Description of Ten New Lobophora Species from the Bismarck Sea (Papua New Guinea)

Phycological Research 2019; 67: 228–238 doi: 10.1111/pre.12372 ...... Description of ten new Lobophora species from the Bismarck Sea (Papua New Guinea)

Christophe Vieira ,1* Olivier De Clerck,1 Laurent Millet2 and Claude E. Payri2 1Phycology Research Group and Center for Molecular Phylogenetics and Evolution, Ghent University, Ghent, Belgium and 2UMR ENTROPIE (IRD, UR, CNRS), Institut de Recherche pour le Développement, Nouméa, Nouvelle-Calédonie, France

...... 2005). The two districts (referred to as localities hereafter) SUMMARY where the expeditions took place are located 670 km apart, and are under the influence of different currents. Situated on Sampling in the framework of the research program called ‘La the northeastern mainland coast of Papua New Guinea, ’ Planète Revisitée in Kavieng and Madang (Papua New Madang experiences oceanic influences from the Bismarck Guinea) brought thus far unknown diversity of the brown algal Sea, and major currents from the Dampier Strait (exchanging genus Lobophora to the surface. DNA-assisted alpha taxonomy allowed identifying the presence of 16 species Lobophora water with the Solomon Sea). While, Kavieng, located in the fl from these two localities, which only share four species in north east of New Ireland Island, experiences oceanic in u- common. Ten species are newly described, including four, ences from both the Bismarck Sea and Western Pacific which are only known to the Bismarck Sea. A more exhaustive Ocean. sampling across the Bismarck Sea, and more largely across The marine flora of the Bismarck Sea is still poorly docu- the Coral Triangle, will very likely unveil an even greater diver- mented with a total of 305 species records of seaweeds, iden- sity. The present study underscores the fragmentary nature of tified based on morphological characters (Coppejans & Millar our knowledge of macroalgal diversity in this region. 2000; Coppejans et al. 2001a, b). Prior Vieira et al. (2016, 2017), a single species of Lobophora, namely L. variegata, Key words: Bismarck Sea, cox3, diversity, Lobophora, New was recorded by Coppejans et al. (2001a). The presence of Ireland, Papua New Guinea, psbA, rbcL. the latter in Papua New Guinea is nevertheless questionable, ...... as it has only been confirmed to occur in the Greater Carib- bean, based on molecular data.

INTRODUCTION DNA sequencing accelerated species discovery to the level MATERIALS AND METHODS where taxonomists have difficulties to keep up with the description of new taxa. For many taxa this results in a grow- Taxon sampling ing gap between the numbers of molecular entities identified Lobophora specimens were collected during two expeditions in and the number of species formally described (Padial et al. northern Papua New Guinea led by the last author as part of 2010). Indeed, one witnesses a shift in macroalgal taxonomy the research program ‘La Planète Revisitée’ (‘Our Planet towards more informal systems of naming taxa, where taxon Reviewed’): Madang (November 2012) and Kavieng (August– identifiers are used to communicate biological information September 2014) (Fig. S1 in the Supporting Information). and species are no longer formally described (De Clerck et al. A total of 87 specimens from Madang and 30 specimens from 2013). The brown seaweed Lobophora (Dictyotales, Phaeo- Kavieng were collected by SCUBA. Plant material was pre- phyceae), distributed in tropical to warm-temperate regions, served in silica for molecular analyses and ethanol or 5% form- forms a good example to underscore the problems associated aldehyde for morphological analyses. Herbarium specimens with historically underestimated alpha-diversity in seaweeds. were either deposited in the Herbarium of the Muséum national Until 2012 only six species were recognized in Lobophora. d’Histoire Naturelle in Paris (PC) or in the IRD New Caledonia Recent DNA-based studies revealed the presence of over Herbarium (NOU) (Table S1 in the Supporting Information). 100 evolutionary taxonomic units from across the globe in tropical to warm-temperate regions (Schultz et al. 2015; Vieira et al. 2016, 2017). Today, less than a quarter of cur- Molecular and morphological analyses rently recognized Lobophora biodiversity is formally described Total genomic DNA was extracted from tissue samples dried (Vieira et al. 2016). in silica gel using a cetyl-trimethyl ammonium bromide The present paper attempts to bridge this gap partly by extraction method (De Clerck et al. 2006). Sequences were describing the species of Lobophora resulting from the research program ‘La Planète Revisitée’ conducted in Madang and Kavieng in Papua New Guinea, which surveyed the ...... fl marine fauna and ora in 2012 and 2014. The Bismarck Sea *To whom correspondence should be addressed. is located in the southwestern Pacific Ocean, northeast of the Email: [email protected] island of Papua New Guinea and south of the Bismarck Archi- Communicating Editor: Wendy Nelson pelago and the Admiralty Islands (Fig. 1; Green & Mous Received 29 August 2018; accepted 9 January 2019.

Fig. 1. Bayesian chronogram of the genus Lobophora, adapted from Vieira et al. (2017), based on the concatenated alignment of cox3 (610 bp) + psbA (919 bp) + rbcL (1360 bp). In green the species occurring only in Madang (M), in blue only in Kavieng (K), and in orange in both localities (M+K), shown in the map on the background. Posterior probabilities are indicated by colored circles (black: full support, dark grey: [95:99], light grey [90:94]). [Color figure can be viewed at wileyonlinelibrary.com] generated from the mitochondrial encoded cytochrome c oxi- Vieira et al. (2014) and two others belong to the ‘L. obscura- dase III gene (cox3), the chloroplast encoded ribulose-1,- complex’ (L. obscura3 and L. obscura8). Lobophora com- 5-biphosphate carboxylase (rbcL) and the photosystem II plexes are defined as monophyletic clades composed of multi- protein D1 (psbA) genes. For details on the molecular ana- ple cryptic species. The genuine L. obscura (Dickie) C.W. lyses, including the species delineation and phylogenetic ana- Vieira, De Clerck & Payri was linked to L. obscura8 based on lyses, we refer to Vieira et al. (2016) and Vieira et al. (2017). a sequence of the type (Vieira et al. 2016). The remaining In short, we applied three DNA-based methods to delimit spe- 12 lineages from Madand and Kavieng correspond to unde- cies, namely the Maximum Likelihood implementation of the scribed species: L. sp5, L. sp16, L. sp18, L. sp28, L. sp29, GMYC model (Pons et al. 2006; Reid & Carstens 2012), the L. sp30, L. sp33, L. sp41, L. sp50, L. sp61 and L. sp82. Automatic Barcode Gap Discovery (ABGD; Puillandre et al. 2012) and the Poisson Tree Processes model (PTP; Zhang Taxonomic results et al. 2013) on the cox3 dataset. Morphological observations of Lobophora species included analyses of the external and We formally describe 10 of the 13 undescribed lineages internal structures of the specimens following Vieira (including L. obscura3) from the Bismarck Sea, species for et al. (2014). which we have collected several specimens: L. sp16, L. sp18, L. sp24, L. sp28, L. sp29, L. sp30, L. sp33, L. sp41, L. sp50 and L. obscura3 (Table 1, Table S2). We pay tribute to the scientific expeditions from the era of the ‘European and American RESULTS voyages of scientific exploration’ between 1764 and 1899 by A Lobophora species tree, inferred from the concatenated naming these new species after some of the ships from these alignment of rbcL, cox3 and psbA sequences, presented with expeditions. Bayesian posterior probability values, is given in Figure 1. Six- teen Lobophora lineages were identified from the Bismarck Sea, including twelve in Madang and eight in Kavieng (Fig. 1, DISCUSSION Fig. S2). As expected, none of the species is conspecific with L. variegata. In line with previous studies (Vieira et al. 2016, Species diversity, distribution and endemism 2017) we regard these lineages as distinct species. Two species were previously described, L. abscondita C.W.Vieira, This is first major taxonomic study within the coral triangle Payri & De Clerck, L. petila C.W.Vieira, Payri & De Clerck in focusing on the genus Lobophora. In line with other studies

Table 1. Descriptions of Lobophora species from the Bismarck Sea, Papua New Guinea, including descriptions of the new species.

Lobophora abscondita C.W.Vieira, Payri & De Clerck Distribution: New Caledonia, Papua New Guinea Specimens: Pointe Bovis, Noumea, New Caledonia, 15 March 2012, leg. C. Vieira (IRD10198); Brun, Noumea, New Caledonia, 15 March 2012, leg. C. Vieira (CV3058); Ilot Laregnere, Noumea, New Caledonia, 15 March 2012, leg. C. Vieira (CV3060); Crouy, Noumea, New Caledonia, 15 March 2012, leg. C. Vieira (CV3076); Pointe Bovis, Noumea, New Caledonia, 15 January 2013, leg. C. Vieira (CV3088); Pointe Bovis, Noumea, New Caledonia, 15 January 2013, leg. C. Vieira (CV3091); Pointe Bovis, Noumea, New Caledonia, 15 January 2013, leg. C. Vieira (CV3096); Pointe Bovis, Noumea, New Caledonia, 15 January 2013, leg. C. Vieira (CV3097); Pointe Bovis, Noumea, New Caledonia, 15 January 2013, leg. C. Vieira (CV3098); Aquarium, Noumea, New Caledonia, 15 January 2013, leg. C. Vieira (IRD11057); Pointe Bovis, Noumea, New Caledonia, 6 May 2013, leg. C. Vieira (IRD11122); Kanumera, Ile des Pins, New Caledonia, 2 May 2013, leg. C. Payri (IRD7919); Sirup Island, Madang, Papua New Guinea, 16 November 2012, leg. C. Payri (PAP509); Sirup Island, Madang, Papua New Guinea, 16 November 2012, leg. C. Payri (PAP513); Kavieng, Papua New Guinea, 12 August 2014, leg. C. Payri (IRD11078); Kavieng, Papua New Guinea, 20 August 2014, leg. C. Payri (IRD11097).

Lobophora astrolabeae C.W.Vieira & Payri sp. nov. [This species corresponds to L. obscura3 in Vieira et al. (2017)] Figs 2a, 3a, 4a, Table S2. Description: Large flabellate thallus, up to 8 cm wide and 4.5 cm tall, thick and leather-like, strongly adherent to the substratum across the whole of the ventral surface by rhizoids, margin irregular, surface smooth, dark brown to grey in color with irregular longitudinal gray bands. Thallus composed of single- to double-cell-layered medulla, five to nine- and three to five-cell-layered cortex on the dorsal and ventral side respectively. The thallus was 164–203 μm thick and composed of 10–12-cell-layers. The species was distinguished from its related species by its thickness and in having longitudinal gray bands, and by the distinctive DNA sequence IRD11138. Type locality: Duad Island, Madang, Papua New Guinea Distribution: China, Papua New Guinea, Salomon Islands. Holotype: IRD11138, collected 15 November 2012, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant common growing on hard substratum mostly dead corals and large rubbles in various habitats including fringing reefs, barrier reef and lagoon; in shallow waters not easily observed because of the color that melts with the dead corals and the strong adhesion to the substratum Etymology: The specific epithet refers to one of the two ships (L’Astrolabe) from the round-the-world expedition La Boussole and L’Astrolabe (1785–1788), ordered by Louis XVI and led by the Chief of expedition Jean-François de Galaup, compte de Lapérouse (1741–1788). The French ship L’Astrolabe disappeared in the Solomon Islands, at Vanikoro, during a violent storm, together with the ship La Boussole. L’Astrolabe explored the coasts of New Guinea in 1827. Specimens: Tab Island, Madang, Papua New Guinea, 12 November 2012, leg. C. Payri (IRD10233); Tab Island, Madang, Papua New Guinea, 12 November 2012, leg. C. Payri (IRD11158); Tab Island, Madang, Papua New Guinea, 12 November 2012, leg. C. Payri (PAP185); Duad Island, Madang, Papua New Guinea, 15 November 2012, leg. C. Payri (IRD11138); Duad Island, Madang, Papua New Guinea, 15 November 2012, leg. C. Payri (IRD11137); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. C. Payri (IRD10237); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. C. Payri (IRD10238); Sek Island, Madang, Papua New Guinea, 19 November 2012, leg. C. Payri (PAP662); Masas Island, Madang, Papua New Guinea, 12 November 2012, leg. C. Payri (PCT41); Riwo Island, Madang, Papua New Guinea, 14 November 2012, leg. H. Verbruggen (PHV419); Malaita, Salomon Islands, 17 July 2004, C. Payri & A. N’Yeurt (IRD1394); Xidao Island, Sanya, Hainan, 16 December 2008, leg. J. Yao & Z. Sun (SAP109518); Tab Island, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3528); Tab Island, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD 4020); Sek Island, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD4072); Sek Island, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD4083; Kavieng, Papua New Guinea, 18 August 2014, leg. C. Payri (IRD11091); Kavieng, Papua New Guinea, 18 August 2014, leg. C. Payri (IRD11092).

Lobophora boudeuseae C.W.Vieira & Payri sp. nov. [This species corresponds to L. sp33 in Vieira et al. (2017)] Figs 2b, 3b, 4b, Table S2. Description: Thallus reniform, up to 3 cm wide and 2 cm tall, predominantly procumbent, spreading out on the substratum or arranged in tiles, dark golden brown in color. Attached to the substratum by ventral rizhoids. Margin entire. Thallus composed of single-cell-layered medulla, two-cell-layered cortex on both sides, dorsal and ventral. The thallus was 64–88 μm thick and composed of 5-cell-layers. The species was distinguished from its related species in the distinctive DNA sequences IRD10234. Type locality: Sirup Island, Madang, Papua New Guinea. Distribution: Papua New Guinea. Holotype: PC0063038 (IRD10234), collected 16 November 2012, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant growing on hard substratum in the shallow part of the lagoon, forming patches common on coral, coralline algae or on bloc near the shore. Etymology: The speciﬁc epithet refers to one of the two ships (La Boudeuse) from the round-the-world expedition La Boudeuse and L’Étoile, between 1766–1769, ordered by the French King Louis XV and led by the Chief of expedition Louis Antoine Bougainville (1720–1773), which led to the ‘discovery’ of Tahiti. The two ships sailed in 1767 along the coasts of New Britain and New Guinea. Specimens: Sirup Island, Madang, Papua New Guinea, 16 November 2012, leg. C. Payri (IRD10234); Sek Island, Madang, Papua New Guinea, 20 November 2012, leg. C. Payri (IRD10232).

Lobophora boussoleae C.W.Vieira & Payri sp. nov. [This species corresponds to L. sp28 in Vieira et al. (2017)] Figs 2c, 3c, 4c, Table S2. Description: Thallus reniform, up to 2 cm wide and 2 cm tall, predominantly procumbent, dark brown in color. Thallus closely attached to the hard substratum by ventral rizhoids; margin free. Thallus composed of single-cell-layered medulla, two- to three- and one- to two-cell-layered cortex on the dorsal and ventral side respectively. The thallus was 78–118 μm thick and composed of 4–6-cell-layers. The species was distinguished from its related species in the distinctive DNA sequence IRD10242. Type locality: Sek Island, Madang, Papua New Guinea.

Table 1. Continued

Distribution: Egypt, Malaysia, Papua New Guinea, Philippines. Holotype: PC0063035 (IRD10242), collected 22 November 2012, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant growing on hard substratum, mostly on dead corals or on the dead part of coral colonies mixed with other macroalgae species; in shallow coastal reefs near Enhalus seagrass bed. Etymology: The speciﬁc epithet refers to one of the two ships (La Boussole) from the round-the-world expedition La Boussole and L’Astrolabe (1785–1788). See Lobophora astrolabeae. Specimens (7): Sek Island, Madang, Papua New Guinea, 22 November 2012, leg. C. Payri (IRD10242); Marsa Alam, Egypt, leg. H. Verbruggen (HV03432); Marsa Alam, Egypt, leg. H. Verbruggen (HV03441); Marsa Alam, Egypt, leg. H. Verbruggen (HV03489), Port Dickson, Pulau Rawa, Malaysia, 17 June 2008, leg. P.E. Lim (d5187); Tambaron Island, Oriental Mindoro, Philippines, leg. G.W. Saunders (GWS025837); Balatasan, Oriental Mindoro, Philippines, leg. G.W. Saunders (GWS025866).

Lobophora challengeriae C.W.Vieira sp. nov. [This species corresponds to L. sp16 in Vieira et al. (2017)] Figs 2d, 3d, 4d, Table S2. Description: Large flabellate thallus, lobbed, up to 4 cm wide and 3 cm tall, predominantly ruffled, light to dark brown in color. Thallus base narrow, attached to the substratum by a basal rizhoids. Mount of hairs at the base. Margin entire. Thallus composed of single-cell-layered medulla, three- to five- and two- to four-cell-layered cortex on the dorsal and ventral side respectively. The thallus was 106–188 μm thick and composed of 6–10-cell-layers. The species was distinguished from its related species in the distinctive DNA sequences IRD11086. Type locality: Kavieng, Papua New Guinea. Distribution: Kenya, Papua New Guinea, Oman, Sri Lanka, Tanzania. Holotype: PC0063047 (IRD11086), collected 13 August 2014, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant obvious forming large clumps mixed with other large algae; growing in shallow waters near mangrove habitat. Etymology: The specific epithet refers to the ship His Majesty’s Ship Challenger (HMS Challenger) from the round-the-world expedition Challenger Expedition (1773–1776), organized by the Royal Society in collaboration with the University of Edinburgh. The Challenger expedition sampled extensively in along the north coast of Papua New Guinea (February 1875). Specimens: Raaha Bay, Mirbat, Dhofar, Oman, 11 February 2001, leg. G. Richards & H. Ford (MW0709); Mirbat, Dhofar, Oman, 5 May 2001, leg. G. Richards & H. Ford (MW1009); Mirbat, Dhofar, Oman, 19 September 2001, leg. G. Richards & H. Ford (MW1909); Mirbat, Dhofar, Oman, (DHO0052); Mirbat, Dhofar, Oman, (DHO0214); Kalpitiya, Sri Lanka, leg. E. Coppejans (HEC15982); Diani Beach, Kenya, leg. O. De Clerck (ODC1511); Pongwe, Zanzibar, Tanzania (TZ0752); Paje, Zanzibar, Tanzania (TZ0809); Paje, Zanzibar, Tanzania (TZ0847); Nungwi, Zanzibar, Tanzania (TZ538); Kavieng, Papua New Guinea, 13 August 2014, leg. C. Payri (IRD11084); Kavieng, Papua New Guinea, 13 August 2014, leg. C. Payri (IRD11086).

Lobophora coquilleae C.W.Vieira sp. nov. [This species corresponds to L. sp29 in Vieira et al. (2017)] Figs 2e, 3e, 4e, Table S2 Description: Thallus flabellate, up to 3.5 cm wide and 2.5 cm tall, predominantly procumbent, light golden brown in color with iridescent lines underwater. Weakly attached to the substratum, margin entire and free, fronds overlapping each other. Thallus composed of single-cell-layered medulla and a single-cell-layered cortex on both sides, dorsal and ventral. The thallus was 30–70 μm thick and composed of 3-cell-layers. The species was distinguished from its related species in the distinctive DNA sequences IRD9853. Type locality: Tausch Island, Madang, Papua New Guinea. Distribution: New Caledonia, Papua New Guinea. Holotype: PC0063036 (IRD9853), collected 21 November 2012, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant growing on hard substratum, on coralline algae or Peyssonnelia; common on outerslope, forming large patches in deep water from 50 m to 20 m. Etymology: The specific epithet refers to the ship La Coquille from a round-the-world expedition (1822–1825), led by the French naval officer and explorer Louis Isidore Duperrey. The scientific expedition through to the Pacific Islands started on August 1822. After reaching Tahiti, the expedition called in at Louis de Bougainville’s Port Praslin in New Britain in August 1823, and then sailed along the New Guinean coast on the way to the Maluku Islands. The expedition called in again in New Guinea along the northern coast in July 1824. The ship was later renamed L’Astrolabe in April 1826. Specimens: Baie de Koue, New Caledonia, 10 March 2005, leg. C. Payri (IRD7672); Kranket Island, Madang, Papua New Guinea, 11 November 2012, leg. C. Payri (IRD9849, PAP089); Kranket Island, Madang, Papua New Guinea, 11 November 2012, leg. C. Payri (IRD 9850, PAP098); Sirup Island, Madang, Papua New Guinea, 16 November 2012, leg. C. Payri (IRD 9851, PAP446); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. C. Payri (IRD10236); Sek Island, Madang, Papua New Guinea, 19 November 2012, leg. C. Payri (IRD 9852, PAP646); Tausch Island, Madang, Papua New Guinea, 21 November 2012, leg. C. Payri (IRD9853); Tausch Island, Madang, Papua New Guinea, 21 November 2012, leg. C. Payri (IRD 9854, PAP802a); Masas Island, Madang, Papua New Guinea, 14 November 2012, leg. H. Verbruggen (PHV369); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. H. Verbruggen (PHV594); Sek Island, Madang, Papua New Guinea, 19 November 2012, leg. H. Verbruggen (PHV646); Tausch Island, Madang, Papua New Guinea, 21 November 2012, leg. H. Verbruggen (PHV825)

Lobophora endeavouriae C.W.Vieira sp. nov. [This species corresponds to L. sp18 in Vieira et al. (2017)] Figs 2f, 3f, 4f, Table S2. Description: Large ﬂabellate thallus, up to 7 cm wide and 4 cm tall, predominantly procumbent, surface smooth, thallus thick leathery in aspect, light to dark green in color. Presence of rizhoids patches randomly distributed on the ventral side. Margin entire. Thallus composed of single-cell-layered medulla, two- and one- to two-cell-layered cortex on the dorsal and ventral side respectively. The thallus was 152–210 μm thick and composed of 8–9-cell-layers. The species was distinguished from its related species in the distinctive DNA sequences CV3258. Type locality: Pointe Bovis, Noumea, New Caledonia. Distribution: Australia, Egypt, Israel, Japan, Juan de Nova, Mayotte, New Caledonia, Palau, Papua New Guinea.

Table 1. Continued

Holotype: PC0063048 (CV3258), collected 5 December 2013, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant growing in the lagoon part, within branches of hard coral (branched Porites), forming large thick lobes. Etymology: The speciﬁc epithet refers to the ship His Majesty’s Bark Endeavour (HMS Endeavour) from the 1768–1771 scientiﬁc expedition to Australia and New Zealand, organized by the Royal Society in collaboration with the University of Edinburgh to observe the transit of Venus across the Sun. The Endeavour cruised across the Torres Strait between Australia and New Guinea in 1770. Specimens: Pointe Bovis, Noumea, New Caledonia, 5 December 2013, leg. C. Vieira, (CV3258); Paradise Beach, Coral Bay, Western Australia, Australia, 18 May 2013, leg. J. Costa, V. Marcelino, L. Stavrias & H. Verbruggen (HV03585); Paradise Beach, Coral Bay, Western Australia, Australia, 18 May 2013, leg. J. Costa, V. Marcelino, L. Stavrias & H. Verbruggen (HV03591); Lotties, Coral Bay, Western Australia, Australia, 19 May 2013, leg. S. Berrin, J. Costa & H. Verbruggen (HV03621); Coral Bay, Western Australia, Australia, 20 May 2013, leg. V. Marcelino, L. Stavrias & H. Verbruggen (HV03652); Coral Bay, Western Australia, Australia, 20 May 2013, leg. V. Marcelino, L. Stavrias & H. Verbruggen (HV03653); Paradise Beach, Coral Bay, Western Australia, Australia, 18 May 2013, leg. H. Verbruggen, V. Marcelino, J. Costa & L. Stavrias (JFC0025); Coral Bay, Western Australia, Australia, 20 May 2013, leg. J. Costa & S. Berrin (JFC0071); Bills Bommie, Coral Bay, Western Australia, Australia, 21 May 2013, leg. J. Costa, L. Stavrias & S. Berrin (JFC0096); Coral Bay, Western Australia, Australia, 21 May 2013, leg. J. Costa, L. Stavrias & S. Berrin (JFC0121); Five Fingers Lagoon, Coral Bay, Western Australia, Australia, 22 May 2013, leg. H. Verbruggen, J. Costa, V. Marcelino & L. Stavrias (JFC0136); Juan de Nova Island, 11 December 2013, leg., M. Zubia (JN091); Coral Beach, Eilat, Israel, leg. H. Verbruggen (HV02912); Coral Beach, Eilat, Israel, leg. H. Verbruggen (HV02919); Coral Beach, Eilat, Israel, leg. H. Verbruggen (HV03056); Bablomekang Island, Palau (PALBB062); Ishigaki Island, Okinawa, Japan, 19 January 1998 (PDI002); Ishigaki Island, Okinawa, Japan (ZoD1); Nakanojima, Japan, 22 November 2010, leg. Z. Sun (MBMD00101); Naama Bay, Sharm el Sheikh, Egypt, 7 May 2012, leg. T. Sauvage, W. Schmidt & D. Gabriel (LAF05802); Naama Bay, Sharm el Sheikh, Egypt, 7 May 2012, leg. T. Sauvage, W. Schmidt & D. Gabriel (LAF05803); El Tor, Egypt, 8 May 2012, leg. T. Sauvage, W. Schmidt & D. Gabriel (LAF05851); El Tor, Egypt, 8 May 2012, leg. T. Sauvage, W. Schmidt & D. Gabriel (LAF05856); Nabq, Egypt, 9 May 2012, leg. T. Sauvage, W. Schmidt & D. Gabriel (LAF05978); Mayotte, leg. L. Mattio (IRD11073); Kavieng, Papua New Guinea, leg. C. Payri (IRD11079).

Lobophora esperanceae C.W.Vieira sp. nov. [This species corresponds to L. sp30 in Vieira et al. (2017)] Figs 2g, 3g, 4g, Table S2. Description: Thallus reniform, spreading out on the substratum, forming patches up to 8 cm wide, predominantly procumbent, dark brown in color. Firmly attached to the substratum by ventral rhizoids. Margin entire. Thallus composed of single-cell-layered medulla, one- to two and one-cell-layered cortex on the dorsal and ventral side respectively. The thallus was 36–66 μm thick and composed of 3–4-cell-layers. The species was distinguished from its related species in the distinctive DNA sequences IRD9857. Type locality: Sirup Island, Madang, Papua New Guinea. Distribution: Papua New Guinea. Holotype: PC0063037 (IRD9857), collected 17 November 2012, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant growing on hard substratum, forming patches common in the shallow part of the outerslope of barrier reef. Etymology: The speciﬁc epithet refers to the frigate L’Espérance commanded by captain Jean-Michel de Kermadec (1748–1793). A two ships (La Recherche and L’Espérance) expedition was launched on September 1971 to search for the missing navigator Lapérouse Jean François de Galaup, comte de Lapérouse, and to make scientiﬁc discoveries and surveys on the way. The expedition sailed between June and September 1792 the seas to the north of New Guinea. Specimens: Sirup Island, Madang, Papua New Guinea, 17 November 2012, leg. C. Payri (IRD9857); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. C. Payri (IRD9858); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. C. Payri (IRD9856); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. C. Payri (IRD9855); Tab Island, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3655); Malamal passage, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3735); Malamal passage Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3755); Tab Island, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3918); Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD4097).

Lobophora etoileae C.W.Vieira sp. nov. [This species corresponds to L. sp41 in Vieira et al. (2017)] Figs 2h, 3h, 4h, Table S2. Description: Large thallus round-shaped, up to 4 cm wide and 3.5 cm tall, predominantly rufﬂed, light golden brown to light green in color. Mount of hairs at the base. Margin entire. Ventral side heavily epiphytized. Thallus composed of single-cell-layered medulla and a single cell-layered cortex on both sides, dorsal and ventral. The thallus was 34–60 μm thick and composed of 3-cell-layers. Sporangia clustered, forming patches all over the thallus. Male sexual reproductive organs unknown. The species was distinguished from its related species in the distinctive DNA sequences IRD9859. Type locality: Sek Island, Madang, Papua New Guinea. Distribution: Papua New Guinea, Palau. Holotype: PC0063069 (IRD9859), collected 20 November 2012, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant growing on hard substratum, in lagoon and bay habitat. Etymology: The speciﬁc epithet refers to one of the two ships (L’Étoile) from the round-the-world expedition La Boudeuse and L’Étoile, between 1766–1769, ordered by the French King Louis XV and led by the Chief of expedition Louis Antoine Bougainville (1720–1773), which led to the ‘discovery’ of Tahiti. The two ships sailed in 1767 along the coasts of New Britain and New Guinea. Specimens: Kranket Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD 9860, PAP002); Sek Island, Madang, Papua New Guinea, 20 November 2012, leg. C. Payri (IRD9859); Kranket Island, Madang, Papua New Guinea, 10 November 2012, leg. H. Verbruggen (PHV046); Sek Island, Madang, Papua New Guinea, 22 November 2012, leg. H. Verbruggen (IRD 9861, PHV891); Palau (PALNG021); Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3467).

Table 1. Continued

Lobophora petila C.W.Vieira, Payri & De Clerck Distribution: French Polynesia, New Caledonia, Papua New Guinea, Tuvalu Specimens: Nuku Hiva, French Polynesia, 27 November 2011, leg. C. Payri (IRD8917); Redika, New Caledonia, 4 June 2006, leg. C. Payri (IRD7657); Canala, New Caledonia, 21 April 2012, leg. C. Payri (IRD7877); Kranget Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD9831); Kranget Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD9832); Kranget Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD9833); Kranget Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD9834); Kranget Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD9835); Kranget Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD9836); Kranget Island, Madang, Papua New Guinea, 10 November 2012, leg. C. Payri (IRD9837); Duad Island, Madang, Papua New Guinea, 15 November 2012, leg. C. Payri (IRD11139); Malamal anchorage, Madang, Papua New Guinea, 17 November 2012, leg. C. Payri (IRD9838); Malamal anchorage, Madang, Papua New Guinea, 17 November 2012, leg. C. Payri (PAP557b); Malamal anchorage, Madang, Papua New Guinea, 17 November 2012, leg. C. Payri (IRD9839); Malamal anchorage, Madang, Papua New Guinea, 18 November 2012, leg. C. Payri (IRD9840); Sek Island, Madang, Papua New Guinea, 19 November 2012, leg. C. Payri (IRD9841); Sek Island, Madang, Papua New Guinea, 20 November 2012, leg. C. Payri (IRD9842); Sek Island, Madang, Papua New Guinea, 20 November 2012, leg. C. Payri (IRD9843); Kanaman pt., Madang, Papua New Guinea, 21 November 2012, leg. C. Payri (IRD9844); Sek Island, Madang, Papua New Guinea, 22 November 2012, leg. C. Payri (IRD9845); Sek Island, Madang, Papua New Guinea, 22 November 2012, leg. C. Payri (PAP900); Tab Island, Madang, Papua New Guinea, 23 November 2012, leg. C. Payri (PAP950); Riwo Island, Madang, Papua New Guinea, 14 November 2012, leg. H. Verbruggen (PHV385); Riwo Island, Madang, Papua New Guinea, 14 November 2012, leg. H. Verbruggen (PHV394); Duad Island, Madang, Papua New Guinea, 15 November 2012, leg. H. Verbruggen (PHV440); Malamal anchorage, Madang, Papua New Guinea, 17 November 2012, leg. H. Verbruggen (PHV551); Sek Island, Madang, Papua New Guinea, 20 November 2012, leg. H. Verbruggen (PHV773); Sek Island, Madang, Papua New Guinea, 22 November 2012, leg. H. Verbruggen (PHV900); Kranket Island, Madang, Papua New Guinea, November 2012, leg. F. Leliaert (MAD3160); Kranket Island, Madang, Papua New Guinea, November 2012, leg. F. Leliaert (MAD3173); Panap Damon Point, Madang, Papua New Guinea, November 2012, leg. F. Leliaert (MAD3825); Kranket Island, Madang, Papua New Guinea, November 2012, leg. F. Leliaert (MAD4033); Kavieng, Papua New Guinea, 12 August 2014, leg. C. Payri (IRD11074); Kavieng, Papua New Guinea, 12 August 2014, leg. C. Payri (IRD11075); Kavieng, Papua New Guinea, 12 August 2014, leg. C. Payri (IRD11076); Kavieng, Papua New Guinea, 12 August 2014, leg. C. Payri (IRD11077); Kavieng, Papua New Guinea, 13 August 2014, leg. C. Payri (IRD11083); Kavieng, Papua New Guinea, 13 August 2014, leg. C. Payri (IRD11087); Kavieng, Papua New Guinea, 19 August 2014, leg. C. Payri (IRD11093); Kavieng, Papua New Guinea, 20 August 2014, leg. C. Payri (IRD11094); Kavieng, Papua New Guinea, 22 August 2014, leg. C. Payri (IRD11100); Funafuti, Tuvalu, 18 November 2014, leg. C. Payri (IRD11146).

Lobophora providenceae C.W.Vieira sp. nov. [This species corresponds to L. sp50 in Vieira et al. (2017)] Figs 2i, 3i, 4i, Table S2. Description: Thallus flabellate shaped, up to 4 cm wide and 3 cm tall, predominantly ruffled, dark brown in color weakly attached to the hard substratum. Thallus base narrow, attached to the substratum by a basal rizhoids. Margin entire. Thallus composed of single-cell-layered medulla, one- to two-cell-layered cortex on both sides, dorsal and ventral. The thallus was 70–112 μm thick and composed of 3–5-cell-layers. The species was distinguished from its related species in the distinctive DNA sequences IRD11145. Type locality: Funafuti, Tuvalu. Distribution: Papua New Guinea, Tuvalu. Holotype: PC0063040 (IRD11145), collected 17 November 2014, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Plant growing in shallow lagoon, forming numerous lobes within branched corals or laying on the floor weakly attached to the substratum Etymology: The specific epithet refers to the ship His Majesty’s Ship Providence (HMS Providence) commanded William Bligh, which returned to the Royal Botanic Gardens Kew with 1283 plants. Specimens: Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3045); Kavieng, Papua New Guinea, 12 August 2014, leg. C. Payri (IRD11080); Funafuti, Tuvalu, 17 November 2014, leg. C. Payri (IRD11145); Funafuti, Tuvalu, 19 November 2014, leg. C. Payri (IRD11155).

Lobophora rechercheae C.W.Vieira sp. nov. [This species corresponds to L. sp5 in Vieira et al. (2017)] Figs 3j, 4j, Table S2. Description: Thallus formless to reniform, up to 2 cm wide and 1 cm tall, predominantly crustose, light to dark green in color. When obvious, the margins were irregularly lobed. Thalli tightly attached to hard substratum by basal rhizoids on the entire ventral surface, but can be removed easily from the substratum. Entire margin. Thallus surface lumpy due to its thinness and as it embraces the irregularities of the substratum. Thallus composed of single-cell-layered medulla, two- to five- and one- to three-cell-layered cortex on the dorsal and ventral side respectively. The thallus was 84–160 μm thick and composed of 4–9-cell-layers. Sexual reproductive organs were unknown. The species was distinguished from its related species in being very proliferative and able to cover important surface area, and the distinctive DNA sequences IRD7883. Type locality: Ilot Ouao, Touho, New Caledonia Distribution: Japan, New Caledonia, Papua New Guinea Holotype: PC0063022 (IRD7883), collected 15 April 2012, deposited in the Herbarium of the Muséum national d’Histoire Naturelle in Paris (PC). Habitat: Grew abundantly, proliferating on dead corals and coralline algae forming large patch of a mosaic of thalli tightly applied to the substratum, 10–18 m deep. Etymology: The specific epithet refers to the frigate La Recherche commanded by captain Antoine Bruni d’Entrecasteaux (1737–1793). A two ships (La Recherche and L’Espérance) expedition was launched on September 1971 to search for the missing navigator Lapérouse Jean François de Galaup, comte de Lapérouse, and to make scientific discoveries and surveys on the way. The expedition sailed between June and September 1792 the seas to the north of New Guinea. Specimens: Ilot Ouao, Touho, North Province, New Caledonia, 15 April 2012, leg. C. Vieira (IRD7883); Masas Island, Madang, Papua New Guinea, 10 November 2012, leg. H. Verbruggen (PHV241); Malamal anchorage, Madang, Papua New Guinea, 17 November 2012, leg. H. Verbruggen (PHV567); Rio Waters, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3496).

Table 1. Continued

Lobophora sp10 Distribution: Curaçao, Juan de Nova, New Caledonia, Papua New Guinea, Saint Vincent, Tuvalu Specimens: Klein Curaçao, Curaçao, 9 February 2017, leg. C. Vieira (CWV0642); Juan de Nova 2 December 2013, leg. M. Zubia (JN046); Chesterfield, New Caledonia, 11 November 2015, leg. C. Payri (IRD11202); Chesterfield, New Caledonia, 13 November 2015, leg. C. Payri (IRD11205); Chesterfield, New Caledonia, 15 November 2015, leg. C. Payri (IRD11208); Kavieng, Papua New Guinea, 13 August 2014, leg. C. Payri (IRD11082); Saint Vincent, 26 April 2015, leg. C. Payri (CP15174); Tuvalu, 19 November 2014, leg. C. Payri (IRD11149).

Lobophora sp61 Distribution: Papua New Guinea Specimens: Yabob Island, Madang, Papua New Guinea, November 2012, leg. L. Le Gall and F. Leliaert (MAD3999); Kavieng, Papua New Guinea, 20 August 2018, leg. C. Payri (IRD11095), Kavieng, Papua New Guinea, 20 August 2018, leg. C. Payri (IRD11096).

Lobophora sp82 Distribution: Papua New Guinea Specimens: Sek Island, Madang, Papua New Guinea, 22 November 2012, leg. C. Payri (PAP866). from the same time period, Coppejans et al. (2001a) reported common ancestor has a Central Indo-Pacificorigin(Vieiraet al. the presence of a single species of Lobophora from Papua 2017). Lobophora sp30 is sister species to L. coquilleae also New Guinea. Making use of gene sequence data here we occurring in the Bismarck Sea and in New Caledonia, making report the presence of 16 lineages, that we equate to species- this clade a Melanesian clade. The clade possibly diverged in level, from two localities only in the Bismarck Sea. Twelve the northeastern part of Papua New Guinea and dispersed to species are found in Madang (Fig. 1, highlighted in green) New Caledonia. Lobophora sp61 does not have close sister spe- and eight in Kavieng (Fig. 1, highlighted in blue). Only four cies, but likely originated from the Central Indo Pacific(Vieira species, however, are shared between the two localities, et al. 2017). Finally, L. boudeuseae is sister species to the L. abscondita, L. petila, L. providenceae and L. sp61 (Fig. 1, clade composed of L. sp31 (restricted to Marquesas) – L. sp32 highlighted in orange). Given the current taxon sampling four (Marquesas) – L. sp5 (distributed in Japan, New Caledonia, Bis- species are so far only known from the Bismarck Sea (L. marck Sea). The clade L. boudeuseae – L. sp31 – L. sp32 – L. esperanceae, L. boudeuseae, L. sp61 and L. sp82; Fig. 1). sp5 has a Central Indo-Pacific origin, and the sub-clade L. Three of them are only occurring in Madang, and one (L. sp31 – L. sp32 settled and diverged in French Polynesia. sp61) in both localities. At least four of the species These data do not vouch in favor of sympatric speciation (L. boussoleae, L. challengeriae, L. endeavouriae and L. for the genus Lobophora in the central Indo-Pacific where it is sp10) from this region are widely distributed species (up to the most diverse. On the contrary, this finer scale analysis of the western Indian Ocean). With a total of 16 species, identi- the diversity seems to indicate the importance of allopatric fied from only two localities, the Bismarck Sea appears to har- speciation by founder events. Nevertheless, a more extensive bor a rich Lobophora diversity. This number will likely sampling in the region is necessary to conclude on the evolu- increase with a more thorough sampling across the region. For tionary processes at play. comparison, New Caledonia, a well-sampled group of Islands further south of the Coral Triangle, contains over 30 species (Vieira et al. 2014). CONCLUSION Sixteen species of Lobophora have now been documented fi from the Bismarck Sea. Thirteen species are currently for- Diversi cation mally described and three remain undescribed (L. sp10, L. Admitting that we have only sampled a fraction of the diver- sp61 and L. sp82). It is recommended that DNA sequence sity from this region, it is difficult to make solid conclusions data are used for species identification in this region. Present with regard to the diversity patterns, biogeographic ranges results support the idea that the coral triangle or more broadly and evolutionary processes of this group in the region. Lobo- the Central-Indo Pacific is the cradle of diversity of the genus phora species from the Bismarck Sea do not trace back to a Lobophora. A more thorough sampling is still needed within single ancestor (Fig. 1) from which one could conclude that this region to fully expose the scope of this diversity, which they radiated in the coral triangle. Rather dispersal events we expect to be higher. More generally, this study highlights and subsequent speciation are more likely scenarios to how little we know about the diversity of the marine flora in account for the diversity observed. The only sister species this region of the coral triangle. The series of papers to come found are L. coquilleae – L. esperanceae, which could indi- on the marine flora from the Bismarck Sea, resulting from the cate speciation within the same region. The four species only research program ‘La Planète Revisitée’, will contribute to known from this region may have resulted from multiple bridge this knowledge gap. founder events. Lobophora sp82 is sister species to the cluster L. abscondita – L. sp83, with L. abscondita distributed in the Bismarck Sea ACKNOWLEDGMENTS (Madang and Kavieng) and in New Caledonia, and L. sp83 known from Gloriosio Islands (located north of Madagascar). We acknowledge the collectors who have contributed to this The clade L. sp82 – L. abscondita – L. sp83 most recent study: F. Leliaert, L. Le Gall and H. Verbruggen. The Madang

Fig. 2. In situ photographs of the Bis- marck species that are newly described. (a) Lobophora astrolabeae sp. nov. (IRD11138); (b) Lobophora boudeuseae sp. nov. (IRD10234); (c) Lobophora boussoleae sp. nov. (IRD10242); (d) ex situ photograph of Lobophora challengeriae sp. nov. (IRD11086); (e) Lobophora coquilleae sp. nov. (IRD9853); (f) Lobophora endeavouriae sp. nov. (CV3258); (g) Lobophora esperanceae sp. nov. (IRD9857); (h) Lobophora etoileae sp. nov. (IRD9859); (i) Lobophora providenceae sp. nov. (IRD11145). [Color ﬁgure can be viewed at wileyonlinelibrary.com]

expedition specimens were obtained during the ‘Our Planet EDF, Stavros Niarchos Foundation and Entrepose Contracting, Reviewed’ Papua Niugini expedition organized by Muséum and in-kind support from the Divine Word University (DWU). National d’Histoire Naturelle (MNHN), Pro Natura Interna- The expedition operated under a permit delivered by the tional (PNI), Institut de Recherche pour le Développement Papua New Guinea Department of Environment and Conserva- (IRD) and University of Papua New Guinea (UPNG), Principal tion. The Kavieng Lagoon Biodiversity Survey (Principal Inves- Investigators Philippe Bouchet, Claude Payri and Sarah tigators: Philippe Bouchet, Jeff Kinch) was part of the Our Samadi. The organizers acknowledge funding from the Total Planet Reviewed expeditions organized jointly by Muséum Foundation, Prince Albert II of Monaco Foundation, Fondation National d’Histoire Naturelle (MNHN), Pro-Natura

Fig. 3. Holotype herbarium specimens of the nine newly described species. (a) Lobophora astrolabeae sp. nov. (IRD11138); (b) Lobophora boudeuseae sp. nov. (IRD10234); (c) Lobophora boussoleae sp. nov. (IRD10242); (d) Lobophora challengeriae sp. nov. (IRD11086); (e) Lobophora coquilleae sp. nov. (IRD9853); (f) Lobophora endeavouriae sp. nov. (CV3258); (g) Lobophora esperanceae sp. nov. (IRD9857); (h) Lobophora etoileae sp. nov. (IRD9859); (i) Lobophora providenceae sp. nov. IRD11145); (j) Lobophora rechercheae sp. nov. (IRD7883). [Color ﬁgure can be viewed at wileyonlinelibrary.com]

Fig. 4. Longitudinal (upper picture) and transverse (lower picture) sections of the holotypes of the newly described species. (a) Lobophora astrolabeae sp. nov. (IRD11138); (b) Lobophora boudeuseae sp. nov. (IRD10234); (c) Lobophora boussoleae sp. nov. (IRD10242); (d) Lobophora challengeriae sp. nov. (IRD11086); (e) Lobophora coquilleae sp. nov. (IRD9853); (f) Lobophora endeavouriae sp. nov. (CV3258); (g) Lobophora esperanceae sp. nov. (IRD9857); (h) Lobophora etoileae sp. nov. (IRD9859); (i) Lobophora providenceae sp. nov. (IRD11145); (j) Lobophora rechercheae sp. nov. (IRD7883). [Color ﬁgure can be viewed at wileyonlinelibrary.com]

International (PNI) and Institut de Recherche pour le Dévelop- Paciﬁque, and CNRS’ Institut Ecologie et Environnement pement (IRD), with support from Papua New Guinea’s (INEE). The expedition was endorsed by the New Ireland Pro- National Fisheries Authority. The organizers acknowledge sup- vincial Administration and operated under a Memorandum of porting funding from the Total Foundation, the Laboratoire Understanding with University of Papua New Guinea (UPNG). d’Excellence Diversités Biologiques et Culturelles (LabEx This work makes use of resources and facilities provided by BCDiv, ANR-10-LABX-0003-BCDiv), the Programme Investis- UGent as part of the Belgian contribution to EMBRC-ERIC sement d’Avenir (ANR-11-IDEX-0004-02), the Fonds (FWO GOH3817N).

REFERENCES Schultz, N. E., Lane, C. E., Le Gall, L. et al. 2015. A barcode analysis of the genus Lobophora (Dictyotales, Phaeophyceae) in the west- Coppejans, E. and Millar, A. 2000. Marine red algae from the north ern Atlantic Ocean with four novel species and the epitypification coast of Papua New Guinea. Bot. Mar. 43: 315–46. of L. variegata (J.V. Lamouroux) E.C. Oliveira. Eur. J. Phycol. Coppejans, E., De Clerck, O. and Leliaert, F. 2001a. Marine brown 50:1–20. algae (Phaeophyta) from the north coast of Papua New Guinea, Vieira, C., Camacho, O., Sun, Z. et al. 2017. Historical biogeography with a description of Dictyota magneana sp. nov. Cryptogamie of the highly diverse brown seaweed Lobophora (Dictyotales, Algol. 22:15–40. Phaeophyceae). Mol. Phylogenet. Evol. 110:81–92. Coppejans, E., Leliaert, F., Dargent, O. and De Clerck, O. 2001b. Vieira, C., Camacho, O., Wynne, M. J. et al. 2016. Shedding new light Marine green algae (Chlorophyta) from the north coast of Papua on old algae: matching names and sequences in the brown algal New Guinea. Cryptogamie Algol. 22: 375–443. genus Lobophora (Dictyotales, Phaeophyceae). Taxon 65:689–707. De Clerck, O., Guiry, M. D., Leliaert, F., Samyn, Y. and Vieira, C., D’hondt, S., De Clerck, O. and Payri, C. E. 2014. Toward Verbruggen, H. 2013. Algal taxonomy: a road to nowhere? an inordinate fondness for stars, beetles and Lobophora? Species J. Phycol. 49: 215–25. diversity of the genus Lobophora (Dictyotales, Phaeophyceae) in De Clerck, O., Leliaert, F., Verbruggen, H. et al. 2006. A revised clas- New Caledonia. J. Phycol. 50: 1101–19. sification of the Dictyoteae (Dictyotales, Phaeophyceae) based on Zhang, J., Kapli, P., Pavlidis, P. and Stamatakis, A. 2013. A general rbcL and 26S ribosomal DNA sequence analyses. J. Phycol. 42: species delimitation method with applications to phylogenetic 1271–88. placements. Bioinformatics 29: 2869–2876. Green, A. and Mous, P. J. 2005. Delineating the Coral Triangle, its ecoregions and functional seascapes. Report Based on an Expert Workshop Held at the TNC Coral Triangle Center, Bali, Indonesia (April–May 2003), and on Expert Consultations Held in June and SUPPORTING INFORMATION August. The Nature Conservancy, Coral Triangle Center and Global Marine Initiative, Indo-Pacific Resource Centre, Bali, Indonesia Additional Supporting Information may be found in the online and Brisbane, Australia; Feb Report No.: Version 3.1. version of this article at the publisher’s web-site: Padial, J. M., Miralles, A., De La Riva, I. and Vences, M. 2010. The Fig. S1. Map of sampling localities in Bismarck Sea. In red, integrative future of taxonomy. Front. Zool. 7: 16. the Coral Triangle according to Green and Mous (2005). Pons, J., Barraclough, T.G., Gomez-Zurita, J. et al. 2006. Sequence- based species delimitation for the DNA taxonomy of undescribed Fig. S2. Lobophora Bayesian species tree based on cox3 based insects. Syst. biol. 55: 595–609. from Vieira et al. (2016, 2017). The values shown at each Puillandre, N., Lambert, A., Brouillet, S. and Achaz, G. 2012. ABGD, node represent Bayesian posterior probabilities. Automatic Barcode Gap Discovery for primary species delimitation. Mol. Ecol. 21: 1864–1877. Table S1. Origin of the specimens from the Bismarck Sea used Reid, N. M. and Carstens, B. C. 2012. Phylogenetic estimation error in this study and in Vieira et al. (2016) and their GenBank can decrease the accuracy of species delimitation: a Bayesian accession numbers for the markers cox3, psbA and rbcL. implementation of the general mixed Yule-coalescent model. BMC Table S2. Comparison of morphological characters among spe- Evol. Biol. 12: 196. cies of Lobophora.