Palabras clave islas. las en poblaciones y/o sus de recuperación conservación de yproponer planes Canarias de marinos ecosistemas los en futuros cambios predecir para abordados ser conocimiento que necesitan de vacíos aquellos identifica también síntesis Esta archipiélago. del marinas bentónicas comunidades las en protagonismo su y mostrar de sintetizar fin el con Canarias islas las en sobre publicados previos estudios los todos revisar es trabajo este de objetivo El décadas. últimas las en declive marcado en poblaciones ycon hábitats de formadoras endémicas, ellas de algunas por 16 especies, yO Pterocladiaceae Gelidiellaceae, , genéticamente: reconocidas familias por cuatro constituido que está Florideophyceae clase ala perteneciente rojas orden algas de un es Gelidiales R and O Pterocladiaceae Gelidiellaceae, Gelidiaceae, families: four recognized genetically comprises and Florideophyceae class to the belongs order which algae ared is Gelidiales A lidiellaceae, Pterocladiaceae, herbario. Pterocladiaceae, lidiellaceae, herbarium. Pterocladiaceae, Keywords populations. for their plans and/or recovery conservation suggest and ecosystems marine the in changes future to predict addressed be to need that gaps knowledge those identifies also review This archipelago. of the communities marine benthic to the species of these I Canary the in Gelidiales the into studies previous all examine is to article this of aim The decades. four last over the decline unforeseen in populations with canopy-forming and endemic are some of which I thogonacladiaceae. bstract esumen GEL P R IDIALES ( R : canopy-forming species, endemism, phenology, Gelidiellaceae, Gelidiaceae, evista Scientia I Scientia evista EV Beatriz A Beatriz IO : endemismo, especies formadoras de hábitat, fenología, Gelidiaceae, Ge Gelidiaceae, fenología, hábitat, de formadoras especies : endemismo, GEL US S rthogonacladiaceae. Es un orden numeroso caracterizado en Canarias Canarias en caracterizado orden numeroso un Es rthogonacladiaceae. t is a copious order characterized in the Canary I Canary the in order acopious characterized t is ES IDIA T RHODOPHYTA U DOI T lfonso*, Sansón Marta and Sangil Carlos LES ( LES DIO nsularum U slands, in order to synthesize and demonstrate the relevance relevance the demonstrate and order to synthesize in slands, : https://doi.org/10.25145/j.S: DI S P RHODOPHYTA Universidad de La Laguna La de Universidad R ES A ES EV , 2; diciembre 2019, diciembre , 2; pp. 153-181; I IO S YPE N D FU ) I ) EN L ) EN R SPEC N T T A TI U S I HE C HE I .2019.02.07 V R SL A E PE S FU A S C A T A N U R N SSN: e-2659-6644 SSN: RA SPEC AR ARIA S Y I slands by 16 species, slands S:

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Revista Scientia Insularum, 2; 2019, pp. 153-181 153 Revista Scientia Insularum, 2; 2019, pp. 153-181 154 pin pin goza 2000; Mercado et al. 2000; goza Puentes 1993; and (Juanes 1993; Melo Neushul ces and Servière-Zara and Scrosati servi ecosystem provide and biodiversity important high harbor that species nant domi canopy-forming and several out are there other rhodophytes among stand as et al. 2017, 2018; 2015; et al. Schmidt A 2015, 2013,2012; 2014, 2018; Borja et al. 2014;et al. et al. Stroobant Quintano Fredriksen and R and Fredriksen ( biotechnology industries and cosmetic food, the in used mainly are which agarose, and agar of high-quality 2019). source Guiry out due anatural economic stand to their interest as Gelidiales world’s the inhabiting species template and (Guiry regions of numerous macroalgal T “ so-called pattern, sporegermination (7) aspecific and spermatangia; divided sally transver (6) fertilization; after cells nutritive linking filaments gonimoblast nates origi that carpogonia (5) tetrasporophytes; intercalary and morphic gametophytes (3) wall; simplecell covered pit (4) connections; iso diplohaplophasic with cycle life componentas the (2) agar planes; three of in branched filament axial tion, the with organiza (1) other algae: red among uniaxial pseudoparenchymatous recognized them enablethat be to characters set of morphological aspecific display also diales bruggen Ver Yoon 2006; 1990; Wilcox Woelkerling 2000; and 1990;et al. et al. Graham ( stages life all centriolesandin complete of absence flagella 1), sporophyte (Figure carposporophyte) and (gametophyte, cycle (6) and life phasic ofsence pit (5) (protoplasmic adjacent connections cells; connections) tri between cytoplasm; thethe (4) pre in (3) plastid; starch the in floridean thylakoids tacked (blue)] (2) of thylakoids; uns phycocyanobilin the on surface form clusters the that (red) and (1) are: dophyta phycobilins [phycoerythrobilin pigments named accessory 2010, 2016).andthe reproductive R of attributes cytological The morphological, charya plants) (Bhatta terrestrial and algae (green organisms green and Glaucophyta with super group the of monophyletic A within lineage eukaryotic R the 2019). Guiry and of the (Guiry entirety with share Gelidiales to the species These enerife, Canary I Canary enerife, et al. 2005, 2006; Scariot et al. Scariot 2005, 2006; hodophyta a combination of specific attributes that outline them as a distinct distinct as a them that outline hodophyta acombination attributes of specific et al. A (Florideophyceae, Kylin R The orderGelidiales d * 2016). Nevertheless, the ecological role of the Gelidiales also makes them them makes also 2016). role of Gelidiales the ecological the Nevertheless, 2010; 2014;et al. Hurd et al. 2015). Yang et al. However, order the Geli mongst the 7262 documented species, approximately species, 233 7262mongst the belong documented red algae -type” (Figure 2) Fredericq and (Figure 1988). (Hommersand -type” epartamento de Botánica, Ecología y Fisiología Vegetal. Universidad de La Laguna, Laguna, La de Universidad Vegetal. yFisiología Ecología Botánica, de epartamento 2004; A 2004; slands, Spain. Corresponding author, [email protected]. Corresponding Spain. slands, ueness 1989;ueness A dl et al. dl 2001; et al. Bouza R ao and Kaliaperumal 1983, 1987; Kaliaperumal and ao 1987a, Santelices b; 2005; Yang et al. Yang 2005; 1. I 2012; Simioni et al. N 1995; Bouzon 1994; 1995;rmisén Bouzon Freile-Pelegrin et al. 2017a, et al. lfonso 2018; O TROD UC 2006; D 2006; 2005, 2016; 2014; Burki Yoon et al. TIO 2014a, b; et al. Boo N íez et al. íez hodophyta) constituted is D rchaeplastida, together together rchaeplastida, ixon 1973; Gabrielson 2012; Polifrone et al. 2018). et al. taíza 2016a; Fili 2016a; ho ------

to identify some genera ( to identify ment (Perrone system andtheand morphology ontogeny filaments) the attach of rhizoidal thick-walled (internal of presence to rhizines the proposed have been owning characters nostic diag some species, vegetative Gelidiales many uncommon in are gametophytes tile (carposporophyte protected morphology. by apericarp) fer as tocarp Nevertheless, development to related carposporophyte cys are and families the to separate used T 1998; 2006; Perrone et al. (Freshwater andBailey described previously thefamilies verified genetically and O family latest the described studies, molecular and phological (2 Boo 4species). et genera, Miller et al. Gaillon, Boo Le Perrone et Felicini (3 24 species) genera, cladiaceae O and (7 158 genera, Kützing (2 Fan 27 species), genera, species), Ptero Gelidiellaceae the O in occurs as parent branch to the angles right at nearly arising pinnules and pinnae the some alternately, irregularly, with cases oris branched, pinnately distichously filaments complexor haptera(Perrone filaments et al. of either independent consisting rhizoidal substratum, to stolons the gled attached of entan system from aprostrate arise Gelidiales of all axes Erect species. llaceae R Prostrate R Espermatangia eproductive structures eproductive Branching pattern T O Cystocarps Cystocarps R hizines etrasporangia hizoidal filaments hizoidal Location Location Haptera rganization rganization N. 2016b). et al. (Boo rthogonacladiaceae R o species Currently, there are four accepted families in the Gelidiales; Gelidiaceae Gelidiaceae Gelidiales; the in Currently, families accepted four are there I system

n relation to the vegetative characters, the branching pattern in Gelidiales Gelidiales in pattern branching the characters, n relation vegetative to the TA T HE F HE BLE 1. M BLE or pinnate or I Present A parallel rows parallel I A Bilocular tion. Endogenous tion. I Present rregular, alternate rregularly or in in or rregularly ndependent forma A pical sori sori pical pical sori pical Gelidiaceae M I et al. L AI I 167 ES I ES N DIA T able 1). 2006). 2006). N GEL GN ronchin and Freshwater 2007). The main attributes Freshwater attributes and ronchin 2007).main The - O IDIA chous or pinnate or chous I A Stichidia V-like rows V-like in or parallel I A sori protrude and teral Unknown Unknown lar and exogenous and lar Unicellu mation. I A S rregular, disti rregularly, in in rregularly, ndependent for Gelidiellaceae pical sori or la or sori pical bsent bsent TI I n addition, the location of sporangia also helps also n addition, location of the sporangia C M LES ( LES 33 OR ADA PH - 2006). The Gelidiellaceae attachment 2006). The Gelidiellaceae P - - O T - L E O D F hizines are only absent Gelidie in are hizines pinnate I Present A V-like rows V-like in or parallel I A ovoid-triangular or Unilocular thick sheath sheath thick a within coalescent Endogenous Present G Pterocladiaceae rregular or or rregular rregularly, in in rregularly, pical sori pical pical sori sori pical I ROM B C A L CH 29 2016 et al. OO (2016b), combining mor (2016b), combining ARA rthogonacladiaceae Boo, rthogonacladiaceae C T rthogonacladiaceae rthogonacladiaceae E main axis main with angle right in nearly I Present A in parallel rows rows parallel in I A Unilocular Unilocular within a thick sheath sheath athick within coalescent Endogenous branched. Extensively Present O rregular or pinnate pinnate or rregular rregularly or R pical or lateral sori lateral or pical pical or lateral sori lateral or pical rthogonacladiaceae S OF A ). 4

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Revista Scientia Insularum, 2; 2019, pp. 153-181 155 Revista Scientia Insularum, 2; 2019, pp. 153-181 156 conservation and recovery plans for their populations suggested. populations for their plans recovery and conservation and predicted be will ecosystems marine to the changes future on studies, previous conclusionscombination and in with identified based be will that gaps knowledge I Canary the in into Gelidiales the on studies previous by drawing archipelago the in communities marine benthic to the Gelidiales the of importance and the relevance highlight to therefore,aims article, This concern. of is major populations reduction their severe in the so islands, shores the in rocky producers on2017b). exposed primary important are macroalgae these Furthermore, 2016b,(Perrone et al. Boo 2006; c, d) et al. filaments rhizoidal the surrounding sheath mucilaginous a develop gonocladiaceae and O Exceptionally, Pterocladiaceae cells. from superficial secondarily and exogenously originates system of attachment the cortication basal the whilst cells, I function. and of morphology different origin, cells with complex system prostrate contrary, O and Pterocladiaceae Gelidiaceae, O theircells. mother with connectionin open that remain cells superficial fromstolon arising filaments rhizoidal of independent consists unicellular system truding in the latest family. A latest the in truding O whereas sori, develop apical Pterocladiaceae and sori. Gelidiaceae olateral apical in rangia or bispo display tetrasporangia 2016a). families et al. Boo The remaining 2006; 1961; (Fan stichidia named reproductive branchlets Boo et són 1999; A (Sansón decades over last the ning decli have been that populations with endemic, canopy-forming and are which ( accepted 2016a). et al. Boo 2002; 1977; (Santelices unknown is R reproductive structure female the llaceae, 1997). Hommersand and (Santelices ovoid I or unilocular, triangular are A branch. tile of fer the on surfaces both emerging carposporangia with Gelidiaceae in circular of Gelidiellaceae, 9 Gelidiaceae and 2 Pterocladiaceae ( 2Pterocladiaceae and 9Gelidiaceae of Gelidiellaceae, 4species are there Nowadays, changes. nomenclatural recently some with of which Nortonson and 1971). (1927) Børgesen 12 registered of species, the already had (Montagne 1840; Picconeislands 1884, 1886; Vickers 1896; 1927; Børgesen Law carried out onthe studies on were phytobenthos first the since changed ficantly n these families, rhizoids have an endogenous origin and arise from the cortical cortical from the arise endogenous and have an origin rhizoids families, n these 2. G 2. ( The reproductive structures in Gelidiellaceae are localized in specialized in specialized are localized Gelidiellaceae in The reproductivestructures Gelidiellaceae is represented by three genera in these islands, Millerella islands, these genera represented in is by three Gelidiellaceae I The number of species documented for the Canary I The Canary the fordocumented number of species M. tinerfensis n the Canary I Canary n the A rthogonacladiaceae and Gelidiellaceae may have apical or lateral sori pro or lateral have may apical Gelidiellaceae and rthogonacladiaceae el 2016), 2014; 2016d;et al. some of Gallardo et al. fonso-Carrillo Boo fonso-Carrillo 2014; Guiry and Guiry 2019). 2014; Guiry and fonso-Carrillo Guiry versely, in Pterocladiaceae and O and versely, Pterocladiaceae in idia b les , M. pannosa M. slands occurrence of 16 species of Gelidiales is currently currently is of 16 occurrence of Gelidiales species slands iodi fter fertilization the cystocarps grow bilocular and and grow bilocular cystocarps the fertilization fter ve ), ), 2013; Martínez 2013;et al. Martínez r Parviphycus s it y i . n t rthogonacladiaceae developrthogonacladiaceae a more rthogonocladiaceae the cystocarps cystocarps the rthogonocladiaceae Santelices ( Santelices Ca he R ico ico slands. There exist certain certain exist There slands. A 2015;et al. et al. n fonso-Carrillo and San and fonso-Carrillo ar P. antipae slands has not signi has slands 2002; Perrone 2002; et al. y Isl A a , lfonso lfonso n P. setaceus n Gelidie d et al. ico s n the n the rtho et al. Boo Boo ) ------

was based both on both Montagne’s based was (1840) var. (as corneum study G. not observe nor collect any specimen in the Canary Canary the in specimen any nornot collect observe on statement (1927). the bybased made Børgesen (1927) Nonetheless, Børgesen did et al. G. arbuscula G. J.V. Lamouroux: genetic analyses. so it is very likely to amisidentification. be likely itso very is archipelago, the in macroalga about of presence this the known are references ther A capillacea Pterocladiella tified species of Huismaniella species tified rrillo Gil- G. canariense G. as species the ted article 33.3article of I the However, combination the canariensis G. I Canary to the species (1979) combination new the proposed by Seoane-Camba until endemic anew as Norton and 1971) (Lawson cartilagineum G. (Haroun versicolor G. or as ( Cruz riensis Islands. the Canary from endemic species an as statement, arbuscula therefore G. proposing to accept Price var. T in collected barium combinationthe arbuscula of G. to attributed Bory. often is cies However, (1927), Børgesen it was who published arbuscula was G. islands the in registered species first The pectinatumG. cies was recognized synonymous with Millerella with synonymous tinerfensis recognized was cies (1977) I ne-Camba considered endemic and Canary to the ( la Cruz de Puerto at collected on specimens based described first Sangil Sangil antipaecus and mous with mous with (1995), R tenuissimadiella Millerella 2019), genera new to the them recent have assigned phylogenetic studies although s a result of this, Børgesen (1927) Børgesen of probably this, s aresult No fur as rare. corneum to referred G. nereideum by Montagne (1840). D R Huismaniella (1988), John et al. et al. (2016d) et al. (2007). More recently, uniden Boo acurrently reported et al. odríguez, D odríguez, by Grunow in Piccone by in Grunow (1884) de la at Puerto collected on specimens based T Gelidiella as 2003; et al. enerife) ( Gelidiaceae is represented in the Canary I represented Canary is the in Gelidiaceae R Gelidium canariense ico et al. ico , egarding egarding and and Parviphycus (1988) later John and et al. or was registered in the archipelago in 1992 in archipelago ( the in registered was Millerella pannosa Millerella Echinocaulon , G. pusillum G. , listed for the Canary I for Canary the , listed A ( (2002) Gil- and íaz de Castro et Prud’homme et de Castro íaz fonso-Carrillo 2003b). A fonso-Carrillo Huismaniella Huismaniella G. corneum G. (2002) valida (2002) et al. Haroun hence, 200) CBN (Greuter et al. G. corneum G. named aspecimen coincided with which enerife, slands [ slands 1988). et al. (Price upon Lyngbye herbarium the also ) and (2004), A ( , Huismaniella G. spathulatum G. G. cartilagineum var. cartilagineum G. as described originally was A (as (as , fonso-Carrillo and Sansón 1999; Guiry and Guiry 1999; Guiry and Sansón and Guiry fonso-Carrillo from the Canary I Canary from the G. canariensisG. , this species was included in the catalogues by Price catalogues included the in was species , this ), P. while setaceus G. canarienseG. (Grunow in Piccone) Seoane-Camba ex Haroun, Haroun, ex (Grunow Piccone) in Seoane-Camba , including individuals deposited in the Bory Her Bory deposited the in individuals , including Gelidiella pannosa Gelidiella sp.). fonso-Carrillo (2014) et al. fonso-Carrillo Gallardo and R odríguez et al. odríguez O (2004) agreed on the lack of evidence of that of of that evidence on lack the agreed (2004) riginally, these species were all described as as described were species all these riginally, slands by A slands espite also being cited in northeast A cited northeast in being espite also (Boo (Boo was not correctly done in accordance with with done not accordance in correctly was fterwards, the entity was referred only as only as referred was entity the fterwards, G. spinosum G. and (Grunow Piccone) in Seoane-Camba]. , G. corneum G. et al. . slands ( slands was documented by A was ) by John (2003), considered synony was 2016d). Gelidiella tinerfensis 2016d). slands by 9 species of by Gelidium 9species slands udiffred udiffred (1985), et al. Betancort I slands, thereupon his reference thereupon his slands, A 1992; 1992; et al. fonso-Carrillo . The authority of this spe this . The authority of T , slands. R slands. (Boo et al. (Boo enerife) as result of their ofenerife) their result as G. crinale G. et al. (Gallardo et al. (Gallardo (2004). (2004). T capillaceum enerife) by Seoa ecently this spe this ecently , 2016d). Geli 2016d). G. microdon G. et al. fonso-Ca Parviphy (2016), (2016), 2016). 2016). 1984), 1984), cana , now frica, frica, was was ------,

Revista Scientia Insularum, 2; 2019, pp. 153-181 157 Revista Scientia Insularum, 2; 2019, pp. 153-181 158 species registered in the Canary I Canary the in registered species Montagne (1840), aforementioned the Thesecond with species. synonymous today (1927)ned by Børgesen in var. capillaceum citation the corneum of who stated G. absent Pterocladia in are cavity. to the attributes emerging These radially carposporangia and axis a central to Pterocladia assigned viously (1997) Hommersand and telices Pterocladiella created of genera Pterocladiella the species two tum tum I Canary florathe of 2018),sen include we themarine confirmation, in needs it species this although so R the first time in T time first the of macroalgae in the Canary I Canary the in of macroalgae group this in of diversity the knowledge order in abetter needed to are reach dies A characteristics. morphological based solely on correctly to identify difficult makes changes, seasonal the with ther group (1988). toge this of species, et al. characterizes that complex The plasticity Finally, Finally, with with I Canary the of lists macroalgae from omitted the been has pectinatum G. Surprisingly, again. ria. llum llum I Canary the in performed dies 1993)eyes et al. ( was reported from a single specimen and never identified and specimen from asingle reported Gelidium pectinatum was and and were documented by Vickers (1896) Cana Gran in collected from specimens Gelidium bipectinatum Eventually, the family Pterocladiaceae in the Canary I Canary the in Pterocladiaceae Eventually, family the The six remaining species of Gelidium species remaining The six G. spinosum G. G. spinulosum G. slands for several decades, probably problem due decades, to anomenclatural for several slands 1992; 1992; 1992; 1990s the in enerife Pinedo (Elejabeitia et al. et al. T slands. Subsequently,slands. (1912) Sauvageau mentioned- spathula G. was recorded for the first time in the Canary I Canary the in time recorded for first the was able 2). (now synonymous with with (now synonymous Pterocladiella capillacea Pterocladiella type . The species (Furnari et al. slands. slands. Gelidium crinale slands. but display cystocarps with nutritive cells around around nutritive with cells but cystocarps display dditional morphological, chemical and genetic stu genetic and chemical morphological, dditional Pterocladiella melanoidea Pterocladiella slands, ( A 1999; T fonso-Carrillo and Sansón 1999). Sansón and San fonso-Carrillo were cited in the first phycological stu phycological were first cited the in G. microdon G. askin and Wynne 2013; and askin A , to include the species preto include species the G. pectinatumG. ) as collected in in collected ) as slands is depicted is by slands was first mentio first was was recorded for was slands in Price in slands G. pusi G. and T enerife. enerife. nder ------J.V. Lamouroux Gelidium Gelidiaceae Families and genera TA

t Ca he BLE 2. Fa 2. BLE n ar m y Isl i lin) P.C. Silva lin) (S.G. Gme spinosum G. G. microdon Kützing microdon G. Bornet G. spathulatum (Kützing) Le Jolis Le (Stackhouse) pusillum G. Montagne pectinatum G. T ex (Hare crinale G. J.V. Lamouroux (Hudson) corneum G. Prud’home van R van Prud’home D guez, Haroun, Gil- Haroun, ex Seoane-Camba G. canariense (Grunow) Borgesen et Saint-Vincent de Bory arbuscula G. l urner) Gaillon urner) i es a , gene n íaz de Castro et et Castro de íaz Species d s . Refe R ra a ra odrí eine - r n

to e to ences d G - el 2016. et al. Gallardo A 2004; et al. John 2003; et al. Price A et al. 1999; Haroun Sansón Haroun Haroun Guadalupe-González 2003; John et al. John 2003; 2002; Gil- et al. 1999; Haroun Sansón 1988; A et al. Price 1840; Montagne 2016. et al. Gallardo A 2004; et al. John 2003; et al. et al. 1927; 1912; et al. Price Børgesen Sauvageau A D et al. 1992; R Price 1981; D so-Carrillo 1927; Gil- 1896; Børgesen Vickers 1981; 1988. al et Price so-Carrillo 1927; Gil- 1896; Børgesen Vickers 2016. et al. A A 1988; 1927; et al. Price Børgesen 1840; Montagne A 2004; et al. et al. Sangil 2002; 1992; A 1927; Price et al 1988; Haroun et al. 1988;1927; Haroun al et Price 1912; Børgesen 1896; Sauvageau Vickers 1840; Montagne 1979; Haroun 1979; Haroun 1971; Norton 1927; and Seoane-Camba gesen Lawson 1912; Bør 1884; Sauvageau Piccone 1840; Montagne Haroun Pinedo 1988; et al. 1984; Price 1971; et al. Norton and Haroun 1927; 1912; Lawson Børgesen 1896; Sauvageau Vickers 2016. et al. Gallardo A 2004; et al. John 2003; et al. 2016. et al. 2014; Gallardo fonso-Carrillo 2016. et al. 2014; Gallardo fonso-Carrillo 2016 et al. 2014; Gallardo fonso-Carrillo 2004; 2004; 1999; et al. John Sansón and fonso-Carrillo íaz-Villa idia 2003; Gil- 2002; Gil- et al. 1992; Elejabeitia et al. 1988; Pinedo et al. 1988; A et al. 1992; A et al. 1994; González- et al. eyes et al. 1999; Haroun Sansón and fonso-Carrillo spec les 2002; Gil- et al. 2003; Gil- et al. Sangil 2002; et al. a et al. spec ch 2016. et al. 2014; Gallardo fonso-Carrillo et al. 2004; John et al. John 2004; R R 2004; A 2004; odríguez et al. odríguez odríguez et al. odríguez 2003; Gil- 2003; 1984; Price 1984; Price i 1984; A et al. elgado do es fonso-Carrillo and Sansón 1999; Sansón and fonso-Carrillo R i fonso-Carrillo and Sansón 1999; Sansón and fonso-Carrillo 1995; A et al. ar es eferences R fonso-Carrillo 2014; Gallardo 2014; Gallardo fonso-Carrillo 2002; A 2002; 2003; Sangil Sangil 2003; et al. odríguez cumen R e i odríguez et al. odríguez 2003; Sangil et al. Sangil 2003; 2003; John et al. John 2003; 2004; R 2004; et al. nclu fonso-Carrillo 2014; fonso-Carrillo fonso-Carrillo 2014; fonso-Carrillo fonso-Carrillo 2014; fonso-Carrillo R R R Gil- 2002; ldanondo- R odríguez and A and odríguez A and odríguez 1995 b; a, et al. uiz 1988; Pinedo 1988; Pinedo fonso-Carrillo and and fonso-Carrillo fonso-Carrillo and and fonso-Carrillo t R e eyes et al. eyes d et al. odríguez udiffred 1985; udiffred 1988; Haroun Haroun 1988; d i e d 2003; John John 2003; A n R ristizabal ristizabal odríguez odríguez odríguez odríguez 2004; 2004; 2003; 2003; 2005; 2005; et al. fon fon - - -

Revista Scientia Insularum, 2; 2019, pp. 153-181 159 Revista Scientia Insularum, 2; 2019, pp. 153-181 160 the Gelidiales of the Canary I of Canary the Gelidiales the cula known about their relevance to the marine communities of the islands ( of islands the communities marine to the relevance about their known is little albeit of cespitose assemblages, part as primarily appear species Gelidiales Bouza García-Jiménez A and ( species the of diversity genetic and ecophysiology phenology, and reproductive the vegetative with dealt havemainly studies shores. These rocky 2002; R lices et Hommersand et lices Pterocladiella Pterocladiaceae G.H. Boo et S.M. Boo et G.H. Boo Huismaniella Parviphycus et S.M.Boo Millerella G.H.Boo Gelidiellaceae Pterocladiella capillacea Pterocladiella and fonso-Carrillo 1986; García- and Sosa fonso-Carrillo et al. The dominant and habitat forming species Gelidiumand habitatspecies canariense, forming The dominant ancel 2008).ancel

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2001; R 1999; 2002; Bouza, Mercado et al. et al. - f G et Hommersand et Santelices Bornet) ex boe (Schous melanoidea P. mersand Hom et Santelices lin) (S.G.Gme capillacea P. Huismaniella sp. T et C. Sangil M. Sansón, J. A (Feldmann) setaceus P. Santelices B. P. antipae (Celan) R J.M. et S.M. Boo ne-Camba) (Seoa tinerfensis M. G.H. Boo et L.Le Gall L.Le et G.H. Boo M. pannosa omínguez-Álvarez et al. omínguez-Álvarez . D el fonso-Carrillo, fonso-Carrillo, o íaz-Villa idia g ico i c (Feldmann) (Feldmann) a i les slands due to their important ecological role on the ecological due important to their slands l have been the target for most studies performed on performed for most studies target the have been a - n n t - - - d molecul d Pinedo Pinedo 2016. et al. 2014; Gallardo rrillo 2004; A et al. John 2003; et al. Sangil 2003; 2002; Gil- et al. 1999; Haroun Sansón Sangil Sangil 1995; A et al. 1993; Lawson 2016. et al. 2014; Gallardo so-Carrillo 2002; Gil- et al. 1999; Haroun zález Elejabeitia 1927; D Børgesen 1896; Vickers 1886; 1884, Piccone 1840; Montagne 2016d et al. Boo A A Sangil A 2004; et al. John 2003; et al. Sangil 2003; et al. 2016. et al. Gallardo 2002; Gil- et al. 1999; Haroun Elejabeitia 2018.Woelkerling et al. et al. 2014; Gallardo so-Carrillo Haroun zález zález 1977; et al. Price Seoane-Camba 2016a. et al. 2016; Boo et al. 2004; Santelices 2004; jabeitia A tizabal et al. 1999; Haroun Sansón and Ca he udiffred et al. 1985;Price udiffred 2016. et al. 2014; Gallardo fonso-Carrillo fonso-Carrillo et al. 2011; Polifrone et al. et al. et al. et al. 2004; Santelices 2004; 2004; Santelices 2004; et al. John 2003; et al. R et al. 1992; R et al. 1992; Elejabeitia et al. 2002; R 2002; et al. 1998; 1992, 1998; eina 1993; et al. Sosa 1995; Lawson et al. 1995; Lawson et al. 1995; A n 2003; John et al. John 2003; 2003; Gil- 1992; A et al. 1992; Betancort et al. 1992; Betancort ar 1992; ar s ar 1992; 1992; et al. 1984; Pinedo et al. elgado 2007; A 2007; et al. y Isl fonso-Carrillo and 1999; Sansón fonso-Carrillo R R eyes eyes A 2002; Gil- et al. ico t odríguez odríguez fonso-Carrillo 2014; Gallardo 2014; Gallardo fonso-Carrillo 1988; Pinedo et al. 1988; Pinedo a u fonso-Carrillo and Sansón Sansón and fonso-Carrillo et al. fonso-Carrillo and Sansón Sansón and fonso-Carrillo 2004; R 2004; 1995; A n di 2002; A 2002; 1988; Guadalupe-Gon d 2016; Boo et al. 2016; Boo 1994; Guadalupe-Gon R R D 1995; A fonso-Carrillo 2014; fonso-Carrillo es 2012). The other et al. 2003; 2003; et al. odríguez 2003; 2003; et al. odríguez s R arias- 2005; 2005; et al. ico fonso-Carrillo and and fonso-Carrillo ancel 2008; A 2008; ancel

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to R tose (Polifrone entire et al. year the ter respectively. I P. capillacea P. of gametophytes male and female T the in winter in only appear arbuscula of G. gametophytes female Fertile cycles. life their in seasonality a clear these factors on factors capillacea P. these or I siteyear studied. the during irradiance and temperature in due be may to literature changes the in bant ( mic species (Piccone species 1884;mic 1927; Børgesen 1979; Seoane-Camba R ende reproductive and morphologyble of about vegetative this information the theavaila widens study This individuals. non-fertile ramified less and to smaller corresponds other the whereas individuals, gametophytes) and (tetrasporophytes fertile in number occur of and ahigher apices by with longer branches racterized canariensein G. morphotypes different two More recently, A population. of level the in recruitment ahigh (0-5 length), suggest may in cm which individuals small of 15 abundant more consisting than length in cm individuals I ( year the found be throughout can phases cycle life all dium canariense 33:1 T in so-Carrillo 1986; et al. Lindgren so-Carrillo 7:1 T in detected was 1998)Canaria (Lindgren found. I have been et al. frone frone ofratios 3:1:1 ( ( capillacea lla Hommersand and Fredericq and Hommersand 1988). 1963; Kamura and (Chihara become prominent when they cystocarps tilization, fer only evident after are but of ends branches they at terminal arise also pogonia sori. Similarly, or car lateral apical formed in are alike spermatangia and rangia female). (male and spo the develop into Both gametophytes they spores, dioecious to generate haploidmeiosis occurs spores. A 1). ofalternation generations (Figure D Phenological studies Phenological T T n Gran Canaria, Betancort and González (1991) González and apopulation Betancort with Canaria, characterized n Gran :F:M) T in ) in comparison with female (F) and male (M) gametophytes. I gametophytes. (M) male and (F) female with comparison ) in ancel (2008) and contrary to previous studies (Fredriksen and R and (Fredriksen studies to previous (2008) contrary and ancel Pterocladiella melanoidea Pterocladiella 2014).documented ratios tetrasporophyte in et al. evidentdifferences These et al. Studies into spatio-temporal variations in populations of endemic populations the Geli in into variations spatio-temporal Studies O Phenological studies on Gelidium on canariense studies Phenological Gelidiales show a Gelidiales enerife (Polifroneenerife et al. and and arbuscula Gelidium other hand, n the 2012) 4:1:1 and T in show a clear dominance in terms of populations of tetrasporophytes of of populations terms tetrasporophytes in dominance show aclear enerife (Polifroneenerife et al. T have agreed on the absence of seasonality of fertile individuals. Thus, Thus, individuals. of fertile of on absence seasonality the have agreed :F:M) ( n both species fertile tetrasporophytes can be observed throughout throughout observed be can tetrasporophytes fertile species n both (2017a) et al. lfonso of coexistence the time for first the described enerife and Gran Canaria (Sosa and García- (Sosa and Canaria Gran and enerife (2014) suggest an important effect (2014)of effect important et al. Stroobant an n fact, suggest D arias- Polysiphonia tetrasporophyte ratios. tetrasporophyte develops female gametophytes year-round according according year-round gametophytes female develops 2012; et al. Stroobant enerife and Gran Canaria populations, while fertile fertile while populations, Canaria Gran and enerife R 2012), P.capillacea in whereas enerife ( enerife odríguez and A and odríguez 1998; Polifrone et al. 2012) 14:1 and ( -type life cycle characterized by isomorphic characterized cycle life -type A iploid where sporophytes form sporangia lfonso et al. lfonso fter attachment and germination of the germination and attachment fter populations. O populations. can be found in summer and win and summer found be in can fonso-Carrillo 1986), 12:3:1fonso-Carrillo (Poli G. arbuscula n G. Pterocladiella capillacea Pterocladiella and Pterocladie and arbuscula , G. 2014). I 2017a) 4:1 and ( T D 2012; A :F) in Gran Canaria (Stroo Canaria Gran :F) in arias- ne morphotype is cha is morphotype ne n addition, cespi the R the ratio was 133:1:1 was ratio the odríguez and A and odríguez lfonso et al. lfonso R G. canariensen G. , aT ico et al. ico eina 1992)eina and ueness 1990).ueness T :F) in Gran Gran :F) in :G ratio of:G ratio 2017a). 2017a). display display 2005). fon ------,

Revista Scientia Insularum, 2; 2019, pp. 153-181 161 Revista Scientia Insularum, 2; 2019, pp. 153-181 162 shore of T in in kins Haw 2011; of phytobenthos et al. the (Häder survival and distribution tor the in species. of this survival to the crucial are tides low emersionthat during periods outcome This suggests levels. synthetic photo same the reaches never it although conditions, sheltered under recover to only 15 exposed is it minutes, arbuscula able is G. When algae. to the nent damage of continuous exposition P to both of DI Häder Häder photoinhibition 2018).and Figueroa 2012; and (Hanelt et al. Harb A damage photosynthetic photo-tolerance, photosensitivity, to related frequently is it of organisms, or ability dissipation demand photochemical the exceeds energy (P radiation active photosynthetically and radiation UV to associated the energy by is determined macroalgae of ability The photosynthetic 2001). et al. (Häder radiation ultraviolet(UV) filtered and light incident natural to individuals conditions by exposing under natural populations Canaria Gran this. to confirm experiments particular needs species bon this acquisition in cula of Gelidium canariense incidentand on activity photosynthetic light the content older with specimens. compared agar of in 20% increase have an viduals November. in indi young Furthermore amaximum reaches July in and increase content Gelidiales these agar the of that laboratory.in authors determined These synthetized be cannot and of Gelidiales matrix intercellular and walls of cell the component anatural is which industries, laboratory and cosmetic food, in used and content arbuscula Gelidium in of agar the et al. Cardell studies. of different physiological studies Physiological ment of C would favor uptake the environ to their species of level. Hence, adaptation these the irradiance of light the 2 namism that allows the water to maintain high C high water to the maintain allows that namism by hydrody astrong characterized are shores of northern islands thermore, the the 2011), Fur incidencethe thelight surface. to et al. most (Martín affecting thus, the where clouds tend to shores accumulate of islands the bit rocky northern the Populations rates. of canariense G. tosynthetic pho to factor their a limiting was light the environment from the that actively and et al. 1997, (Mercado et al. macroalgae in 1998, 2001; 2005), Mercado Giordano et al. *s -1 and and arbuscula G. canariense, Gelidium in air and 400 μmol*m 400 and air in Pterocladiella capillacea Pterocladiella and (2001) demonstrated that these species had a low capacity of use HC of (2001) use alow had capacity species these that demonstrated et al. C when the light incidence low. is level light C when the car of However,strategy specific the (2001)et al. P the Not only is the incident light, but also the desiccation tolerance a key fac akey tolerance desiccation Not the incident the but only is light, also The photosynthetic inhibitionThe photosynthetic of Mercado Mercado I Canary the in Canopy-forming Gelidiales 2016). D enerife. The results show a clear effect of desiccation tolerance to vertical todesiccation vertical oftolerance effect show clear a Theenerife. results et al. omínguez-Álvarez et al. omínguez-Álvarez (2001) studied the effect of dissolved inorganic carbon ( carbon (2001) inorganic dissolved of effect the studied values during the experiment reached 2000 μmol*m 2000 reached experiment the during values AR -2 *s -1 at a depth of 4-6 m. The results after 30 minutes 30 after m.The results of at adepth 4-6 . A O and filtered UV radiation revealed perma revealed radiation UV filtered and AR mong the different strategies of carbon uptake uptake carbon of strategies mong different the 2 by diffusion rather than by the active uptake uptake theactive by than rather by diffusion Pterocladiella capillacea Pterocladiella (2011) to tolerance emersion the studied Gelidium arbuscula arbuscula Gelidium , G. canarienseG. (1977) studied the annual variation (1977) variation annual the studied P. capillacea P. and arbuscula G. O 2 slands have also been the focus focus the been have also slands concentrations independently independently concentrations AR . A ). However, when this gar is aphycocolloid is gar was studied in the the in studied was on the northern northern the on ccording to ccording , O G. arbus G. 3 – ( inha C) DIC) DI C) C) ------order (Chihara and Kamura 1963; Kamura Fredericq and and 1988). Hommersand order (Chihara García-Jimé Gelidium so-called the is pattern germination their and res, tetraspores. uninucleate generate tetrasporangia the whereas res, produce binucleate can haploidbisporangia bispores diploid or uninucleate bispo to Suneson (1982) other rhodophytes, the ding several and algae some in coralline dium canariense of Geli individuals in tetrasporangia divided cruciately, irregularly and decussately R tetrasporangia. cruciate typical despitethe andsomein spores Gelidiales, of sporangia types mation of different R and R (Maggs even species or amongfamilies differences uponced certain descriptions by Kylin (1923).detailed Yet pla been attention has recently, particular studies development early and Biological species. of this metabolism the in compounds of these function particular the to assess studies of need further the authors the admitted Nevertheless, tions. arbuscula G. though even algae, to the damage or response cellular no clear was there that ( studied was compounds volatile these the conditionsdark) on productionand light of of different light red light, (white herbivores 2007). (Fink against defenses and antioxidants ( sulfide dimethyl ethylene and as such pounds imply and/or emission the reception of com macroalgae in ductive mechanisms 2013).competition to related Various interspecific or repro processes physiological capillacea P. in of tetrasporangia maturation to the pounds usually only appear in immature sporangia (Maggs and R and (Maggs sporangia immature in only appear usually canariense G. found in change. of climate productionsis scenario under the predicted andtolerance or on temperature desiccation photosynthe irradiance of bined effect ( changes to environmental adaptability its high upper demonstrates the sublittoral, which or pools in infralittoral in buted species worldwide over grows other two the and capillacea of humidity. P. Lastly, maintenance favors the species morphologyclumped of this sion. I the emer after most affected the indeed was species, below other two sublittoral the The endemic canariense species. G. ofdistribution these 1992; Fierst et al. 1992; Callow (Fletcher and factors Johnson and Brawley by several threatened cess suc their with of macroalgae, benthic cycle life the in stages vulnerable and tical ico ico et al. G. arbuscula of G. case n the General features of the life cycle of Gelidiales are well known since the first first the since known well are of Gelidiales cycle of life the features General A R demonstrated the fastest rehydration and recovery. This species is distri is recovery. rehydration and species This fastest the demonstrated D ttachment and germination of diaspores to the substrate are the most cri the are substrate to the of diaspores germination and ttachment ecent studies have demonstrated the connection of connection some the have demonstrated com volatile ecent studies produced high concentrations of dimethyl sulfide under all light condi light all under sulfide concentrations of dimethyl produced high 2005; Simioni et al. Simioni 2005; omínguez-Álvarez et al. omínguez-Álvarez deposited T in 2005). I mature binucleate of genus, other species the in mature spores, which n Gelidiales, the diaspores are tetraspores and carpospo and tetraspores are diaspores the n Gelidiales, FC (Herbarium of Universidad de La Laguna). A of Laguna). UniversidadFC (Herbarium de La 2014a; A , water loss of the thallus was reduced, although the the although reduced, was water of loss thallus the ico et al. ico 2011). However, about com the known is little A lfonso et al. lfonso (2005) described bisporangia along with with along (2005) bisporangia described lstyne and Puglisi 2007). The result was 2007).was The result Puglisi and lstyne D MS), abiogenic compound with 2018). O , which grows in the upper the in grows , which I G. arbuscula n G. ico 1991).ico et al. (García-Jiménez ne example is the for the is ne example -type, specific to the to specific -type, R ico ico (2001) (2001) et al. ,

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Revista Scientia Insularum, 2; 2019, pp. 153-181 163 Revista Scientia Insularum, 2; 2019, pp. 153-181 164 from populations in T in from populations arbuscula G. Gelidium of canariense tetrasporophytes and gametophytes female of phoresis in in high as twice populations were However, populations. among between differences tic differentiation. Gelidium canarienseand geneof andpatterns systems mating their in differ species both showed that results (1998) diploid the only used subpopulation. and Their data previous the re-evaluated et al. Sosa mind, in With this apossible on results. bias the authors the suggested so diploid the in subpopulation used (tetrasporophytes), those than greater times four haploid the in number were subpopulation of the used genes cies, (gametophytes) spe dominant these of contribution knowledge populationfirst of the dynamics to contrary, no significant differences were found between tetrasporophytes and game and tetrasporophytes werebetween differences found contrary, no significant haploidin diploidfrequency and allele O populations. different refore displays nation is that the clonal growth and genetic drift is higher in in higher is drift genetic and growth clonal the that nation is García- and Genetic studies phylogenetic and needed. are cycles life onvation their experiments andculti studies further species, cespitose Gelidiales all among differences logical branch. of fertile the pletely surfaces covered both Millerella pannosa of Millerella tetraspores from cultivated Canaria. Canaria. rosa genus genus the in character considered adiagnostic was of absence gametophytes tially, the populations and even colonize new substrates. new colonize even and populations their of would spores, contribute the mination to maintain notably ability to their Hurd (Santos D and system attachment perennial from the arising axes erect I tions. of sporement the would germination favorpopula of recruitment and the their attach Therefore,thefaster hydrodynamism. strong shores inhabit with that cies 2014a). for spe et al. strategy adaptive an (Simioni be could behavior This species ( release hafter 2-4 between 2). began (Figure species nation pattern these Surprisingly, in spore the germination A More recently,began. axis the anderect filament formation the of rhizoidal which antibiotic solution an in were cultivated for 55 tetraspores the after days, released, ment canariense G. in Canary I Canary nez nez lfonso lfonso G. canariense G. in than arbuscula G. (Santelices 1997). (Santelices Subsequently, R et al. 2014). This characteristic, along with the earlier attachment andger attachment earlier the 2014).along with et al. characteristic, This Gelidiella n addition, Gelidiales species also have modular-clonal growth, with new new with growth, have modular-clonal also species n addition, Gelidiales Genetic studies focusing on Gelidiales of the Canary I of Canary the on Gelidiales focusing studies Genetic The life history of most of the cespitose Gelidiales is still incomplete. I still is cespitosetheof Gelidiales most of history life The et al. (1999) protocol cultivation for canariense axenic G. developed an slands, so that the species could be exploited as an agar source. A source. agar exploited be an could species as the that so slands, O nly plants cultivated at 20ºC developed spermatangial sori which com sori which developed spermatangial at 20ºC cultivated plants nly R (2018) described in detail the initial stages of the tetraspore develop of tetraspore the (2018) stages initial the detail in described eina (1992, electro eina by isoenzyme 1993) variability genetic analyzed Gelidiella ace in Gelidiella sori description of first spermatangial the until and agreed with the Gelidium the with agreed and arbuscula G. and A lfonso et al. lfonso enerife and Gran Canaria. D Canaria. Gran and enerife . A ico ico ccording to the authors, to one the possibleccording expla et al. 2018), 2hbefore other nearly Gelidium had significant differences differences significant had arbuscula G. (2002) obtained male gametophytes gametophytes male obtained (2002) Gelidiella tenuissima Gelidiella (as T o better understand the bio the understand o better slands are scarce. Sosa Sosa scarce. are slands G. arbuscula arbuscula G. espite being the espite the being -type germi -type uarte 1996;uarte fter being being fter ) of Gran ) of Gran from the from the and the and n the n the and and ni ------

Boo Boo of genera Onikusa the species with together Gelidium of European clade to the based on five genetic markers. R on canariense,G. markers. fivegenetic based G. spinosum to related closely G. cultivated by R cultivated I Canary and2016). Hoban (Krueger-Hadfield history of life the phases toant consider all it complex import is with cycles, biological of macroalgae genetic diversity and ture conclusions any Hence, on population before struc the stating gametophytes. female of thevariability genetic albeitanalyzing only islands, different of isolation between alevel established they Furthermore, populations. between variability genetic the (1993),ina reproduction to role on of important sexual the do agree they although andGarcía- Sosa of findings to the contrary populations, among diversity genetic populations of natural diversity species. this in reproductive system main the canariense G. in frequencies allele tophytes Gelidium canariense Gelidium from material D phism polymor amplified random analyzing Palma) La Gomera and (La islands western ( islands shores of central rocky of northern the ties lla tenuissimalla by R sequence genetic analyzed the that synonymy.also concludedthe orderstudy in required This to is confirm P. setaceus to similar very teristics niella differentinto divided five groups ( thefamily of classification (2016d), They proposed a criteria. new molecular morphological and combining Canary I Canary and and arbuscula were by dominated Gelidium islands western and tral of cen coasts exposed northern the in communities macroalgae the that observed of endemic the M.to tinerfensis that although genetic analyses are needed to support this. to needed support this. are analyses genetic although antipae Parviphycus between similarity morphological (2016d) the Finally, et al. Boo archipelago. highlighted from this species-complex (2007). However, authors, to D the according et al. , Parviphycus Several phylogenetic studies of Gelidiales have included species from the have from the included species of phylogenetic Gelidiales studies Several et al. (2002) Bouza Bouza and The phylogeny of the Gelidiellaceae was also re-analyzed by Boo et al. Boo by re-analyzed also was The phylogenytheGelidiellaceae of The first studies into the phytobenthos of the Canary I Canary the into studies the phytobenthos of first The N (1995) Freshwater et al. slands. T and Huismaniella Huismaniella as referred is slands (2014, including phylogeny 2016b) molecular of the Gelidiales, analyzed 4. E 4. A ( Millerella pannosa of (currently Millerella synonym considered as c RA o ueness and Fredriksen. R Fredriksen. and ueness o f G l P

o D adnatus el ) marker variation. Their results demonstrated high levels of levels high demonstrated Their results variation. ) marker gy Parviphycus setaceus idia , di, , Millerella s G. attenuatum , G. i les tri of of collected in T in collected Gelidium canariense bu is part of aspecies-complex part is pusillum G. , while . A n t (2006) also contributed of study to genetic also the (2006) n tio dditional studies are necessary to clarify this this to clarify necessary are studies dditional and (2002) and identified as Gelidie (2002) identified and et al. ico Ca he G. canariense G. that establish esults , concluding that sexual reproduction is sexual that , concluding Perronella gracilis Perronella as described by described as , Suhria sp G. pulchellum G. and a N G. pusillum G. and enerife exhibiting morphological charac morphological . exhibiting (2002) include D et al. ronchin G. canariense G. that indicate esults n n analysis of the type material of material of type the A analysis d c d T ar and Gelidium and enerife and Gran Canaria) and and Canaria) Gran and enerife Millerella, of Millerella, species the and y Isl onse . They focus on eight locali focus . They a A ). Asample from the r n fonso-Carrillo fonso-Carrillo Gelidiella v slands had already already had slands d . n atio s . A G. canariense.G. ), is analogue ), analogue is collected and and collected t a later date, t alater date, ,

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Revista Scientia Insularum, 2; 2019, pp. 153-181 165 Revista Scientia Insularum, 2; 2019, pp. 153-181 166 2013; 2012; A at 9to 14 et al. over years, Sansón the (Polifrone cm et al. long.cm O in 2013;et al. A 1979) (Polifrone cm 20 (Seoane-Camba cm from 30 et al. than to less canariensein G. is described decline greatest The of Gelidium areduction length in have indicated the throughoutyears. occurred changes assess interestare great to of studies These theirpopulations. of cartography digital a including thesitesislands, different of and and arbuscula of G. population size on and distribution data vided rina rina and canariense G. arbuscula, G. community G2-zonated and G1- the noted as Gelidiales, by dominated these communities the including archipelago, the of upper in the sublittoral assemblages mon macroalgae G. canariense G. and tacantha abies-marina (as of yellow Trep band the beneath just walls belt adense on rocky the forming tions (1912)Sauvageau Norton and (1971) Lawson and popula of these each described R 3) (Figure et al. (Wildpret Canaria T Gomera, La shores Palma, of northern La at wave-exposed ral canariense A Pinedo and A and guez ( pools rocky or infralittoral in arbuscula above Gelidium located capillacea by Pterocladiella belt red anew characterized highlighted dies rife) Norton presented and by (1971) Lawson stu New decades. two after changed ( cal not economi was compilation transport the abandoned since and was practice this 1960 until when industries sent and to mainland air-dried was material cally, this Gelidium 2014). 1950, Since 40-50up weight haveto collected fishermenof tonsdry of local Porse and 2011; Bixler (McHugh 2004; medium to produce culture try Hurd et al. biotechnology indus the in used be only can one the and that quality of highest the ( of agar T of shores. rocky exposed northern in of sublittoral the beginning at the located species these with zonation pattern documented this also Norton and (1971) Lawson intermingled. appeared where species platforms both of canariense G. band darker and it athicker developed neath arbuscula by G. uppermostthe red-purple characterized belt was 2008). et al. odríguez G. arbuscula G. 1987) gave rise to its exploitation as a natural resource 1987) resource to et al. its exploitation rise anatural gave as (Wildpret enerife A (as fonso-Carrillo 2003a, b). 2003a, fonso-Carrillo Cystoseira abies-marina Besides differences in abundance and vertical distribution, recent studies distribution, studies recent and vertical abundance in differences Besides Wildpret Wildpret (N of T la Cruz de Puerto in distribution macroalgae of The vertical of Gelidium arbuscula abundance great The A per year from natural populations that were uprooted that populations by Periodi waves. from natural year per fonso-Carrillo 2003a). I fonso-Carrillo are dominant and canopy-forming species of the rocky upper sublitto of rocky the canopy-forming species and dominant are fonso-Carrillo 1986; Betancort and González 1991; González 1986; and Betancort Pinedofonso-Carrillo et al. Pterocladiella capillacea contrary, Pterocladiella n the 2017b). et al. lfonso (1927) Børgesen of 14 individuals cm described fonso-Carrillo 1994). capillacea P. fonso-Carrillo Nowadays, (2012) 12 by Polifrone studied than were those et al. less , while indicates the location of most com the the 1980s. the indicates in study This (1987) et al. arbuscula of distribution Gelidium documented the Cystoseira abies-marina ). T wenty years later, years wenty R 1987; A nterestingly, the agar obtained from Gelidium obtained nterestingly, agar the fonso-Carrillo 2003b; Sangil et al. 2003b; Sangil fonso-Carrillo specimens in the last four decades. decades. four last the in specimens , whose individuals have decreased have decreased , whose individuals ). A G. canariense G. and have exhibited similar length length similar have exhibited ccording to Sauvageau (1912), to Sauvageau ccording odríguez et al. odríguez , except in abrupt rocky abrupt rocky in , except Treptacantha abies-ma , G. G. and arbuscula G. enerife and Gran Gran and enerife G. arbuscula belt arbuscula G. , whereas under whereas , G. canarienseG. (2008a,b) pro D 2012; Sansón in the north north the in arias- , generally , generally R 2004; 2004; 1992; 1992; lfonso lfonso odrí ene in in is is ------

bution and ecological role on the islands needs further research. research. further needs role onbution islands the ecological and diales of the Canary I of Canary the diales on Geli the studies further in techniques combine molecular morphological and we unless not detected be will that species cryptic new indicate some could species spinosum G. losum, pectinatum G. (i.e. records corneum, doubtful Gelidium on and historic based are islands Some documented to these species uncertain. 2019). (Bertolini when decline sensitive species stages succession early in species habitat-forming as role overlookedmight These key play a species important. also is known, still is little which of cespitosethe Gelidiales, Thestudy endemic of species. of the risk extinction the and status conservation of their assessment allow tions will of popula distribution the and on abundance data current and of historical parison order in com The to populations. recover the necessary are strategies management and plans conservation thus, to changes, vulnerable are that Gelidiales threatened canopy-formingthe with especially concerngraveis of rhodophytes. This of these diversity, the in morphology, changes help ecology physiology future to and predict I Canary the in diales dalupe-González dalupe-González melanoidea diella 1995a, b; Guadalupe-González et al. 1995a, b; Guadalupe-González I Canary from the cespitose Gelidiales ral T R en Silvestres de Especies Listado the species” in “vulnerable consideredrecently as canarienseG. have been and arbuscula G. abundance, population and morphology A change. due to climate changing also are that variables species, these of growth the variables on different of environmental effects 2013; A I period. same the temperature over surface sea in increase a with significant significantly structures and in arbuscula G. and number of reproductive length theindividual in decline asignificant strated ( BCM and T et al. beitia (Eleja archipelago the across cespitose assemblages the in most common species Catálogo Español de Especies Amenazadas (O Amenazadas Especies de Español yCatálogo de Protecciónégimen Especial FC ( EC/596/2019, B EC/596/2019, 2017b). deposited specimens since Thestudy of herbarium 20151970 until in et al. D lfonso et al. lfonso The accurate diversity of Gelidiales species in the Canary Canary the in species diversity Gelidiales of Theaccurate Geli of ecology and the in gaps biology knowledge identifies review This D pto. Botánica, Ecología yFisiología Ecología Vegetal; ULL) de Ciencias; Facultad pto. Botánica, n contrast, these changes are not detected in P.capillacea not in are detected changes these n contrast, ifferent studies on intertidal and subtidal communities seve document communities and subtidal intertidal on ifferentstudies 1992; Pinedo D pto. Biología; Facultad de Ciencias del Mar; ULPGC) have demon ULPGC) pto. Mar; del de Ciencias Biología; Facultad 5. F have been only occasionally reported (Seoane-Camba 1977; (Seoane-Camba reported Gua only have occasionally been et al. O u 2017b). Experimental studies are necessary in order to verify the the order in to verify 2017b). necessary are studies Experimental ). The apparent high morphological plasticity characteristic of characteristic ). morphological plasticity The apparenthigh E 2019). E t slands. 1995; et al. Sangil u slands. Certain aspects need further investigation and will will and investigation further need aspects Certain slands. r et al. e pe G. canariense G. 1992; r spec 1995), Millerella whereas tinerfensis R eyes eyes ti 2003; R 2003; o ves Gelidium pusillum Gelidium slands. et al. . I n addition, this trend also correlates correlates trend also n addition, this s a result of these sharp changes in in changes sharp of these s aresult R González- 1994, 2005; n G ancel 2008) and their real distri real their and 2008) ancel el , G. sphatulatum, G. spinu G. sphatulatum, G. idia les (Sansón et al. I is one is of the slands is still still is slands or Pterocla or uiz uiz et al. rden rden ------

Revista Scientia Insularum, 2; 2019, pp. 153-181 167 Revista Scientia Insularum, 2; 2019, pp. 153-181 168 Conceptualization: B Universidad de La Laguna and the funding entity CajaSiete granted to B. A CajaSiete granted entity funding the and Laguna Universidad de La Canary Islands. Canary of the ecosystems marine the in changes order tools in future to predict useful and capillacea lla worldwide distributed ( species otherceae), introduced are there while R O macroalgae assemblages at the eastern A eastern at the assemblages macroalgae rella tinerfensis O archipelago. the in tem services from the Canary I Canary from the levels. population and individual on changes environmental of effects the to foresee it difficult makes information lack of This to yet discovered. be are Gelidiales ment many in rates recruit reproductive and or phenology, successful vegetative shifts spatio-temporal incomplete.distribution, The still is cycle life their and missing are gametophytes whose female of reference Gelidiellaceae species to in the particularly required are eview and edition of the final draft: all authors. all draft: edition of final and the eview riginal draft: B draft: riginal This work has been possible thanks to the predoctoral agreement between agreementbetween the to predoctoral possiblethanks been work has This T R o conclude, it is pivotal to improve our knowledge on Gelidiales species species o conclude, it to improve pivotal on is Gelidiales knowledge our egarding the vegetative and reproductive and morphology, vegetative the findings additional egarding ). A , Gelidium canariense ll these attributes make these species model macroalgae to study to study model macroalgae species these make attributes these ll A slands owing to the fact that some of contribute that them fact to to the ecosys owing slands , MS. A . Au Ackn t hor thers are endemic and vulnerable species ( species vulnerable endemic and are thers o s , wle c G. arbuscula G. R ecibido o tlantic ( tlantic n d tri gment : enero de 2020; aceptado 2020; de : enero bu Gelidium pusillum Gelidium ) or on common species cespitose tio s n and Gelidiella and : febrero de 2020 de : febrero Pterocladie lfonso. Mille - - - - - Betancort Bertolini A A A A A A A A Afonso-Carrillo Afonso-Carrillo Afonso-Carrillo Afonso-Carrillo Afonso-Carrillo Afonso-Carrillo A udiffred lfonso rmisén ndersen lstyne lfonso lfonso ldanondo- dl , S.M., Simpson , S.M., , B., Sansón , K.L. &Puglisi , K.L. , R , B., Hernández , B., Sangil tana, nes de Gelidium canariensis de nes Marine 841-849. on diversity. Biodiversity 49: algae of understory ción General de Cantabria. España. 27 p. 27 España. Cantabria. de General ción J. Juanes, 69: 394-402. 69: Sci. Aquat. impacts. ecological and function tribution, cies of Gelidiales. Libro de resúmenes I resúmenes de Libro of Gelidiales. cies Bol. Inst. Esp. Oceanogr. 19: 165-170. Oceanogr. Esp. Inst. Bol. Fuerteventura. de isla la en ción a asociada algal sidad marine algae, algae, marine Canary I Canary Gelidium canariense species endemic threatened the 111-114. 61: 50: 119-127. 50: Mar. Bot. distribution. and I Canary the from algae marine I Canary the 254 p. Laguna. La de Universidad Publicaciones de Servicio analítica. Clave rias. Sociedad Española de Ficología. España. 64 p. 64 España. Ficología. de Española Sociedad 4: 14-27. yHumanidades Ciencias de Revista Catharum: Canarias. en marina nica 173-184. 15: Canaria (1951-1966). Cruz la de Etnografía de Puerto agar. de ción Cuaderno Pajar. El 52: 399-451. 52: biol. classification of eukaryotes with emphasis on the taxonomic of protists. of protists. taxonomic the on emphasis with eukaryotes of classification S.S., , C. 2019. Can secondary species maintain a primary role? Consistent inter-regional effects effects inter-regional Consistent role? aprimary maintain species 2019., C. secondary Can , R , P. , M.J. &González . 1994. Productos derivados del Gelidium del derivados . 1994. Productos . A A A 16: 51-58. 16: Brugerolle , N., D ristizabal . 2018. R . 1985. Marine algae of El Hierro (Canary I (Canary Hierro of El . 1985. algae Marine Bot. Mar. 60: 543-553. Mar. Bot. slands. A , J., , J., Pinedo , J. &Sansón , J. 2014. Lista actualizada de algas marinas de las islas Canarias. I Canarias. islas las de marinas algas de , J. 2014. actualizada Lista , J. 2003b. Bajíos y algas marinas de Puerto de la Cruz: una historia de la botá la de historia una Cruz: la de Puerto de marinas y algas Bajíos , J. 2003b. , J. 2003a. 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D , R ’ íaz-Villa : producción, estructura y aplicaciones. En: En: yaplicaciones. estructura : producción, O , S. &Gil- , S. : de los recursos a las aplicaciones. D aplicaciones. alas recursos los : de ia . rbigny, 1840, una lapa en peligro de extin de peligro en rbigny, lapa 1840, una A etraspore germination of two vulnerable vulnerable of two germination etraspore ., A tlantic O tlantic slands). Vieraea slands). s 2005. The new higher level level higher new The 2005. et al. , S., derson (Gelidiales, R (Gelidiales, ( R , T hodophyta, Gelidiales). R . 2007. New records of benthic of benthic . 2007. records New odríguez cean): morphology, taxonomy morphology, cean): , O . R 14: 157-183. 14: ., Barta hodophyta) from the the from hodophyta) , M.C. 2003. D 2003. , M.C. J. Eukaryot. Micro Eukaryot. J. , J. nforme de la la de nforme R ., Bowser slas Cana slas Bot. Mar. Mar. Bot. 56: 6. 6. 56: iputa iver is ------,

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Cardell Burki Elejabeitia D D D Chihara Brawley D D Delgado Fredriksen Fan Filipin Fletcher Fink Fierst Fredriksen arias- íez íaz-Villa ixon omínguez-Álvarez University of California Publications in in Publications California of University Gelidiales. of the 1961., K.C. studies Morphological Mar. Freshw. Freshw. Mar. systems. aquatic in compounds organic of volatile , P. functions 2007. Ecological , Cold Spring Spring Cold perspective. phylogenomic global from a life of tree , F. 2014.eukaryotic The , J.L., TerHorst , J.L., I Biology of the Rhodophyta the of , P.S. 1973. Biology , E.P., Bouzon ., R , S.H. & , S.H. , C.E., Esteban , C.E., algas marinas bentónicas de El Hierro ( Hierro El de bentónicas marinas algas A de zona la de lógica lles y D del y aislamiento a016147. 6: Biol. Perspect. Harb. 233-252. 27: J. Phycol. Br. algae. the in reproduction on sexual tive 68: 117-124. 68: Madrid de Botánico dín Hidalgo, T Hidalgo, T from of Gelidiales species of three zonation and versicolor logia, Estuar. Coast. Shelf Sci. 99:108-120. Sci. Shelf Coast. Estuar. change. mate cli to relationship potential their and Sea Cantabrian Eastern the in changes assemblage R net) comb. nov. (Gelidiales, R nov.net) comb. (Gelidiales, 539-546. 32: Mar. Bot. availability. nitrogen and temperature density, flux ( Botany 41: 238-249. 41: Phycol. J. histories. life complex in dominance of phase patterns 40: 155-168. 40: Phy. Behav. floridanum of Gelidium res tetraspo the of germination and adhesión realese, the on effects of salinity 2016. Evaluation 27: 303-329. 27: J. phycol. Br. spores. algal , M. &Kamura , M. Muguerza , E., González , E., odríguez , R R esúmenes V Simposio I VSimposio esúmenes Gelidium var cartilagineum Goill. (L.), , T hodophyta) from Norway. Growth and nitrogen storage in response to varying photon varying to response in storage nitrogen and Norway. Growth from hodophyta) , Y., R 34: 303-308. , S. &Rueness , S. , S. &Rueness , S. .L. &Callow .L. ., Tronholm 3: 69-74. 3: 32: 315-368. Johnson (S. Gmel.) Lamouroux ( (S. Gmel.) Lamouroux eyes , J.M. &Afonso-Carrillo, J.M. enerife ( enerife , N., Santolaria , Z.L., O , Z.L., , S., R , S., , J. &Afonso-Carrillo , C.P., Kubler , M.N. &Jorge , M.N. , C. &Fernández , C. , S. 1963., S. O L.E. 1992. Gametogenesis, gametes and zygotes: an ecological perspec ecological an zygotes: and gametes 1992. Gametogenesis, L.E. (Schousboe (Schousboe melanoidea on Pterocladia studies , J. 1990. Culture , A , M.E. 1992. The settlement, attachment and establishment of marine marine of establishment and attachment 1992.settlement, The , M.E. , J. 1989. Culture studies of Gelidium latifolium (Grev.), J. 1989. studies Thur et Born Culture I ico -manitol en dos algas de las I las de algas dos en -manitol 21: 1-28. 21: Vieraea Canarias). slas uriques ., Afonso-Carrillo 12-13: 97-110. 12-13: Macaronésica Bot. Canaria). (Gran rinaga R &Gil- , J.M. bérico del Bentos Marino. Puerto de la Cruz. p. 85. Cruz. la de Puerto Marino. Bentos del bérico ( R , J.E. &D , J.E. n the germination of tetraspores of Gelidiella acerosa of tetraspores germination n the hodophyta, Florideophyceae). hodophyta, , L., Pereira , L., Phycologia 29: 182-190. Phycologia hodophyta). , A , D R ., . New York. New (US hodophyta) en el Puerto de la Cruz ( Cruz la de Puerto el en hodophyta) Ganzedo , L. 1977. Curvas anuales del contenido en agar-agar agar-agar en contenido del 1977. anuales , L. Curvas . 1984. Contribución al estudio de la vegetación fico vegetación la de estudio al . 1984. Contribución udgeon I odríguez , J. 1992. A , J. 1986. R 32: 167-176. 32: Vieraea Canarias). slas . , D canariensis , J. &Sansón . T , U. & ., Braum , S. Bory Gelidium Bory arbuscula Canarias: slas , M.C. 2011. Photosynthetic response 2011., M.C. response Photosynthetic enerife, Canary I Canary enerife, A R lgas marinas bentónicas de Punta del del Punta de bentónicas marinas lgas eproducción y fenología de Gelidium de yfenología eproducción ). Hafner Press. 251 Press. p.). Hafner . 2005. Fertilization success can drive drive can success Fertilization . 2005. Gorostiaga Grun. Grun. , F., Schmidt M. 2004. A 2004. M. 28: 2925-2938. 28: Phycol. Appl. J. Ann. Inst. Bot. A.J. Cavani A.J. Bot. Inst. Ann. , J.M. 2012. Seaweed 2012. Seaweed , J.M. slands. Anales del Jar slands. , E.C. &Simioni, E.C. diciones a la flora de flora ala diciones T enerife). Libro de de enerife). Libro . ex Phyco Bor , C. , C. ------

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Woelkerling Wildpret Woelkerling Vickers Tronchin Taskin Suneson Verbruggen T Stroobant Sosa Sosa Sosa Simioni Simioni Seoane-Camba ronchin , P. , P. , P. , E. &Wynne , E. A , C., Schmidt , C., R , C., A & A , , S. 1982. The culture of bisporangial plants of Dermatolithon litorale plants of bisporangial 1982., S. culture The Consejería de A de Consejería canario. litoral del marinas fanerógamas de ypraderas algas de campos los de cartografía 80: 1-5. 80: algarum Notulae status. R (Gelidiaceae, ne-Camba R ving the red algal tree of life. BMC Biol Evol. of life. tree algal red the ving 2010. D cens Aphanta pachyrrhiza Aphanta R capillaceacladiella (Gelidiales, G.Furnari ( G.Furnari bipectinatum dium ( Lemoine 769-772. 2: Life Med. Coast. 45: 548-558. R (Gelidiales, licher Onikusa genera the in species of reclassification 10: 279-284. Phycol. App. J. of results. Are-evaluation species. dium 29: 118-124. 29: Phycol. J. electrophoresis. byisozyme determined Mar. Biol. 113: Biol. 679-688. Mar. electrophoresis. byisozyme ned gametophytes in populations of Gelidium arbuscula populations in gametophytes 90: 1050-1060. 90: Photobiol. chem. Photo ultrastructure. and performance photosynthetic carotenoids, pigments, tosynthetic ridanum of Gelidium flo tetraspore of the tubeformation germ and system endomembrane A on the Kreusch radiation (UV radiation Chow 99-112. 5: Malac. Bot. Acta las. A ., Valero . &García- , W., , E.M. & Freshwater , E.M. , E.M., Freshwater , E.M., ed A ed . 1896. Contribution à la flore algologique des Canaries. Canaries. des algologique flore à la . 1896. Contribution , M., Polifrone , M., and P. psammophila and , H., Maggs García- , W.J. 1990. A , W.J., Furnari over lgae. Cambridge University Press. Cambridge, UK. pp. 1-6. UK. Cambridge, Press. University Cambridge lgae. , F., R , J. 1979. Sobre algunas gelidiáceas nuevas o poco conocidas de las costas españo costas las de conocidas opoco nuevas gelidiáceas , J. 1979. algunas Sobre Gil- ata mining approach identifies research priorities and data requirements forresol requirements data and priorities research identifies approach mining ata ( , M., Pereira , M., , M., Batista , M., R R , T R 50: 577-586. 50: Phycol. J. Florideophyceae). hodophyta, , M.J. 2013. Proposal of Gelidium, M.J. 2013. serra Proposal Br. phycol. J phycol. Br. Corallinaceae). hodophyta, R odríguez amlov , E.C., Felix , E.C., R ., Schmidt A eina eina +UVB) on young gametophytes of Gelidium floridanum gametophytes on young +UVB) gricultura y Pesca, Gobierno de Canarias. I Canarias. de Gobierno yPesca, gricultura , C. , M. &Viera-, M. , G. 1992. Genetic variability and differentiation of sporophytes and sporophytes of differentiation and variability 1992., G. Genetic n introduction. I n introduction. , F., Maraschin , G. 1993. Genetic variability of Gelidium canariensis ( 1993., G. variability Genetic gen. et sp. nov., et gen. Gelidium profundum Bot. Mar. Mar. Bot. data. morphological and on molecular based hodophyta) A , G., Guardia , G., , D ., Saunders , M.C. & , M.C. , D , F.J. &Gonz , D sp. nov. Phycologia , C., Felix , C., .W., Bolton , M. hodophyta): nomenclatural validity, tipification and taxonomic taxonomic and tipification validity, nomenclatural hodophyta): .W. ( 2007. Gelidiales Four . T ., O R R 68: 21-23. 68: Taxon. Geograph. Plant. J. Webbia: hodophyta). hodophyta) at Canary I Canary at hodophyta) ., Polo uriques R Afonso-Carrillo odríguez , G.W., Le Gall , R n: Cole, K.M. and Sheath, R Sheath, and K.M. Cole, n: , M. , M. &Bouzon, M. a , J.J. &A . &Perrone lez-P , L.K., R , L.K., R , C. &Bouzon, C. ., Polo 46: 325-348. 46: e . 10: 1-16. , M. rez A nderson over (S.G. Gmel.) Nov. Geli comb. replace to A , M. katsuka and Suhria , L.K., dos Santos , L.K., . 2014. R . 17: 107-116. 17: . (Gelidiaceae: R , L., Yoon , L., , C. 2018., C. Gelidiella tinerfensis R , J. 1987. Evaluación cuantitativa y , J. 1987. cuantitativa Evaluación , T , Z.L. 2014b. Effects of ultraviolet ultraviolet of 2014b. Effects , Z.L. A hodophyta) new to southern A southern to new hodophyta) . 1998. Genetic structure of Geli structure . 1998. Genetic slands (Spain, A (Spain, slands ., Kreusch , Z.L. 2014a. Effects of brefeldin of brefeldin 2014a. Effects , Z.L. Ann. Sci. Nat. (Bot.) (Bot.) Nat. Sci. Ann. , R sp. nov., caerules Pterocladiella nforme técnico. técnico. nforme reassessment and and Areassessment .J. 2002. eproductive pattern of Ptero pattern eproductive , H.S. &D , H.S. .G. (eds.) Biology of the of the .G. (eds.) Biology hodophyta) determi , M., Pereira , M., (Suneson) Hamel et et Hamel (Suneson) J. A , R : growth rate, pho rate, : growth tlantic O tlantic ., Costa ex ex rgardh e Clerck R 4: 293-306. 4: hodophyta) hodophyta) J. J. cean). , G.B., , G.B., , D Seoa frica, frica, - End , O . T ., ------.

Revista Scientia Insularum, 2; 2019, pp. 153-181 177 Revista Scientia Insularum, 2; 2019, pp. 153-181 178 Yoon Yoon Yoon Yang Yang Yang , S., D , S., , E.C., Boo , E.C., , E.C., Kim , E.C., , H.S., Nelson , H.S., , H.S., Zuccarello , H.S., , H.S., Müller , H.S., D Springer I Springer D Chapman, M., Melkonian, L., lis, D I algae. red major lineages of red algae ( algae of red lineages major tent. 7: 2394-2406. Evol. Biol. Genome Florideophyceae. of the algae red multicellular midt Sci. Rep. 6: 21361. Rep. Sci. algae. red florideophyte & . 2016. R ordrecht D ordrecht oolittle Yoon PNAS , W.E., Fredericq , K.M., Kim , K.M., , S.M., Bhattacharya , S.M., , H.S. 2016., H.S. D nternational Publishing Switzerland. 45 p. Switzerland. Publishing nternational , W., Lindstrom n: Seckbach, J. & Chapman, D J. &Chapman, Seckbach, n: 102: 373-378. hodophyta. I hodophyta. , K.M., Sheath , K.M., , R eidelberg London New York. New pp. 27-42. London eidelberg .F. &Bourne , G.C. &Bhattacharya , G.C. , S.Y., Lee ivergence time estimates and the evolution of major lineages in the the in lineages of major evolution the and estimates time ivergence , S. et al. , S. n: A n: R , R , S.C., Boo , S.C., rchibald, J.M., Simpson, A Simpson, J.M., rchibald, 42: 482-492. 42: Phycol. J. hodophyta). , P.E. 2005. Phylogeny determined by protein domain con domain byprotein determined , P.E. Phylogeny 2005. , J.M., Boo , J.M., , D .G., O 2015. Highly conserved mitochondrial genomes among among genomes mitochondrial 2015. conserved Highly ., Saunders .J. and Corliss, J. Corliss, .J. and tt , S.M., Pueschel , S.M., , F. , G.H., Lee , G.H., . 2010. Evolutionary history and taxonomy of taxonomy and history . 2010. Evolutionary .J. (eds.) R D . &Bhattacharya , G.W., Knoll ed algae and genomic age. Springer Springer age. genomic and algae ed O , J.H., Nelson . (eds.) Handbook of the Protists. Protists. of the . (eds.) Handbook , C., Qiu , C., - Margu C.H., .G.B., Slamovits, , A ., Fredericq, S., Graf S., ., Fredericq, , D , H. &Bhattacharya , H. . 2006. D . 2006. , W. A ., Yi efining the efining , G., Sch , G., , L. , L. - - , Meiosis ( Meiosis Male gametophytes develop spermatangia merged in whitish sori (a) located in the apical region of region apical the in (a) sori located whitish in merged spermatangia develop gametophytes Male in the outermost cortical cells (b), each of them release one tiny spermatium. Fertile female game female Fertile spermatium. one tiny release of them (b), each cells cortical outermost the in cells when release. D release. when cells tophytes (c) have a whitish longitudinal line at the terminal region of last order branches. I branches. order of last region terminal the at line longitudinal (c) awhitish tophytes have O - throu acarpospore release will carposporangium Each carposporangia. multiple inside holds gh the cystocarp ostiole and give rise to a tetrasporophyte. D atetrasporophyte. to rise give and ostiole cystocarp the gh tetrasporangia (f) located in sori at ends of last order branches and immerse in the cortex (g). (g). cortex the in immerse and branches order of last ends at sori in located (f) tetrasporangia lateral branches. I branches. lateral A nce carpogonia are fertilized a prominent and bilocular mature cystocarp (e) develops which (e) which develops cystocarp mature bilocular and aprominent fertilized are carpogonia nce Figure 1. Life cycle of Gelidium canariense cycle 1. Life Figure fter syngamy (S!) each procarp will develop into a diploid carposporophyte (2n) (arrows). carposporophyte adiploid into develop will procarp (S!) each syngamy fter procarp. complex a constitute that filaments nutritive and carpogonia several are there R !) occurs in each tetrasporangium to generate four haploid (n) tetraspores leaving empty empty leaving (n) haploid tetraspores four generate to tetrasporangium each in !) occurs phases of the cycle. I cycle. of the phases originated cells mother from formed are spermatangia section n transverse ashed line indicates the haploid phases and continuous line indicates diploid diploid indicates line continuous and phases haploid the indicates line ashed mages obtained and adapted from A from adapted and obtained mages f igu (n) dioecious. haploid and are . Gametophytes r es iploid tetrasporophyte (2n) form iploid tetrasporophyte (2017). et al. lfonso nside nside -

Revista Scientia Insularum, 2; 2019, pp. 153-181 179 Revista Scientia Insularum, 2; 2019, pp. 153-181 180 by a cell wall and separated from the original spore (arrow). (4) 24 hours after tetraspore release. release. tetraspore (arrow). spore after (4) hours 24 original the from separated and wall by acell migration in content protoplasm the from develop to begin tube (2) spore: germination of the attachment and release after 4hours 2to 3) (2 Between and wall. cell lacking still settlement surrounded is tube (arrow); (3) germination protrude the is tube germination (red color) and Figure 2. Gelidium 2. Figure Multicellular organism showing elongate distal cell (dc) and primary rhizoid (arrow). rhizoid (dc) primary and cell distal elongate showing organism Multicellular -type germination pattern germination -type Scale 20 µ 20 Scale m. A m. dapted from A from dapted and release after immediately . (1) tetraspore (2018) et al. lfonso . Figure 3. (1) Habitat forming Gelidiales in a mixed community in the north of T north the in community amixed in 3. (1) Gelidiales Figure forming Habitat Below, detail of the species: (2) Gelidium canariense species: of the Below, detail Pterocladiella capillacea (4)and Pterocladiella , (3) Gelidium arbuscula . enerife.

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