A combined barcode and morphological approach to the systematics and biogeography of Laurencia pyramidalis and Laurenciella marilzae (Rhodophyta)

Figures & data

Table 1. Laurencia complex species reported from Lusitanian Macaronesia, with sources of information.

Species	Azores	Salvagens	Madeira	Canary Islands
Chondrophycus glandulifer (Kützing) Lipkin & P.C. Silva		John et al. (2004)	Neto et al. (2001) (as L. glandulifera); John et al. (2004)	Haroun et al. (2002) (as Laurencia glandulifera); John et al. (2004)
Laurencia brongniartii J. Agardh				Haroun et al. (2002); Gil-Rodríguez et al. (2012)^1
Laurencia chondrioides Børgesen				Haroun et al. (2002); Gil-Rodríguez et al. (2012)^1
Laurencia caduciramulosa Masuda & Kawaguchi				Cassano et al. (2008)
Laurencia canariensis Montagne ex Kützing				Afonso-Carrillo & Sansón (1999); John et al. (2004); Gil-Rodríguez et al. (2012)^1
Laurencia catarinensis Cordeiro-Marino & M.T. Fujii				Haroun et al. (2002) (as L. intricata); Machín-Sánchez et al. (2012a)
Laurencia dendroidea J. Agardh				Haroun et al. (2002) (as L. majuscula); Cassano et al. (2012b)
Laurencia flexilis Setchell		Parente et al. (2000)	Neto et al. (2001)	Haroun et al. (2002); Gil-Rodríguez et al. (2012)^1
Laurencia glandulifera (Kützing) Kützing			Neto et al. (2001)	Haroun et al. (2002)
Laurencia intricata J.V. Lamouroux		Parente et al. (2000)		Afonso-Carrillo & Sansón (1999); Machín-Sánchez et al. (2012b)^1
Laurencia majuscula (Harvey) A.H.S. Lucas		John et al. (2004)	Neto et al. (2001)	Haroun et al. (2002);^a Cassano et al. (2012a)^1
Laurencia microcladia Kützing	Tittley et al. (2009)	Parente et al. (2000)	Neto et al. (2001)	Haroun et al. (2002)
Laurencia minuta Vandermeulen, Garbary & Guiry				Haroun et al. (2002); Gil-Rodríguez et al. (2012)^1
Laurencia nidifica J. Agardh			John et al. (2004)
Laurencia obtusa (Hudson) J.V. Lamouroux	Neto (1994)	John et al. (2004)	Neto et al. (2001)	Haroun et al. (2002); Gil-Rodríguez et al. (2012)^1
Laurencia tenera C.K. Tseng				Haroun et al. (2002); Gil-Rodríguez et al. (2012)^1
Laurencia viridis Gil-Rodríguez & Haroun	Gil-Rodríguez & Haroun (1992)	Gil-Rodríguez & Haroun (1992); Parente et al. (2000)	Gil-Rodríguez & Haroun(1992); Neto et al. (2001)	Gil-Rodríguez & Haroun (1992); Haroun et al. (2002)
Laurenciella marilzae (Gil-Rodríguez, Sentíes, Díaz-Larrea, Cassano & M.T. Fujii) Gil-Rodríguez, Sentíes, Díaz-Larrea, Cassano & M.T. Fujii				Gil-Rodríguez et al. (2009) (as Laurencia marilzae)
Osmundea hybrida (De Candolle) K.W. Nam	Neto (1994) (as Laurencia hybrida)	Audiffred & Weisscher (1984) (as L. hybrida); John et al. (2004)	Levring (1974) (as L. hybrida); Neto et al. (2001)	Gil-Rodríguez & Afonso-Carrillo (1980a) (as L. hybrida); Haroun et al. (2002); Machín-Sánchez et al. (2012b)^1
Osmundea osmunda (S. Gmelin) K.W. Nam & Maggs			Neto et al. (2001); John et al. (2004)
Osmundea pinnatifida (Hudson) Stackhouse	Neto (1994) (as Laurencia pinnatifida)	John et al. (2004)	Neto et al. (2001)	Haroun et al. (2002); Machín-Sánchez et al. (2012b)
Osmundea truncata (Kützing) K.W. Nam & Maggs		Parente et al. (2000); John et al. (2004)	Neto et al. (2001)	Haroun et al. (2002); Machín-Sánchez et al. (2012b)
Palisada corallopsis (Montagne) Sentíes, M.T. Fujii & Díaz-Larrea		Parente et al. (2000) (as Chondrophycus corallopsis); John et al. (2004) (as C. corallopsis)	Neto et al. (2001) (as C. corallopsis)	Haroun et al. (2002) (as C. corallopsis); Gil-Rodríguez et al. (2012)^1
Palisada flagellifera (J. Agardh) K.W. Nam				Gil-Rodríguez et al. (2010)
Palisada patentiramea (Montagne) Cassano, Sentíes, Gil-Rodríguez & M.T. Fujii		Parente et al. (2000) (as Chondrophycus patentirameus)		Audiffred & Prud’homme van Reine (1985) (as Laurencia paniculata); Gil-Rodríguez et al. (2012)^1
Palisada perforata (Bory de Saint-Vincent) K.W. Nam		Parente et al. (2000) (as Chondrophycus perforatus); John et al. (2004) (as C. perforatus)	Neto et al. (2001) (as C. perforatus)	Haroun et al. (2002) (as Chondrophycus papillosus and as C. perforatus); Cassano et al. (2009)
Palisada thuyoides (Kützing) Cassano, Sentíes, Gil-Rodríguez & M.T. Fujii				Gil-Rodríguez & Haroun (1993) (as Laurencia paniculata); Gil-Rodríguez et al. (2012)^1
Yuzurua poiteaui (J.V. Lamouroux) Martin-Lescanne		Parente et al. (2000) (as Chondrophycus poiteaui); John et al. (2004) (as C. poiteaui)		Haroun et al. (2002) (as C. poiteaui); Gil-Rodríguez et al. (2012)^1

¹These papers made changes to the taxonomy of the species. Record regarded as doubtful.

Fig. 1. Map of Atlantic Ocean showing the Macaronesian archipelagos. Sites where Macaronesian specimens were collected in the present study are in the Azores: Pico (PI: Pocinho-Monte Candelaria 38.49686466° N/ 28.53992863° W; Barca-Madalena 38.53988167° N/ 28.52058390° W; Prainha do Norte 38.47720139° N/ 28.20443794° W; Lajes do Pico-Poça de Baleia 38.38998195° N/ 28.25144459 W; Lajes do Pico-Fábrica de Baleia 38.38862491° N/ 29. 19430601° W; Santa Cruz Ribeiras 38.404° N/ 28.1872° W), São Miguel (SM: Cerco da Caloura-Baía 37.7570843° N/ 25.81716330° W; Ferraria 37.8579856° N/ 25.85265462° W; Mosteiros 37.8992112° N/ 25.82103420° W), Santa Maria (SM^a: Boca de Ribeira Seca 36.94337327° N/ 25.16456403° W; Emissores 36.99720678° N/ 25.17678807° W; Anjos-Este 37.00542551° N/ 25.16444382° W; Anjos-Ponta dos Frades 37.00788999° N/ 25.15019092° W; Anjos-Piscinas 37.00458430° N/ 25.15727202° W); in Madeira (MA: Seixal-Praia da Laje 32.82554110° N/ 17.11529253° W; Porto Moniz-Piscinas 32.86802811° N/ 17.17135116° W; Ponta de Sao Jorge-Casi 32.8357° N/ 16.9053° W); in the Canary Islands: Fuerteventura (FV: Garcey 28.345492° N/ 14.178111° W; El Cotillo 28.7013° N/ 14.0182° W), Lanzarote (LZ: Arrecife 28.957972° N/ 13.544525° W; Pechigueras 28.855217° N/ 13.872631° W), La Gomera (LG: El Charco de las Condesas 28.0839772° N/ 17.33672469° W; El Charco del Conde 28.05150° N/ 17.20269° W; Punta de la Dama 28.031° N/ 17.183° W), Tenerife (TF: El Pris 28.50981317° N/ 16.42174616° W; Puerto de la Cruz 28.4175° N/ 165462° W; Punta del Hidalgo 28.5739° N/ 16.5462° W).

Figs 2–5. Laurencia pyramidalis: type specimens from the herbarium of the Naturalis Biodiversity Center (section NHN), Leiden (L). 2. Herbarium sheet L 0820668 bearing lectotypes and syntypes (Herbarium Kützing). 3. Syntype, enlarged from Fig. 2. 4. Syntype, enlarged from Fig. 2. 5. Lectotype, enlarged from Fig. 2.

Fig. 6. Phylogram generated using neighbour-joining analyses from COI-5P sequences (A) and phylogenetic tree generated using Bayesian inference inferred from rbcL sequences (B). Voucher numbers (in bold) and the geographical origin of each specimen for which sequences were generated in the current study are indicated. FR: France, AA: Azores archipelago, MA: Madeira, CI: Canary Islands, MX: Mexico, BR: Brazil. Bootstrap values > 60% are indicated above nodes and Bayesian posterior probabilities are indicated under nodes. Bold branches indicate strongly supported nodes (bootstrap values > 95% and Bayesian probability > 0.99).

Figs 7–14. Laurencia pyramidalis from the Lusitanian Macaronesian region. 7. Habit. The main axes have sparse branching in the lower portions and first order laterals that decrease in length upwards and bear three further orders of branching in the same arrangement, resulting in a thallus that is pyramidal in outline. Scale bar = 2 cm. 8. Basal anchorage crust with stoloniferous branches. Scale bar = 1 mm. 9. Spiral branching with three or four orders of whorled branches. Branches issued from the first-order branches curved markedly towards the main axis, becoming almost parallel to it. Scale bar = 2 mm. 10. In surface view, cortical cells contain one or two corps en cerise (cc) in living specimens (arrows). Scale bar = 30 µm. 11. Branches with truncate tips showing dense hyaline trichoblasts with one corps en cerise per cell in living specimens (e.g. arrow). Scale bar = 50 µm. 12. Surface view showing polygonal cortical cells connected to each other by secondary pit connections (arrows). Scale bar = 30 µm. 13. Transverse section of the upper portion of a branch showing an axial cell (a) with four pericentral cells (p). Scale bar = 30 µm. 14. Transverse section of a thallus showing medullary cells with thickened walls. Note pericentral cells (p) with annular thickenings (arrow). Scale bar = 50 µm.

Figs 15–22. Laurencia pyramidalis from the Lusitanian Macaronesian region: reproductive structures. 15. Female gametophyte with cystocarps (arrows) located on the penultimate branches; they are subapical, sessile and prominent. Scale bar = 1 mm. 16. Longitudinal section showing a slightly pyriform cystocarp with a non-protuberant ostiole and clavate carposporangia. Scale bar = 200 µm. 17. Male gametophyte with spermatangial receptacles (arrows) located on the ultimate fertile branchlets. Scale bar = 500 µm. 18. Longitudinal section of a cup-shaped spermatangial receptacle. An axial cell (a) is discernible at the base bearing a fertile branch with many ovoid spermatangia. Scale bar = 50 µm. 19. Tetrasporangial plants with cylindrical branchlets. Scale bar = 2 mm. 20. Surface view of a tetrasporangium (te) with two presporangial cover cells (pr). Scale bar = 30 µm. 21. Transverse section near the apex of axial tetrasporangial segments with an axial cell (a), two vegetative pericentral cells (p1, p2) and two fertile pericentral cells (arrows). Scale bar = 30 µm. 22. Longitudinal section through a tetrasporangial branchlet showing the parallel arrangement of the tetrasporangia. Each fertile pericentral cell (arrows) cuts off presporangial cover cells (pr) distal to the initial tetrasporangium (te). Scale bar = 50 µm.

Figs 23–25. Laurenciella marilzae from the Azores. 23. Habit. Scale bar = 5 mm. 24. In surface view, cortical cells contain one corps en cerise (cc) in living specimens. Scale bar = 30 µm. 25. Longitudinal section through a branch showing corps en cerise in cortical and medullary cells. Scale bar = 10 µm.

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