INTRODUCTION
The recent literature checklist (Tsuda & Walsh, Reference Tsuda and Walsh2013) of the marine benthic algae in central Polynesia shows that previous algal floristics on the marine Rhodophyta (red algae), Phaeophyceae (brown algae) and Chlorophyta (green algae) in the contiguous South Pacific islands of Tonga (108 species), Niue (3 species) and Cook Islands (106 species) between 158° and 178°W longitude are wanting. The low algal species numbers (excluding Cyanobacteria) in these island groups are in contrast to the adjacent Samoan Archipelago (Samoa and American Samoa) to the north with 359 species (Skelton & South, Reference Skelton and South1999, Reference Skelton and South2004, Reference Skelton and South2007) and the adjacent Melanesian islands of Fiji to the west with 434 species (N'Yeurt et al., Reference N'Yeurt, South and Keats1996; South & Skelton, Reference South and Skelton2003). N'Yeurt & Payri (Reference N'Yeurt and Payri2006, Reference N'Yeurt and Payri2007, Reference N'Yeurt and Payri2010) include 309 taxa of brown, green and red algae in French Polynesia which lies east of the Cook Islands.
The expansive Tongan Archipelago (Figure 1) is the southernmost island group in central Polynesia and comprises some 170 islands scattered over 748,000 km2; some 36 islands are inhabited with a total population of about 102,000 (London Times, 1997; National Geographic Society, 2005). Previous records of 108 documented Tongan algal species and one Cyanobacteria species were reported in Harvey & Bailey (Reference Harvey and Bailey1851), Martens (Reference Martens1866), Grunow (Reference Grunow1874, Reference Grunow1915), Bailey & Harvey (Reference Bailey, Harvey and Gray1874), Dickie (Reference Dickie1876), Yamada (Reference Yamada1930), Howe (Reference Howe1932), Hillis (Reference Hillis1959), Taylor (Reference Taylor1964, Reference Taylor1966), Ducker (Reference Ducker1967), Papenfuss (Reference Papenfuss1967), Skelton & South (Reference Skelton and South2007) and N'Yeurt & Payri (Reference N'Yeurt and Payri2007). Aside from the few incidental records of Tongan algae in the latter two references on Samoa and French Polynesia, respectively, the previous 12 references were published more than 46 years ago.
Map of the Tongan Archipelago, showing the main island groups and localities.
Seven additional species, with type specimens credited to the islands of Tonga, were not listed in Tsuda & Walsh (Reference Tsuda and Walsh2013). The species include three Rhodophyta, i.e. Callophycus seriatus (Harvey ex Kützing) P.C. Silva, Reference Silva1957, Bostrychia radicans (Montagne) Montagne, Reference Montagne1842a [= Bostrychia bipinnata Harvey ex J. Agardh, Reference Agardh1863] and Bostrychia tenella (J.V. Lamouroux) J. Agardh, Reference Agardh1863 [= Bostrychia terrestris (J. Agardh) Harvey ex Zanardini, Reference Zanardini1872], and four Chlorophyta, i.e. Bornetella capitata (Harvey ex E. Wright) J. Agardh, Reference Agardh1887, Bornetella nitida Munier-Chalmas ex Sonder, Reference Sonder and Mueller1880, Caulerpa chemnitzia (Esper) J.V. Lamouroux, Reference Lamouroux1809 [=Caulerpa peltata J.V. Lamouroux, Reference Lamouroux1809] and Caulerpa nummularia Harvey ex J. Agardh, Reference Agardh1873 [=Caulerpa stellata Harvey ex J. Agardh, Reference Agardh1873; =Caulerpa peltata var. stellata (Harvey) Weber-van Bosse 1898 (Belton et al., Reference Belton, Prud'homme van Reine, Huisman, Draisma and Gurgel2014)].
In April and May 2005, the first co-author was invited and afforded the opportunity to examine 1,232 Pacific algal specimens from the British Natural History Museum (BM), London, of which 343 specimens, many unidentified, were collected from Tonga. The algal specimens comprised 64 species, i.e. 1 Cyanobacteria, 20 Rhodophyta, 5 Phaeophyceae and 38 Chlorophyta. A database search of Tongan algae at the Bishop Museum (BISH), Honolulu, by the second co-author and the examination of 43 algal specimens from Tonga, predominantly unidentified, yielded 30 species, i.e. 7 red algae, 4 brown algae and 19 green algae.
MATERIALS AND METHODS
This study is based on two sets of algal specimens identified independently by each of the two co-authors. The bulk of the Tongan specimens deposited at the BM was collected predominantly by William H. Harvey in 1855 and 1874 during his voyage to the Pacific Ocean, and were much later identified by the first co-author in 2005. The second co-author identified the Tongan algal specimens deposited in BISH which were collected by Clyde T. Imada, Bishop Museum, in July 1988, by Winona P. Char in July 1995 and Timothy J. Motley in July 1995. Specimen numbers on few of the BISH specimens were preceded by ‘IA’ and represented specimen numbers of the late Isabella A. Abbott. Where synonyms of currently accepted names were previously used in references recording Tongan algae, these are given in brackets in the Results section. Silva et al. (Reference Silva, Basson and Moe1996) and Guiry & Guiry (Reference Guiry and Guiry2014) were used to identify references, higher algal classification and current species names. Island names for the Tonga Group follow those as listed in Motteler (Reference Motteler2006), while the authors of plant names follow the guidelines of Brummit & Powell (Reference Brummitt and Powell1992).
RESULTS
Thirty-four new algal records for Tonga, i.e. 8 Rhodophyta and 26 Chlorophyta are presented below.
SYSTEMATICS
Phylum RHODOPHYTA
Class FLORIDEOPHYCEAE
Order NEMALIALES
Family GALAXAURACEAE
Dichotomaria obtusata (J. Ellis & Solander) Lamarck, Reference Lamarck1816;
N'Yeurt & Payri, Reference N'Yeurt and Payri2010
SPECIMENS EXAMINED
BM 841094, Tonga, coll. W.H. Harvey, 1855; BM 841098, Vava‘u Group, Vava‘u, coll. W.H. Harvey, 1855.
Family LIAGORACEAE
Liagora viscida (Forsskål) C. Agardh, Reference Agardh1822
SPECIMEN EXAMINED
BM 701498, Tonga, coll. and det. W.H. Harvey, 1855.
Trichogloea requienii (Montagne) Kützing, Reference Kützing1847;
N'Yeurt & Payri, Reference N'Yeurt and Payri2010
(Figure 2A)
New records of marine algae for Tonga: (A) Trichogloea requienii (Montagne) Kützing (BM 000841014;); (B) Titanophora pikeana (Dickie) Feldmann (BISH 692759); (C) Caulerpa plumulifera Zanardini (BM 000841481); (D) Codium extricatum P.C. Silva (BM 000840921); (E) Udotea glaucescens Harvey ex J. Agardh (BM 840810); (F) Valoniopsis pachynema (G. Martens) Børgesen (BISH 550188). Scale bars: A, 10 mm; B, 20 mm; C–F, 10 mm.
SPECIMENS EXAMINED
BM 841012, 841015 and 841016, Ha‘apai Group, Uoleva, coll. W.H. Harvey, coral reef pools, 1855.
Order CORALLINALES
Family CORALLINACEAE
Jania pumila J.V. Lamouroux, Reference Lamouroux1816;
Abbott, Reference Abbott1999
[= Corallina pumila (J.V. Lamouroux) Kützing, Reference Kützing1858]
SPECIMEN EXAMINED
BISH 529618. Tongatapu Group, ‘Eua I., epiphytic on Laurencia sp., reef flat off ‘Ohomua Harbor, coll. C. Imada (s.n.), 26 July 1988.
Order NEMASTOMATALES
Family SCHIZYMENIACEAE
Titanophora pikeana (Dickie) Feldmann, Reference Feldmann1942;
Abbott, Reference Abbott1999
(Figure 2B)
SPECIMEN EXAMINED
BISH 692759. Vava‘u Group, Pangaimotu Island, Lisa Bay, coll. W. Char (IA 22494), 8 July 1995, det. I.A. Abbott, 11 November 2003.
REMARKS
This specimen represents the first record of Titanophora pikeana within central Polynesia outside of Hawaii (Abbott, Reference Abbott1999). The species is also reported from the neighbouring western Melanesian islands of Fiji (South & Skelton, Reference South and Skelton2003). A second species, Titanophora weberae Børgesen, was reported from Fiji by N'Yeurt et al. (Reference N'Yeurt, South and Keats1996), Upolu, Samoa by Skelton & South (Reference Skelton and South2007) and from Tahiti, Society Islands by N'Yeurt & Payri (Reference N'Yeurt and Payri2010).
Order RHODYMENIALES
Family LOMENTARIACEAE
Ceratodictyon repens (Kützing) R.E. Norris, Reference Norris1987
SPECIMEN EXAMINED
BM 840695, Vava‘u Group, Vava‘a, coll. W.H. Harvey 1855.
Order CERAMIALES
Family RHODOMELACEAE
Amansia rhodantha (Harvey) J. Agardh, Reference Agardh1841
SPECIMENS EXAMINED
BM 841369, 841371 and 841394, Ha‘apai Group, Lifuka, coll. W.H. Harvey, 1855.
Tolypiocladia glomerulata (C. Agardh) F. Schmitz,
in Schmitz & Falkenberg, Reference Schmitz, Falkenberg, Engler and Prantt1897
SPECIMENS EXAMINED
BM 841009, 841010 and 841011, Vava‘u Group, Vava‘u, coll. W.H. Harvey, 1855.
Phylum CHLOROPHYTA
Class ULVOPHYCEAE
Order ULVALES
Family ULVACEAE
Ulva clathrata (Roth) C. Agardh, Reference Agardh1811;
N'Yeurt & Payri, Reference N'Yeurt and Payri2007
[= Enteromorpha clathrata (Roth) Greville, Reference Greville1830]
SPECIMEN EXAMINED
BISH 529619, Tongatapu Group, ‘Eua I., reef flat off Ohonua Harbor, coll. C. Imada (s.n.), 26 July 1988.
Ulva flexuosa Wulfen, Reference Wulfen1803;
N'Yeurt & Payri, Reference N'Yeurt and Payri2007
[Enteromorpha flexuosa (Wulfen) J. Agardh, Reference Agardh1883]
SPECIMEN EXAMINED
BISH 692762, Tongatapu Group, Tongatapu Island, south-east side, Emeline Beach, coll. W. Char (IA 22513), 11 July 1995.
Ulva lactuca Linnaeus, Reference Linnaeus1753;
N'Yeurt & Payri, Reference N'Yeurt and Payri2007
SPECIMENS EXAMINED
BM 770090, Tongatapu Group, Tongatapu Island, coll. W.H. Harvey, date unknown; BISH 692784, Tongatapu Group, Tongatapu Island, south-east side, Emeline Beach, coll. W. Char (IA 22508), 11 July 1995.
Order BRYOPSIDALES
Family BRYOPSIDACEAE
Bryopsis pennata J.V. Lamouroux, Reference Lamouroux1809;
Abbott & Huisman, Reference Abbott and Huisman2004
SPECIMENS EXAMINED
BISH 529620, Tongatapu Group, ‘Eua I., reef flat off ‘Ohonua Harbor, coll. C. Imada (s.n.), 26 July 1988; BISH 692764, Tongatapu Island, Emeline Beach, coll. W. Char (IA 22515), 11 July 1995; BISH 692769 and 692770, Vava‘u Group, Foilifuka Island, coll. T. Motley, 28 July 1995.
Family CAULERPACEAE
Caulerpa brachypus Harvey, Reference Harvey1860
SPECIMENS EXAMINED
BM 515866, 515867 and 515868, Ha‘apai Group, Lifuka, coll. W.H. Harvey, date unknown.
Caulerpa cupressoides (M. Vahl) C. Agardh, Reference Agardh1817;
Abbott & Huisman, Reference Abbott and Huisman2004
[= Caulerpa mamillosa Montagne, Reference Montagne1842b]
SPECIMENS EXAMINED
BM 701588, 701589 and 701592, Vava‘u Group, Vava‘u, coll. W.H. Harvey, date unknown; BISH 692795 and 692796, Vava‘u Group, specific island unknown, coll. W. Char (IA 22503), 10 July 1995; BISH 692797, Tongatapu Group, Tongatapu Island, Emeline Beach, coll. W. Char (IA 22507), 11 July 1995.
Caulerpa fastigiata Montagne, Reference Montagne1837
SPECIMENS EXAMINED
BM 515892, Vava‘u Group, Vava‘u, coll. W.H. Harvey, date unknown; BM 841587 and 841591, Tongatapu Group, Tongatapu Island, coll. H. Moseley, July 1874.
Caulerpa plumulifera Zanardini, Reference Zanardini1878
(Figure 2C)
SPECIMENS EXAMINED
BM 841481, Ha‘apai Group, Lifuka, coll. W.H. Harvey, 1855, det. A. Weber-van Bosse; BM 841487, Tongatapu Group, Tongatapu Island, coll. E. Graeffe, 1869; BM 841550, Ha‘apai Group, Lifuka, coll. W.H. Harvey, date unknown, det. A. Weber-van Bosse. This is a currently accepted name in AlgaeBase, and also reported from Sri Lanka in Silva et al. (Reference Silva, Basson and Moe1996). However it was not recollected from Tonga since the 19th Century, and new collections as well as a molecular study on this species would be desirable to ascertain its taxonomic status.
Caulerpa sertularioides (S.G. Gmelin) M. Howe, Reference Howe1905;
Abbott & Huisman, Reference Abbott and Huisman2004
SPECIMENS EXAMINED
BM 701530, 841473 and 841477, Vava‘u Group, Vava‘u, coll. W.H. Harvey, 1855; BISH 692793 and 692794, Vava‘u Group, specific island unknown, coll. W. Char (IA 22500), 10 July 1995.
Caulerpa taxifolia (M. Vahl) C. Agardh, Reference Agardh1817
SPECIMENS EXAMINED
BM 701533, 701534 and 701535, Tonga, coll. W.H. Harvey, date unknown.
Caulerpa urvilleana Montagne, Reference Montagne, Hombron and Jacquinot1845
SPECIMENS EXAMINED
BM 701587, 701590 and 701591, Tonga, coll. W.H. Harvey, date unknown.
Caulerpa verticillata J. Agardh, Reference Agardh1847
SPECIMENS EXAMINED
BM 515895, 841524 and 841525, Tongatapu Group, Tongatapu Island, mangrove, coll. W.H. Harvey, 1855.
Caulerpa webbiana Montagne, Reference Montagne1837
SPECIMENS EXAMINED
BM 515896, 515897 and 515898, Ha‘apai Group, Lifuka, coll. W.H. Harvey, 1855.
Family CODIACEAE
Codium arabicum Kützing, Reference Kützing1856
SPECIMENS EXAMINED
BM 659654 and 840916, Tonga, coll. W.H. Harvey, 1855.
Codium extricatum P.C. Silva, Reference Silva1959
(Figure 2D)
SPECIMENS EXAMINED
BM 840910, Tongatapu Group, Tongatapu Island, coll. H. Moseley, ca. 1870; BM 840921, Tonga, coll. W.H. Harvey, 1855.
Codium geppiorum O.C. Schmidt, Reference Schmidt1923;
N'Yeurt & Payri, Reference N'Yeurt and Payri2007
[Codium bulbopilum Setchell, Reference Setchell1924]
SPECIMENS EXAMINED
BM 840911, Tongatapu Group, Tongatapu Island, fringing reef, coll. H. Moseley, July 1874, det P.C. Silva; BM 840913, Tongatapu Group, Tongatapu Island, coll. H. Moseley, July 1874, det. P.C. Silva as Codium bulbopilum. BM 840918, Tongatapu Group, Tongatapu Island, coll. E. Graeffe, date unknown, det. P.C. Silva; BM 840922, Tonga, coll. W.H. Harvey, 1855, det. P.C. Silva; BISH 692788, Tongatapu Group, Tongatapu Island, south-east side, Emeline Beach, coll. W. Char (IA 22520), 11 July 1995.
Family HALIMEDACEAE
Halimeda borneensis W.R. Taylor, Reference Taylor1975;
Kooistra et al.,
Reference Kooistra, Coppejans and Payri2002
SPECIMENS EXAMINED
BM 701496 and 841152, Tongatapu Group, Tongatapu Island, coll. H. Moseley, 1874; BM 841075, Tongatapu Group, Tongatapu Island, coll. H. Moseley, date unknown; BISH 753200, Vava‘a Group, Hunga Island, coll. T. Motley, 12 July 1995.
REMARKS
Halimeda simulans in the Indo-Pacific region is now recognized as Halimeda borneensis (Kooistra et al., Reference Kooistra, Coppejans and Payri2002).
Halimeda discoidea Decaisne, Reference Decaisne1842;
Hillis, Reference Hillis1959
SPECIMENS EXAMINED
BM 701508 and 840959, Tonga, coll. W.H. Harvey, 1855; BM 840951, Tongatapu Group, Tongatapu Island, coll. H. Moseley, 1874; BISH 692787, Tongatapu Group, Tongatapu Island, south-east side, W. Char (IA 22506), 11 July 1995.
Family UDOTEACEAE
Udotea glaucescens Harvey ex J. Agardh, Reference Agardh1887
(Figure 2E)
SPECIMENS EXAMINED
BM 515944, 515945, 840806, 840807, 840809, 840810. Tongatapu Group, Tongatapu Island, coll. W.H. Harvey, 1855.
Order CLADOPHORALES
Family ANADYOMENACEAE
Microdictyon umbilicatum (Velley) Zanardini, Reference Zanardini1862
SPECIMENS EXAMINED
BM 841210 and 841212, Tongatapu Group, Tongatapu Island, coll. H. Moseley, July 1874; BM 841209, Tonga, coll. W.H. Harvey, 1855
Family BOODLEACEAE
Boodlea composita (Harvey) F. Brand, Reference Brand1904
SPECIMENS EXAMINED
BM 701628, Ha‘apao Group, Lifuka, coll. W.H. Harvey, date unknown; BM 701629, Vava‘u Group, Vava‘u, coll. W.H. Harvey, date unknown; BM 840740, Tongatapu Group, Tongatapu Island, fringing reef, coll. H. Moseley, date unknown.
Family CLADOPHORACEAE
Chaetomorpha linum (O.F. Müller) Kützing, Reference Kützing1845
SPECIMEN EXAMINED
BM 841384, Tonga, collector and date unknown, in Herb. Dickie.
Family PITHOPHORACEAE
Dictyosphaeria versluysii Weber-van Bosse, Reference Weber-van Bosse1905;
Skelton & South, Reference Skelton and South2007
[= Dictyosphaeria australis Setchell, Reference Setchell1926]
SPECIMEN EXAMINED
BISH 692778, Vava‘u Group, Foilifuka Island, coll. T. Motley, 28 July 1995.
Family SIPHONOCLADACEAE
Boergesenia forbesii (Harvey) Feldmann, Reference Feldmann1938;
Skelton & South, Reference Skelton and South2007
SPECIMENS EXAMINED
BM 637958, Tonga, coll. W.H. Harvey, date unknown, det. F. Leliaert (7 April 2002); BM 841679, Tonga, coll. W.H. Harvey, no date; BISH 692780, Tongatapu Group, Tongatapu Island, south-east side, Emeline Beach, coll. W. Char (IA 22517), 11 July 1995.
Family VALONIACEAE
Valonia ventricosa J. Agardh, Reference Agardh1887;
Kraft, Reference Kraft2007
[= Ventricaria ventricosa (J. Agardh) Olsen & J.A. West, Reference Olsen and West1988]
SPECIMENS EXAMINED
BISH 529613. Tongatapu Group, ‘Eua Island, reef flats off Ohonua Harbor, coll. C. Imada (s.n.), 26 July 1988, det. P. Skelton; BISH 692775, Vava‘u Group, Foilifuka Island, coll. T. Motley, 28 July 1995.
Valoniopsis pachynema (G. Martens) Børgesen, Reference Børgesen1934;
Kraft, Reference Kraft2007
(Figure 2F)
SPECIMEN EXAMINED
BISH 550188. Tongatapu Group, Tongatapu Island, Nukualofa Reef, coll. H.E. Parks and W.A. Setchell (UC Herb. 17103), early June 1926.
DISCUSSION
Eight species of Rhodophyta and 26 species of Chlorophyta are reported for the first time from the marine waters of Tonga in the central South Pacific Ocean. The 34 new species records from Tonga represent 26 (41%) of the 64 species from Tonga examined at BM and 15 (50%) of the 30 species of Tongan algae which were deposited at BISH. Seven species of the newly recorded Chlorophyta were present in both the BM and BISH collections.
One species of Rhodophyta, i.e. Trichogloea requienii, and four species of Chlorophyta, i.e. Caulerpa plumulifera, Codium extricatum, Udotea glaucescens and Valoniopsis pachynema represent the first records from central Polynesia. The Tongan Archipelago may thus be considered a ‘stepping stone’ area for the distribution of marine algae between eastern Melanesia (Fiji) and western Polynesia (the Cook Islands). The red alga Titanophora pikeana was only reported previously from the adjacent western Melanesian islands of Fiji (South & Skelton, Reference South and Skelton2003) and the isolated Hawaiian Islands to the north (Abbott, Reference Abbott1999). The other 28 algal species are widely distributed marine species around tropical and subtropical islands in the Pacific.
ACKNOWLEDGEMENTS
The first author extends his appreciation to the British Natural History Museum for the invitation to reevaluate its holdings of Southern Pacific marine algae and to the courtesy extended by Jennifer A. Bryant, Curator of Algae, Botany Department during his visit from 18 April to 18 May 2005. The second author extends his appreciation to the three vascular plant taxonomists, i.e. Clyde T. Imada, the late Winona P. Char and the late Timothy J. Motley, for collecting the algal specimens in Tonga. We acknowledge Nicole M. Shun and Benjamin K. Brown, Bishop Museum, for providing the initial images of BISH 550188 and 692759. The identifications of the Tongan algal specimens deposited in BISH were partially supported by a National Science Foundation, Biological Research Collections Pacific Basin Grant DB1-1057453 to the Bishop Museum (Shelley A. James, Principal Investigator). Any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors and do not necessarily reflect the views of the National Science Foundation. This paper is Contribution 2014-002 of the Pacific Biological Survey, Bishop Museum.