Open Access

Hydroids (Cnidaria, Hydrozoa) from marine environments in Taiwan

  • Li-Chun Tseng1,
  • Cheng-Han Wu1,
  • Wen-Hung Twan2,
  • Zhi-Can Tang3 and
  • Jiang-Shiou Hwang1Email author
Zoological Studies201453:29

DOI: 10.1186/s40555-014-0029-z

Received: 17 February 2014

Accepted: 19 May 2014

Published: 10 June 2014

Abstract

Background

Hydroids were identified from extensive surveys of benthos, and especially from worm tubes, in the Danshuei River estuary (DRE), in the northeast Taiwan Strait (NETS), and on a reef in Nanwan Bay (NWB), Taiwan.

Results

A total of 33 species (12 of them identified to generic and 1 to family rank only), referable to 22 genera, 12 families, and 2 orders, were distinguished. The majority of them were referable to order Leptothecata (84.8%) with the remainder being assigned to order Anthoathecata (15.2%). The only species found at NETS was Monoserius pennarius (Linnaeus, 1758). Plumularia habereri (Stechow, 1909) was recorded at both DRE and NWB. The known range of Dynamena nanshaensis (Tang, 1991), recorded for the second time, is extended further north. Six species (Aglaophenia latecarinata, Plumularia floridana, Diphasia palmata, Dynamena brevis, Dynamena obliqua, Synthecium elegans) are recorded for the first time in waters adjacent to the Chinese mainland.

Conclusions

Species composition and abundances varied considerably from one environment to another and especially between DRE and NWB. This study provides the first taxonomic account of the hydroid fauna inhabiting the eastern Taiwan Strait region.

Keywords

Cnidaria Hydroids Reef Worm tube Taiwan East China Sea South China Sea

Background

Hydroids are largely sessile epibenthic cnidarians that inhabit all marine habitats from shallow to abyssal waters (Vervoort [1966]). Since hydroids feed on plankton, they likely play an important role in marine ecosystems (Gili and Hughes [1995]). Hydroids are often found growing on other marine organisms, including sponges (Calder [1991b]), other hydroids (Tang [1991b]), deep-sea corals (Henry [2001]), molluscs (Kubota et al. [1999]), arthropods (Genzano [2002]), and macroalgae (Fraschetti et al. [2006]; Cunha and Jacobucci [2010]; Oliveira and Marques [2007]). Except for the well-known association between proboscidactylid polyps and sabellid polychaetes (Schuchert [2009]), relatively, little attention has been given to worm tubes as substrates for hydroids.

Several studies on hydroids from temperate and tropical waters of the northwest Pacific have been carried out over the past few decades. These include the fauna of Japan to the north (e.g., Hirohito [1988], [1995]), the Yellow Sea (Tang and Huang [1986]), the East China Sea (Tang and Xu [1978]; Liu and Li [2002]), the Taiwan Strait (Kubota et al. [1999]; Xu and Huang [2004]), the Philippine Islands (Hargitt [1924]), and the South China Sea (Tang [1991a], [b], [1998]; Yeh [1995]; Zhao [1998]). Liu ([2008]) and Huang and Lin ([2012]) provided checklists and illustrations of Hydrozoa in waters adjacent to mainland China. A number of earlier reports explored the rich hydroid fauna of the western Pacific Ocean (for some references, see Vervoort [1995]). Literature is scanty on hydroids from waters of northeast Taiwan (Jäderholm [1903]), from the coast of southern Taiwan (Yeh [1995]), around Taiwan (Hwang and Shao [1998]), in the middle part of the western coast of Taiwan (Kubota et al. [1999]), in the Taiwan Strait (Xu and Huang [2004]), and from coral reef areas of the country (Fontana et al. [2012]). So far, distribution and species composition of hydroids have not been explored in the eastern Taiwan Strait. The present study was undertaken to establish a baseline of information on the group in waters of Taiwan.

Methods

Field sampling and samples treatment

Investigations on hydroids were undertaken in three environmentally distinct regions: (1) the Danshuei River estuary (DRE, 25°10′ N, 121°24′ E) in north Taiwan, (2) the northeastern Taiwan Strait (NETS, 24°48′ N, 119°55′ E), and (3) the reef zone in Nanwan Bay (NWB, 21°56′ N, 120°46′ E), south Taiwan (Figure 1, Table 1). Samples from DRE and NETS were collected by bottom trawl onboard the Ocean Research Vessel II and from a fishery vessel, respectively. At NWB, hydroid sampling on a coral reef was undertaken at 12 to 15 m depth using self-contained underwater with breathing apparatus (SCUBA) and at 100 m depth using a remotely operated vehicle (ROV). A total of 24 worm tube specimens collected at the coral reef site by SCUBA diving, and two additional ones using the ROV, were examined for hydroid associates. Specimens were preserved in seawater with 5% to 10% buffered formalin immediately after collection.
Figure 1

Map of Taiwan and vicinity, showing the three sampling locations. DRE, Danshuei River estuary; NETS, northeastern Taiwan Strait; NWB, Nan-Wan Bay.

Table 1

Sampling location, date, depth, and methods

Sampling area

Date

Depth (m)

Methods

Estuary of Danshuei River

21 Aug. 2009

10 to 15

ORII CR-1662, bottom trawling

Northeastern Taiwan Strait

14 Mar. 2009

50 to 60

Fishery boat, bottom trawling

Reef zone of Nan-Wan Bay

22 Dec. 2009

5 to 10

SCUBA diving

Reef zone of Nan-Wan Bay

Aug. 2008

100

Remotely operated vehicle

ORII, Ocean Research Vessel II.

Hydroids identification

In the laboratory, each sample for taxonomic identification was dissected on a glass slide with the help of a dissecting microscope. Identification of species was undertaken using recent taxonomic keys and relevant references, including Vervoort ([1966]), Millard ([1975]), Rees and Vervoort ([1987]), Calder ([1988], [1991a], [1997], [2012]), Hirohito ([1988], [1995]), Cornelius ([1995a], [1995b]), Schuchert ([1996], [1998], [2001], [2003], [2004]), Marques ([2001]), and Vervoort and Watson ([2003]).

Results

Hydroids from DRE and NETS, collected by bottom trawl, were often found together with rocks. However, those actually attached to rock at DRE represented less than 30% (N > 100) of the total number of specimens. At NETS, hydroids were found tangled in the trawl net because most colonies were fragmented and longer than 40 cm. Worm tubes, an important substrate at NWB, supported a substantial growth of sessile organisms including hydroids, soft corals, ascidians, and red algae (Figure 2). As for hydroids, they were associated with worm tubes at the rate of 100% (N = 24). Most hydroids attached to middle and distal surfaces of the tube, while fewer were present at the base.
Figure 2

Photographs of worm tubes in field (a) and colony size (b) taken from the coast of Nan-Wan Bay.

In our samples, 22 species were identified from riverbed environments at DRE, 11 epizoic species were found on worm tubes at NWB, and 1 species occurred in samples from NETS (Table 2). Overall, a total of 33 hydroid species (12 of them identified solely to generic level and 1 to family level) referable to 22 genera, 12 families, and 2 orders, were discovered. Most of the species belonged to order Leptothecata (84.8%), with the remainder being referable to order Anthoathecata (15.2%). Taking samples from all study areas together, hydroids of the family Sertulariidae (comprising 14 species, with 3 identified to generic level only) contained the largest number of species. Six species (Aglaophenia latecarinata Allman, 1877; Plumularia floridana Nutting, 1900; Diphasia palmata Nutting, 1905; Dynamena brevis (Fraser, 1935); Dynamena obliqua Lamouroux, 1816; and Synthecium elegans Allman, 1872) are recorded here for the first time from waters adjacent to mainland China (Table 2).
Table 2

Taxonomic list of hydroids collected from north and south Taiwan

Scientific classification

Present study

Recorded in Liu ([2008])

Hydrozoa

  

 Order Anthoathecata Cornelius, 1992

  

  Bougainvilliidae Lütken, 1850

  

   Bimeria sp.

NWB

-

  Eudendriidae L. Agassiz, 1862

  

   Eudendrium sp. 1

DRE

-

   Eudendrium sp. 2

DRE

-

   Eudendrium sp. 3

NWB

-

  Pennariidae McCrady, 1859

  

   Pennaria disticha Goldfuss, 1820

NWB

ECS, SCS, CTR-STR, MEDIT

 Order Leptothecata Cornelius, 1992

  

  Aglaopheniidae Marktanner-Turneretscher, 1890

  

   Aglaophenia cupressina Lamouroux, 1816

NWB

XSHA, NSHA, SAFRI, EAFRI, INMA, AU

   Aglaophenia latecarinata Allman, 1877a

DRE

-

   Macrorhynchia philippina Kirchenpauer, 1872

DRE

SCS, CTR-STR

   Monoserius pennarius (Linnaeus, 1758)

NETS

ECS, SCS, IWPAC

  Campanulariidae Johnston, 1836

  

   Campanulariidae indet.

DRE

-

   Obelia dichotoma (Linnaeus, 1758)

DRE

YS, ECS, SCS, COSMOP

  Halopterididae Millard, 1962

  

   Halopteris sp.

NWB

-

  Hebellidae Fraser, 1912

  

   Hebella sp.

DRE

-

  Kirchenpaueriidae Stechow, 1921

  

   Kirchenpaueria sp.

DRE

-

  Lafoeidae Hincks, 1868

  

   Zygophylax sp.

DRE

-

  Plumulariidae Agassiz, 1862

  

   Nemertesia sp.

DRE

-

   Plumularia habereri Stechow, 1909

DRE, NWB (ROV)

SCS, ID, JP, CARIB

   Plumularia floridana Nutting, 1900a

NWB

-

  Sertulariidae Lamouroux, 1812

  

   Diphasia digitalis (Busk, 1852)

DRE

SCS, CTR-STR

   Diphasia palmata Nutting, 1905a

DRE

-

   Dynamena brevis (Fraser, 1935)a

DRE

-

   Dynamena crisioides Lamouroux, 1824

NWB

ECS, SCS, CTR-STR

   Dynamena disticha (Bosc, 1802)

DRE

ECS, SCS, ID, JP, BM, BR (Synonym, Dynamena cornicina McCrady, 1859)

   Dynamena nanshaensis Tang, [1991]

NWB (ROV)

NSHA

   Dynamena obliqua Lamouroux, 1816a

DRE

--

   Dynamena quadridentata (Ellis and Solander, 1786)

DRE

ZJ, FJN, HK, TKG, CTR-STR

   Idiellana pristis (Lamouroux, 1816)

DRE

ECS, SCS, CTR-STR

   Salacia sp.

NWB

-

   Sertularella diaphana (Allman, 1885)

DRE

ECS, CTR-STR

   Sertularella sp.

NWB

-

   Sertularia loculosa Busk, 1852

DRE

SCS, WAFRI, SAFRI, LK, ID, JP (Synonym, Sertularia ligulata Thornely, 1904)

   Sertularia sp.

DRE

--

  Syntheciidae Marktanner-Turneretscher, 1890

  

   Synthecium elegans Allman, 1872a

DRE

--

aFirst record of the species in waters adjacent Mainland China. DRE, Danshuei River estuary; NETS, northeastern Taiwan Strait; NWB, Nan-Wan Bay. Distributions of certain species as recorded in Liu ([2008]). AU, Australia; BM, Bermuda; BR, Brazil; CARIB, Caribbean; COSMOP, Cosmopolitan; CTR-STR, Circumtropical-Subtropical; EAFRI, East Africa; ECS, East China Sea; FJN, Fujian; HK, Hong Kong; ID, Indonesia; INMA, Indo-Malaysia; IWPAC, Indo-West Pacific; JP, Japan; LK, Sri Lanka; MEDIT, Mediterranean; NSHA, Nansha Islands; SAFRI, South Africa; SCS, South China Sea; TKG, Tonkin Gulf; WAFRI, West Africa; XSHA, Xisha Islands; YS, Yellow Sea; ZJ, Zhejiang.

Hydroid species composition and abundance varied considerably between DRE and NWB. Only one species, Plumularia habereri Stechow, 1909, was found at both locations. Species found in the deep zone at NWB by the ROV were P. habereri and Dynamena nanshaensis Tang, [1991a]. At NETS, only one species was found, the robust aglaopheniid Monoserius pennarius (Linnaeus, 1758). Its colony had a single stem, with pinnate side branches.

Discussion

Historical records of hydroids in coastal waters around Taiwan are limited. Yeh ([1995]) reported seven species of Plumulariidae collected in the southern part of the island. Hwang and Shao ([1998]) described two venomous hydroids from the same location. Kubota et al. ([1999]) provided the first record of Eugymnanthea japonica (Yamada, 1950) (Leptothecata, Eirenidae), from western Taiwan, where it was associated with two bivalve species (Crassostrea gigas (Thunberg, 1793), Perna viridis (Linnaeus, 1758)). Recently, Fontana et al. ([2012]) reported what was believed to be a cosmopolitan and possibly genus-specific association between Zanclea spp. and its coral hosts, based in part on hydroids from Acropora corals on reefs in Kenting and the Penghu Islands, Taiwan. Previous literature provided li`mited data of hydroids in this area, as shown in the extensive review on hydroid species in Chinese waters by Liu ([2008]). His report recorded 615 species of Hydrozoa in waters adjacent to mainland China, including the East China Sea, western Taiwan Strait, and South China Sea. Shortly after, Huang and Lin ([2012]) provided illustrations of 829 species of Hydrozoa in waters adjacent to mainland China. However, hydroids in the eastern Taiwan Strait were still inadequately known, and our study is the first to document the hydroid fauna in waters of the region.

The stinging hydroid Aglaophenia cupressina Lamouroux, 1816 is widely distributed on the coral reefs of the Bunaken area (Ricciardi [2007]). Tang ([1991a]) noted that the sting of A. cupressina, frequent on reefs in the Indo-West Pacific area, produces an itchy rash in humans. Hwang and Shao ([1998]) reported A. cupressina and the feather hydroid Halocordyle disticha (a synonym of Pennaria disticha Goldfuss, 1820), both venomous to humans, in Taiwan. Eldredge and Smith ([2001]) noted that the sting of P. disticha causes mild irritation in humans. The two species were recorded on worm tubes from NWB in our study.

The colonial hydroid P. habereri was found in a riverbed environment at DRE and by ROV in NWB. This species is widely distributed in the South China Sea, Indonesia, and Japan (Liu [2008]) at varied depth ranges, e.g., 10 to 35 m (Schuchert [2003]) and 1 to 4 m (Calder and Kirkendale [2005]). In our study, the species was collected at a depth of 100 m, the deepest ever recorded. In addition, waters in the estuarine environment at DRE have a wide range of salinities. We conclude that P. habereri is a euryhaline species.

A noteworthy discovery in our samples was D. nanshaensis, collected from hard substrates on the reef at NWB. This is only the second record of the species since its original description from the Nansha Islands (Tang [1991a]). Thus far, D. nanshaensis is known only from the South China Sea, specifically from the type locality (Nansha Islands, about 10° N) and now from our study in Nanwan Bay (about 21°56′ N), south Taiwan, the northernmost record of the species.

The macrobenthic hydroid M. pennarius was especially abundant at Nansha Island (Zhao [1998]; Tang [1991b]). Its dense population there establishes an ecological environment described as an ‘imitative steppe on sea floor’ (Zhao [1998]) and a ‘sea-floor prairie’ (Tang [1991b]). M. pennarius is widely distributed in the Indo-Malayan and Sino-Japanese subregions of the Indo-West Pacific region (Tang [1991b]), including India, Ceylon, Indonesia, Philippines, Taiwan, southern Japan, Palau, and New Zealand (Schuchert [2003]). It is a species of warm waters and high salinity in the East China Sea (Liu and Li [2002]). Common macrobenthic communities there include: Callianassa spp., M. pennarius-Stellaster equestris (Tang and Xu [1978]), and S. equestris, M. pennarius-Callianassa japonica (Liu and Li [2002]). Our material from NETS confirmed that colony length of M. pennarius ranges from 30 to 100 cm and that it inhabits muddy-sandy seabeds at depths of 50 to 100 m (Tang [1991b]; Schuchert [2003]; Vervoort and Watson [2003]).

Samples from DRE included the cosmopolitan species Dynamena disticha (Bosc, 1802). Liu ([2008]) reported D. disticha from the East China Sea, South China Sea, Indonesia, Japan, Bermuda, and Brazil. Puce et al. ([2009]) reported that D. disticha can be found in association with algae, hydroids, barnacles, rocks, and concretions in the Mediterranean Sea. This suggests high adaptability to diverse environments, and indeed, D. disticha has a broad distribution. Some hydroids form communities in association with other co-occurring marine organisms. Fraschetti et al. ([2006]) reported D. disticha and Obelia dichotoma (Linnaeus, 1758) in association with brown algae Cystoseira amentacea (C.Agardh) Bory de Saint-Vincent, 1832 in the Mediterranean Sea. Both hydroid species were recorded from DRE in this study. Cunha and Jacobucci ([2010]) reported 16 species of hydroids on fronds of the alga Sargassum cymosum C.Agardh, 1820 in subtropical Brazil. Among species in their list, A. latecarinata, D. disticha, and O. dichotoma were also found in our study. Gravier-Bonnet and Bourmaud ([2006]) recorded 95 hydroid species from a reef zone in the southwest Indian Ocean, with P. disticha, A. cupressina, Macrorhynchia philippina, and Dynamena crisioides being common to abundant. Three of these species (P. disticha, A. cupressina, and D. crisioides) were also identified from samples in the reef zone at NWB.

Notably, six species (A. latecarinata, P. floridana, D. palmata, D. brevis, D. obliqua, and S. elegans) identified from DRE have never been recorded previously in seas of China (Liu [2008]). This reveals that the assemblage of hydroids in the eastern Taiwan Strait differs significantly from those in coastal waters of mainland China. Shao ([1998]) estimated that the number of marine species around Taiwan represented 10% of the total number known worldwide. The island of Taiwan, located in the west Pacific Ocean, has a total of 1,566 kilometers of coastline (including the Penghu Islands) (Tung [2006]), and complex current systems influence the island (Tseng et al. [2013a], [b]). Diverse water masses of the East China Sea, South China Sea, China Coastal Current, and Kuroshio Current have a pronounced effect on the endemic marine biota. Varied marine habitats promote a highly diversity biota around Taiwan Island (Shao [2009]). Expanding the geographic scope and frequency of marine surveys would potentially show that the indigenous hydroid fauna in waters around Taiwan is even richer.

Conclusions

The study documents a unique and heterogeneous assemblage of hydroids found in three quite dissimilar marine environments in Taiwan (DRE, NETS, and NWB). The DRE riverbed system supports a comparatively richer hydroid fauna than a group of species associated with worm tubes from NWB. The three sampling sites differed significantly in faunal composition and numbers of species. Differences are attributed primarily to disparities in substrate types and salinity characteristics among the sites. Studies on developmental stages of hydroids under laboratory conditions are needed to better understand their life cycles in nature.

Declarations

Acknowledgements

This manuscript was completed thanks in large part to Professor Z.-C. Tang, who passed away on 18 August 2011 during the study. His efforts and dedication to the taxonomic component of the work are greatly appreciated. Thanks to Mrs. Tang (Mei-Jun Huang) for assisting in screening hydroid samples. Thanks are due to Professor Santosh Kumar Sarkar (Department of Marine Science, University of Calcutta, India), whose comments and advice improved the paper. We thank two anonymous referees for their editing and suggestions that substantially improved the quality of the manuscript. We are grateful for financial support from the Ministry of Science and Technology of Taiwan via projects NSC 100-2611-M-019-010; NSC 101-2611-M-019-011 and NSC 102-2611-M-019-003 to J.-S. Hwang. Our appreciation is extended to members of J.-S. Hwang's laboratory and to the captain and technicians of Ocean Research Vessel II for their assistance during field sampling of hydroids in Danshuei River estuary.

Authors’ Affiliations

(1)
Institute of Marine Biology, College of Life Sciences, National Taiwan Ocean University
(2)
Institute of Life Science, National Taitung University
(3)
Institute of Oceanology, Academia Sinica

References

  1. Calder DR: Shallow-water hydroids of Bermuda: the Athecatae. R Ont Mus Life Sci Contrib 1988, 148: 1–107.Google Scholar
  2. Calder DR: Shallow-water hydroids of Bermuda: the Thecatae, exclusive of Plumularioidea. R Ont Mus Life Sci Contrib 1991, 154: 1–140.Google Scholar
  3. Calder DR: Association between hydroid species assemblages and substrate types in the mangal at Twin Cays, Belize. Can J Zool 1991, 69: 2067–2074. 10.1139/z91-288View ArticleGoogle Scholar
  4. Calder DR: Shallow-water hydroids of Bermuda: superfamily Plumularioidea. R Ont Mus Life Sci Contrib 1997, 161: 1–107.Google Scholar
  5. Calder DR: On a collection of hydroids (Cnidaria, Hydrozoa, Hydroidolina) from the west coast of Sweden, with a checklist of species from the region. Zootaxa 2012, 3171: 1–77.Google Scholar
  6. Calder DR, Kirkendale L: Hydroids (Cnidaria, Hydrozoa) from shallow-water environments along the Caribbean coast of Panama. Caribb J Sci 2005, 41: 476–491.Google Scholar
  7. Hargitt CW: Hydroids of the Philippine Islands. Philipp J Sci 1924, 24: 467–505.Google Scholar
  8. Cornelius PFS: North-west European thecate hydroids and their medusae. Part 1: Introduction, Laodiceidae to Haleciidae. Synopses Br Fauna (ns) 1995a, 50: 347.Google Scholar
  9. Cornelius PFS: North-west European thecate hydroids and their medusae. Part 2: Sertulariidae to Campanulariidae. Synopses Br Fauna (ns) 1995b, 50: 386.Google Scholar
  10. Cunha AF, Jacobucci GB: Seasonal variation of epiphytic hydroids (Cnidaria: Hydrozoa) associated to a subtropical Sargassum cymosum (Phaeophyta: Fucales) bed. Zoologia 2010, 27: 945–955. 10.1590/S1984-46702010000600016View ArticleGoogle Scholar
  11. Eldredge LG, Smith CM: A guidebook of marine introduced species in Hawaii. Bishop Mus Tech Rep 2001, 21: 1–80.Google Scholar
  12. Fraschetti S, Terlizzi A, Bevilacqua S, Boero F: The distribution of hydroids (Cnidaria, Hydrozoa) from micro- to macro-scale: spatial patterns on habitat-forming algae. J Exp Mar Biol Ecol 2006, 339: 148–158. 10.1016/j.jembe.2006.07.007View ArticleGoogle Scholar
  13. Fontana S, Keshavmurthy S, Hsieh HJ, Denis V, Kuo CY, Hsu CM, Leung JK, Tsai WS, Wallace CC, Chen CA: Molecular evidence shows low species diversity of coral-associated hydroids in Acropora corals. PLoS ONE 2012, 7: e50130. 10.1371/journal.pone.0050130View ArticleGoogle Scholar
  14. Genzano GN: Associations between pycnogonids and hydroids from the Buenos Aires littoral zone, with observations on the semi-parasitic life cycle of Tanystylum orbiculare (Ammotheiidae). Sci Mar 2002, 66: 83–92. 10.3989/scimar.2002.66n183View ArticleGoogle Scholar
  15. Gili J, Hughes RG: The ecology of marine benthic hydroids. Oceanogr Mar Biol Annu Rev 1995, 33: 351–426.Google Scholar
  16. Gravier-Bonnet N, Bourmaud C: Hydroids (Cnidaria, Hydrozoa) of coral reefs: preliminary results on community structure, species distribution and reproductive biology in Juan de Nova Island (Southwest Indian Ocean). West Indian Ocean J Mar Sci 2006, 5: 123–132.Google Scholar
  17. Henry L-A: Hydroids associated with deep-sea corals in the boreal north-west Atlantic. J Mar Biol Ass U K 2001, 81: 163–164. 10.1017/S0025315401003502View ArticleGoogle Scholar
  18. The hydroids of Sagami Bay. (Part 1. Athecata). Publs. Biol. Lab., ImpHousehold, Tokyo; 1988.Google Scholar
  19. The hydroids of Sagami Bay II.. Thecata. Publs. Biol. Lab., Imp. Household, Tokyo; 1995.Google Scholar
  20. Huang Z, Lin M: An illustrated guide to species in China’s Seas (volume 3) - Animalia (1): Porifera, Cnidaria Plathyelminthes, Nemertinea, Nematoda, Acanthocephala, Rotifera, Gastrotricha, Kinorhyncha, Priapulida, Annelida, Sipuncula, Echiura. In The living species and their illustrations in China’s Seas (Part II). China Ocean Press, Beijing, China; 2012. in ChineseGoogle Scholar
  21. Hwang DF, Shao KT: Illustrations of toxic fishes and shellfishes in Taiwan. Cheng Chung Book Co., Ltd. Taipei; 1998.Google Scholar
  22. Jäderholm E: Aussereuropäischen Hydroiden im schwedischen Reichsmuseum. Arkiv Zoologi 1903, 1: 259–312.Google Scholar
  23. Kubota S, Dai C-F, Lin C-L, Ho J-S: The first occurrence of the paedomorphic derivative hydrozoan Eugymnanthea (Leptomedusae, Eirenidae) from Taiwan, with a report of a new host. Publ Seto Mar Biol Lab 1999, 38: 219–222.Google Scholar
  24. Liu R: Checklist of marine biota of China Seas. Science Press, Academia Sinica, Beijing; 2008.Google Scholar
  25. Liu L-S, Li X-Z: Distribution of macrobenthos in spring and autumn in the East China Sea. Biodiv Sci 2002, 10: 351–358.Google Scholar
  26. Marques AC: Simplifying hydrozoan classification: inappropriateness of the group Hydroidomedusae in a phylogenetic context. Contr Zool 2001, 70: 175–179.Google Scholar
  27. Millard NAH: Monograph on the Hydroida of southern Africa. Ann S Afr Mus 1975, 68: 1–513.Google Scholar
  28. Oliveira OMP, Marques AC: Epiphytic hydroids (Hydrozoa: Anthoathecata and Leptothecata) of the world. Check List 2007, 3: 21–38.View ArticleGoogle Scholar
  29. Puce S, Bavestrello G, Di Camillo CG, Boero F: Long-term changes in hydroid (Cnidaria, Hydrozoa) assemblages: effect of Mediterranean warming? Mar Ecol 2009, 30: 1–14. 10.1111/j.1439-0485.2009.00283.xView ArticleGoogle Scholar
  30. Rees WJ, Vervoort W: Hydroids from the John Murray Expedition to the Indian Ocean, with revisory notes on Hydrodendron, Abietinella, Cryptolaria and Zygophylax (Cnidaria: Hydrozoa). Zool Verh Leiden 1987, 237: 1–209.Google Scholar
  31. Ricciardi F: First record of a color variation of Cuthona diversicolor (Tergipedidae: Aeolidina) feeding on the hydroid, Aglaophenia cupressina , on a North Sulawesi (Indonesia) coral reef. Zool Stud 2007, 46: 473.Google Scholar
  32. Schuchert P: The marine fauna of New Zealand: athecate hydroids and their medusae. NZ Oceanogr Inst Mem 1996, 106: 1–159.Google Scholar
  33. Schuchert P: How many hydrozoan species are there? Zool Verh Leiden 1998, 323: 209–219.Google Scholar
  34. Schuchert P: Hydroids of Greenland and Iceland (Cnidaria, Hydrozoa). Medd Grønland, Biosci 2001, 53: 1–184.Google Scholar
  35. Schuchert P: Hydroids (Cnidaria, Hydrozoa) of the Danish expedition to the Kei Islands. Steenstrupia 2003, 27: 137–256.Google Scholar
  36. Schuchert P: Revision of the European athecate hydroids and their medusae (Hydrozoa, Cnidaria): families Oceanidae and Pachycordylidae. Rev Suisse Zool 2004, 111: 315–369.View ArticleGoogle Scholar
  37. Schuchert P: The European athecate hydroids and their medusae (Hydrozoa, Cnidaria): Filifera Part 5. Rev Suisse Zool 2009, 116: 441–507.View ArticleGoogle Scholar
  38. Shao KT: Marine ecology. National Press Company, Ming Wen Book Co., Ltd, Taipei, Taiwan; 1998.Google Scholar
  39. Shao KT: Marine biodiversity and fishery sustainability. Asia Pac J Clin Nutr 2009, 18: 527–531.Google Scholar
  40. Tang ZC: On a collection of hydroida from the Nansha Islands, Hainan Province, China. Researches on the marine biology of Nansha Islands and neighboring waters collected papers (I):25–36. Ocean Publishing Press, Beijing; 1991a.Google Scholar
  41. Tang ZC: Monoserius pennarius assemblage and its ecological and geographical studies on the continental shelf of waters around the Nansha islands, South China Sea. In Researches on the marine biology of Nansha Islands and neighboring waters collected papers (II). Ocean Publishing Press, Beijing; 1991b:255–261. In Chinese with English abstractGoogle Scholar
  42. Tang ZC: A new species of the genus Filellum (Hydrozoa: Lafoeidae) from the waters around Nansha Islands, South China Sea. Studies on marine fauna and flora and biogeography of the Nansha Islands and neighbouring waters III:42–46. Ocean Publishing Press, Beijing; 1998.Google Scholar
  43. Tang ZC, Huang MJ: A new species of the genus Symplectoscyphus (Hydroida) from the Huanghai Sea. Chin J Oceanol Limnol 1986, 4: 317–318.View ArticleGoogle Scholar
  44. Tang ZC, Xu FS: Preliminary analysis of the distribution and community of macrobenthos from the continental shelf of the East China Sea. In Transactions of the Continental Shelf of the East China Sea. Institute of Oceanology, Chinese Academy of Sciences, Qingdao; 1978:156–164. In ChineseGoogle Scholar
  45. Tseng L-C, Dahms H-U, Chen Q-C, Hwang J-S: Geospatial variability of autumn community structure on epipelagic zooplankton in the upper layer of the northern South China Sea. Zool Stud 2013a, 52: 2. 10.1186/1810-522X-52-2View ArticleGoogle Scholar
  46. Tseng L-C, Hung J-J, Chen Q-C, Hwang J-S: Seasonality of the copepod assemblages associated with interplay waters off northeastern Taiwan. Helgoland Mar Res 2013b, 67: 507–520. 10.1007/s10152-012-0339-7View ArticleGoogle Scholar
  47. Tung C-Y: Prospects of the Taiwan Strait west coast economic zone. In Trading arrangements in the Pacific Rim: ASEAN and APEC (New York: Oceana Publications, May 2006), Document III. C Edited by: Davidson P. 2006, 1–8.Google Scholar
  48. Vervoort W: Bathyal and abyssal hydroids. Galathea Rep 1966, 8: 97–174.Google Scholar
  49. Vervoort W: Bibliography of Leptolida (non-siphonophoran Hydrozoa, Cnidaria). Works published after 1910. Zool Verh Leiden 1995, 301: 1–432.Google Scholar
  50. Vervoort W, Watson JE: The marine fauna of New Zealand: Leptothecata (Cnidaria: Hydrozoa) (Thecate hydroids). Natl Inst Water Atmos Res (NIWA) Biodivers Mem 2003, 119: 1–538.Google Scholar
  51. Xu Z, Huang J: On new species and record of Laingiomedusae and Leptomedusae (Cnidaria, Hydrozoa, Hydroidomedusae) in the Taiwan Strait. J Xiamen Univ (Nat Sci) 2004, 43: 107–114.Google Scholar
  52. Yeh Y-C: Morphology of Plumulariidae hydroida in Taiwan. National Sun Yat-Sen University, Kaohsiung, Taiwan; 1995.Google Scholar
  53. Zhao H: Type and evolution of landscapes of Nansha Islands. Chin Geogr Sci 1998, 8: 144–151. 10.1007/s11769-997-0028-7View ArticleGoogle Scholar

Copyright

© Tseng et al.; licensee Springer. 2014

This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly credited.