INTRODUCTION
A red alga occurring on the North Atlantic Ocean of Morocco, belonging to the family Delesseriaceae, has been identified as Polyneura bonnemaisonii (C. Agardh) Maggs & Hommersand; it is reported for the first time in Morocco, as well as for the first time in Africa.
The genus Polyneura J. Agardh includes six species worldwide and one species (Polyneura californica J. Agardh) with an uncertain taxonomic status (Guiry & Guiry, Reference Guiry and Guiry2015). Polyneura is one of the most limited distributed genera, with four endemics species (Wynne, Reference Wynne2014; Guiry & Guiry, Reference Guiry and Guiry2015). Whereas, the type species Polyneura bonnemaisonii has been reported from the North Atlantic Ocean, but only in Europe. Among the seven taxa, only two species have been reported from Africa, they are Polyneura denticulata Feldmann from Senegal (John et al., Reference John, Prud'homme van Reine, Lawson, Kostermans and Price2004) and Polyneura venosa (Harvey) Papenfuss from South Africa (Wynne, Reference Wynne2014).
Recently, two studies were published by Riadi et al. (Reference Riadi, Salhi, Bouksir, Moussa and Hassoun2013) and Hassoun et al. (Reference Hassoun, Salhi, Bouksir, Moussa, Riadi and Kazzaz2014) concerning the marine phycoflora of Morocco, but none of the reported taxa belongs to the family Delesseriaceae. In this publication, we add a new Delesseriaceae species for Moroccan and African marine phycoflora.
MATERIALS AND METHODS
The specimens examined in this study were collected from Hyayda on the north-west coast of Morocco (35°05′15.7″N 6°12′51.4″W), on the shores of the Atlantic Ocean. Samplings were performed in 2013 and 2014, in the upper-subtidal (0.5–1 m depth) and lower-intertidal zones of wave-exposed rocky shores. Fresh samples were transported to the laboratory in plastic bags with seawater. Studies were carried out on samples preserved in 5% formalin-seawater solution; anatomical sections were made by hand with a razor blade. The specimens studied were deposited in the Phycological Herbarium of the Phycology and Mycology Laboratory, Faculty of Sciences, Abdelmalek Essaâdi University, Tetouan, Morocco (HTET 879). The photomicrographs were taken with a Panasonic Lumix DMC-FS16 digital camera (Osaka, Japan) coupled with a Nikon microscope. For nomenclatural purposes, the following taxonomic databases were used: Index Nominum Algarum (Silva, Reference Silva2015) and AlgaeBase (Guiry & Guiry, Reference Guiry and Guiry2015).
RESULTS AND DISCUSSION
SYSTEMATICS
Family DELESSERIACEAE
Genus Polyneura J. Agardh
Polyneura bonnemaisonii (C. Agardh) Maggs & Hommersand (Figure 1)
Basionym Delesseria bonnemaisonii C. Agardh
Synonym Polyneura hilliae (Greville) Kylin
Polyneura bonnemaisonii (C. Agardh) Maggs & Hommersand: (A) habit; (B, C) surface view of blade showing elongate-polygonal cells and microscopic veins; (D) dentate margin of blade; (E) apex showing the growth of thallus by transverse division of a prominent apical cell; (F) transverse section of the blade, which is monostromatic; (G) detail of the cells in surface view; (H) tetrasporangial sori; (I) mature tetrasporangia, divided tetrahedrally. Scale bars: A, 1 cm; B–E, 150 µm; F–G, 50 µm; H, 250 µm; I, 100 µm.
DISTRIBUTION
This species, which has a type locality of Add Finisterre Armoricae, Brittany (Agardh, Reference Agardh1822), has a limited distribution in the North-Eastern Atlantic Ocean: Europe (Cullinane & Whelan, Reference Cullinane and Whelan1983; Bárbara et al., Reference Bárbara, Cremades and López Rodríguez1992; Maggs & Hommersand, Reference Maggs and Hommersand1993; Cires Rodríguez & Cuesta Moliner, Reference Cires Rodríguez and Cuesta Moliner2010; Wynne, Reference Wynne2014).
DESCRIPTION
Red-pink delicate plants, 20–50 mm in height and to 40 mm in width. Thalli leafy, membranous and translucent, consisting of one or more blades attached to the substrate by a small basal disc. Blade simple to deeply cleft into wedge or fan-shaped segments. Margin undulate and with rounded apices. Venation palmate; no distinct midrib; macroscopic veins present, restricted to the basal parts of blades and well evident in transparence; microscopic veins are present, but only in younger parts of blade. The margin of the thallus is cut by numerous multicellular denticulations or fine teeth.
Apex showing the growth of thallus by transverse division of a prominent apical cell. The blade is mostly monostromatic, about 50 µm thick, but increasing in thickness near stipe, since the blades become polystromatics. Cells at the surface of blade are rectangular to polygonal, strongly pigmented, and measured 30–75 µm long and 20–45 µm wide.
Tetrasporangial sori are grouped, rounded, or irregularly coalescing, up to 500 µm in diameter, disposed on the side of the blades. Tetrasporangia tetrahedrally divided, spherical, 45–53 µm in diameter.
HABITAT AND PHENOLOGY
Tetrasporophytic plants of Polyneura bonnemaisonii were collected in March 2014, sterile plants in December 2013. They were found in the low- to mid-intertidal zone.
REMARKS
Our specimen of Polyneura bonnemaisonii is in agreement with the generic criteria and the general vegetative features described by Gayral (Reference Gayral1966) and Cabioc'h et al. (Reference Cabioc'h, Floc'h, Le Toquin, Boudouresque, Meinesz and Verlaque1992) as Polyneura hilliae (Greville) Kylin, and conforms in all aspects with previous descriptions made by Maggs & Hommersand (Reference Maggs and Hommersand1993).
The Moroccan specimens are very similar to Erythroglossum laciniatum (Lightfoot) Maggs & Hommersand, a species already reported from the Atlantic Ocean of Morocco (Benhissoune et al., Reference Benhissoune, Boudouresque, Perret-Boudouresque and Verlaque2003). However, E. laciniatum differs from our plants in the location of the tetrasporangial sori, which are developing in discontinuous lines inside the margins of large blades, paired on either side of midrib in narrow marginal outgrowths, or between veins of small blades and near apices (Maggs & Hommersand, Reference Maggs and Hommersand1993). Besides that, the blade of E. laciniatum has a variable number of triangular apical cells along apices and margins; furthermore, this blade measures up to 100 µm in thickness.
Other related species, Haralidiophyllum bonnemaisonii (Kylin) A.D. Zinova and Haraldia lenormandii (Derbès & Solier) Feldmann, show, along with Moroccan species, the following features: entire or dentate margins; blade thin, membranous and monostromatic when young (Maggs & Hommersand, Reference Maggs and Hommersand1993). However, both species can be distinguished from P. bonnemaisonii by several characters, such as: neither species have microscopic and macroscopic veins. Also, a distinctive feature of Haraldia lenormandii is that the tetrasporangial sori are arranged in the median region of the blade, in a series of discontinuous sori (Maggs & Hommersand, Reference Maggs and Hommersand1993; Bárbara, Reference Bárbara2009). In Haralidiophyllum bonnemaisonii, there are no microscopic veins, only some macroscopic nerves in the basal region. The blade margins are smooth and growth is by means of marginal initials, undergoing oblique divisions, whereas Polyneura grows by means of discrete apical cells, which undergo transverse divisions (Wynne, Reference Wynne2014).
Our plant somewhat resembles Polyneura denticulata Feldman, already reported in the Atlantic Ocean of Senegal (John et al., Reference John, Prud'homme van Reine, Lawson, Kostermans and Price2004), but is essentially different as one compares the tetrasporangial structure of both species. In Polyneura denticulata, the tetrasporangial sori cover the blades, whereas, the position of the tetrasporangial sori in Polyneura bonnemaisonii are near the branch apices and they cover most of the branch width according to Maggs & Hommersand (Reference Maggs and Hommersand1993). Another difference can be seen in the blade margins; in Polyneura denticulata they are ‘denticulate’, whilst in Polyneura bonnemaisonii the teeth are much smaller (Bárbara, Reference Bárbara2009).
At the macromorphological level, our specimens (Polyneura bonnemaisonii) present several similarities with Cryptopleura ramosa (Hudson) L. Newton, studied by González-García & Conde-Poyales (Reference González-García and Conde-Poyales1994), Gómez-Garreta et al. (Reference Gómez-Garreta, Gallardo, Ribera, Cormaci, Furnari, Giaccone and Boudouresque2001) from the Mediterranean Sea of Morocco; Navarro & Gallardo (Reference Navarro and Gallardo1989) and Benhissoune et al. (Reference Benhissoune, Boudouresque, Perret-Boudouresque and Verlaque2003) from the Atlantic Ocean of Morocco. But C. ramosa has veins that are only one-cell wide, situated just behind the apices and smooth blade margins, without dentation. In addition, the tetrasporangia of C. ramosa measures up to 95 µm in diameter, whereas, in Polyneura bonnemaisonii it never exceeds 65 µm (Maggs & Hommersand, Reference Maggs and Hommersand1993).
Nitophyllum dentatum Bornet, a species already reported from Morocco (Wynne, Reference Wynne2014), shares some similarities with our alga – mainly the small teeth situated along the blade margins. However, N. dentatum is clearly distinguished from Polyneura bonnemaisonii by the absence of any microscopic veins (Maggs & Hommersand, Reference Maggs and Hommersand1993).
It is worth mentioning that the thalii of Polyneura bonnemaisonii measure up to 20 cm in height, but our alga (collected in winter) did not exceed 6 cm in height; according to Maggs & Hommersand (Reference Maggs and Hommersand1993) and Gayral (Reference Gayral1966), small specimens are typical of winter, and the thalli reach their largest dimensions during spring; during summer, the blades show scattered holes.
This algal species can also provide important information in characterizing the turbidity present in water, and according to Miossec et al. (Reference Miossec, Soudant and Le Stum2012), it is a good indicator for low turbid waters.
ACKNOWLEDGEMENTS
The authors are indebted to Dr Michael J. Wynne (University of Michigan) for confirming the identity of the species and supplying us with some hard-to-find literature. We are also grateful to Hajar Bouksir and Mohamed Kadiri for their contributions and an anonymous referee who helped improve the paper.