidae) from waters off the coast of Pakistan, northern Arabian Sea. Check List, 11(1551):1-4.
  • GUL, S.; MORANDINI, A.C. & MOAZZAM, M. 2015b. First record of the crowned jellyfish Netrostoma coerulescens (Cnidaria: Scyphozoa) from Pakistani waters. Marine Biodiversity Records , 8(e156):1-4.
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  • LOTAN, A.; BEN-HILLEL, R. & LOYA, Y. 1992. Life cycle of Rhopilema nomadica: a new immigrant scyphomedusan in the Mediterranean. Marine Biology , 112:237-242.
  • OMORI, M. & KITAMURA, M. 2004. Taxonomic review of three Japanese species of edible jellyfish (Scyphozoa: Rhizostomeae). Plankton Biology and Ecology, 51:36-51.
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  • YAHIA, M.N.D.; YAHIA, O.K.D.; GUEROUN, S.K.M.; AISSI, M.; DEIDUN, A.; FUENTES, V. & PIRAINO, S. 2013. The invasive tropical scyphozoan Rhopilema nomadica Galil, 1990 reaches the Tunisian coast of the Mediterranean Sea. BioInvasions Records, 2(4):319-323.

Publication Dates

  • Publication in this collection
    Dec 2016

History

  • Accepted
    17 July 2016
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E-mail: einicker@usp.br